A Review of the Genus Eunice - Smithsonian Institution Libraries

* 

A Review of the Genus Eunice 

(Polychaeta: Eunicidae) 

Based upon Type Material 

KRISTTAN FAUCHALD 

m 

i 

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 523

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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 5 2 3 




Kristian Fauchald 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1992

ABSTRACT 

Fauchald, Kristian. A Review of the Genus Eunice (Polychaeta: Eunicidae) Based upon Type 

Material. Smithsonian Contributions to Zoology, number 523,422 pages, 117 figures, 53 tables, 

1992.—The genus Eunice has been widely reported in all kinds of marine environment. A total 

of 286 names have been applied to taxa originally described in, or subsequently referred to, the 

genus. All such taxa published prior to 1985 have been accounted for below. Of these, some or 

all types of about 175 are available and are described below. Twenty-five species are 

incompletely known; types are missing and original descriptions (or subsequent descriptions of 

types) are insufficient to relate these species to the rest. The descriptions of these species have 

been briefly summarized. A key to species includes nearly all species for which types were 

available. Species considered incompletely known are excluded from the key, but are included 

when possible in a set of tables comparing similar species. Forty-seven named taxa are 

indeterminable. Twenty-two have been referred to other genera, including some new 

combinations. In some cases the type lot included two species; some of these types could be 

referred to other, known species; however, four species are described as new. Seven species have 

been given new names for nomenclatural reasons. 

Attempts to group the 206 reasonably well-characterized species into supra-specific taxa 

using cladistic techniques did not lead to interpretable results. The reasons might include the 

very large number of species compared to the number of characters. A detailed analysis of the 

characters has been started using freshly collected material in an attempt to expand the number 

of characters and measure the variability of currently used characters. Similar reviews are 

planned for the remaining eunicean genera. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is 

recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral 

Montastrea cavernosa (Linnaeus). 

Library of Congress Cataloging-in-Publicalion Data 

Fauchald, Kristian. 

A review of the Genus Eunice (Polychaeta-Eunicidac) based upon type material / Kristian Fauchald. 

p. cm.—(Smithsonian contributions to zoology ; no. 523) 

Includes bibliographical references. 

1. Eunice (Annelida). I. Title. II. Series. 

QU.S54 no. 523 

[QL391.A6] 

59 s—dc20 91-19518 

[595.1'47] 

CIP 

® The paper used in this publication meets the minimum requirements of the American 

National Standard for Permanence of Paper for Printed Library Materials Z39.48— 1984.

Contents 

Page 

Introduction 1 

Nomenclatural Notes 2 

Publication Dates 2 

Family Name 2 

Generic Names 2 

Type Species 3 

Scope of the Study 4 

Limitations of the Study 5 

Acknowledgments 5 

Materials and Methods 6 

Microscopes and Computer 6 

Discussions of Individual Species 7 

Synonyms 7 

Material Examined 7 

Comments on the Material Examined 8 

Descriptions 8 

Unknown Morphological Features 8 

Expected States of Selected Unknown Features 8 

Character States Unknown 8 

Assumed States Used in Preparing Key 8 

Remarks 8 

Illustrations 8 

Tables 9 

Comparison Tables 9 

Morphological Features of the Eunicids 9 

Overall Body Features and Posterior End 9 

Prostomium and Peristomium 11 

Jaw Apparatus 12 

Branchiae 13 

Parapodia 13 

Setae 14 

Cladistic Analysis 15 

Selection of Outgroup Taxa 15 

Character List 18 

Plesio- and Apomorphic States of the Characters 18 

Size Characters 18 

Prostomium and Peristomium 18 

Eyes 19 

Antennae 19 

Peristomium 19 

Branchiae 20 

Parapodia 21 

Pectinate Setae 22 

Falcigers and Spinigers 22 

Aciculae 23 

Subacicular Hooks 23 

Results 24 

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

Overview of the Species 24 

Eunice Cuvier, 1817 27 

Key to the Species of Eunice 28 

Description of Species 40 

Eunice aciculata (Treadwell, 1922) 40 

Eunice adriatica Schmarda, 1861 42 

Eunice aedificatrix (Monro, 1933) 42 

Eunice aenea Blanchard, 1849 45 

Eunice aequabilis Grube, 1878 45 

Eunice afra Peters, 1854 47 

Eunice qfuerensis Hartman, 1944 49 

Eunice americana Hartman, 1944 51 

Eunice amoureuxi Rullier, 1974 51 

Eunice amphiheliae Marin in Filhol, 1885 53 

Eunice anceps Pruvot, 1930 54 

Eunice annulicornis Johnston, 1865 54 

Eunice antarctica Baird, 1869 56 

Eunice antennata (Lamarck, 1818) 57 

Eunice antennata aedificatrix Monro, 1933 60 

Eunice antennata gracilis Grube, 1866 60 

Eunice antillensis Ehlcrs, 1887 60 

Eunice aphroditois (Pallas, 1788) 62 

Eunice aphroditois djiboutiensis Gravier, 1900 64 

Eunice aphroditois punctata Fishelson and Rullier, 1969 64 

Eunice arcturi (Treadwell, 1928) 64 

Eunice arenosa Kinberg, 1865 64 

Eunice argentinensis (Treadwell, 1929) 65 

Eunice armillata (Treadwell, 1922) 67 

Eunice articulata Ehlers, 1887 69 

Eunice articulata (Hoagland, 1920) 71 

Eunice atlantica Kinberg, 1865 71 

Eunice attenuata Grube, 1866 72 

Eunice aucklandica Averincev, 1974 72 

Eunice auriculata Treadwell, 1900 72 

Eunice australis Quatrefages, 1866 74 

Eunice badia Grube, 1878 74 

Eunice balfouriana (Mclntosh, 1885) 74 

Eunice barvicensis Mclntosh, 1885 76 

Eunice bassensis Mclntosh, 1885 77 

Eunice bellii Audouin and Milne Edwards, 1833 77 

Eunice benedicti (Verrill, 1885) 78 

Eunice bertolonii (Chiaje, 1828) 80 

Eunice biannulata Moore, 1904 81 

Eunice biannulata mexicana Fauchald, 1970 81 

Eunice bicirrata Rullier, 1964 81 

Eunice biformicirrata (Treadwell, 1922) 83 

Eunice bilobata Treadwell, 1906 84 

Eunice binominata Quatrefages, 1866 85 

Eunice bipapillata Grube, 1866 86 

Eunice bitorquata Grube, 1870 86 

Eunice borneensis Grube, 1878 88 

Eunice bottae Quatrefages, 1866 88 

Eunice bowerbanki Baird, 1869 91 

Eunice brasiliensis Kinberg, 1865 93

NUMBER 523 

Eunice brevis (Ehlers, 1887) 93 

Eunice bucciensis (Treadwell, 1921) 94 

Eunice burmeisteri Grube, 1878 96 

Eunice caeca Shisko, 1981 96 

Eunice capensis Schmarda, 1861 98 

Eunice capensis (Kinberg, 1865) 98 

Eunice cariboea Grube, 1856 98 

Eunice cedroensis Fauchald, 1970 101 

Eunice challenged Mclntosh, 1885 101 

Eunice cincta (Kinberg, 1865) 103 

Eunice cingulata Claparede, 1868 103 

Eunice cirribranchis Grube, 1870 104 

Eunice cirrobranchiata Mclntosh, 1885 104 

Eunice claparedii Quatrefages, 1866 106 

Eunice coccinea Grube, 1878 106 

Eunice coccinioides Augener, 1922 107 

Eunice collaris Grube, 1869 109 

Eunice collini Augener, 1906 

Ill 

Eunice complanata Grube, 1877 113 

Eunice concinna (Verrill, 1900) 113 

Eunice congesta Marenzeller, 1879 113 

Eunice conglomerans Ehlers, 1887 114 

Eunice contingens (Chamberlin, 1919) 116 

Eunice crassitentaculata (Treadwell, 1922) 117 

Eunice culebra Treadwell, 1901 117 

Eunice curticirrus Knox, 1960 119 

Eunice denticulata Webster, 1884 119 

Eunice depressa Schmarda, 1861 121 

Eunice dilatata Grube, 1877 121 

Eunice djiboutiensis (Gravier, 1900) 123 

Eunice dubia Woodworth, 1907 124 

Eunice dubitata Fauchald, 1974 124 

Eunice ebranchiata Quatrefages, 1866 126 

Eunice edwardsi Mclntosh, 1885 127 

Eunice ehlersi Gravier, 1900 127 

Eunice eimeorum, new species 129 

Eunice elegans (Verrill, 1900) 130 

Eunice elseyi Baird, 1869 132 

Eunice enteles (Chamberlin, 1918) 132 

Eunice equibranchiata Mclntosh, 1885 133 

Eunice eugeniae, new species 133 

Eunice excariboea, new species 134 

Eunice fasciata (Risso, 1826) 136 

Eunice fauveli Gravier, 1900 136 

Eunice fijiensis Baird, 1869 138 

Eunice filamentosa Grube, 1856 138 

Eunice fimbriata Grube, 1870 140 

Eunice flaccida Grube, 1869 141 

Eunice flavapunctata (Treadwell, 1922) 143 

Eunice flavocuprea Grube, 1869 144 

Eunice flavofasciata Grube, 1878 144 

Eunice flavopicta Izuka, 1912 146 

Eunice floridana (Pourtales, 1867) 146 

Eunice franklini Monro, 1924 148


Eunice frauenfeldi Grube, 1866 148 

Eunice fucata Ehlers, 1887 150 

Eunice fuscafasciata (Treadwell, 1922) 152 

Eunice fusicirris Grube, 1878 152 

Eunice gagzoi Augener, 1922 153 

Eunice gaimardi Quatrefages, 1866 155 

Eunice gallica (Lamarck, 1818) 155 

Eunice gigantea auctores 155 

Eunice goodei, new name 156 

Eunice goodsiri (Mclntosh, 1885) 156 

Eunice gracilicirrata (Treadwell, 1922) 157 

Eunice gracilis Grube, 1866 159 

Eunice gracilis Moore, 1903 160 

Eunice gracilis (Crossland, 1904) 160 

Eunice gravieri Fauvel, 1911 160 

Eunice grubei Gravier, 1900 161 

Eunice grunwaldi (Risso, 1826) 161 

Eunice guanica (Treadwell, 1921) 161 

Eunice guildingi Baird, 1869 163 

"Eunice" gunneri (Storm, 1881) 165 

Eunice guttata Baird, 1869 165 

Eunice harassii Audouin and Milne Edwards, 1833 166 

Eunice havaica Kinberg, 1865 168 

Eunice hawaiensis Treadwell, 1906 169 

Eunice heterochaeta Quatrefages, 1866 169 

Eunice hirschi, new name 171 

Eunice hispanica (Lamarck, 1818) 173 

Eunice imogena (Monro, 1924) 173 

Eunice impexa Grube, 1878 174 

Eunice incerta (Hansen, 1882) 176 

Eunice indica Kinberg, 1865 176 

Eunice interrupta Treadwell, 1906 177 

Eunice investigatoris Fauvel, 1932 179 

Eunice jagori Grube, 1878 179 

Eunice januarii Grube, 1881 180 

Eunice japonica, new name 180 

Eunice jeffreysii Mclntosh, 1903 182 

Eunice johnsoni Hartman, 1954 182 

Eunice kerguelensis Averincev, 1974 183 

Eunice kinbergi Ehlers, 1868 185 

Eunice kinbergi (Webster, 1884) 186 

Eunice kobiensis Mclntosh, 1885 186 

Eunice langi (Treadwell, 1943) 188 

Eunice laticeps Ehlers, 1868 188 

Eunice laurillardi Quatrefages, 1866 190 

Eunice leptocirris Grube, 1870 192 

Eunice leucodon Ehlers, 1901 194 

Eunice leuconuchalis Benham, 1900 194 

Eunice leucosticta Grube, 1878 194 

Eunice levibranchia (Hoagland, 1920) 194 

Eunice limosa Ehlers, 1868 196 

Eunice lita (Chamberlin, 1919) 197 

Eunice longicirrata (Kinberg, 1865) 198 

Eunice longicirrata Webster, 1884 198

NUMBER 523 

vii 

Eunice longicirris Grube, 1869 198 

Eunice longicornis Grube, 1866 200 

Eunice longiqua Kinberg, 1865 200 

Eunice longisetis Webster, 1884 200 

Eunice lucei Grube, 1856 201 

Eunice macrobranchia Schmarda, 1861 203 

Eunice macrochaeta Schmarda, 1861 204 

Eunice madeirensis Baird, 1869 204 

Eunice magellanica Mclntosh, 1885 204 

Eunice magnified Grube, 1866 206 

Eunice makemoana (Chamberlin, 1919) 207 

Eunice manihine Longbottom, 1972 207 

Eunice manorae Aziz, 1938 209 

Eunice marenzelleri Gravier, 1900 209 

Eunice margaritacea Williams, 1853 210 

Eunice margaritacea (Verrill, 1900) 210 

Eunice margariticaeca Fischli, 1900 210 

Eunice marovoi Gibbs, 1971 211 

Eunice martensi Grube, 1878 213 

Eunice maxima Quatrefages, 1866 214 

Eunice medicina Moore, 1903 214 

Eunice megabranchia Fauchald, 1970 216 

Eunice megalodus Grube, 1878 217 

Eunice mexicana Fauchald, 1970 217 

Eunice microprion Marenzeller, 1879 219 

Eunice mindanavensis Mclntosh, 1885 221 

Eunice minuta Grube, 1850 222 

Eunice modesta Grube, 1866 222 

Eunice monilifer (Chamberlin, 1919) 224 

Eunice mossambica Peters, 1854 224 

Eunice mucronata Moore, 1903 224 

Eunice multicylindri Shisko, 1981 227 

Eunice multipectinata Moore, 1911 228 

Eunice murrayi Mclntosh, 1885 . 230 

Eunice mutabilis Gravier, 1900 230 

Eunice mutilata Webster, 1884 232 

Eunice mutilata samoae Hartmann-Schroder, 1965 233 

Eunice mutilatoides Augener, 1922 233 

Eunice narconi Baird, 1869 235 

Eunice nesiotes (Chamberlin, 1919) 235 

Eunice nicidioformis Treadwell, 1906 237 

Eunice nigricans Schmarda, 1861 238 

Eunice northioidea Moore, 1903 240 

Eunice norvegica (Linnaeus, 1767) 241 

Eunice notata (Treadwell, 1921) 243 

Eunice (Marphysa) novaehollandiae (Kinberg, 1865) 244 

Eunice oerstedii Stimpson, 1854 244 

Eunice oliga (Chamberlin, 1919) 244 

Eunice oliga papeetensis (Chamberlin, 1919) 245 

Eunice ornata Andrews, 1891 245 

Eunice ovalifera Fauvel, 1936 246 

Eunice pacifica Kinberg, 1865 246 

Eunice palauensis Okuda, 1937 248 

Eunice (Eriphyle) paloloides Moore, 1909 249


Eunice panamena (Chamberlin, 1919) 249 

Eunice papeetensis (Chamberlin, 1919) 250 

Eunice parasegretata Hartmann-SchrOder, 1965 252 

Eunice parca Grube, 1878 253 

Eunice parva Hansen, 1882 253 

Eunice parvibranchis Grube, 1870 255 

Eunice paucibranchis Grube, 1866 256 

Eunice paupera Grube, 1878 256 

Eunice pauroneurata (Chamberlin, 1919) 256 

Eunice pectinata Grube, 1869 258 

Eunice pelamidis Quatrefages, 1866 260 

Eunice pellucida Kinberg, 1865 261 

Eunice pennata (Miiller, 1776) 263 

Eunice perimensis Gravier, 1900 264 

Eunice perrieri Gravier, 1900 266 

Eunice petersi, new name 267 

Eunice philocorallia Buchanan, 1893 267 

Eunice pinnata (Miiller, 1779) 269 

Eunice plicata Baird, 1869 269 

Eunice polybranchia (Verrill, 1880) 271 

Eunice prayensis Kinberg, 1865 273 

Eunice procera Grube, 1866 274 

Eunice prognatha Mclntosh, 1885 274 

Eunice pruvoti, new name 276 

Eunice pulvinopalpata Fauchald, 1982 277 

Eunice punctata (Risso, 1826) 279 

Eunice punctata Peters, 1854 279 

Eunice punctata Grube, 1856 279 

Eunice purpurea Grube, 1866 279 

Eunice pycnobranchiata Mclntosh, 1885 281 

Eunice quadrioculata Grube, 1856 284 

Eunice quinquefida Moore, 1903 284 

Eunice quoya Quatrefages, 1866 284 

Eunice reducta Fauchald, 1970 285 

Eunice rissoi Quatrefages, 1866 285 

Eunice rosaurae Monro, 1939 287 

Eunice roussaei Quatrefages, 1866 288 

Eunice rubella Knox, 1951 288 

Eunice rubra Grube, 1856 291 

Eunice rubrivittata (Treadwell, 1921) 292 

Eunice rubrocincta Ehlers, 1868 294 

Eunice rullieri, new name 294 

Eunice samoae Hartmann-Schroder, 1965 295 

Eunice savignyi Grube, 1878 297 

Eunice schemacephala Schmarda, 1861 297 

Eunice schizobranchia Claparede, 1870 298 

Eunice scombrinis Quatrefages, 1866 299 

Eunice sebastiani Nonato, 1965 299 

Eunice segregata (Chamberlin, 1919) 303 

Eunice semisegregata Fauchald, 1969 305 

Eunice siciliensis Grube, 1840 307 

Eunice simplex Peters, 1854 307 

Eunice sonorae Fauchald, 1970 307 

Eunice splendida Grube, 1856 308 

Eunice spongicola (Treadwell, 1921) 308

NUMBER 523 

ix 

Eunice stanleyi, new species 310 

Eunice stigmatura (Verrill, 1900) 311 

Eunice stragulum Grube, 1878 313 

Eunice subdepressa Grube, 1866 313 

Eunice suviensis (Treadwell, 1922) 313 

Eunice taenia Claparede, 1864 313 

Eunice tahitana Kinberg, 1865 314 

Eunice tentaculata Kinberg, 1865 314 

Eunice tentaculata Quatrefages, 1866 316 

Eunice tenuicirrata (Verrill, 1900) 316 

Eunice tenuis (Treadwell, 1921) 316 

Eunice teretiuscula Schmarda, 1861 317 

Eunice thomasiana Augener, 1922 317 

Eunice tibiana (Pourtales, 1867) 319 

Eunice torquata Quatrefages, 1866 319 

Eunice torresiensis Mclntosh, 1885 321 

Eunice triantennata (Risso, 1826) 322 

Eunice tribranchiata Mclntosh, 1885 323 

Eunice tridentata Ehlers, 1905 324 

Eunice tristriata Grube, 1870 325 

Eunice tubicola (Treadwell, 1922) 325 

Eunice tubifex Crossland, 1904 327 

Eunice unidentata Rioja, 1962 329 

Eunice unifrons (Verrill, 1900) 330 

Eunice valenciennesii Grube, 1878 332 

Eunice valens (Chamberlin, 1919) 332 

Eunice valida Gravier, 1900 333 

Eunice validissima Grube, 1866 333 

Eunice validobranchiata Monro, 1937 333 

Eunice violacea Grube, 1856 335 

Eunice violaceomaculata Ehlers, 1887 335 

Eunice viridis auctores 337 

Eunice vittata (Chiaje, 1829) 337 

Eunice vittatopsis Fauchald, 1970 340 

Eunice vivida Stimpson, 1854 340 

Eunice wasinensis, new name 340 

Eunice websteri Fauchald, 1969 342 

Eunice woodwardi Baird, 1869 343 

Eunice zonata Chiaje, 1841 345 

Tables 19-53 345 

Appendix A. Character List for Eunice 375 

Appendix B. Characters and Character States for Each Species As Setup for Use with 

DELTA 378 

Appendix C. Important Publications on Eunice Issued Since 1985 and New 

Taxa Described 415 

Literature Cited 416 

TABLES 

Variability within a Species 

1. Eunice aedificatrix 44 

2. Eunice antennata 59 

3. Eunice cariboea 99 

4. Eunice dubitata 126 

5. Eunice floridana 147 

6. Eunice lucei 203


7. Eunice marovoi 213 

8. Eunice mexicana 219 

9. Eunice mucronata 226 

10. Eunice muUicylindri 227 

11. Eunice nicidioformis 238 

12. Eunice ornata 246 

13. Eunice polybranchia 273 

14. Eunice pycnobranchiata 283 

15. Eunice torresiensis 322 

16. Eunice validobranchiata 335 

17. Eunice violaceomaculata 336 

18. Eunice vittata 339 

Comparison among Species 

19. Overview of group A-l 345 

20. Group A-l with articulated peristomial cirri 346 

21. Group A-l without articulated peristomial cirri 347 

22. Overview of groups A-2 and A-4 348 

23. Group A-2 348 

24. Overview of group B-l 349 

25. Group B-l with antcnnal articulations 350 

26. Group B-l without antennal articulations 350 

27. Overview of group B-2 351 

28. Group B-2 without antennal articulations 353 

29. Group B-2 with cylindrical antcnnal articulations and all nolopodial cirri 

articulated 354 

30. Group B-2 with cylindrical antcnnal articulations and notopodial cirri 

articulated anteriorly only 355 

31. Group B-2 with cylindrical antcnnal articulations but without notopodial 

articulations 356 

32. Group B-2 with drop-shaped or moniliform antcnnal articulations 357 

33. Overview of groups B-3 and B-4 358 

34. Groups B-3 and B-4 with articulated antennae 359 

35. Groups B-3 and B-4 with articulated antennae and pectinate branchiae .... 360 

36. Groups B-3 and B-4 with articulated antennae and palmate or simple 

branchiae 360 

37. Groups B-3 and B-4 without antennal articulations 361 

38. Group B-4 without antennal articulations; with branchiae and with median 

antennae longer than other antennae 362 

39. Group B-4 without antennal articulations, with branchiae and all antennae 

equally long 363 

40. Abranchiate species 364 

41. Overview of group C-l 365 

42. Group C-l with cylindrical antennal articulations 366 

43. Group C-l with distally drop-shaped or moniliform articulations 367 

44. Group C-l with all articulations drop-shaped or moniliform 367 

45. Group C-l without antennal articulations 368 

46. Overview of group C-2 369 

47. Group C-2 with all notopodial cirri articulated 370 

48. Group C-2 with only anterior notopodial cirri articulated 371 

49. Group C-2 without notopodial articulations 371 

50. Overview of Group D 372 

51. Group D 372 

52. Overview of species with unusual kinds of setae 373 

53. Species with unusual kinds of setae 373




Kristian Fauchald 

Introduction 

Eunicean polychaetes are uniquely defined by the presence 

of laterally arranged maxillae and paired ventral mandibles 

mounted on an eversible, muscular U-shaped structure (Ehlcrs, 

1868; Dales, 1962; Clark, 1964; Fauchald, 1974a). Traditionally 

the group was considered as a single family, Eunicidae, 

with a series of subfamilies (Fauvel, 1923; Day, 1967). Lately 

most authors recognize the group as a distinct order, Eunicida 

(e.g., Fauchald, 1977; Pettibone, 1982), with a series of families. 

The family Eunicidae Berthold (1827) is the nominal 

family. Other families include Onuphidae Kinberg (1865), 

Lumbrineridae Malmgren (1867), Harunaniellidae Imajima 

(1977), Iphitimidae Fauchald (1970), Arabellidae Hartman 

(1944), Lysaretidae Kinberg (1865), Oenoniidae Kinberg 

(1865), and Dorvilleidae Chamberlin (1919a). The parasitic 

families Histriobdellidae Vaillant (1890) and Ichthyotomidae 

Eisig (1906) and the mesopsammic family Dinophilidae Remane 

(1932) are usually considered as members of the order, 

but are sometimes relegated to a satellite status. Relationships 

between the Eunicidae and the other families are outlined 

below in the section on cladistics. Formal definitions of the 

families can be found in Fauchald (1977), Pettibone (1982), 

and Colbath (1989). 

Eunicoid polychaetes are known from fossilized jaws present 

in Ordovician fossil beds (Kielan-Jaworowska, 1966). 

Members of the order are present in all marine environments, 

most are free-living; parasitic taxa are found in part on other 

polychaetes, in part on fish or crustaceans. In some families, all 

or nearly all species are tubicolous; in other families most 

species lack tubes. Eunicids sensu stricto are present in all 

marine benthic environments. They are especially common in 

tropical shallow seas, in coral reef rubble, and in mangrove 

Kristian Fauchald, Department of Invertebrate Zoology, National 

Museum of Natural History, Smithsonian Institution, Washington DC. 

20560. 

swamps. Some species form complex tubes (e.g., Hanley, 

1986:215); most tubicolous species form less-complicated 

structures (Ehlers, 1887), but many eunicids lack tubes completely. 

Most species live in cracks and crevices in mixed 

rubble, rock, and sand environments in shallow water. Many 

species drill in dead coral rubble (Hutchings, 1981) and are 

sufficiently common to be of considerable ecological importance. 

The Atlantic and Pacific palolo-worms are known for 

their brief, lunar periods of spawning (e.g., Mayer, 1900; 

Hofmann, 1974). The Atlantic and Pacific palolos belong to 

different genera, Eunice and Palolo, respectively. The habit of 

having brief, intense spawning periods may be more common 

in the family than currently appreciated. 

The first described eunicid species was originally placed in 

Nereis. Nereis norvegica Linnaeus (1767) was described from 

an ahermatypic coral reef in Norway. Nereis aphroditois was 

described from a hermatypic coral reef in Sri Lanka (Ceylon) 

by Pallas (1788). Other species were added by O.F. Miiller 

from Norway (Muller, 1776, 1779). 

Cuvier (1817:524) named a new genus, Eunice, to contain 

these and related taxa (a more detailed discussion of nomenclatural 

problems is given below). Audouin and Milne Edwards 

(1832), Kinberg (1865), Quatrefages (1866), Malmgren (1867), 

Ehlers (1868), and Grube (1870b, 1878a,b) reviewed early 

work and added numerous new species. Mclntosh (1885) and 

Chamberlin (1919a) added new taxa based on materials from 

the Challenger and Albatross cruises, respectively, many from 

deep-water habitats. Treadwell (1921, 1922) added many new 

species from coral reefs in the Caribbean Sea and the Pacific 

Ocean, respectively. He also attempted to expand the set of 

features used to characterize species to include color patterns; 

he was not followed in this attempt by any later worker. The 

live color is, however, very characteristic for several species 

and may be useful as a character in a well-studied geographical 

area. Fauvel (1917, 1919) reviewed many of the tropical species, 

especially from the Indian Ocean, and established numer-


ous synonyms. He failed to examine many available types, 

perhaps because of the political situation in Europe at the time. 

Augener (1922a,b, 1923) reviewed some early types and established 

other synonyms. Hartman (1944) reviewed, without 

access to many types, all species described from the Americas; 

later Hartman (1948,1956) reviewed types of species described 

by Kinberg and Treadwell, respectively. In her 1944 publication 

Hartman codified the characters used to separate genera 

and species in the family and, following suggestions by Ehlers 

(1868), proposed a scheme of informal species groups within 

the genus Eunice for the American members of the genus. This 

scheme was expanded by Fauchald (1970) to include all species 

listed in Hartman (1959, 1965). He added a codification of the 

branchial distribution to the scheme, originally suggested by 

Grube (1878a), resulting in a more detailed set of groups than 

the original Ehlers-Hartman scheme. Miura (1986) detailed the 

branchial distribution of several species, suggesting that this 

distribution would add a valuable character to the description of 

eunicids. 

NOMENCLATURAL NOTES 

PUBLICATION DATES.—Some crucial publications are frequently 

misquoted in the literature. First of these is a set of 

French publications involving Savigny, Lamarck, and Cuvier. 

These three gentlemen based their publications on the collections 

of Musee National d'Histoire Naturelle, Paris. Savigny 

worked up the material collected during Napoleon's travels in 

Egypt and other polychaete material deposited in the Parisian 

collections. He probably had a manuscript finished before 1815 

(the plate including the eunicids was engraved by 1812). This 

manuscript was not published until 1820 as a separate issue 

(preprint) of volume 1, part 3 of the "Description of Egypt." 

The publication date is printed on the back page of the preprint, 

which was separately paginated. Volume 1(3) was later repaginated 

as part of the complete volume and the publication 

date 1809 was printed on the frontispiece. Only volume 1(1), 

'The Fishes of the Nile," was issued in 1809; the publication 

date for Savigny's study of the annelids was 1820 and Savigny's 

plates were not issued until 1826. Sherbourn (1897:287) 

claimed that volume 1(3) was issued in 1822, quoting a review 

in Gottingische Gelehrte Anzeigen, 2:695, as his source. Quite 

correctly, a review of Savigny's study starts on that page, but 

the publication date of volume 1(3) is not mentioned. On the 

next page, 1822 is mentioned as a publication date but only for 

parts of volume 2; Sherbourn must have misunderstood the 

rather complicated review. 

Cuvier issued in 1817 volume 2 of Le Rigne animal.... in 

which the name Eunice was first used (p. 524). Cuvier 

(1817:525) listed the same previously published species that 

were later included in the genus in Savigny's publications, but 

added no new taxa. In the next calendar year (1818) Lamarck 

published volume 5 of his Histoire naturelle Lamarck 

quoted Savigny's manuscript extensively; he described Leodice 

and in addition nearly all species Savingy had included in 

his manuscript, quoting Savigny's manuscript in his synonymy 

lists for all. 

Three important publications from the mid-1860s have 

caused confusion. Kinberg in a series of papers published in 

Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, 

Stockholm, reported on the polychaetes collected during the 

Eugenie expedition. The euniceans were included in the number 

10 of the journal for the year 1864, this number was 

published in 1865. Other parts of this series was published 

earlier and later, but do not concern the euniceans. Quatrefages' 

large Histoire des Annel£s bears the publication date 1865 on 

the title page of the first volume; as pointed out by Wright 

(1867:578; see also Fauchald, 1986:252), this volume was not 

issued until 1866. Thus, new euniccan taxa proposed by Kinberg 

antedates those by Quatrefages, and Kinberg's names have 

priority in cases where the two gentlemen described the same 

species. Malmgren first treated eunicids in his 1867 publication; 

note that both of Malmgren's major publications (1865 

and 1867) were issued as separately paginated reprints as well 

as in a journal. 

FAMILY NAME.—The family Eunicidac, as a concept, was 

first used by Lamarck (1818). This mention is not available for 

purposes of synonymy because there is no reference to the 

genus Eunice in Lamarck's text and the name is spelled Eunicae. 

Savigny (1820) used the family name Eunicae, but the 

generic name Eunice is listed only as a synonym of Leodice. 

Such mention of a family name is invalid for priority purposes 

according to the International Code of Zoological Nomenclature 

(ICZN; also referred to herein as "the Code"). Latreille 

(1825:239) defined the family and listed Eunice, with Leodice 

as a synonym; however, the name form used is invalid. The first 

valid mention of the family is by Berthold (1827:227-228), a 

translation of Latreille (1825) to German. 

GENERIC NAMES.—The oldest generic name applied to a 

member of this group (other than the Linnean Nereis) is Tibiana 

in Lamarck (1816). The specimen examined by Lamarck 

was the tube of a deep-water species from the Indian Ocean 

(Lamarck, 1816:148). Lamarck believed he was describing a 

coral, perhaps an octocoral, and the name Tibiana has not been 

used as a generic name for polychaetes. According to the Code, 

articles 12b(8), 23f(III), and 72c(I), the name is available for 

purposes of priority (ICZN, 1985). In a description of a tube 

under the specific name Marphysa tibiana by Pourtales (1867) 

(now known as Eunice tibiana), a specific reference to Tibiana 

as the name of a eunicid-tube was made. The name has otherwise 

not been used. I have prepared an application to the 

International Commission on Nomenclature to have the name 

suppressed. 

Cuvier (1817:525) defined Eunice and Lamarck (1818:321- 

323) diagnosed Leodice. The two taxa have been used more or 

less as synonyms since they were both first named. 

Savigny worked up the collections of eunicid polychaetes in 

the museum in Paris (MNHN). His descriptions in Savigny

NUMBER 523 

(1820) are exemplary as are the plates for that volume. Cuvier's 

(1817) and Lamarck's (1818) volumes were both based at least 

in part on Savigny's manuscript, but because of the publication 

dates, Savigny cannot be quoted as author for Leodice nor for 

the new species included in the "Description of Egypt" included 

by either one or the other of the two earlier authors. 

From Audouin and Milne Edwards in 1833 until the present, 

previous workers have attempted to rescue Savigny as author of 

the many genera and species described in Savigny (1820). The 

Code, however, is very clear on the point of priority of publication; 

and the publication dates are sufficiently well established 

that Cuvier and Lamarck end up with the honors. 

Savigny recognized the separation between what is now 

called Eunice and Marphysa. Part of his definition, the presence 

or absence of peristomial cirri, is the only valid character 

separating the two genera. 

Audouin and Milne Edwards (1833:208), in their section on 

the classification of the eunicids, commented: 

La famillc dont nous faisons ici l'hisloire a etc ciablic sous le nom d'Eunices 

par M. Savigny, pour rcccvoir, outre les Eunices de M. Cuvier, auxquelles il 

donne le nom de Liodices. trois genres nouveaux, les Lysidic.es, les Aglawes et 

les Oenones. 

The family about which we are writing, was established by Savigny to receive, 

other than M. Cuvier's Eunice to which he gave the name Leodice, three new 

genera, Lysidice, Aglaura, and Oenone. (K.Fd translation, italicization as in the 

original.] 

Audouin and Milne Edwards thus synonymized Leodice 

with Eunice as did Grube (1850). Note that Audouin and Milne 

Edwards recognized Savigny as author of the family; this 

problem has been discussed above. 

The generic name Eunice has been considered preoccupied 

by Eunice Rafinesque (1815); this is incorrect, because Rafinesque's 

use is a nomen nudum. Verrill (1900) considered 

Eunice preoccupied by Eunica Hiibner, 1816, thus preferring 

the name Leodice. Hiibner's name is spelled differently; Eunice 

Cuvier, 1817, cannot be considered validly preoccupied by 

Hubner's name according to the Code. 

The valid name for the genus is here considered to be Eunice 

Cuvier, 1817. 

Other generic names proposed for the group include Nereidonta, 

erected by Blainville (1828:476) for his species N. 

paretti. This species may be a Palola according to Ehlers 

(1868:353). The question will be addressed in a future review 

of that genus. Mayeria was proposed by Verrill (1900:650) for 

Staurocephalus gregaricus Mayer, 1900, without any description 

of Mayer's material or any other material. Mayer's species 

was referred to Eunice schemacephala Schmarda (1861) by 

Augener (1925:29); the latter is here considered an indeterminable 

species of Eunice. Mayer left no specimens of his species 

in any collections; thus his species is indeterminable. The 

genus Mayeria lacks a clear definition and is here considered 

indeterminable. 

Eriphyle Kinberg (1865), erected for E. capensis Kinberg 

(1865), is here considered a junior synonym of Eunice as first 

proposed by Malmgren (1867; see below). The species for 

which Eriphyle was erected was renamed Eunice kinbergi by 

Ehlers (1868; see below). 

The generic name Nicidion, originally proposed as a subgenus 

of Eunice by Kinberg (1865:564), has been used by a 

variety of authors at the generic level. The original single 

identifying feature of the taxon, the absence of branchiae, does 

not appear to be a valid generic character. Nicidion is here 

treated as a synonym of Eunice. 

Other generic names used in the family pertain to species to 

be treated in the future. 

TYPE SPECIES.—Cuvier (1817:525) listed in a footnote a 

series of species he transferred to his new genus, Eunice: 

Terebella aphroditois. Nereis pinnata, Nereis norwegica (note 

spelling), Nereis tubicola, and Nereis cuprea. The last two are 

onuphids, Hyalinoecia tubicola and Diopatra cuprea, respectively 

(e.g., Fauvel, 1923). Terebella aphroditois, originally 

described as Nereis by Pallas (1788) and moved to Terebella by 

Gmelin (1788), is a valid species of Eunice (see below). Nereis 

pinnata and pennata were originally both described by O.F. 

Miiller (1776; see also 1779); of these, N. pinnata is indeterminable 

(see below) and N. pennata is a valid species of Eunice 

(see below). Nereis norwegica, originally spelled norvegica, 

was described by Linnaeus (1767). I cannot determine from the 

text how many of these species Cuvier had seen. He made the 

following comment (Cuvier, 1817:525): 

J'en connais une de la mer des Indes, de plus de quatre pieds de long. 

I know one [specimen] from the Indian Ocean that is more than 4 feet long. 

[K.Fd translation.] 

Cuvier probably examined the collection in Paris in some 

detail, but he explicitly stated that he had seen at least one 

specimen. Traditionally, his statement has been considered as 

referring to E. aphroditois, originally described from Sri 

Lanka; this species can become very large. However, Cuvier 

did not specifically name the large specimen he had seen; no 

other valid species is named by Cuvier (1817). 

Lamarck (1818:321-323) quoted Savigny's manuscript extensively. 

He listed (for Leodice) L. gigantea, L. antennata, L. 

gallica, L. norwegica, L. pinnata, L. hispanica, L. opalina, and 

L. sanguinea and gave a brief Latin diagnosis for each. 

For L. gigantea Lamarck listed the following synonyms: 

An terebella aphroditois Gmel. p. 3114 

Eunice.Cuv.Regne anim. 2 p.525 

Leodice gigantea Sav. MSS. 

This list, with original italization indicated, is an excerpt 

from a longer list of synonyms published by Savigny two years 

later; it is quoted here to demonstrate that as originally published, 

the name Leodice gigantea was tied to the species name 

aphroditois. Savigny's (1820:49) synonymy list for Leodice 

gigantea reads (French name forms omitted): 

1. Leodice gigantea. 

Nereis aphroditois Pall. Nov. Act. Petrop. torn II, pag. 229, tab. 5, fig. 1-7.


-Terebella aphroditois Gmclin Syst. nat torn I part. 6. p«g. 3114. no. 9. 

= Variete d'age ou espece ires-voisine. 

Nereis giganlea Collect, du Mus. 

Eunice giganlea Cuv. Collect et Regn. anim. torn II. pag. 525. 

The implication is that Savigny included all forms listed as 

synonyms of Leodice gigantea. Furthermore, it is implied that 

the species had been named by Cuvier (1817:525), as mentioned 

above; this is not the case. Cuvier may have labeled the 

specimen in the collections by the name gigantea, but that 

name is nowhere mentioned in his publication. The earliest 

valid use of the name Leodice gigantea is in Lamarck's (1818) 

publication. Because Pallas' description of Nereis aphroditois 

is valid, Savigny established Leodice (= Eunice) gigantea 

Lamarck (1818) as a junior synonym of Pallas species. This 

discussion establishes only a valid "name" for this species, but 

not a valid type species for the genus Eunice; a valid "concept" 

was established by Ehlers (1868; see below). 

Malmgren (1867:64-65), in commenting on what he considered 

Kinberg's incorrect use of generic names, made the following 

statement: 

Fasthlllande sasom princip, an lita ett genusnamn alltid Itfolja den art. hvilken 

genusnamnets uppsia'Uarc haft tillfalle all sjclf undersdka cllcr foretradesvis 

afscit, anser jag de gamla genusnamnen inom fam. Eunicidae bora rattast 

anvandas pi foljande satt. Namnet Eunice gafs af CUVIER it Eunice giganlea 

CUV. Regn. Anim. Tom II p.525. nouv. edit. Tom III. p. 199. hvarfore detu 

namn omojligen kan anvandas for nagon annan grupp inom familjen an dit 

Eunice gigantea CUV. hor. Nar KINBERG nyligen uppstallt en nytt genusnamn 

Eriphyle for en grupp, dit nan sjelf afven raknar Eunice gigantea CUV., 

och begagnat namnet Eunice CUV. i en betydelse, for hvilken SAVIGNY's 

gamla namn Leodice (sens, str.) hade varit del lampligaste och rattaste uttryck, 

maste namnet Eriphyle KNBG, fltminstone i dess narvarande betydelse sisom 

fullkomligt ofverflodigt och synonymt med CUVIER *s Eunice heltochh&llet 

forsvinna, och namnet Leodice SAV. s. str. bdr begagnas for den grupp, sisom 

KINBERG benamnt Eunice, ty Leodice antennata SAV. Descr. d. l'Egypte 

Tom. XXI p. 380, som bdr anses for typ for Leodice SAV. hor afven till 

KINBERGS genus Eunice. 

Maintaining as a principle to let a generic name always follow the species that 

the author of the generic name had the opportunity to examine himself, or 

preferentially designated, I consider that the old generic names in the family 

Eunicidae should be used in the following fashion. The name Eunice was given 

by Cuvier to Eunice gigantea, Cuvier, Regn. Anim., Tom. II. p. 525, new 

edition, Tom. Ill, p. 199, hence this name cannot possibly be used for any other 

group of the family than the one to which Eunice gigantea Cuvier belongs. 

When Kinberg recently erected a new generic name, Eriphyle, for a group to 

which he himself ranks Eunice gigantea Cuvier, and applies the name Eunice 

Cuvier in a sense for which Savigny's old name Leodice (sensu stricto) had 

been the most appropriate and most correct name, the name Eriphyle Kinberg, 

at least in its current sense, should altogether disappear as being completely 

unnecessary and synonymous with Cuvier's Eunice, and the name Leodice 

Savigny sensu stricto should be applied to the group that Kinberg named 

Eunice, because Leodice antennata Savigny, Descr.d.l'Egypte, Tom. XXI, 

p.380, which must be considered the type for Leodice Savigny, belongs to 

Kinberg's genus Eunice. [K.Fd translation; capitalization and italicization as in 

the original.] 

This rather complicated uttering boils down to three major 

theses: 

1. Malmgren believed that Cuvier named his genus for the 

species gigantea, but including possibly other species. However, 

as indicated above, the name gigantea is nowhere mentioned 

on the page of Cuvier's book Malmgren cited. 

2. Malmgren declared Eriphyle Kinberg a junior synonym of 

Eunice. 

3. Malmgren designated antennata as type for the genus 

Leodice, but considered the latter as a synonym of Eunice. 

Verrill (1900:638) quoted Malmgren as saying: "Malmgren 

... restricted Eunice to the type of E. gigantea" Verrill's 

statement, although a misunderstanding of Malmgren's intent, 

is a clear designation of type for Eunice and is valid according 

to the Code. Moreover, it is the first correct designation and 

takes precedence over all other designations. 

The type species has often been quoted as Nereis aphroditois 

Pallas, 1788, with type designation by Hartman (1959:308). 

Both Cuvier and Savigny cited Nereis aphroditois Pallas, 1788, 

as a synonym for Eunice (or Leodice) gigantea Cuvier. However, 

the first valid designation of type for the genus in the 

sense of the Code is Verrill (1900), fixing the type as Eunice 

gigantea Cuvier, 1817. This binomen is invalid in the sense of 

the Code, but there is a valid binomen available (Code recommendation 

67B). Ehlers (1868:306-310) gave a detailed definition 

of E. aphroditois and in a discussion suggested that 

Cuvier's £. gigantea was a synonym as earlier authors had 

done. Ehlers listed Leodice gigantea Savigny and E. gigantea 

Milne Edwards and E. gigantea Quatrefages as synonyms. The 

former reference is to the last edition of Cuvier's Le Rigne 

animal..., the so-called student's edition; the latter is 

Quatrefages (1866). The two names probably refer to the same 

species, but both are without types and cannot be defined 

accurately without access to material from the type areas. 

Cuvier's material came from the Indian Ocean, presumably one 

of the French possessions, the Seychelles, Madagascar, or the 

Mascarenes. Pallas' material was from the coast of Sri Lanka. 

The two species are considered synonymous in this paper. 

The type species should be cited as Leodice gigantea Lamarck 

(1818 = Nereis aphroditois Pallas, 1788) by subsequent 

designation (Verrill, 1900:688). 

SCOPE OF THE STUDY 

Most species treated in this review were described either as 

Eunice or Leodice. Species of the Linnean genus Nereis described 

prior to the erection of Eunice and Leodice and considered 

members of Eunice by Hartman (1959, 1965) are also 

included. The few species in the latter category either come 

from well-circumscribed type localities (e.g., E. norvegica), or 

a well-defined tradition has developed associating a morph 

with a name (£. aphroditois). I reviewed descriptions of other 

species of Nereis described prior to 1817 to seek out additional 

eunicids, but found none. Species named in Eriphyle Kinberg, 

1865, and Nicidion Kinberg, 1865, have been included, because 

both genera are usually treated as subgenera of Eunice 

(cf. Hartman, 1959) if they are recognized at all. Species 

originally described as Eunice (Marphysa) also have been

NUMBER 523 

included, but not species described and exclusively recognized 

in Marphysa, except for a few that have been transferred from 

that genus to Eunice. 

Only species published before 1985 have been included in 

the study. Appendix C lists publications describing new taxa 

added since that time. 

LIMITATIONS OF THE STUDY 

During a study of the clastic effects of polychaetes on coral 

reefs, Pat Hutchings of the Australian Museum collected a very 

large collection of eunicids, including masses of small specimens. 

She requested my assistance in getting them identified to 

species. I realized that without a review of types of previously 

described species, the task would be impossible. I first reviewed 

(Fauchald, 1986) species described from Australia and 

New Zealand. This paper is an expansion of that effort to 

include species described from other areas. 

Studies of types alone are inadequate for clarification of 

systematic problems in any group; however, without a description 

of all available types, all other taxonomic studies will lack 

foundation. 

The family Eunicidae is poorly known, even if a few members 

have been intensively studied. We know, for example, 

very little about longevity and numbers of (annual) reproductive 

episodes, and whether the females grow between these 

bouts. Perhaps, most importantly from a systematise s point of 

view, we know little about how morphological features vary 

with size or sex. Some species become sexually mature at about 

10-15 mm in length (e.g., Eunice marovoi Gibbs, 1971; and 

examination of Gibbs' types below). On the other hand, very 

large individuals, 3-4 m in length, still appear to be reproductively 

active judging from the presence of large eggs in the 

body cavity (e.g., E. aphroditois, E. sebastiani). Several authors 

(e.g., Ehlers, 1868:309-311; Fauvel, 1917:209-232; 

Hartman, 1944:98) have wrestled with the problem of identifying 

smaller specimens of species that become very large, but 

their conclusions are unconvincing. 

Newly metamorphosed eunicid juveniles may have 5-15 

setigers (Akesson, 1967) and grow, as usual among annelids, 

by addition of segments from a pre-pygidial growth zone. 

Unless anterior regeneration has taken place, the oldest segments 

are those immediately behind the peristomium. Parapodia, 

branchiae, and setae of each segment bear a characteristic 

relation to the relative position of the segment in the body. 

Branchiae start and end at a specifiable range of segments; 

subacicular hooks start within a limited range of segments. 

Thus ontogenetic changes in the structure and shape of the 

parapodia must take place. Because the distribution of features 

is relatively stable, the underlying ontogenetic changes must be 

structured. The complement of structures present on each segment 

varies with a segment's relative position, so all segments 

change in shape and structure during the life of the organism. 

For example, early juveniles of a species related to Eunice afra 

have branchiae represented by single filaments in segments 

5-8 when the whole body consists of 20-30 setigers; adults of 

the same species with 120-130 setigers have branching branchiae 

starting at setiger 18-20 and continued to the posterior 

end (Fauchald, personal observations). Ontogenetic changes in 

soft-body features have been poorly documented, and a study 

of the types will yield limited information about such changes. 

Changes in setal structures with size have been better documented. 

During the early juvenile life, the setal complement of 

the setigers change in an orderly pattern; eventually the adult 

complement of setae is established (Akesson, 1967:157). Such 

orderly changes have been more frequently explored in the 

closest relatives of the eunicids, the onuphids (Blake, 1975:55; 

Hsieh and Simon, 1987:200-201; Paxton, 1986:20-21) and 

have been found to follow fairly restricted patterns of transformation. 

The obvious implication is that the setal complement of 

each segment is a well-regulated feature. The setal complement 

and distribution at any given time or over time may show 

species-specific patterns. These patterns may be related to the 

distribution of other morphological features or show independent 

patterns. In other polychaetes, such as maldanids (Green, 

1987) and nephtyids (Rainer, 1984), even the number of different 

kinds of setae in each segment is considered a speciesspecific 

feature. Particular ontogenetic sequences may be unique 

to a single species, or characteristic of larger groups. 

For these reasons, I started a study of the morphological 

variability of species available in shallow water at Carrie Bow 

Cay, Belize. Preliminary results (Fauchald, 1991) indicate that 

various length measures and some numerically variable morphological 

features are very precisely controlled in each taxon. 

Some features appear size related; others are size independent 

The species studied appear to have their own patterns; simple, 

generalized patterns have yet to emerge. Thus, without considerably 

more detailed studies of ontogenesis in many more 

species, one can only suggest possible synonyms from a study 

of the types only. For this reason, I have made only a few 

suggestions about new synonyms. I expect that the study of 

variability will lead to many more, but I also suspect that some 

current names hide more than one taxon. 

ACKNOWLEDGMENTS 

Many types came from the collections of the National Museum 

of Natural History, Smithsonian Institution (with the 

collection designation of USNM for collections of the former 

United States National Museum), but most were found in other 

collections. 

I would like to thank my colleagues in many museums for 

lending me types and for helping me track down types no 

longer deposited in the obvious collections. Just as important, 

some colleagues informed me that types of some species were 

no longer in existence or had never existed. These minutiae 

were needed to identify many of the species. The specific 

persons and museums involved include Ms. Leslie Harris and


Ms. Susan Williams, Allan Hancock Foundation, University of 

Southern California, Los Angeles (AHF); Dr. Welton L. Lee, 

California Academy of Sciences (CAS); Ms. Ardis Johnston, 

Museum of Comparative Zoology, Harvard University (MCZ); 

Dr. Willard Hartman, Peabody Museum, Yale University 

(YPM); Dr. Ernest Kirsteuer, Mr. Harold Feinberg, and Mr. 

Jerry Thurmond, American Museum of Natural History, New 

York (AMNH); Dr. Arthur E. Bogan, Academy of Natural 

Sciences, Philadelphia (ANSP); Dr. Vivianne Solis Weiss and 

Dr. Maria Elena Caso, University of Mexico; Drs. Paulo da 

Cunha Lana, E. Nonato, and A.E. Migotto, Brazil; Ms. Inger 

Winsnes and Dr. Kari Andersen, Zoological Museum, University 

of Oslo; Dr. Endre Willassen, Zoological Museum, University 

of Bergen (ZM Bergen); Dr. Tor StrOmgren, University of 

Trondheim, Museet, Trondheim; Mr. Roy OlerOd, Riksmuseet, 

Stockholm (RM); Ms. Mary Petersen and Dr. JOrgen Kirkegaard, 

Zoological Museum, Copenhagen (ZMC); Dr. Gesa 

Hartmann-Schroder, Zoologisches Museum und Institut, Hamburg 

(ZMH); Dr. G. Buzhinskaya, Akademia Nauk, Leningrad; 

Dr. G. Hartwich, Zoologisches Museum, Berlin (ZMB); Dr. 

Jadwiga Ludwig, Wroclaw; Dr. Erich Kritschcr, Zoologisches 

Museum, Wien (ZM Wien); Dr. Cl. Vaucher, Museum d'Histoire 

Naturelle, Geneve, Switzerland. Dr. Jeanne Renaud- 

Mornant, Musee National d'Histoire Naturelle, Paris (MNHN, 

Paris); Dr. J. David George and Mr. Alex Muir, British Museum 

(Natural History) (BM(NH), now the Natural History 

Museum, London); Dr. Louis Amoureux, Rennes, France; Dr. 

Y. Gillet, Universite Catholique d'Angers, France; Dr. Adriana 

Giangrande, Dipartimento di Biologia, Universita degli studi di 

Lecce, Italy; Dr. Minoru Imajima, National Science Museum, 

Tokyo; Dr. Tomoyuki Miura, Faculty of Fisheries, Kagoshima 

University, Japan; Mr. Tony Saville, Canterbury Museum, 

Christchurch, New Zealand. 

Of these, Dr. Jeanne Renaud-Mornant, Mr. Alex Muir, and 

Dr. G. Hartwich were particularly helpful tracking down specimens. 

The first two also put up with visits from me. 

The study was prompted by a request from Dr. Pat Hutchings 

of the Australian Museum for identification of eunicid species 

she had been finding on the Great Barrier Reef. This request 

lead me to spend some time in Sydney as a visiting curator at 

the Australian Museum. I am grateful for the funding provided 

through Dr. Des Griffin, Director of the Australian Museum, 

for this visit. 

Dr. Klaus Rutzler provided funding for the study of variability 

of the eunicid species associated with the reefs off Belize 

through the I MS WE, SWAMP, and CCRE programs and thus 

provided the wherewithal for assessing, if preliminarily, validity 

of the many types I have examined. 

Dr. Mary E. Rice, Smithsonian Marine Station at Link Port, 

made it possible for me to undertake extensive investigations of 

Kinbergonuphis simoni, used as one of the outgroups in the 

cladistic analysis. 

Discussions with my colleagues Drs. Meredith L. Jones, 

Marian H. Pettibone, W. Duane Hope, Mary E. Rice, and 

Frederick M. Bayer clarified several taxonomic and nomenclatural 

concepts. Talks with Drs. David E. Russell and J. Kirk 

Fitzhugh, at the time both graduate students at George Washington 

University, gave me a much needed sounding board for 

hare-brained schemes. Discussions with Drs. Richard O'Grady, 

Dan Brooks, and Vicki Funk gave me insight into the thought 

processes and theory associated with phylogenctic analysis. 

Ms. Linda Ward demonstrated great skill and patience in many 

ways in putting the study together, not least with proofing all 

the long data entry tables and preparing and checking the 

citations. 

I dedicate this study to Dr. Leonard P. Hirsch partly for his 

constant support and encouragement, but mostly for his insistent 

prodding on methodological issues. 

MATERIAL AND METHODS 

The types of nearly 200 species of Eunice arc available. The 

German collections are especially valuable, both the Berlin and 

Hamburg museums have types of many species described by 

Grube. Additional types are in the holdings of the museum in 

Wroclaw (Breslau) now in Poland, where Grubc lived and 

worked for many years. The British Museum (Natural History), 

now called the Natural History Museum, and the Musde National 

d'Histoire Naturelle, Paris, have large holdings as docs 

the National Museum of Natural History, Smithsonian Institution, 

and American Museum of Natural History, New York. 

Some authors, for example Claparcde and Rioja, never deposited 

any types. 

I have designated neotypes and lectotypes for a few species. 

Neotypes have been designated only for poorly defined, widely 

dispersed species such as E. norvegica. Most early authors did 

not designate holotypes; if they had more than one specimen 

available, these have here been considered syntypes. Only in 

cases where confusion could arise as to the identity of a species, 

as when the type lot(s) contained two or more species, have 

lectotypes and paralectotypes been designated. 

MICROSCOPES AND COMPUTER.—I used a Wild M8 

stereomicroscope and a Zeiss Universal compound microscope 

with interference optics, both equipped with camera lucida. The 

paper was prepared on an IBM PS2-80. Word-processing packages 

included NotaBene and WordPerfect 4.1 through 5.1. 

Tables and calculations were prepared using SYSTAT 4.0, 

SuperCalc4 1.0, and PlanPerfect 3.0. For the cladistic analysis 

I used PAUP 1.1. DELTA 1.1 (Dallwitz and Paine, 1986) was 

used for the preparation of the text and key and for the data 

matrix for one of the PAUP runs. The rather idiosyncratic 

punctuation used in DELTA natural language descriptions was 

modified for publication. Details of the preparation of the key 

and the cladistic analysis are given elsewhere. 

Several species have been moved to other genera. I give only 

brief comments for each of these, including a reference to the 

author responsible for the current generic disposition. New 

combinations are noted. 

Some species for which types arc missing were poorly 

described originally and have rarely if ever been found by

NUMBER 523 

subsequent workers. They may have been listed in comprehensive 

works (e.g., Fauvel, 1923:451); such mention cannot be 

considered a use of the name in the sense of the Code. These 

species are here characterized as indeterminable and are briefly 

described, with references to the original descriptions and the 

first author to declare them invalid. A few nomina nuda are also 

included. 

All other species are as extensively described and illustrated 

as possible. Where no types were available, original descriptions 

(or redescriptions of the original material) have been 

paraphrased. Type material is missing for some frequently 

reported species, such as Eunice antennata and E. vittata. In 

these cases specimens from the type locality (or as close to the 

type locality as practical) were examined in lieu of types. 

Some recently described species, for which types are missing, 

have not been reported since their original description. 

These species are characterized as best as possible from the 

original illustrations and description and are considered incompletely 

known. In most cases these species are too poorly 

known to be included in the keys or the cladistic analysis. All 

species described (other than those declared indeterminable or 

moved to other genera) are included in the tables and can thus 

be compared to related taxa. 

DISCUSSIONS OF INDIVIDUAL SPECIES.—The species are discussed 

in alphabetic order, and the treatment of each is separated 

into several sections. Lightfaced centerheads identify 

species that have been transferred to other genera; boldfaced 

centerheads identify species that I consider as belonging to 


A species number, arbitrarily designated by me, precedes 

boldfaced species centerheads. These numbers, rather than 

species names, are used in the comparison tables, making the 

tables less sprawling than they otherwise would be. Species of 

Eunice are listed in Appendix B, using the species numbers. In 

the first line following the heading are given references to 

illustrations and to tables demonstrating variability of the type 

material of the species. 

Synonyms: The synonymy lists contain references only to 

the original descriptions, to redescriptions of the type material, 

and to important changes in generic assignment. The two 

generic names Eunice and Leodice have been used synonymously 

since about 1820 and various switches from one to the 

other and back again have been omitted as trivial. A brief check 

of materials in various museums revealed that up to 50% of the 

specimens had been misidentified. Without a careful check of 

the specimens on which a given record has been based, inclusion 

of such records in the synonymy lists may perpetuate old 

errors. 

Several species have been victims of what I call a serial 

synonymy. A serial synonymy may arise as follows: Author A 

synonymizes species Y with species X. Later, author B synonymizes 

species Z with species Y. Finally, author C roots out 

this combination of synonyms from the literature and proceeds 

to lump all three named species under what she (or he) perceives 

to be the oldest available name. Very different morphs 

have been paraded under the same species name, often for 

many years, without anybody examining the types or even 

examining the types of one species and comparing them to 

descriptions of other species. Authors of widely used faunal 

surveys sometimes take serial synonyms into account when 

writing descriptions (e.g., Fauvel, 1923). The result is that 

species may become so poorly characterized that delimitation 

of any species in the genus becomes difficult. In other cases the 

descriptions encompass only one section of the species listed in 

the synonymy lists (e.g., Fauvel, 1953). Without direct comparison 

of the types of all named taxa included in one of these 

series, it is impossible to decide what parts (if any) of such a 

chain of synonyms are valid. References to obvious serial 

synonyms have not been included. 

Most beginning students of polychaetes will use one of the 

major keys or handbooks as his or her major source of information. 

As I know from practical experience (Fauchald, 1977), no 

one scientist can handle all families equally well when writing 

a handbook or a key. One has to depend on available published 

sources. In cases where the original literature is poor, the result 

is often the introduction of unjustified synonyms into the 

literature. Keys include, obviously and trivially, only those 

species considered by the author when constructing it. Species 

not considered by the author may very well key out without any 

problems and the identification still be incorrect. 

The secondary literature may cause some additional problems. 

These problems are here pointed out by the following 

example. Please note that both Fauvel (1953) and Hartman 

(1968, 1969) pointed out clearly and accurately the sources of 

their information in the publications. Problems arise in assuming 

taxonomic accuracy in these general works. 

Using Fauvel (1953) to identify polychaetes from India, the 

purpose stated in the title of Fauvel's book, may create some 

problems. For many species he assumed to be widely dispersed 

geographically, Fauvel used illustrations already issued in his 

two volumes on the French fauna (Fauvel, 1923, 1927). The 

descriptions in Fauvel (1953) are very brief. He may have 

based his descriptions on Indian specimens or he may have 

abbreviated the descriptions prepared for the Atlantic fauna to 

cover Indian records. 

Using Hartman's (1968,1969) atlas of Californian polychaetes 

poses a related problem. Nearly all Hartman's descriptions 

were based on direct examination of California material. Most 

of the illustrations, however, were copied or redrawn from 

earlier publications, usually from the original descriptions. In 

several cases, the descriptions and illustrations do not agree 

(K.Fd, personal observation). 

As it is usually not clear what aids were used for identification 

in later records of euniceans, the lists of synonyms have 

been limited to those that identify the species and the material 

used and a few well-documented synonyms. At a later date I 

will try to cope with all sources of synonyms (corrector not) for 

the group. 

Material Examined: In this section, type status of the

8 

specimens examined, as well as museum reference numbers, 

locality information, and number of specimens is given. Locality 

information is given as originally spelled either on the labels 

or in the original description. Where necessary, equivalent 

modern names have been added in parentheses. Depths of 

capture have been recalculated to the metric system. 

Comments on the Material Examined: This section includes 

comments on the state of preservation of the specimens; 

any specimens referred to other species are named, as are 

specimens specifically described or illustrated if the quality of 

the specimens differ. I have also included comments on missing 

type materials. 

Descriptions: The descriptions have been standardized, 

with all features named and described in the same sequence, 

beginning with overall body characters. Included are state of 

completeness of the specimen; sex (if identifiable); numbers of 

setigers present, total length, maximum width, length through 

setiger 10, width at setiger 10. All measurements expressed are 

in mm. In addition, the setiger at which the maximal width is 

first reached is noted. The shape of the body in general appearance 

is briefly noted. Descriptions of pygidium and anal cirri 

are given in this section. 

Next are named and described all features associated with 

prostomium and peristomium. The jaw structure, often limited 

to a maxillary formula, is given in a separate section and the 

branchiae are described in a separate paragraph. 

In the paragraph describing the parapodia, each feature is 

mentioned for anterior, median, and posterior setigers when 

possible. The ventral cirri are considered part of the neuropodia 

and are included in the description of the neuropodia. The 

notopodial cirri, in the literature referred to as the dorsal cirri, 

are here considered extensions of the notopodia and are for that 

reason described separately. 

In the section on setae, the different kinds of setae are 

described in the following order: limbate setae, pectinate setae, 

compound hooks (and compound spinigers when present), 

aciculae, and finally subacicular hooks. 

I have attempted to account for features not included in a 

description, either in terms of actual absence or because the 

type material is incomplete or in too poor condition to discern 

a given structure. 

Several types have fragmented over time or were originally 

collected in several pieces. I have consistently assumed that the 

anterior end represents the individual originally described and 

that all other fragments, although they probably belong to the 

species, may belong to other individuals or even species collected 

at the same time. The problem is particularly serious 

when material has been collected by dredging, where one may 

get representatives of several species, and it may be difficult to 

associate anterior and posterior ends (cf. Eunice torresiensis 

and E. tribranchiata below). If the type has been cut into pieces 

that clearly match, I have assumed that the pieces are part of a 

single specimen. 

Caution was taken to avoid damaging specimens. Single 


type specimens were not dissected to examine the jaws, even if 

the descriptions thereby remain incomplete. 

The parapodia used for illustrations were, as much as possible, 

appended to the types in small glass vials to be available 

for future study. 

The following four items are listed separately from the 

description to ensure that they are not considered part of the 

description itself. 

Unknown Morphological Features: In this section is given 

a brief summary of major morphological features left undescribed, 

usually because of the fragmentary nature of the types. 

Expected States of Selected Unknown Features: For some 

species a few predictions of the expected state of various 

unknown characters can be given. These predictions were made 

to stimulate a closer examination of specimens. 

The next two features indicate the status of each character, 

using the character table. 

Character States Unknown: "Inappropriate characters" are 

those characters that cannot under any circumstances be scored 

for the species in question; for example, all features associated 

with branchiae in an abranchiate species. "Unknown characters" 

arc those characters that should have been scored but 

which had to be left out, usually because of the incomplete state 

of the type. 

Assumed States Used in Preparing Key: Many species 

were described from anterior fragments only; some characters, 

such as the posterior branchial distribution, are thus unavailable. 

Because knowledge of the branchial distribution is crucial 

for creating a reasonably accurate key, I have assumed a state 

for the characters listed. The assumed states cannot be verified 

on the types; a study of additional material from the type area 

may test the likelihood of the assumption. In theory, as the 

holotype is the only specimen that is the carrier of a particular 

name, verification is impossible, but for all biological, rather 

than nomenclatural purposes, the character states can be 

demonstrated, based on the study of fresh material. Most of the 

assumed states follow traditions in the literature; where nothing 

else is stated, I follow suggestions in Hartman (1944) or 

Fauchald (1970). 

Remarks: This section includes a few comments on similar 

species and reference to the appropriate comparison table(s). 

Where appropriate I have also specified differences between 

the original and the current descriptions. Remarks about each 

species have been kept brief because most of the comparisons 

to similar species can be done through the tables. 

Illustrations: The anterior ends are drawn in lateral view. I 

adopted the convention of not drawing appendages from the far 

side of the worm in order to avoid cluttering illustrations of 

species with long notopodial cirri and branchiae. Parapodial 

and setal features are illustrated; where necessary, parapodia 

from both the anterior and median-posterior end were removed 

and illustrated. The parapodia were drawn in full anterior view 

as mounted on a microscopic slide. I have adopted the following 

conventions when drawing setal forms in the parapodial

NUMBER 523 

illustrations. Seial length is indicated as accurately as possible; 

the number of setae drawn bears no relation to the number of 

different setae present. Limbate setae are rendered as single 

lines, pectinate setae as short single lines in the same fascicle as 

limbate setae. In compound falcigers and spinigers, the joints 

between appendages and shafts are indicated. 

In poorly preserved specimens only the setae are illustrated; 

soft parts are usually so badly distorted that illustrations would 

be misleading. 

The jaws, when illustrated, were drawn in dorsal view as 

flattened on a microscopic slide. The shapes of supporting 

plates are thereby distorted, but I preferred on this occasion to 

follow the convention first apparently adopted by Schmarda 

(1861) in illustrating eunicid jaws. 

Tables: When the type material consisted of at least three 

specimens, the variability of all characters is presented in a 

series of tables. Each table consists of two parts. The upper part 

gives ranges, means, and standard deviations of measurements 

and presents variability in numerically characterized features. 

The lower half is a list of invariable features. Where possible, 

both incomplete and complete specimens have been included. 

Where only incomplete specimens were available, the only 

length measure used was the length through setiger 10 ("Length 

through 10"). 

Comparison Tables: Because the cladistic analysis gave 

no readily interpretable results, I decided to include a series of 

tables comparing "similar" species as aids for identification. 

These tables are based on species groupings first suggested by 

Ehlers (1868) and codified in Hartman (1944) and Fauchald 

(1970). I would like to emphasize that even though eventual 

cladistic analysis may show that some (or all) of these groups 

represent phylogenetic lines, phylogenetic conclusions cannot 

be drawn from membership in a group at this time. The groups 

are artificial aids for the identification of the species, nothing 

more, nothing less. 

The comparison tables are grouped hierarchically so that 

each major group table (Tables 19, 22, 24, 27, 33, 40, 41, 46, 

50, and 52) has been broken down into suitable smaller tables. 

The characters treated in the subordinate tables are exclusively 

presence-absence characters. 

Not all characters included in the character table were included 

in the comparisons, and the characters used vary from 

one table to the next. Characters are listed in the left-hand 

column. The other columns show the character state for each 

character of a single species; each column is headed by a 

species number. The character numbers and states are listed in 

the character table. As noted above, each species was given a 

species number (another consequence of the use of the DELTA 

format). The species number for each species precedes the 

name in the systematic treatment. The species numbers included 

in each table are also listed at the bottom of each 

comparison table. 

Note that if one of the characters used in the breakdown of 

the genus is unknown for a species, this species is listed in all 

tables to which it might belong. Some anomalies arise as a 

consequence: Eunice prayensis has been listed in four subtables 

and two major tables: poorly known species will turn up 

in more tables than well-known ones. 

Morphological Features of the Eunicids 

This review of eunicid morphology, with special reference to 

the nominal genus Eunice, is divided into sections corresponding 

to major body features. Characters and character states used 

in the cladistic analysis and in preparation of the key are based 

on the morphological structures described herein, but are detailed 

in the section on cladistic analysis. The main structures 

are diagrammed in Figure 1. The left- and right-hand diagrams 

show the same "specimen" in dorsal and left lateral view; the 

three median diagrams illustrate a pre-branchial, a branchial, 

and a postbranchial parapodium of the same "specimen" in 

anterior view. 

OVERALL BODY FEATURES AND POSTERIOR END 

Eunicids are relatively slender-bodied polychaetes with short 

parapodia. The parapodia gradually change shape along the 

body and no major segmental body regions are recognized. 

Three different body shapes may be recognized, based on 

observations in full dorsal view. In all three, the anterior end 

tapers very little and the prostomium appears truncate. 

1. The widest part of the body is at about setiger 10, 

corresponding to the position of the posterior end of the inverted 

jaw apparatus; the body tapers evenly and very gradually 

towards the posterior end from that point. This body shape is 

illustrated in Figure 1. 

2. The maximum body width, reached at about setiger 10, is 

retained through most of the rest of the body; body tapering is 

relatively abrupt in a posterior end consisting of perhaps 20 

segments. 

3. Maximum body width occurs in a wide, usually flattened 

region in the posterior one-third of the body. The rest of the 

body is cylindrical or slightly tapering from the posterior end of 

the inverted jaw apparatus at about setiger 10. The latter body 

shape is most common in species of Marphysa, but is present in 

several species of Eunice. 

The anterior end is always very muscular. The muscles are 

mainly used to support and move the large, protrusile jaw 

apparatus (for more detail see below). In cross section each 

anterior segment is usually distinctly convex dorsally. The 

ventrum is less distinctly convex than the dorsum or it may 

appear flattened or the ventral nerve cord may be located in a 

groove between paired longitudinal muscles. The paired ventrolateral 

longitudinal muscles usually form distinct longitudinal 

bands continuing at least through the first one-third of 

the body. The bands fade posteriorly and are indistinct in the 

posterior half of the body in most species. The middle part of 

the body region is less muscular than the anterior end. In cross 

section the middle body appears flattened, most usually ven-

10 


First branchia 

Presetal lobe 

Acicular lobe 

Notopodial cirrus 

Postsetal lobe 

Ventral ctrrus 

Antenna 

J 

Ceratostyle 

Ceratophore 

Anterior ring 

•Posterior ring 

Pensiomial cirrus 

First branchia- 

PARAPODIUM 2 

Branchial tilament 

Branchia - 

Branchial stem 


Last branchia 

Notopodial acicula 

Neuropodiai acicula 

Limbale and pectinate setae 

Compound talciger 

Last branchiar 

ree tip ot ventral cirrus 

PARAPODIUM 12 

Inflated base of ventral cirrus 


Acicula 

Subacicular hook 

•Limbate and pectinate setae 

Compound falciger 

Pygidium 

Ventral cirrus 

Superior (short) anal cirrus- 

PARAPODIUM 30 

Infenor (long) anal cirrus - 

FIGURE 1.—Diagram of Eunice showing location of various morphological features. 

trally, but often also dorsally. The posterior end is often nearly 

circular in cross section, but especially in species of body type 

3 can be strongly dorsoventrally flattened. 

The color of live specimens is usually highly characteristic. 

The most frequent color pattern, especially among species 

similar to Eunice aphroditois, is a dark, purplish red, with 

white or pale gray dots scattered over the surface. One, 

sometimes two, anterior sctigers, most usually including setiger 

4, has a transverse white dorsal band. Large specimens may be 

purplish black with a greenish iridescence and the light-colored 

spots or bands present on small to medium-sized specimens 

may be difficult to see. Species related to E. antennata are brick 

red, with minute white dots, but may also be uniformly bright 

scarlet without any markings. Species related to E. vitiata often 

have a white base color with yellow or ocher cross bars and 

spots, but rarely have the white dots and bars that are present in 

other species. Colors fade rapidly in preservation and have been 

poorly described for most species. Treadwell (1921) gave good 

color notes for all species observed by him near the Dry 

Tortugas Islands in the Gulf of Mexico. All of his types are now

NUMBER 523 11 

uniformly yellowish gray. 

Colors and color patterns are clearly inconvenient as 

systematic characters, but appear to show very distinct 

systematic patterns. Color notes will be taken in the ongoing 

study of variability of eunicids, and color patterns will be 

introduced formally as systematic characters at a future date. 

Most species consist of 100-200 segments as adults (defined 

as individuals containing recognizable sexual products), but 

adults with as few as 35 and as many as 1500 segments are 

known. In specimens with few segments, each segment is 

usually just slightly wider than long, in dorsal view. In large 

specimens, segments are often much wider than long. In 

species with an expanded posterior end, segments are usually 

very much wider than long, appearing crowded in the posterior 

one-third of the body. 

Sexually mature individuals vary in length from about 5 mm 

to about 6000 mm. Most specimens examined during this study 

arc 50-150 mm in length. 

Eunicids have two pairs of anal cirri. The upper pair, actually 

located laterally on the pygidium, is usually shorter than the 

width of the pygidium and filiform in all species. The lower 

pair, whose base forms the ventrolateral rim of the anus, varies 

in length. In some species the lower anal cirri are short, barely 

reaching beyond the posteriormost five setigers and the 

pygidium proper; in other species they may outreach 15-20 

setigers. The anal cirri are incomplete in most types examined, 

so the length of the anal cirri may be considerably more 

variable than indicated herein. Long anal cirri are always 

basally inflated. Distally, cirri are perhaps most frequently 

tapering, but may also be medially inflated or digitiform, with 

or without articulations. Articulations, when present, may be 

cylindrical or moniliform. Species with moniliform articulations 

in the anal cirri also have moniliform articulations in the 

ceratostyles. The dorsal edge of the pygidium may be smooth 

or crenulated. Far too few observations have been made to give 

any particular systematic or taxonomic significance to this 

feature at this time. 

PROSTOMIUM AND PERISTOMIUM 

Most eunicid prostomia consist of two lobes in tandem. The 

bilobed nature of the prostomium may be visible only as a 

ventral longitudinal groove or suture, but is usually also visible 

dorsally as a median sulcus of varying length. In some species 

(e.g., Eunice kinbergi, Figure 54d), each prostomial lobe is 

separated longitudinally into a narrow, high medial ridge and 

paired wider, lower, lateral regions. Palps may be marked off 

by shallow, transverse or circular grooves on the anteroventral 

side of the prostomium. The palpal grooves are far more visible 

in preserved than in live material. Even when not distinctly set 

off from the rest of the prostomium, palpal regions are usually 

raised above the rest of the anteroventral surface of the 

prostomium. 

In small specimens, the prostomium may be just slightly 

narrower than the peristomium, but is usually distinctly 

narrower ("distinctly narrower" defined as one-third narrower 

than the peristomium). The prostomium may be partially 

withdrawn under an anterior peristomial fold on the dorsal side; 

this is most obvious in forms with a narrow prostomium. The 

prostomium is usually distinctly shorter than the peristomium, 

when viewed dorsally, but the two may be of similar length. 

The prostomium is usually held more or less horizontally, but 

in some species the orientation appears to have shifted, and so 

the two prostomial lobes slope strongly ventrally. The 

prostomial orientation has rarely been mentioned in the 

literature and has yet to be adequately documented. 

The dorsoventral dimension of the prostomia is here defined 

as the prostomial depth. Some species have very shallow 

prostomia as viewed from the side, being less than half as deep 

as the peristomium. In other species, the prostomium may be 

much deeper and may be very nearly as deep as the 

peristomium. The depth of the peristomium has consequences 

for distortions in shape associated with protrusion of the jaws. 

In species with a shallow prostomium, the jaws are protruded 

very nearly directly anteriorly underneath the prostomium; in 

species with a deeper prostomium, the prostomium is tilted 

back to allow eversion of the jaws. The jaws in these cases are 

everted anteroventrally, rather than strictly anteriorly. The 

difference in direction of eversion may have consequences for 

feeding behavior and burrowing activity. 

Eyes, when present, are found posterolaterally on the dorsal 

side of the prostomium. They are paired, ovate, and vary in 

color from bright red through purple to black; the most 

common color is dark purple. Eye colors fade in preservation, 

making them useful characters only when recorded from fresh 

specimens. The eyes may be located in line with (between) the 

antennae, or distinctly outside the line of the antennae. In some 

species the eyes are found on the ceratophores of A-I or A-II 

(see below for a definition of these terms). All eyes are located 

directly on the dorsal surface of the anterior part of the cerebral 

ganglion, overlain by translucent tissue. Four small eyespots 

have been reported present in a few species (e.g., Eunice 

americana. Figure 8a; E. gravieri, not illustrated). Eyes are 

absent in several species. 

The five antennae are situated in an occipital crescent or in a 

transverse row. The outer lateral antennae (A-I) are often 

shorter than the other antennae. The inner lateral antennae 

(A-II) and the unpaired median antenna (A-III) may be similar 

in length, but, perhaps most frequently, A-III is longer than all 

other antennae. In some species all antennae are short, barely 

reaching the tip of the prostomia, with thick, sausage-shaped 

styles. The spacing of the antennae and the thickness of the 

antennal styles (ceratostyles) vary among species. The antennal 

bases (ceratophores) are short and ring-shaped in most species, 

but may also be short, cylindrical structures confluent with the 

prostomial surface. The ceratophores lack articulations except 

in a single species (Eunice unidentata, not illustrated). The 

shape and structure of the ceratophores appear to be stable

12 


features within any given species. 

The ceratostyles may have cylindrical or moniliform 

articulations, or may lack articulations. The kinds of articulations 

present may vary among the antennae and from base to tip 

in the same antenna. The shape of what is herein referred to as 

a "moniliform articulation" may vary from drop-shaped 

through rounded quadrangular, to nearly triangular with the 

widest edge distally. Whether or not these differences arc 

fixation artifacts has yet to be determined. Casual observations 

on live material indicate that shapes do not change upon 

fixation, but documentation is incomplete. 

The peristomium forms a fold covering the base of the 

prostomium dorsally (see above). The pocket formed by this 

fold is deepest near the midline in live animals. The bases of 

antennae and eyes may be covered by the pcristomial fold, both 

in live and preserved specimens. Laterally, the peristomial fold 

terminates in an ear-shaped fold. Ventral to this fold, the 

separation between the prostomium and peristomium is 

indistinct externally for a short distance. The antcrovcntral part 

of the peristomium is a more or less scoop-shaped lower lip. 

The lip may consist of paired, inflated, strongly muscular 

cushions, distinctly set off from the rest of the peristomium by 

a shallow groove. The cushions are usually separated in the 

ventral midline by a frontal notch and a mid-ventral, poorly 

muscularized region. The lower lip may also be relatively 

poorly muscularized; if this is the case, the peristomium will 

taper anteriorly. 

The peristomium is nearly always separated into two rings. 

The rings are usually separated by dorsal and ventral grooves or 

by a groove encircling the peristomium. The separation may be 

visible only on either the dorsal or the ventral side; in a few 

cases, the separation is marked only as shallow grooves anterior 

to the bases of the peristomial cirri. The rings are ontogenetically 

presegmental in origin and do not represent fused 

segments (Akesson, 1967). 

Paired, dorsolateral peristomial cirri are located near the 

anterior edge of the posterior peristomial ring. The cirri vary in 

shape from short, ovate structures barely outreaching the 

posterior peristomial rings, to long, slender, tapering structures 

reaching well beyond the tip of the prostomium. They may be 

articulated, usually with cylindrical articulations. The peristomial 

cirri are articulated only in species in which the 

ceratostyles are articulated, but are not articulated in all species 

with articulated ceratostyles. 

JAW APPARATUS 

The eversible jaw apparatus consists of paired ventrally 

located mandibles and four pairs of lateral maxillae in addition 

to an unpaired plate on the left side (seen from above); a fifth 

pair of sclerotinized plates is present; these plates nearly always 

lack teeth, but may have a single sharpened ridge in a few 

species. The maxillae are diagrammed as seen slightly 

compressed in dorsal view in Figure 2. 

Mx- 

Left 

Right 

/Mx-IV 

/ yMx-V 

Maxillary 

carrier 

FlOURE 2.—Diagram of maxillae of Eunice showing the numbering system of 

the various pans. Mx-VI is absent in most species. The maxillary fonnula for 

this set of jaws would be 1+1. 8+9. 12+0.6+11. 1+1. and 1+1. Mx-III in this 

instance is long and located behind left Mx II. 

The mandibles are narrow and expanded anteriorly as a 

cutting edge, which may be impregnated with CaCO 3 . In the 

genus Palola the mandibles are scoop-shaped and enclose the 

maxillae when retracted. In other genera, mandibles are flat, but 

may be tilted in relation to each other to form a shallow V. 

The maxillae are attached to longitudinal, muscular ridges 

arranged on both sides of the eversible pharynx. Most of the 

muscle mass of the pharyngeal bulb is associated with the 

maxillae. Relatively less bulky muscles are used in retracting 

the apparatus. Protrusion is apparently mainly a function of a 

contraction of the body muscles of the whole anterior end (cf. 

Clark, 1964; Wolf, 1976). The dorsalmost pair of maxillae 

(Mx-I) are large, curved structures in a forceps-shaped 

arrangement. They are basally attached a pair of wide, thin, 

posteriorly tapering, usually short, maxillary carriers. Remaining 

jaw elements are numbered in order progressing ventrally 

and anteriorly from Mx-I. Each second maxilla (Mx-II) is a 

large plate, the base of which is folded at approximately right 

angles over a muscular ridge. The teeth of Mx-II are located on 

the convex edge and curve posteriad. The number of teeth 

varies from three to about 15; the number of teeth is largely 

independent of the size of the specimens. An Mx-III is present 

only on the left side. It may be located directly below Mx-II or 

under and in front of left Mx-II. Mx-III either forms a transition

NUMBER 523 13 

to the frontal jaws (Mx-IV and V) or is part of an arc formed by 

the frontal jaws. When located behind Mx-II it resembles Mx-II 

in that it is an angled plate with teeth along the free edge. When 

located in front of Mx-II, the Mx-III base plate is smaller and 

the outer edge is curved, usually fitting into the curve of the left 

Mx-IV. The number of teeth is usually much lower in the 

second kind and the base plate is often tucked underneath the 

left Mx-IV. The shape of Mx-III and its relation to other 

maxillae are probably taxonomically informative; the topic is 

currently being pursued. Left and right Mx-IV consist of 

angled, comma-shaped plates with teeth along the cutting edge. 

The right Mx-IV usually has more teeth than the left one. In 

cases where Mx-III forms part of a distal arc with Mx-IV, the 

combined number of teeth in Mx-III and left Mx-IV often 

approximates the number present in right Mx-IV. The number 

of teeth vary from 2 or 3 to about 15 in each of Mx-III and IV. 

Mx-V is a small plate lateral to each Mx-IV and usually with a 

single tooth; in a few cases several teeth may be present. Mx-V 

may be asymmetric. Lateral to Mx-V may be found another 

sclcrotinized piece, which, when present, may have a small 

tooth and is then referred to as Mx-VI. 

BRANCHIAE 

Branchiae are present in most species. They usually are 

located on the dorsal edge of the notopodia near the base, but 

may emerge, especially in posterior setigers, from the body 

wall dorsal to the notopodial bases. Branchiae are readily 

differentiated from notopodial cirri by the presence of a 

vascular loop, visible in most cases with the use of a high 

power stereo microscope in situ. Minute branchial capillaries 

loop between the epidermal cells and are visible in optical 

section as minute punctae lined up just below the surface of the 

branchiae. 

Branchiae may consist of single filaments, or they may be 

more or less branched. Where best developed, branchiae are 

pectinate, with long, usually tapering stems and more than 40 

filaments arranged in a single comb on the dorsolateral side of 

the shaft Miura (1986) demonstrated that in most species the 

maximum number of filaments is seen just posterior to the start 

of branchiae, with the number of filaments tapering off towards 

the posterior end. In some species, an intermediate region of 

low numbers of filaments is present. In a set of diagrams 

showing the numbers of branchiae per segment, Miura 

demonstrated that patterns of branchial distribution are more or 

less species specific. Potential variability in these patterns are 

now under study as part of the study of variability within each 

species. In many species, branchiae are limited to the anterior 

one-third to one-half of the body, but branchiae often are 

present from setiger 5-10 to the posteriormost distinct 

segments. 

The length of the branchial stems varies. When the stems are 

long in relation to the filaments, branchiae appear pectinate; 

when the shafts are short, branchiae have a palmate appearance. 

In species with palmate branchiae, the maximum number of 

filaments is usually only two or three. In some species, the 

branchial stems may be bent, or be slightly coiled, resulting in 

some rather unusual shapes. The basic structure of all branchiae 

are the same: a stem with a single series of filaments attached 

along one side. In some species filaments may themselves be 

branching; these secondary branching patterns are irregular, 

varying from one segment to the next. Only a few species are 

prone to secondary branching (e.g., Eunice johnsoni, Figure 

59j). 

Most species have an anterior and posterior region with 

single filaments, even if the branchiae are strongly pectinate 

elsewhere. The number of setigers with single filaments 

anteriorly and posteriorly appear to vary without any distinct 

patterns in some species; in other species distinct patterns may 

be present. 

In most species the number of branchial filaments decrease 

monotonously after reaching a maximum somewhere in the 

anterior end of the body. In some species with branchiae 

continued to one of the last distinct segments, the number of 

filaments decrease to one or two in a mid-body region and 

increase again, usually to three or four in the posterior one-third 

of the body. The number of filaments decreases to a single 

filament in the last 10-15 prepygidial segments. The presence 

of a mid-body region with reduced branchiae is nearly always 

associated with the presence of tridentate yellow subacicular 

hooks, but not entirely so, and certainly not all species with 

such hooks have reduced number of filaments in a mid-body 

region. 

In some species, I have observed an increase in filament 

length toward the posterior end without an increase in the 

number of filaments. This feature has not been well documented 

on the types and has been omitted in the present study, 

but will be taken up as part of the study of variability mentioned 

above. 

In most species, filaments are digitiform. More rarely, 

filaments are smoothly tapering, whereas in some species, 

especially in those with single branchiae, they may be flattened, 

often nearly foliose with abruptly tapering tips. Fauchald 

(1991) found that single branchial filaments in juveniles often 

are flattened, whereas fully pectinate branchiae of adults of the 

same species have digitiform filaments. 

PARA PODIA 

Eunicid para podia are biramous. The notopodia are represented 

by short bases and notopodial cirri. The notopodia are 

supported by internal aciculae (Stop-Bowitz (1987:128) 

pointed out that the Latin term acicula, plural aciculae, is 

feminine). The notopodia are set off from the cirri by a distinct 

groove in some species (e.g., Eunice pennata, Figure 87p; £. 

petersi, Figure 89e), but in most instances the notopodia are 

separated from the cirri only by the distinctly to slightly 

expanded diameter of the cirri near their junction to the

14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

notopodia. The notopodial aciculae, usually two or three in 

number, are slender, usually slightly yellow to clear, pliable 

rods; in some species one or more of the aciculae may be dark 

brown to black. Notopodial aciculae, being difficult to see in 

most species, have been reported only sporadically, but are 

apparently always present. The notopodial cirri may be 

articulated, especially in the anterior end of the body. The 

articulations are usually cylindrical, more rarely drop-shaped or 

moniliform. The first notopodial cirri are in some cases much 

longer than those in the following setigers (e.g., Eunice 

polybranchia. Figure 91a). More usually, cirri increase in 

length through the first 10-15 setigers and decrease monotonously 

in length posteriorly. In E. aphroditois (Figure 13c,d) 

and similar species, notopodial cirri are strongly inflated 

medially, often appearing rather flaccid. Notopodial cirri of this 

type retain the same length towards the posterior end. Thus, 

because the body narrows, and other parapodial features 

become reduced, these notopodial cirri become the dominant 

parapodial feature near the posterior end. 

The neuropodia are herein described as they appear mounted 

on microscopic slides with the anterior side facing the observer. 

Anterior neuropodial acicular lobes arc usually rounded or 

truncate, supported by or more aciculae. The aciculae may 

emerge near the middle of the acicular lobe or dorsal to the 

midline. In some species, the acicular lobes become progressively 

reduced posteriorly and the aciculae may emerge directly 

from the body wall. In most species, the acicular lobes retain 

roughly the same size relative to the width of the body 

throughout, but become conical or triangular towards the 

posterior end. 

The pre- and postsetal lobes actually comprise a continuous 

structure around the dorsal edge of the neuropodial acicular 

lobes, wrapping around the dorsal edge as a flattened collar, 

covering the bases of the setae. The appearance of the anterior 

and posterior part of this collar is nearly always so different that 

it is most usefully described as if it consisted of separate 

pre- and postsetal lobes. The presetal lobes are usually 

distinctly shorter than the acicular lobes, and form low, usually 

transverse, folds covering the bases of the compound falcigers 

and spinigers. In some species the presetal lobes are as high as 

the acicular lobes and closely follow the outline of the acicular 

lobes. The presetal lobes rarely change appreciably in shape 

along the body. The postsetal lobes are more variable. In some 

species, the anterior postsetal lobes outreach the acicular lobes 

to form a projecting, triangular or rounded lobe. The high part 

of this lobe may be dorsal to, directly behind, or ventral to the 

high point of the acicular lobes. In most species the anterior 

postsetal lobes are either low, transverse folds or follow the 

outline of the acicular lobes closely; in neither case will the 

postsetal lobes be visible in a parapodium mounted in anterior 

view. In median and posterior setigers the postsetal lobes are 

low, transverse folds or follow the outline of the acicular lobes 

closely in nearly all species. 

Anterior, usually prcbranchial, ventral cirri are tapering or 

conical; rarely slender or digitiform. The bases of the ventral 

cirri are inflated and glandular in the next 30-50 setigers in 

nearly all species examined. The distribution, size, and shape of 

the inflated bases vary among species. Most commonly, 

inflated bases are limited to 30-50 seligers and are ovate to 

spherical. The tips of the ventral cirri project as tapering, 

sometimes truncate or digitiform, tips. In posterior setigers, the 

inflated bases usually are gradually reduced. In some species, 

the bases are withdrawn into the body wall and the glandular 

tissues are retained but no longer form a perceptible bulge. 

Posterior ventral cirri usually become longer and more slender 

than those in the antcriormost setigers, but rarely become as 

long as the notopodial cirri in the same setigers. In some 

species the posterior ventral cirri emerge on the posterior face 

of the parapodia and project dorsally, behind the postseial 

lobes. In species in which the inflated bases form thick, 

transverse welts along the ventral edge of the parapodia, the 

inflated bases arc often present also in far posterior setigers. 

SETAE 

All setae arc neuropodial in origin. Limbalc and pectinate 

setae are found in dorso-postcrior fascicles forming bundles; 

compound falcigers and spinigers are in antcrovcntral, flattened, 

often fan-shaped fascicles. Subacicular hooks are also 

present and each ncuropodium is supported by one or more 

aciculae. In a few species (e.g., Eunice afuerensis. Figure 

7h,jjc; E. pelamidis, Figure 86i) compound falcigers are 

replaced by pseudocompound falcigers from setiger 25-50. 

Limbate setae are usually slightly curved, tapering and have, 

when viewed with light microscope, a single, usually narrow, 

limbation. Limbate setae outreach all other setae, and may be 

slender or thick shafted, and marginally smooth or serrated. 

They usually decrease in number from anterior to posterior 

setigers and may be wholly absent in the posterior one-third of 

the body. 

The distal half of each limbate seta consists of a core and an 

asymmetric hood formed by a halo of setal fibers (cf. Kryvi and 

SOrvig, 1990). The inappropriate term "limbate setae" is 

retained for two reasons: it is the appearance of the setae in the 

light-microscope, and it is the term used in the taxonomic 

literature. 

Pectinate setae are present in fascicles of limbate setae. They 

are usually slender and less than J /2 as long as the limbate setae. 

Each pectinate seta consists of a shaft, sometimes flattened, 

which is distally expanded into a variably wide, dentate blade. 

The blade may be distinctly furled (Figure 32d), forming an 

open, shallow scoop, or be flat (Figure 6b,g). In a few species, 

the edge of the blade is slightly oblique, but in most species it 

is at right angles with the shaft. The number of teeth along the 

edge varies between five and 30, the most usual number is from 

10 to 15. The two outermost (marginal) teeth may be thicker 

than the other teeth (Figure 20d), or they may be as thick as the 

other teeth (Figure 6b). One or both of the marginal teeth may

NUMBER 523 15 

be longer than the other teeth (compare Figures 13a, 5h, and 

9c). All teeth may be wide and abruptly tapering distally 

(Figure 21c) or evenly tapering from the bases (Figure 5h). The 

number of pectinate setae usually increases from anterior to 

posterior setigers; in some species, pectinate setae appear to be 

absent in the first 15-20 setigers. 

Compound falcigers are usually distinctly narrower than 

aciculae or subacicular hooks, but in some species they are as 

thick as the latter. Falcigers are composed of shafts and 

appendages. The shaft is usually slender, but often expanded 

distally to a distinct head (compare Figures 23a and 4e). Shaft 

heads may be marginally dentate or serrated. Shafts are usually 

clear, nearly translucent, but may become more strongly 

sclerotinized, especially towards the bases, which then become 

chestnut, brown, or even black in color (e.g., Eunice sebastiani). 

The appendages may be short (about as long as the 

distally expanded head of the shafts, Figure 4e) or they may be 

distinctly longer, becoming in some instances, very long and 

narrow (Figure 8b). The appendages may distinctly taper 

towards the tip (Figure 8b) or have nearly to completely parallel 

sides (Figure 12f). Distally, they arc usually bi- or, rarely, 

tridentate. In bi- and tridentate falcigers (Figures 4e and 5e), the 

teeth are in a row, with one above the other. The teeth may be 

nearly erect or variably curved and differ in size from barely 

noticeable to long and pointed. Reduction in size of the teeth is 

most common in the proximal teeth. Distally the appendages 

are covered by a pair of guards. The guards usually fit closely 

around the appendages, but are very much wider in a few 

species. Each guard consists of a convex membrane, which 

distally may be rounded or bluntly to sharply pointed (Figures 

3b, 4e, 5e). The guards may be equipped with distinct mucros 

(Figure 130- Such mucros are mostly present in forms with 

bluntly to sharply pointed guards, but may also be present in 

forms with rounded guards (Figure 16f,i). The edge facing 

away from the teeth may be slightly thickened and the other, 

cutting edge may be serrated. 

Pseudocompound falcigers (Figure 7h, j,k; Figure 86h,i) have 

thick shafts that are barely inflated near the reduced joint. The 

joint may be recognizable only as a swelling on one side of the 

falciger or be more distinct. The pseudoappendages are short, 

with nearly parallel sides and bidentate. Both teeth are directed 

distally and tapering. 

Shafts of the compound spinigers closely resemble those of 

the falcigers (Figure 57c). Appendages (blades) vary in length 

and are always knife-edged; the narrower edge may be serrated. 

Each neuropodium may be supported by a single acicula; 

perhaps most common are paired aciculae, but up to about 

seven have been recorded (e.g., Eunice manihine, four in Figure 

69e). When multiple aciculae are present, they are arranged in 

a dorsoventral row. The aciculae usually are thicker than all 

other setae. The number and shape may vary from anterior to 

posterior end. Multiple aciculae are more common in anterior 

than in posterior setigers. Most species have tapering, distally 

bluntly pointed, straight aciculae; some species have distally 

hammer-headed or irregularly thickened and truncated aciculae. 

In some species, aciculae are distally bifid. Gently curved, 

or geniculate aciculae are common. Aciculae from a single 

neuropodium may differ in shape. 

Subacicular hooks are exclusively present in the neuropodia 

and are first present from setigers 15-35 in most species. 

Hooks are usually present in all remaining setigers. Hooks 

originate near the base of the aciculae, but form a distinct angle 

relative to the aciculae (Figure 3e) and emerge along the lower 

edge of the neuropodia. Subacicular hooks are usually slightly 

thinner than the aciculae and are often somewhat lighter in 

color. Most species have a single subacicular hook per 

parapodium, but in some species a vertical series of as many as 

5 may be present (e.g., Eunice vittata, three in Figure 114e). 

Distally, subacicular hooks may be bidentate (Figures 3d and 

7c,h), tridentate (Figures 4d, 5c,d) or simple and spine-like 

(Figure 103k). In some species the two distal teeth in the 

tridentate hooks are arranged in tandem (e.g., E. cirrobranchiata, 

Figure 31e; E.flaccida, Figure 46c), but in most cases 

the teeth are crested (i.e., with the tips of the teeth in a single 

plane parallel to the curvature of the setal axis, e.g., Figures 4d, 

5c,d). 

Structurally, aciculae and subacicular hooks are similar 

when viewed with a light microscope; when any internal 

structure is visible, they appear to consist of an inner core of 

dense fibers, covered by a non-fibrous sheath. In some species, 

the separation between core and sheath is distinct, with a 

dark-colored core and a clear, often translucent sheath. In other 

species the separation between core and sheath is indistinct. 

Aciculae and subacicular hooks may be clear and nearly 

colorless, yellow, chestnut colored, various shades of brown, 

and in some instances a very distinct jet black. If paired 

aciculae are present, they are often of different colors. If paired 

subacicular hooks are present, they usually have the same 

color, but replacement subacicular hooks that have yet to 

emerge are often lighter in color. Aciculae and hooks in anterior 

setigers are often lighter in color then those farther back, and 

smaller specimens tend to have more light-colored aciculae 

than do larger specimens. 

Cladistic Analysis 

Species of Eunice have for the last 50 years organized into 

informal species groups (Hartman, 1944; Fauchald, 1970). A 

cladistic analysis was performed to test if these informal groups 

represented monophyletic, possibly definable taxa. The analysis 

was done using PAUP 2.4 on an IBM PS-2/80 and the 

corresponding mainframe version on an IBM 4381. 

SELECTION OF OUTGROUP TAXA 

This initial analysis was intended to justify selection of 

outgroups to be used in this study. A complete analysis of the 

order Eunicoidea is beyond the scope of this study. The


relations proposed here follow the traditional family subdivisions 

of the order and are those that appear most probable to me 

at this point. I invite testing of all assumptions made in the 

following section. 

The eversible pharyngeal structures in polychactcs are 

stomodeal invaginations. Such invaginations may take one of 

two major forms (cf. Dales, 1962). They may be axial structures 

with muscle layers arranged reasonably evenly on all sides or at 

most with narrow dorsal and ventral gaps in the musculature. 

Alternatively, they may be muscularized lower lips, in which 

the dorsal side of the stomodeum is poorly muscularized. The 

euniceans have a distinctive form of the latter construction. 

The jaws of all polychaetes are epidermal structures, formed 

either superficially by the epidermis as part of the cuticle or in 

invaginated epidermal pockets. The main constituents are 

sclero-proteins of varying density impregnated with a variety of 

materials. Eunicoidean jaws often are impregnated with large 

quantities of calcium carbonate (Colbath, 1986, and references 

therein). 

Members of the order Eunicoidca arc uniquely characterized 

by having an eversible, muscular lower lip in the sense of Dales 

(1962), supporting laterally arranged maxillae and a pair of 

ventral mandibles. The muscular part of the stomodeum is 

scoop-shaped with the jaws located in partially isolated pockets 

formed by folds in the scoop. The mandibles are in a midvcntral 

pocket. The maxillae are supported on lateral muscular ridges 

and pockets along the sides of the mouth cavity. The 

mandibles, although probably of great interest, have been 

relatively poorly described in most taxonomic work and are 

herein not considered further for the purposes of selecting an 

outgroup. 

The eunicoidean jaws, although reduced in some, especially 

small or parasitic members, differ from the jaws of other 

polychaetes, especially of the Phyllodocida, in which the jaws 

are formed along the inner lining of an axial, muscular 

eversible pharynx (cf. Dales, 1962; Clark, 1964; Fauchald, 

1974a). Without anticipating a planned analysis of the relations 

among the major groups of the polychaetes, it appears likely 

that the structure of the jaws could be considered a synapomorphy 

for Eunicoidea. Thus, the search for an outgroup for Eunice 

sensu lato is herein limited to the order Eunicoidea. 

As mentioned above, the following families are currently 

recognized in the order Eunicidae, Onuphidae, Lumbrineridae, 

Arabellidae, Iphitimidae, Lysaretidae, Oenoniidae, Hartmaniellidae, 

and Dorvilleidae. The iphitimids were considered part of 

the dorvilleids by Gaston and Benner (1981), a finding that will 

influence the composition of the Dorvilleidae. However, these 

and other taxonomic problems associated with some of the 

families do not influence the selection of outgroups for the 

following reasons. Ehlers (1868:273-274, 280-282) recognized 

two distinct groups of eunicoideans: the prionognaths 

and the labidognaths. The families Eunicidae, Onuphidae, and 

Lumbrineridae belong to the latter; the other families to the 

former group. The two groups differ markedly in the jaw 

structures. 

The prionognaths have (often numerous) jaw pieces arranged 

longitudinally along a single ridge. The carriers either form 

small box-like structures near the base of the postcriormost 

jaws, or arc long, narrow, flexible support structures, completely 

uncovered by muscle in situ (Fauchald, 1970; Wolf, 

1976). These jaws are hardened by very strong sclerotinization 

and a series of metal ions arc present, similar to the condition in 

certain glycerid jaws (Colbath, 1986; see also Voss-Foucart et 

al., 1973). 

In the labidognaths, one pair of jaws are large forceps 

(Hartmann-Schrttdcr, 1967; Wolf, 1976). The jaws are attached 

to a pair of large, wide, usually short maxillary carriers covered 

in life with a thin layer of transverse muscle, making it possible 

for the organisms to lilt the jaws along the medial axis. The 

other jaw pieces include distinct, large, often paired pieces. 

Each piece consists of a curved base and a cutting edge with a 

variable number of teeth along the edge. In addition, the jaws, 

when hardened, are calcified wilh calcitc or aragonitc (Colbath, 

1986). 

As stated above, I assume that the presence of maxillae 

arranged in scries laterally in the ontogenctic derivative of the 

stomodacal invaginalion is a synapomorphy for the order 

Eunicoidca. I further assume thai the group to which the 

eunicids sensu stricto belong is characterized by additional 

synapomorphics associated with the jaw structures; i.e., the 

distinction between prionognaihs and labidognaihs. Juvenile 

jaws of labidognath taxa have jaws lhat resemble the 

prionognath condition (Blake, 1975; Hsich and Simon, 1987). 

For the purpose of this analysis, I have assumed that the two 

sets of jaw characters (or a subset of these characters) can be 

used as synapomorphies characterizing both groups of eunicoidean 

polychaetes. 

The labidognaths have traditionally been separated into two 

groups; one containing the eunicids and onuphids and another 

containing the lumbrinerids. The latter lack the complex 

prostomial structures present in the two former and the 

parapodia are also relatively simple, characters that could easily 

be symplesiomorphies. The lumbrinerids are characterized by 

two synapomorphies associated with the jaws: the jaws are 

symmetrically paired and are always (as far as investigated) 

impregnated with calcite, independent of depth and latitude 

(Colbath, 1986). 

The eunicids and onuphids have asymmetrical jaws, in 

which the left side has an additional jaw piece, usually placed 

behind the large maxillary plate (Mx II) or anterior to, but 

behind that plate, in close association with the distal, smaller 

jaw pieces, when the jaw apparatus is viewed from above. In 

addition, the jaws are impregnated with aragonite, as far as 

known, independent of the origin of the material. Both these 

features are here considered synapomorphies for the eunicids 

and onuphids. 

The onuphids and eunicids differ in the structure of the 

prostomium. In the former the prostomium is a pentagonal to

NUMBER 523 17 

rounded lobe about as wide at the base as the peristomium. A 

pair of small, frontal antennae (frontal palps, sensu Paxton, 

1986) are present, as are five occipital antennae arranged in a 

row or a semicircle near the base of the prostomium. Each 

antenna consists of a ceratophore and a style. The ceratophores 

are often long with numerous articulations, but may also be 

smooth. They are never ring-shaped. The peristomium is a 

single ring with a pair of frontal peristomial cirri (missing in 

several genera). 

Among the eunicids, the prostomium is more or less 

distinctly cleft longitudinally (the cleft is always visible 

ventrally, even among those members of the genus Marphysa 

that are characterized as having rounded prostomia in the 

literature) and the ventral side of the prostomium may be 

equipped with a pair of palpal cushions, more or less distinctly 

set off from the prostomium proper. The frontal palps are 

missing. The antennae are in a transverse row or an open 

horseshoe. Each antenna consists of a ceratophore and a 

ceratostyle. The ceratophore is either ring-shaped or a 

cylindrical extension of the dorsal wall of the prostomium; it is 

ringed only in one described species (£. unidentata). The 

ceratostyles are often ringed and may consist of many 

moniliform articulations. The peristomium consists of two 

rings (not segments, cf. Akeson, 1967) and the peristomial cirri 

are attached near the anterior edge of the second peristomial 

ring. 

The search for an outgroup for the genus Eunice has been 

limited to the families Eunicidae and Onuphidae. Historically, 

all non- Eunice genera of the Eunicidae were formed as subsets 

of Eunice. The sets of characters currently used to recognize 

genera in the family can be summarized fairly simply. Two 

genera {Nematonereis and Lysidice) have one and three 

antennae, respectively. During ontogenesis other members of 

the family go through stages in which they have first one and 

then three antennae before adding the last, outer pair (personal 

observations). Marphysa lacks peristomial cirri, the only 

feature that consistently can be used to separate all species 

assigned to this genus from species of Eunice. Palola lacks 

subacicular hooks and pectinate setae; as an autapomorphic 

feature, the mandible is very large, scoop-shaped, and encloses 

the other jaws. Palola consists of a few species; all live in 

calcareous substrates. Nauphanta Kinberg, 1865, recently 

resurrected (Fauchald, 1986), is characterized by the presence 

of large, flat pectinate setae emerging in rows on the dorsal 

edge of the neuropodia. The genus is poorly known, and several 

additional species of Marphysa may turn out to belong to this 

genus. Additional genera have been described as offshoots of 

Marphysa (Paramarphysa and Heteromarphysa). These are 

poorly known and have been relatively rarely reported. 

Selection of any one of the other eunicid genera, with the 

exception of Palola and Nauphanta, might involve creation of 

a paraphyletic group. Ontogenetically, species of Eunice and 

Marphysa undergo "Nematonereis" and "Lysidice" states 

before they reach the adult configuration of antennae. Furthermore, 

other differences all involve the loss or modification of 

selected setal types. Selection of Palola is possible because a 

distinct apomorphy defines it; however, it is entirely possible 

that the genus will be lumped back into Eunice, as is current 

usage by several authors (e.g., Hofmann, 1974). Nauphanta is 

simply too poorly known to be an effective outgroup. A 

full-fledged cladistic analysis is probably not possible until all 

other genera named have been reviewed. I suspect that although 

there may be definable genera in the family, the current 

distribution is paraphyletic. I undertook this first survey to 

attempt to get an initial idea of possible patterns of relations 

among the species examined. 

I took two routes out of a problematic selection of outgroup. 

The first involved the selection of an onuphid as outgroup. The 

generic subdivision of the onuphids has recently been the 

subject of thorough revisions (Fauchald, 1982a, but especially 

Paxton, 1986). All members of this family are tubicolous and in 

some taxa the anteriormost parapodia are very different from 

those farther back (especially Rhamphobrachium and allied 

genera; see Paxton, 1986; see also Nothria, Paradiopatra, and 

Hyalinoecia). The transition from the modified segments to the 

remainder of the body segments is usually rather abrupt. 

Members of other onuphid genera, such as Onuphis, Kinbergonuphis, 

Paradiopatra, and Mooreonuphis, are less strongly 

cephalized and transitional segments are present over a longer 

section of the body. Eunicids and all other eunicoids appear far 

less cephalized than the onuphids. The anterior parapodia of 

eunicids differ in size more than in structure from those farther 

back. Transitions in the size, shape, and equipment of the 

parapodia are not abrupt. 

Kinbergonuphis simoni Santos, Day, and Rice, 1981, was 

selected as the outgroup onuphid. The level of variability of all 

the taxonomic characters among the onuphids is generally 

unknown. Kinbergonuphis simoni has been the subject of a 

three-year study at the type locality in Florida by Hsiang 

Hwey-Lian (Hsiang and Simon, 1987). She followed in detail 

the larval development and metamorphosis, including setal 

transition during ontogenesis, formation of the jaw apparatus 

(Hsiang and Simon, 1987), and a detailed study of the adult 

population structure at the type locality. Fauchald (1991) has 

studied the variability of adult morphology in this species from 

various localities around the Florida peninsula, but especially 

from a locality near Vero Beach on the east coast, a locality also 

studied by the authors of the species. The species is thus 

comparatively well known and when decisions as to character 

polarity had to be made, reference to this accumulated 

information assisted greatly. 

Even if the onuphids and eunicids are related, a number of 

features with similar relative position in the body are 

consistently very different. For example, the digitate postsetal 

lobes of anterior parapodia of onuphids are very different 

structurally from the transverse, low folds present in the 

eunicids. Similarly, the onuphids have pseudocompound or

18 


simple hooks present in a few anterior setigers; eunicids have 

compound hooks present in all segments (except in Eunice 

afuerensis and E. pelamidis). Further, the hooks appear very 

different and no structural analysis has been done to demonstrate 

their homologies. 

For this reason, a second outgroup was selected. In this case 

a eunicid, Marphysa sanguinea (Montagu, 1815), was selected 

as an outgroup; this species is a member of the genus that most 

closely resembles Eunice. Marphysa sanguinea has been 

reported widely from warm-water areas and the identity of the 

different populations is problematic. For that reason, a single 

specimen from Florida (USNM 17131) was examined when the 

descriptions of the species were too vague to be used in the 

analysis, or when features had not been included in previous 

descriptions. As mentioned above, the presence of peristomial 

cirri is a distinct synapomorphy for all species of Eunice in 

relation to M. sanguinea (or alternatively the absence of 

peristomial cirri in Marphysa may represent an apomorphy for 

that genus; this issue will be explored at a later date). 

CHARACTER LIST 

Originally a character list including, in all, 228 characters 

was developed. The standardized format used in character files 

of DELTA, a program for developing standardized keys and 

descriptions, was originally used to develop the list. The list 

was based on the first 50 species encountered in an alphabetic 

listing of the species. The character states were systematically 

organized from small to large (few to many) and from absent to 

present. Additional character states were included as encountered 

going through the remaining descriptions. Once an 

outgroup had been selected (see below), the character list was 

rescored, using the character state present in the outgroup as the 

plesiomorphic condition. Where a character was missing in the 

outgroup, the character states were ordered as in the original 

list, but were always run as unordered in the analysis. 

The original DELTA-style table included some 228 "characters," 

many of which created serious problems in the cladistic 

analysis. Iterated reductions in the number of characters, 

combining the extremely detailed notations necessary for 

descriptions, led to the development of an analyzable character 

list. The result was a list (Appendix A) of 68 characters, of 

which 38 are multistate and 30 binary. Appendix A also 

includes a series of measurements and numerical counts that 

were excluded from the analysis, because they could not be 

broken down into discrete states. The total number of character 

states are 210, barely adequate for the number of taxa present, 

if all character states had shown perfect behavior in the 

analysis. 

Plesio- and Apomorphic States of the Characters 

All characters are included in this overview, except those 

that identify the specimens, such as species number, name, state 

of specimen completeness, and sex. Characters and character 

states are those included in the preparation of the key; it is a 

printout of the DELTA file CHARS without the formatting 

characters. The characters are grouped by major morphological 

structures. Character state 1 is always plesiomorphic. 

Size Characters 

1. number of setigers 

2. total length mm 

These two characters are mentioned only for complete 

specimens. In all other species they have been scored as 

unknown. Length is measured from the tip of the prostomium 

to the end of the pygidium; antennae and anal cirri are omitted. 

3. maximum width in mm 

4. maximum width first reached at sctigcr 

5. length through setiger 10 in mm 

6. width at sctigcr 10 in mm 

These characters can be scored for all specimens. The 

combined score of characters 3, 4, and 6 can be used to 

characterize the overall shape of the body. Character 5 can give 

a good approximate measure of total size of incomplete 

specimens (Fauchald, 1991). 

Prostomium and Peristomium 

7. prostomium 

1. frontal ly truncate 

2. frontally obliquely truncate 

3. frontally rounded 

8. prostomium 

1. dorsally inflated 

2. dorsally flattened 

3. dorsally excavate with a thickened rim 

9. median sulcus 

1. median sulcus invisible dorsally 

2. median sulcus shallow 

3. median sulcus deep 

These characters define the shape of the prostomium. They are 

observed from the dorsal and lateral side. For characters 7 and 

8, both outgroup species have character state 1. Kinbergonuphis 

simoni lacks a median sulcus wholly; Marphysa 

sanguinea has a shallow, rather wide notch. 

10. prostomium 

1. distinctly shorter than peristomium 

2. about as long as peristomium 


1. about as wide as peristomium 

2. distinctly narrower than peristomium 


1. less than x li as deep as peristomium 

2. as deep as x li of the peristomium or deeper 

These characters define the size of the prostomium in relation 

to the rest of the anterior end. By "distinctly shorter" in 

character 10 is meant "more than 30% shorter than"; the 

break-off point is defined similarly for characters 11 and 12.

NUMBER 523 19 

Both outgroup species have short, wide prostomia and both 

have the prostomium about as deep as one-half of the 

peristomium. 

Eyes 

13. eyes 

1. present 

2. absent 

14. eyes 

1. lateral to the bases of A-I 

2. behind bases of A-I 

3. between bases of A-I and A-I I 

4. behind bases of A-II 

5. on ceratophores of A-I 

6. on ceratophores of A-II 

The presence or absence of eyes is probably very important, but 

because the color fades, and most types are rather old, character 

13 is not featured prominently in any discussions. Both 

outgroup species have character state 1; the other states are 

arranged in order of frequency in the ingroup. 

Antennae 

15. antennae arranged in a 

1. horseshoe 

2. semicircle 

3. straight line 

16. antennae 

1. evenly spaced 

2. with A-I isolated by a gap 

3. with A-I 11 isolated by a gap 

17. antennae 

1. similar in thickness 

2. with A-I slimmer than other three 

3. with A-III slimmer than other four 

4. with A-I thicker than other three 

5. with A-II thicker than other three 

6. with A-III thicker than other four 

The relative position and size of the antennae have never been 

used as systematic characters. The differences observed have 

an unknown value as systematic characters at this point. The 

features have been included here because they are being 

recorded in fresh material and appear to show some interesting 

properties. Kinbergonuphis simoni has antennae in a horseshoe; 

in Marphysa sanguinea A-I is placed just slightly ahead 

of the other antennae. Character 15 was always treated as 

unordered. For character 16 state 1 is by far the most common 

in both onuphids and eunicids and is present in both outgroup 

species. For character 17, state 4 may be the most common 

among onuphids, but is not present in K. simoni, which has 

state 1, as does M. sanguinea. The character was run unordered 

in all runs. 

18. ceratophores 

1. long in all antennae 

2. long in A-I and ring-shaped in other three 

3. ring-shaped in all antennae 


1. articulated 

2. without articulations 

In Kinbergonuphis simoni the ceratophores are cylindrical with 

three or four vaguely indicated articulations; in Marphysa 

sanguinea the ceratophores are short and ring-shaped without 

articulations. 

20. ceratostyles 

1. tapering 

2. medially inflated 

3. digitiform 

4. club-shaped 

The shape of the ceratostyles is usually not considered a strong 

character; a common excuse has been that the styles are often 

lost and thus not worthy of detailed description. Kinbergonuphis 

simoni has tapering styles; Marphysa sanguinea has 

digitiform styles. 




The styles are articulated in Marphysa sanguinea; and lack 

articulations in Kinbergonuphis simoni. 

22. ceratostyle articulations 

1. long or short cylinders 

2. moniliform or drop-shaped distally 

3. moniliform throughout 

Character state 3 may be heterogenous; some "moniliform" 

articulations are medially inflated, shaped more or less as pearls 

on a string. In other forms, the "moniliform" articulations are 

truncated cones with the base distally. It is possible that the 

apparent difference is due to contraction, but this has never 

been well documented. In several species the inner one-third to 

one-fifth of each style lack articulations; in most cases the 

articulations become more distinct distally. 

23. length of 

1. A-II and III similar and A-I shorter 

2. antennae increasing from A-I to A-III 

3. A-II greater than A-I and A-III 

4. all antennae similar (short) 

Length of the antennae is here characterized only as relative 

length in relation to the other antennae. The absolute length is 

important, but was poorly recorded. A-I is shorter than the other 

antennae in character states 1-3 and about as long as the other 

antennae in character state 4. The antennae must differ by at 

least 30% in length for the difference to be considered distinct. 

Peristomium 

24. peristomium 

1. with distinct, muscular lower lip 

2. cylindrical 

I originally had a series of characters describing the shape of the 

peristomium; character 24 is the last remnant of these


characters. It was the only character among them that could be 

characterized whether the jaws were everted or not; it has been 

poorly described in the literature, even if most illustrations in 

ventral view show the distinction rather clearly. Kinbergonuphis 

simoni has character state 2; Marphysa sanguinea has 

character state 1. 

25. separation between peristomial rings 

1. visible dorsally only 

2. visible ventrally only 

3. visible both dorsally and ventrally 

4. present on all sides 

5. absent 

In at least one case, the peristomium appears to be undivided; 

this state may be a juvenile feature. Kinbergonuphis simoni has 

a single peristomial ring on which the peristomial cirri arc 

attached anteriorly, as do all onuphids; M. sanguinea has 

character state 3. The character was run unordered in all runs. 

It is possible that it should be run in order from 5 to 1, but not 

enough is known about variability and ontogeny to decide this 

issue. 

26. anterior ring makes up 

1. V2 of total peristomial length 

2. 2 /3 of total peristomial length 

3. 3 A of total peristomial length 






9. 9 /io of total peristomial length 

This feature is undoubtedly subject to variation due to 

contractions. Differences between the extremes appear to 

reflect valid differences between taxa. 

27. peristomial cirri reach 

1. middle or anterior end of peristomium 

2. middle or front of prostomium 

3. beyond prostomium 

Scoring the length of the peristomial cirri in this fashion avoids 

the use of terms such as "short," "medium," and "long." By 

definition, Kinbergonuphis simoni has character state 2 because 

it lacks the anterior peristomial ring; M. sanguinea lacks 

peristomial cirri, so characters 27 and 28 have not been scored 

for this species. 

28. peristomial cirri 

1. tapering 

2. digitiform 


4. basally inflated 

5. ovate 

Kinbergonuphis simoni has character state 1; the character was 

run unordered. 




Peristomial cirri are not articulated in any onuphid to my 

knowledge. 

Branchiae 

30. branchiae 

1. present 

2. absent 

Branchiae arc present in both outgroup species; they are absent 

in some taxa in both families. They are absent in most other 

euniceans; possibly homologous structures termed branchiae 

arc present sporadically in other groups. 

31. branchiae 

1. pectinate 

2. palmate 

3. single filaments 

Single branchial filaments present in immature specimens with 

only a few segments appear structurally different from those of 

the adults. Some species retain these branchiae; in other species 

with single filaments, the juvenile branchiae are lost. In the 

latter case the structure of adult branchiae appears to differ from 

the juvenile branchiae. 

32. branchiae 

1. distinctly longer than notopodial cirri 

2. about as long as notopodial cirri 

3. distinctly shorter than notopodial cirri 

The definition of "distinctly" follows the pattern established for 

the antennae: one-third longer (or shorter) than the notopodial 

cirri in this case. The three character states proposed here 

represent a first attempt to measure the relative length of the 

two structures. Eventually I expect to have to characterize the 

absolute lengths of each structure; without careful study of the 

variability in a number of specimens, this first step appears to 

show promise. Both outgroup species have character state 1. 

33. branchiae 

1. reduced in mid-body region 

2. not reduced in mid-body region 

Both outgroup species have character state 2 as do most species 

of Eunice. 

34. branchial stems 

1. erect 

2. flexible 

Among species with character state 2 have been included also 

species with distinctly twisted, nearly cork-screwed branchiae. 

The difference between folded and twisted (or cork-screwed) 

shapes may be more due to fixation than to structural 

characteristics. There is, however, a clear-cut distinction 

between the two character states defined above. Kinbergonuphis 

simoni has state 1; Marphysa sanguinea has state 2. 

35. branchiae from setiger number 

36. to setiger number 

37. branchiae 

1. present to near posterior end 

2. terminating well before posterior end

NUMBER 523 21 

By "near posterior end" is here meant within 10 setigers of the 

pygidium in species with 100 or more setigers. Especially in 

species in which the body tapers smoothly towards the 

posterior end, the position of the last branchiae may be difficult 

to determine. 

38. branchiae present on 

1. more than 65% of total number of setigers 

2. less than 55% of total number of setigers 

This feature was included to make comparisons between small 

and large species easier. It is defined as the percentage of 

branchiated setigers in relation to the total numbers of setigers 

in complete specimens. The character corresponds to the 

grouping, suggested already by Grube, into species with 

branchiae terminating in mid-body and those branchiated 

through the posterior end of the body. The break-off point was 

determined by scoring all species for which the branchial 

distribution was known; no species described have branchiae 

on more than 55% but less than 65% of the body. All other gaps 

in distribution are less than 5%. Both outgroup species have 

character state 1 in characters 37 and 38. Characters 37 and 38 

are often, but not necessarily, congruent. 

39. single filaments in number of anterior setigers 

40. single filaments in number of posterior setigers 

41. maximum number of filaments 

42. maximum number of filaments first reached in setiger 

number 

Parapodia 

43. median acicular lobes 

1. distally truncate 

2. distally rounded 

3. triangular or conical 

4. withdrawn into body wall 

5. bilobed 

44. median acicular lobes with 

1. aciculae emerging ventral to midline 

2. aciculae emerging at midline 

3. aciculae emerging dorsal to midline 

The states of these two characters are relatively easily scored in 

carefully mounted parapodia. The character states for character 

43 are clearly unordered. Detailed observations were also made 

on anterior and posterior parapodial lobes, but were excluded 

from use in this study. Many of the types are incomplete 

posteriorly, and in order to make the necessary observations, 

additional parapodia would have to be removed, something 

deemed less than desirable on material consisting of types only. 

45. anterior presetal lobes 

1. follow outline of acicular lobes closely 

2. form low transverse lobes 

3. project as free lobes 

46. median presetal lobes 




47. posterior presetal lobes 




Kinbergonuphis simoni has character state 1 in all three 

characters; Marphysa sanguinea has character state 2. The 

latter is by far the most common state among the eunicids. 

48. anterior postsetal lobes 




49. median postsetal lobes 




50. posterior postsetal lobes 




These character states are determined as parapodia are observed 

from the front as mounted on a microscopic slide. During 

mounting, care has to be taken to avoid twisting the 

parapodium around its long axis. Kinbergonuphis simoni has 

character state 3 for character 48, provided that the digitiform 

postsetal lobes of the onuphids are homologous with the 

transverse folds called state 3 among the eunicids. For 

characters 49 and 50, both outgroup species have character 

state 1. 

51. ventral cirri not inflated in number of anterior setigers 

The first several ventral cirri show a smooth taper from the 

base; then, usually from setigers 5-10, the bases become 

distinctly inflated, whereas the tips retain their taper. This 

character lists the numbers of segments without distinct basally 

inflated ventral cirri. The numbers do not show any distinctly 

separated ranges and the character is not included in cladistic 

analysis. 

52. anterior ventral cirri 

1. tapering from narrow bases 

2. tapering from wide, triangular bases 

3. digiliform 

Both outgroup species have character state 1. 

53. median ventral cirri 


2. without basal inflations 

Tubicolous eunicid species always have inflated bases; 

however, a number of species for which tubes have not been 

reported also have inflated bases. Both outgroup species have 

inflated bases; Kinbergonuphis simoni constructs a tube; 

Marphysa sanguinea lives in a burrow possibly with a thin 

lining. 

54. bases of median inflated ventral cirri 

1. bases ovate or spherical 

2. bases thick, transverse welts


3. bases scoop-shaped 

The scoop-shaped bases are folded around the lower edge of the 

neuropodium and usually enclose the emerging bases of the 

subacicular hooks; the scoop-shaped region is usually angled, 

and so it reaches farther dorsally on the posterior face of the 

parapodium than on the anterior face. 

55. median inflated ventral cirri 

1. narrow tips short and button-shaped 

2. narrow tips tapering 

3. narrow tips digitiform 

4. narrow tips absent 

In nearly all species, median ventral cirri have a distinctly 

marked tip lacking the distinct glands present in the bases; the 

presence of these glands makes it possible to recognize the 

presence of the base, without the narrow tip. 

56. median ventral cirri 

1. tapering 

2. digitiform 

This character is scored only for species in which the bases are 

not inflated. 

57. posterior ventral cirri 


2. without basal inflation 

58. bases of posterior inflated ventral cirri 

1. ovate or spherical 

2. thick, transverse welts 

3. triangular welts 

4. scoop-shaped 

59. posterior inflated ventral cirri with narrow tips 

1. short and button-shaped 

2. tapering 

3. digitiform 

4. absent 

These characters parallel the ones already characterized for 

median setigers. In most species, the posterior ventral cirri lack 

basal inflations; however, when inflated, the shape of the bases 

is often different from that present in median setigers. 

60. posterior ventral cirri 

1. tapering 

2. digitiform 

3. short, nearly tubercular 

4. broadly triangular, nearly scoop-shaped 

61. anterior notopodial cirri 



3. tapering 

4. digitiform 

5. clavate 

62. median notopodial cirri 



3. tapering 

4. digitiform 

5. clavate 

63. posterior notopodial cirri 



3. tapering 

4. digitiform 

5. clavate 

These character states are listed in the same order for all three 

regions of the body; both outgroup species have character state 

1 throughout the body. In most species, the notopodial cirri 

differ in shape from one end of the body to the other. 

64. notopodial cirri 

1. articulated throughout body 

2. articulated in anterior setigers 


Both outgroup species have character state 3. Notopodial cirri 

are rarely articulated in species without articulated ccratostylcs; 

most species with articulated notopodial cirri have articulations 

limited to the anterior end, usually about the first 20 setigers in 

a specimen with 100-120 setigers. 

Pectinate Setae 

65. anterior pectinate setae 

1. furled 

2. flat 


1. distally tapering 

2. distally flaring 

67. median and posterior pectinate setae 

1. furled 

2. flat 


1. distally tapering 

2. distally flaring 

Onuphids often have narrow, furled, tapering pectinate setae 

with relatively few teeth in anterior setigers and wider, flaring 

pectinate setae in median and posterior setigers. A similar 

pattern can be found in some eunicids, but most of them appear 

to have similar pectinate setae in anterior and posterior setigers. 

The median and posterior pectinate setae appear to differ in a 

few cases, but documentation is difficult because these setae 

are small and are often broken in posterior setigers where the 

limbate setae, which protect pectinate setae in anterior setigers, 

are few in number or absent. 

Falcigers and Spinigers 

Compound falcigers differ a great deal in shape from anterior 

to posterior end and from superior to inferior positions in a 

single fascicle. The differences are mainly in the length and 

shape of the appendages, but the shafts may also change along 

the length of the body. In addition to the characters scored here, 

these shape differences should be consulted for additional 

characterizations of the different species.

NUMBER 523 23 

69. dentition of appendages of compound falcigcrs 

1. bidentate 

2. tridentate 

Both outgroup species and most species of Eunice have 

bidentate falcigers; species similar to E. antennata have 

tridentate appendages. 

70. guards distally 

1. mucronate 

2. without mucro 

"Mucronate" is here defined as having a distinct, slender spike 

attached distally to the guard. The two edges of a guard may 

meet at a low angle to form a sharply pointed guard, but this has 

not been coded as mucronate. The shape of the guards was 

originally coded in a more extensive version of the character 

list, but inconsistent scoring made it difficult to maintain the 

identity of the different character states. The issue will be 

pursued in a study of the variability within the genus Eunice. 

71. pscudocompound falcigers 

1. present 

2. absent 

Among the onuphids the anterior falcigcrs are often pscudocompound; 

Kinbergonuphis simoni happens to have only 

compound falcigcrs. Most cunicids, including Marphysa 

sanguinea, lack pseudocompound falcigcrs. 

72. compound spinigers 

1. present 

2. absent 

Among the onuphids, members of the genus Mooreonuphis 

have compound spinigers in some anterior setigers, usually in 

the lowermost position of the fascicles (Fauchald, 1982). 

Compound spinigers, when present among the eunicids, 

replace the compound falcigers and are thus present medially 

anterior to the aciculae. Both outgroup species happen to lack 

compound spinigers; the presence of spinigers is still scored as 

plesiomorphic, because presence of spinigers is widespread in 

both families. 

Aciculae 

The number of aciculae in anterior, median, and posterior 

parapodia may also turn out to be of interest; this character was 

included in earlier PAUP runs, but was omitted in the data 

transfer to this version of the character table. These characters 

are tracked in a current study of variability in the family. 

73. aciculae 

1. light yellow or translucent (clear and colorless) 

2. dark honey-colored to black 

This character may, rarely, be difficult to score; however, in 

most circumstances, there are no problems separating between 

species with brown and light-yellow aciculae. Kinbergonuphis 

simoni has light-yellow aciculae; Marphysa sanguinea has 

brown aciculae. 

74. separation of acicular cores and sheaths 

1. distinct 

2. indistinct 

75. cross section of aciculae 

1. round 

2. flattened and knife-edged 

Character 75 is best observed in posterior parapodia in 

three-quarter view, where the aciculae penetrate the body wall. 

76. aciculae distally 

1. pointed (sharp or blunt) 

2. flattened with rounded tabs 

3. expanded, knurled and knobbed 

4. hammer-headed 

5. bifid (bidentate) 

Most species, including both outgroup species, have sharply or 

bluntly pointed aciculae. The two last character states may not 

be separable at all times: in the hammer-headed forms, both 

projecting knobs are similar in size; in the bidentate forms one 

is very much larger than the other. 

Subacicular Hooks 

77. subacicular hook color 

1. light yellow or translucent (colorless) 


Kinbergonuphis simoni has light-colored subacicular hooks; 

Marphysa sanguinea has brown subacicular hooks. 

78. separation of cores and sheaths 

1. distinct 

2. indistinct 

79. distal ends of hooks 

1. simple and spine-like 

2. bidentate 

3. tridentate with teeth in a crest 

4. tridentate with teeth in tandem 

Character state 1 is often referred to as being unidentate. Both 

outgroup species have bidentate subacicular hooks. Character 

state 4 may appear when the shafts are very much thicker than 

the head: the medial part of the shaft is continued as the distal 

tooth and on each side the two upper ends of the wide shafts 

may be continued as distinct teeth on either side of the "true" 

head. 

80. hooks first present from setiger number 

81. hooks 

1. present in all setigers thereafter 

2. missing in a some setigers 

3. missing in many setigers 

Relative size of the hooks appears to decrease with increasing 

size of the specimens and especially in large specimens of 

species similar to Eunice aphroditois, hooks may be missing 

over long stretches of the body. Kinbergonuphis simoni has 

hooks in all segments; the specimen of Marphysa sanguinea 

used as the other outgroup specimen also has hooks in all 

segments; other specimens referred to Marphysa sanguinea by 

various authors have character state 3. This problem cannot be 

resolved without a detailed review of the species, based on


material from the type area and elsewhere. 

82. hooks 

1. always single (except for replacements) 

2. sometimes paired 

3. paired in most sctigers 

4. in multiples in some setigers 

5. in multiples in most setigers 

Character states 1 and 2 may be difficult to separate; I anticipate 

that this character may be reduced to three states (1 and 2), 3, (4 

and 5). Both outgroup species have character state 1. 

RESULTS 

The analysis included only species for which at least 75% of 

the characters could be scored. Nearly all of the 167 species 

included were scored for at least 90% of the characters. The 

number of unscored characters for each species included is 

indicated in the description of each species. All characters were 

run equally weighted. 

In all runs, more than 100 equally parsimonious trees were 

found; exactly how many trees could be produced was not 

determined. Strict consensus trees were created using both 

outgroups. Under such circumstances, a reduction of the 

number of terminal taxa by considering stable pairs of taxa as 

single taxa will often lead to a reduction in the number of 

equally parsimonious trees. This device reduces the apparent 

number of terminal taxa in relation to the number of characters 

present, making it more probable that a relatively limited 

number of trees can be identified. In the analysis of Eunice, a 

few stable species pairs were established, but attempts to 

reduce the number of taxa through running combinations 

turned out to be futile. 

Species of Eunice have for the last 120 years been separated 

into several groups based on the color and dentition of 

subacicular hooks (Ehlers, 1868; Hartman, 1944; Fauchald, 

1970). These groupings were never considered to have any 

systematic importance: the groupings functioned as sorting 

devices to allow taxonomists to group the species into 

manageable subunits. 

Species with yellow subacicular hooks are grouped together 

in the cladograms. Species with black subacicular hooks are 

organized into a reasonable number of (in most cases 5-6) 

clades; however, sister taxa for any "black" species varies 

widely from one case to the next. Thus, although the number of 

groups in each cladogram was limited, membership of any 

group was extremely varied. Some "black" groups could be 

defined by reversals; other groups were characterized by 

combinations of character states clade. 

Theoretically, these features of the cladograms do not singly 

or severally represent insurmountable problems for interpretation. 

It is possible that Eunice is a single clade with repeated 

reversal from light-colored to dark subacicular hooks and 

aciculae. It is also possible that in this particular case, the way 

in which PAUP handles polytomies made the results difficult to 

interpret. Another problem probably lies in the character 

definitions. The selection of characters must be re-examined; 

other characters must be added and all characters must be 

examined for possible homologies. 

Ontogenctic studies are needed to analyze the distribution of 

different shapes of subacicular hooks. In the present study, 

known ontogenetic differences were excluded from consideration, 

because information was available for so few species. For 

example, it appears as if shape of the subacicular hooks may be 

related to size. The type specimens vary a great deal in size, but 

I was at this time unable to statistically test an association of a 

particular kind of hook with the size of the specimens in which 

they were found. This aspect is being pursued in some detail in 

a study of variability. 

The next step is obvious: study the variability of as many 

characters as possible, including but not limited to those listed 

in the character table. This study will be done within a single 

population, preferably from the type locality, of as many 

species as possible. Ontogenetic changes may reduce the 

number of states of some characters listed above, but may also 

add characters not yet considered. I am also attempting to 

categorize and organize the soft-body shapes into characters so 

that more of them can be included in cladistic analysis. 

Overview of the Species 

Included in this overview arc all species originally described 

in Eunice and all species now or at one time considered as 

members of that genus. Species described in genera usually 

considered synonymous with Eunice, but which have never 

been referred to by that generic name, have been excluded. 

accrescens Hoagland, 1920, as Leodice; referred to Palola; 

previously also listed in Eunice. 

aciculata Treadwell, 1922, as Leodice; referred to Eunice, 

adriatica Schmarda, 1861, as Eunice; referred to Palola 

siciliensis. 

aenea Blanchard, 1849, as Eunice; referred to Marphysa. 

aequabilis Grube, 1878a, as Eunice, 

afra Peters, 1855, as Eunice, 

afuerensis Hartman, 1944, as Eunice, 

americana Hartman, 1944, as Eunice, 

amoureuxi Rullier, 1974, as Eunice, 

amphiheliae Marion in Filhol, 1885, as Eunice, 

anceps Pruvot, 1930, as Eunice; referred to Eunice pruvoti in 

text. 

annulicornis Johnston, 1865, as Eunice, 

antarctica Baird, 1869, as Eunice, 

antennata aedificatrix Monro, 1933, as Eunice; referred to 

Eunice aedificatrix in text. 

antennata Lamarck, 1818, as Leodice; referred to Eunice, 

antillensis Ehlers, 1887, as Eunice, 

aphroditois djiboutiensis Gravier, 1900, as Eunice; referred to

NUMBER 523 25 

Eunice djiboutiensis in text 

aphroditois punctata Fishelson and Rullier, 1969, as Eunice; 

referred to Eunice rullieri in text. 

aphroditois Pallas, 1788, as Nereis; referred to Eunice, 

arenosa Kinberg, 1865, as Eunice. 

argentinensis Treadwell, 1929, as Leodice; referred to Eunice, 

ariculata Treadwell, 1900, as Eunice; error in Hartman, 1959, 

for auriculata. 

armillata Treadwell, 1922, as Leodice; referred to Eunice, 

articulata Ehlers, 1887, as Eunice. 

articulata Hoagland, 1920, as Leodice; referred to Eunice 

hirschi in text. 

atlantica Kinberg, 1865, as Eunice, 

attenuata Grube, 1866b, as Eunice, 

auriculata Treadwell, 1900, as Eunice; referred to Euniphysa 

in text. 

australis Quatrefages, 1866, as Eunice, 

badia Grube, 1878, as Eunice. 

balfouriana Mclnlosh, 1885, as Nicidion; referred to Eunice, 

barvicensis Mclntosh, 1885, as Eunice, 

bassensis Mclntosh, 1885, as Eunice, 

bellii Audouin and Milne Edwards, 1833, as Eunice; referred to 

Marphysa. 

benedicti Verrill, 1885, as Leodice; referred to Eunice, 

biannulata Moore, 1904, as Eunice, 

biannulata mexicana Fauchald, 1970, as Eunice; referred to 

Eunice mexicana in text. 

bicirrata Rullier, 1964, as Eunice. 

biformicirrata Treadwell, 1922, as Leodice; referred to Eunice, 

bilobata Treadwell, 1906, as Eunice, 

binominata Quatrefages, 1866, as Eunice, 

bipapillata Grube, 1866a, as Eunice, 

bitorquata Grube, 1870b, as Eunice; referred to Palola 

siciliensis. 

borneensis Grube, 1878, as Eunice (Eriphyle); referred to 


bottae Quatrefages, 1866, as Eunice, 

bowerbanki Baird, 1869, as Eunice, 

brasiliensis Kinberg, 1865, as Eunice, 

brevis Ehlers, 1887, as Nicidion; referred to Eunice, 

bucciensis Treadwell, 1921, as Leodice; referred to Eunice, 

burmeisteri Grube, 1878, as Eunice, 

caeca Shisko, 1981, as Eunice. 

capensis Schmarda, 1861, as Eunice; referred to Marphysa. 

capensis Kinberg, 1865, as Eriphyle; referred to Eunice 

kinbergi. 

cariboea Grube, 1856, as Eunice, 

cedroensis Fauchald, 1970, as Eunice, 

challengeri Mclntosh, 1885, as Eunice, 

cincta Kinberg, 1865, as Nicidion; referred to Eunice, 

cingulata Claparede, 1868, as Eunice, 

cirribranchis Grube, 1870b, as Eunice, 

cirrobranchiata Mclntosh, 1885, as Eunice, 

claparedii Quatrefages, 1866, as Eunice. 

coccinea Grube, 1878b, as Eunice. 

coccinioides Augener, 1922b, as Eunice. 

collaris Grube, 1869, as Eunice. 

collini Augener, 1906, as Eunice. 

complanata Grube, 1877, as Eunice. 

concinna Verrill, 1900, as Leodice; referred to Eunice. 

congesta Marenzeller, 1879, as Eunice. 

conglomerans Ehlers, 1887, as Eunice. 

contingens Chamberlin, 1919a, as Leodice; referred to Eunice. 

crassitentaculata Treadwell, 1922, as Leodice; referred to 


culebra Treadwell, 1901, as Eunice, 

curticirrus Knox, 1960, as Eunice, 

denticulata Webster, 1884, as Eunice, 

depressa Schmarda, 1861, as Eunice; referred to Marphysa. 

dilatata Grube, 1877, as Eunice, 

dubia Woodworth, 1907, as Eunice, 

ebranchiata Quatrefages, 1866, as Eunice; referred to Palola 

siciliensis. 

edwardsi Mclntosh, 1885, as Eunice, 

ehlersi Gravier, 1900, as Eunice, 

elegans Verrill, 1900, as Leodice; referred to Eunice, 

elseyi Baird, 1869, as Eunice. 

enteles Chamberlin, 1918, as Leodice; referred to Eunice, 

equibranchiata Mclntosh, 1885, as Eunice, 

erithrocephala Risso, 1826, as Leodice; referred to Eunice, 

fasciata Risso, 1826, as Leodice; referred to Eunice, 

fasciculata Lamarck, 1816, as Tibiana; referred to Eunice, 

fauveli Gravier, 1900, as Eunice, 

fijiensis Baird, 1869, as Eunice, 

filamentosa Grube, 1856, as Eunice, 

fimbriata Grube, 1870b, as Eunice, 

flaccida Grube, 1869, as Eunice. 

flavapunctata Treadwell, 1922, as Leodice; referred to Eunice, 

flavocuprea Grube, 1869, as Eunice, 

flavofasciata Grube, 1878b, as Eunice, 

flavopicta Izuka, 1912, as Eunice. 

floridana Pourtales, 1867, as Marphysa; referred to Eunice, 

franklini Monro, 1924, as Eunice, 

frauenfeldi Grube, 1866c, as Eunice, 

fucata Ehlers, 1887, as Eunice. 

fuscafasciata Treadwell, 1922, as Nicidion; referred to Eunice, 

fusicirris Grube, 1878a, as Eunice, 

gagzoi Augener, 1922b, as Eunice, 

gaimardi Quatrefages, 1866, as Eunice, 

gallica Lamarck, 1818, as Leodice; referred to Eunice, 

gigantea Lamarck, 1818, as Leodice; referred to Eunice, 

goodsiri Mclntosh, 1885, as Marphysa; referred to Eunice, 

gracilicirrata Treadwell, 1922, as Leodice; referred to Eunice, 

gracilis Grube, 1866a, as Eunice, 

gracilis Moore, 1903, as Eunice; referred to Eunice japonica in 

text. 

gracilis Crossland, 1904, as Nicidion; referred to Eunice 

wasinensis in text.

26 

gravieri Fauvel, 1911, as Eunice. 

gregaricus Mayer, 1900, as Staurocephalus; referred to 

Eunice; previously also referred to Mayeria. 

grubei Gravier, 1900, as Eunice, 

guanica Treadwell, 1921, as Leodice; referred to Eunice, 

guildingi Baird, 1869, as Eunice. 

gunneri Storm, 1881, as Leodice; referred to Eunice norvegica. 

guttata Baird, 1869, as Eunice. 

hamata Schmarda, 1861, as Eunice; referred to Marphysa. 

harassii Audouin and Milne Edwards, 1833, as Eunice, 

havaica Kinberg, 1865, as Eunice, 

hawaiensis Treadwell, 1906, as Eunice, 

heterochaeta Quatrefages, 1866, as Eunice, 

hispanica Lamarck, 1818, as Leodice; referred to Eunice, 

imogena Monro, 1924, as Nicidion; referred to Eunice, 

impexa Grube, 1878b, as Eunice, 

incerta Hansen, 1882, as Nicidion; referred to Eunice cariboea 

in text. 

indica Kinberg, 1865, as Eunice, 

interrupta Treadwell, 1906, as Eunice, 

investigatoris Fauvel, 1932, as Eunice, 

jagori Grube, 1878a, as Eunice, 

januarii Grube, 1881, as Eunice; referred to Marphysa. 

jeffreysii Mclntosh, 1903, as Eunice; referred to Euniphysa in 

text. 

johnsoni Hartman, 1954, as Eunice, 

kinbergi Ehlers, 1868, as Eunice, 

kinbergi Webster, 1884, as Nicidion; referred to Eunice goodei 

in text. 

kobiensis Mclntosh, 1885, as Eunice, 

langi Treadwell, 1943, as Leodice; referred to Eunice, 

laticeps Ehlers, 1868, as Eunice, 

laurillardi Quatrefages, 1866, as Eunice, 

leptocirris Grube, 1870b, as Eunice. 

leucodon Ehlers, 1901, as Eunice; referred to Palola siciliensis. 

leuconuchalis Benham, 1900, as Eunice; referred to Eunice 

australis. 

leucosticta Grube, 1878a, as Eunice. 

levibranchia Hoagland, 1920, as Leodice; referred to Eunice, 

limosa Ehlers, 1868, as Eunice, 

lita Chamberlin, 1919a, as Leodice; referred to Eunice, 

longicirrata Kinberg, 1865, as Nicidion; referred to Eunice, 

longicirrata Webster, 1884, as Eunice; referred to Eunice 

websteri. 

longicirris Grube, 1869, as Eunice, 

longicornis Grube, 1866b, as Eunice, 

longiqua Kinberg, 1865, as Eunice, 

longisetis Webster, 1884, as Eunice, 

lucei Grube, 1856, as Eunice, 

macrobranchia Schmarda, 1861, as Eunice, 

macrochaeta Schmarda, 1861, as Eunice, 

madeirensis Baird, 1869, as Eunice; referred to Palola. 

madrepora pertusae Gunncrus, 1768, as Nereis; referred to 

Eunice norvegica. 


magellanica Mclntosh, 1885, as Eunice. 

magnifica Grube, 1866a, as Eunice. 

makemoana Chamberlin, 1919a, as Leodice; referred to Eunice. 

manihine Longbottom, 1972, as Eunice. 

manorae Aziz, 1938, as Eunice. 

marenzelleri Gravier, 1900, as Eunice. 

margaritacea Williams, 1853, as Eunice. 

margaritacea Verrill, 1900, as Leodice; referred to Eunice. 

margariticacea Fischli, 1900, as Eunice. 

marovoi Gibbs, 1971, as Eunice. 

martensi Grube, 1878a, as Eunice. 

maxima Qualrefages, 1866, as Eunice. 

medicina Moore, 1903, as Eunice. 

megabranchia Fauchald, 1970, as Eunice. 

megalodus Grube, 1878b, as Eunice. 

microprion Marcnzcllcr, 1879, as Eunice. 

mindanavensis Mclntosh, 1885, as Eunice. 

minuta Grube, 1850, as Eunice. 

modesta Grube, 1866a, as Eunice. 

monilifer Chamberlin, 1919b, as Leodice; referred to Eunice. 

mossambica Peters, 1855, as Eunice; referred to Nauphama; 

previously also referred to Marphysa. 

mucronata Moore, 1903, as Eunice. 

mullicylindri Shisko, 1981, as Eunice. 

multipectinala Moore, 1911, as Eunice. 

murrayi Mclntosh, 1885, as Eunice. 

mulabilis Gravier, 1900, as Eunice. 

mulilata Webster, 1884, as Eunice. 

mutilata samoae Hartmann-Schrfldcr, 1965a, as Eunice; referred 

to Eunice samoae in text. 

mutilatoides Augener, 1922b, as Eunice. 

narconi Baird, 1869, as Eunice. 

nesiotes Chamberlin, 1919a, as Leodice; referred to Eunice. 

nicidioformis Treadwell, 1906, as Eunice. 

nigricans Schmarda, 1861, as Eunice. 

northioidea Moore, 1903, as Eunice. 

norvegica Linnaeus, 1767, as Nereis; referred to Eunice. 

notata Treadwell, 1921, as Leodice; referred to Eunice. 

oerstedii Stimpson, 1854, as Eunice. 

oliga Chamberlin, 1919a, as Leodice; referred to Eunice. 

oliga papeetensis Chamberlin, 1919a, as Leodice; referred to 

Eunice papeetensis in text. 

ornata Andrews, 1891, as Eunice. 

ovalifera Fauvel, 1936, as Eunice. 

pacifica Kinberg, 1865, as Eunice. 

palauensis Okuda, 1937, as Eunice. 

paloloides Moore, 1909, as Eunice (Eriphyle); referred to 

Palola. 

panamena Chamberlin, 1919a, as Leodice; referred to Eunice. 

parasegregata Hartmann-Schroder, 1965b, as Eunice. 

parca Grube, 1878a, as Eunice. 

parva Hansen, 1882, as Eunice. 

parvibranchis Grube, 1870b, as Eunice. 

paucibranchis Grube, 1866a, as Eunice.

NUMBER 523 27 

pauper a Grube, 1878b, as Eunice. 

pauroneurata Chamberlin, 1919a, as Leodice; referred to 


pectinata Grube, 1869, as Eunice, 

pelamidis Quatrefages, 1866, as Eunice, 

pellucida Kinberg, 1865, as Eunice, 

pennata Miiller, 1776, as Nereis; referred to Eunice, 

pentadactylum Schmarda, 1861, as Sphaerodorum; referred to 

Eunice schemacephala. 

perimensis Gravier, 1900, as Eunice, 

perrieri Gravier, 1900, as Eunice, 

philocorallia Buchanan, 1893, as Eunice, 

pinnata Miiller, 1779, as Nereis; referred to Eunice, 

plicata Baird, 1869, as Eunice. 

polybranchia Vcrrill, 1880, as Leodice; referred to Eunice, 

prayensis Kinberg, 1865, as Eunice, 

procera Grube, 1866b, as Eunice, 

prognatha Mclntosh, 1885, as Eunice, 

pulvinopalpata Fauchald, 1982b, as Eunice, 

punctata Risso, 1826, as Leodice; referred to Eunice, 

punctata Peters, 1855, as Eunice; referred to Eunice peter si. 

punctata Grube, 1856, as Eunice; referred to Eunice binominata. 

purpurea Grube, 1866b, as Eunice, 

pyenobranchiata Mclntosh, 1885, as Eunice: 

quadrioculata Grube, 1856, as Eunice; referred to Marphysa. 

quinquefida Moore, 1903, as Eunice, 

quoya Quatrefages, 1866, as Eunice, 

ramosa Lamarck, 1816, as Tibiana; referred to Eunice, 

reducta Fauchald, 1970, as Eunice, 

rissoi Quatrefages, 1866, as Eunice, 

rosaurae Monro, 1939, as Eunice, 

roussaei Quatrefages, 1966, as Eunice, 

rubella Knox, 1951, as Eunice, 

rubra Grube, 1856, as Eunice. 

rubrivittata Treadwell, 1921, as Leodice; referred to Eunice, 

rubrocincta Ehlers, 1868, as Eunice, 

savignyi Grube, 1878b, as Eunice, 

schemacephala Schmarda, 1861, as Eunice, 

schizobranchia Claparede, 1870, as Eunice, 

scombrinis Quatrefages, 1866, as Eunice, 

sebastiani Nonato, 1965, as Eunice. 

segregata Chamberlin, 1919a, as Leodice; referred to Eunice, 

semisegregata Fauchald, 1969, as Eunice, 

siciliensis Grube, 1840, as Eunice; referred to Palola. 

simplex Peters, 1855, as Eunice Palola. 

sonorae Fauchald, 1970, as Eunice, 

splendida Grube, 1856, as Eunice. 

spongicola Treadwell, 1921, as Leodice; referred to Eunice, 

stigmatura Verrill, 1900, as Leodice; referred to Eunice, 

stragulum Grube, 1878b, as Eunice; referred to Marphysa. 

subdepressa Grube, 1866b, as Eunice, 

suviensis Treadwell, 1922, as Leodice; referred to Eunice, 

taenia Claparede, 1864, as Eunice; referred to Palola siciliensis. 

tahitana Kinberg, 1865, as Eunice, 

tentaculata Kinberg, 1865, as Eunice, 

tentaculata Quatrefages, 1866, as Eunice; referred to Eunice 

laticeps. 

tenuicirrata Verrill, 1900, as Leodice; referred to Eunice, 

tenuis Treadwell, 1921, as Leodice; referred to Eunice, 

teretiuscula Schmarda, 1861, as Eunice; referred to Marphysa. 

thomasiana Augener, 1922b, as Eunice, 

tibiana Pourtales, 1867, as Marphysa; referred to Eunice, 

torquata Quatrefages, 1866, as Eunice, 

torresiensis Mclntosh, 1885, as Eunice, 

triantennata Risso, 1826, as Leodice; referred to Eunice, 

tribranchiata Mclntosh, 1885, as Eunice, 

tridentata Ehlers, 1905, as Eunice, 

tristriata Grube, 1870b, as Eunice, 

tubicola Treadwell, 1922, as Leodice; referred to Eunice, 

tubifex Crossland, 1904, as Eunice, 

unidentata Rioja, 1962, as Eunice, 

unifrons Verrill, 1900, as Leodice; referred to Eunice, 

valenciennesii Grube, 1878a, as Eunice; referred to Eunice 

tentaculata Kinberg. 

valens Chamberlin, 1919c, as Leodice; referred to Eunice, 

valida Gravier, 1900, as Eunice; referred to Palola siciliensis. 

validissima Grube, 1866a, as Eunice, 

validobranchiata Monro, 1937, as Eunice, 

violacea Grube, 1856, as Eunice, 

violaceomaculata Ehlers, 1887, as Eunice, 

vittata Chiaje, 1828, as Nereis; referred to Eunice, 

vittatopsis Fauchald, 1970, as Eunice, 

vivida Stimpson, 1854, as Eunice, 

websteri Fauchald, 1969, as Eunice. 

woodwardi Baird, 1869, as Eunice, 

zonata Chiaje, 1841, as Eunice. 

Eunice Cuvier, 1817 

TYPE SPECIES.—Leodice gigantea Lamarck, 1818:322 (junior 

synonym of Nereis aphroditois Pallas, 1788:229-230, pi. 5: 

figs. 1-7), by subsequent designation (Verrill, 1900:638, see 

discussion above). 

SYNONYMS.—Eriphyle Kinberg, 1865; described for E. 

capensis Kinberg, 1865:561 (= Eunice kinbergi Ehlers, 1868, 

see below). Kinberg also listed Leodice gigantea Savigny and 

Nereis aphroditois Pallas as members of his genus. 

Leodice Lamarck (1818), erected for Leodice gigantea (see 

discussion in introductory section of paper). 

Mayeria Verrill, 1900, erected for Staurocephalus gregaricus 

Mayer, 1900; no specimens available, usually considered; 

type species usually considered a junior synonym of 

Eunice schemacephala (see below). 

Nicidion Kinberg, 1865; erected for three species, N. 

longicirrata, N. cincta, and N. gallapagensis (all discussed 

below).

28 


Tibiana Lamarck, 1816; erected for two species, T. 

fasciculata and T. ramosa, both tubes of an unidentifiable 

species of Eunice. 

Kinberg (1865) attempted to subdivide the genus using the 

structure of the prostomium, the number of pairs of maxillae, 

the presence or absence of branchiae and their structure, and the 

presence or absence of pcrislomial cirri. The presence of 

peristomial cirri is used to separate the genus Marphysa (and 

similar genera, see above) from Eunice; the other features have 

not been accepted in the literature as valid characters at the 

generic level. Two of Kinberg's new genera (Nicidion and 

Eriphyle) are here considered junior synonyms of Eunice; the 

two other, Nausicaa and Nauphanta are more closely related to 

Marphysa and will be discussed in an upcoming review of that 

genus. 

The genus Nicidion was erected by Kinberg (1865:564) for 

abranchiate Eunice; the absence of branchiae does not appear to 

be well correlated with other variable features. The genus 

Eriphyle was erected by Kinberg (1865:561) for species with 

quadrilobate prostomia and nine, rather than seven maxillae; 

these features are not well correlated with each other or with 

other features. 

Key to the Species of Eunice 

The key was produced using the auxiliary program, KEY, in the DELTA program. Of 

the 82 characters in the character table, 65 were used in preparing the key; of these 48 were 

directly included in the key. The key includes 179 species; because of uncertainty in 

coding some states, some species will key out more than once. A total of 191 tax a can be 

recognized through the key, including the duplications due to uncertainties. 

None of the numerical measurements or counts were included among the key characters. 

For this reason alone, it is crucially important that use of the key be followed up by a 

careful study of description and illustrations of the named taxon and the comparative tables 

be consulted. 

For the purpose of preparing the key only, certain characters, such as the presence or 

absence of branchiae and the color of the aciculae and the subacicular hooks were 

weighted. The sole reason for this weighting was to sort the characters so that those that 

could be relatively easily observed without dissection were used to break down the key 

into several subsections. The advantage for the user is that once he or she is in a particular 

section of the key, chances are that the specimen under study will belong to one of the taxa 

in that section of the key. Note that the sections of the key do not correspond to the groups 

used as a basis for the comparison tables. These groups are based, at least in part, on the 

distribution of branchiae, a feature that was not included in the key. The advantage for the 

user is that a set of readily characterized features were not used in preparing the key, giving 

some measure of independent confirmation of an identification obtained through the key. 

1(0). Subacicular hooks light yellow or translucent 2 

Subacicular hooks dark honey-colored to black 61 

2(1). Subacicular hooks bidentate [Figure 7j] 3 

Subacicular hooks tridentate with teeth in a crest [Figure 4d] 26 

Subacicular hooks tridentate with teeth in tandem [Figures 41o, 46c] ... 60 

3(2). Pseudocompound falcigers present [Figure 7h,k] afuerensis 

Pseudocompound falcigers absent 4 

4(3). Branchiae present on more than 65% of total number of setigers 5 

Branchiae present on less than 55% of total number of setigers 9 

5(4). Branchiae present to near posterior end 6 

Branchiae terminating well before posterior end 8 

6(5). Peristomial cirri reach middle or anterior end of peristomium [Figure 1001]; 

branchiae distinctly longer than notopodial cirri 7 

Peristomial cirri reach middle or front of prostomium [Figure 106d]; branchiae 

about as long as notopodial cirri 

stigmatura 

Peristomial cirri reach beyond prostomium [Figure 64a]; branchiae distinctly 

shorter than notopodial cirri 

leptocirris 

7(6). Ceratostyles articulated [Figure 15a]; aciculae light yellow or translucent;

NUMBER 523 29 

length of antennae increasing from A-I to A-III; antennae in a transverse row 

armillata 

Ceratostyles without articulations [Figure 1001]; aciculae dark honey-colored to 

black; all antennae similar in length (short); antennae arranged in a 

horseshoe 

schizobranchia 

8(5). Eyes present; ceratostyles club-shaped [Figure 59f]; penstomial cirri digitiform; 

antennae similar in thickness 

johnsoni 

Eyes absent; ceratostyles tapering [Figure 55g]; peristomial cirri tapering; A-I 

slimmer than other three antennae 

heterochaeta 

9(4). Peristomial cirri reach middle or anterior end of peristomium [Figure 55a] 

10 

Peristomial cirri reach middle or front of prostomium [Figure 10a] 17 

Peristomial cirri reach beyond prostomium [Figure 12b] 23 

10(9). Subacicular hooks always single (except for replacements); neuroaciculae 

emerging at parapodial midline 11 

Subacicular hooks paired in some setigers; neuroaciculae emerging dorsal to 

parapodial midline 15 

11(10). Ceratostyles articulated; hoods of compound falcigers distally without mucros; 

prostomium less than l /i as deep as peristomium 12 

Ceratostyles without articulations; hoods of compound falcigers distally 

mucronate; prostomium at least as deep as x li of peristomium 

hawaiensis 

12(11). Ceratostyle articulations long or short cylinders [Figure 61a]; anterior presetal 

lobes form low transverse folds 13 

Ceratostyle articulations moniliform or drop-shaped distally [Figure 86a]; 

anterior presetal lobes follow outline of acicular lobes closely 

pectinata 

13(12). Peristomial cirri articulated; peristomial cirri tapering; notopodial cirri 

articulated in anterior setigers; median acicular lobes triangular or conical 

[Figure 86] 14 

Peristomial cirri without articulations; peristomial cirri digitiform; notopodial 

cirri without articulations; median acicular lobes distally 

rounded 

semisegregata 

14(13). Ceratostyles tapering; anterior postsetal lobes form low transverse folds; 

narrow tips of ventral cirri tapering; anterior notopodial cirri medially 

inflated [Figure 61] 

kobiensis 

Ceratostyles digitiform; anterior postsetal lobes project as free lobes; narrow 

tips of ventral cirri digitiform; anterior notopodial cirri 

tapering [Figure 103] 

segregata 

15(10). Ceratostyle articulations long or short cylinders; inflated bases of ventral cirri 

thick, transverse welts 

valens 

Ceratostyle articulations moniliform or drop-shaped distally; inflated bases of 

ventral cirri ovate or spherical 16 

16(15). Antennae evenly spaced and similar in thickness [Figure 21b]; peristomial cirri 

basally inflated; narrow tips of ventral cirri digitiform biannulata 

Antennae with A-III isolated by a gap; A-I thicker than other three [Figure 87g]; 

peristomial cirri digitiform; narrow tips of ventral cirri tapering 

pennata 

17(9). Ceratostyles articulated; inflated bases of ventral cirri ovate or spherical [Figure 

10b] 18 

Ceratostyles without articulations; inflated bases of ventral cirri thick, 

transverse welts [Figure 69e] 22 

18(17). Ceratostyle articulations long or short cylinders 19 

Ceratostyle articulations moniliform or drop-shaped distally 20


19(18). Anterior notopodial cirri basally inflated edwardsi 

Anterior notopodial cirri medially inflated 

antarctica 

Anterior notopodial cirri tapering 

gracilicirrata 

Anterior notopodial cirri digitiform 

caeca 

20(18). Subacicular hooks always single (except for replacements) 21 

Subacicular hooks paired in some seiigers 

japonica, new name 

Subacicular hooks paired in most setigcrs 

websteri 

Three or more subacicular hooks in most setigers 

mexicana 

21(20). Ceratostyles tapering; peristomial cirri digitiform; narrow tips of ventral cirri 

digitiform; anterior notopodial cirri basally inflated parasegregata 

Ceratostyles digitiform; peristomial cirri tapering; narrow tips of ventral cirri 

tapering; anterior notopodial cirri medially inflated 

tridentata 

22(17). Branchiae pectinate; hoods of compound falcigers distally mucronate; 

subacicular hooks paired in some setigers; eyes present benedicti 

Branchiae palmate; hoods of compound falcigers distally without mucros; three 

or more subacicular hooks in most setigers; eyes absent manihine 

23(9). Ceratostyles articulated; peristomial cirri articulated; pcrisiomial cirri digitiform; 

anterior notopodial cirri digitiform [Figure 12] 24 

Ceratostyles without articulations; peristomial cirri without articulations; 

peristomial cirri tapering; anterior notopodial cirri tapering [Figure 71f-m] 

25 

24(23). Ceratostyle articulations long or short cylinders; ncuroaciculac distally 

flattened with rounded tabs; subacicular hooks paired in some seiigers; 

antennae similar in thickness 

antillensis 

Ceratostyle articulations moniliform or drop-shaped distally; neuroaciculae 

distally pointed (sharp or blunt); subacicular hooks always single (except for 

replacements); antennae with All I thicker than other four . . . articulata 

25(23). Hoods of compound falcigers distally mucronate; neuroaciculae emerging 

dorsal to parapodial midline; narrow tips of ventral cirri tapering; notopodial 

cirri articulated in anterior setigers 

megabranchia 

Hoods of compound falcigers distally without mucros; neuroaciculae emerging 

at parapodial midline; narrow tips of ventral cirri digitiform; notopodial cirri 

without articulations 

validobranchiata 


Branchiae present on less than 55% of total number of setigers .... 40 

27(26). Ceratostyle articulations long or short cylinders [Figure 5a] 28 

Ceratostyle articulations moniliform or drop-shaped distally [Figure 67d] . . . 

lucei 

All ceratostyle articulations moniliform [Figure 58a] 30 

28(27). Peristomial cirri reach middle or anterior end of peristomium; eyes absent; 

peristomial cirri tapering; ceratostyles tapering 

aequabilis 

Peristomial cirri reach middle or front of prostomium; eyes present; peristomial 

cirri digitiform; ceratostyles digitiform 29 

29(28). Appendages of compound falcigers distally bidentate; branchiae about as long 

as notopodial cirri; neuroaciculae emerging dorsal to parapodial midline; 

anterior notopodial cirri tapering 

stigmatura 

Appendages of compound falcigers distally tridentate; branchiae distinctly 

longer than notopodial cirri; neuroaciculae emerging at parapodial midline; 

anterior notopodial cirri basally inflated 

martensi 

30(27). Branchiae pectinate 31 

Branchiae palmate 

interrupta 

31(30). Subacicular hooks always single (except for replacements); prostomium 

distinctly shorter than peristomium 32 

Subacicular hooks paired in some setigers; prostomium about as long as

NUMBER 523 31 

peristomium 

rubra 

32(31). Peristomial cirri reach middle or anterior end of peristomium [Figure 8li] 

33 


33(32). Neuroaciculae distally pointed (sharp or blunt) 34 

Neuroaciculae distally hammer-headed 

panamena 

Neuroaciculae distally bifid (bidentate) 

ornata 

34(33). Notopodial cirri articulated throughout body 35 

Notopodial cirri articulated in anterior setigers 37 

35(34). Appendages of compound falcigers distally bidentate; ceratostyles tapering; 

neuroaciculae emerging at parapodial midline; prostomial median sulcus 

deep 36 

Appendages of compound falcigers distally tridentate; ceratostyles digitiform; 

neuroaciculae emerging dorsal to parapodial midline; prostomial median 

sulcus shallow 

bottae 

36(35). Eyes behind bases of A-I; ceratophores ring-shaped in all antennae; anterior 

ventral cirri tapering from narrow bases; anterior notopodial cirri basally 

inflated 

antennata 

Eyes between bases of A-I and A-II; ceratophores long in all antennae; anterior 

ventral cirri digitiform; anterior notopodial cirri digitiform 

torresiensis 

37(34). Antennae similar in thickness; ceratostyles tapering; peristomial cirri tapering; 

prostomium less than x li as peristomium 38 

Antennae with A-I thicker than other three; ceratostyles digitiform; peristomial 

cirri digitiform; prostomium at least as deep as x li of peristomium 

gracilis 

38(37). Eyes behind bases of A-I; branchiae about as long as notopodial cirri; median 

acicular lobes distally truncate; anterior postsetal lobes form low transverse 

folds 

havaica 

Eyes between bases of A-I and A-II; branchiae distinctly longer than notopodial 

cirri; median acicular lobes distally rounded; anterior postsetal lobes project 

as free lobes 

ornata 

39(32). Notopodial cirri articulated throughout body hirschi, new name 

Notopodial cirri articulated in anterior setigers 

ornata 

Notopodial cirri without articulations 

aedificatrix 

40(26). Branchiae present to near posterior end murrayi 


41(40). Hoods of compound falcigers distally mucronate [Figure 30c] 42 

Hoods of compound falcigers distally without mucros [Figure 17b] .... 53 

42(41). Ceratostyles articulated 43 

Ceratostyles without articulations 50 

43(42). Peristomial cirri reach middle or anterior end of peristomium [Figure 117a] 

44 


Peristomial cirri reach beyond prostomium 

rubrivittata 

44(43). Subacicular hooks always single (except for replacements); narrow tips of 

ventral cirri digitiform 

rubrivittata 

Subacicular hooks paired in some setigers; narrow tips of ventral cirri short and 

button-shaped 

flavocuprea 

Three or more subacicular hooks in most setigers; narrow tips of ventral cirri 

tapering 

woodwardi 

45(43). Ceratostyles tapering [Figure 8a] 46 

Ceratostyles digitiform [Figure 7Id] 48 

46(45). A-II and III similar in length; A-I shorter; peristomial cirri without


articulations; prostomial lobes frontally truncate; prostomial lobes dorsally 

inflated 

47 

Length of antennae increasing from A-I to A-III; pcristomial cirri articulated; 

prostomial lobes frontally rounded; prostomial lobes dorsally flattened . . 

atlantica and indica 

47(46). Antennae in a horseshoe; peristomial cirri tapering; neuroaciculae emerging 

dorsal to parapodial midline; narrow tips of ventral cirri tapering 

cedroensis 

Antennae in a transverse row; peristomial cirri medially inflated; neuroaciculae 

emerging at parapodial midline; narrow lips of ventral cirri digiliform 

americana 

48(45). Neuroaciculae emerging at parapodial midline; posterior ventral cirri without 

basal inflation; anterior notopodial cirri tapering; prostomium about as long 

as peristomium 49 

Aciculae emerging dorsal to midline; posterior ventral cirri basally inflated; 

anterior notopodial cirri digitiform; prostomium distinctly shorter than 

peristomium 

medicine 

49(48). A-I I and III similar in length with A-I shorter; branchiae distinctly shorter than 

notopodial cirri; anterior prcsctal lobes follow outline of acicular lobes 

closely; narrow tips of ventral cirri digiliform 

rubrivittata 

Length of antennae increasing from A-I to A-III; branchiae distinctly longer 

than notopodial cirri; anterior presctal lobes form low transverse folds; 

narrow tips of ventral cirri tapering 

arenosa 

50(42). Subacicular hooks always single (except for replacements); posterior ventral 

cirri without basal inflation; prostomial median sulcus shallow 51 

Three or more subacicular hooks in most setigcrs; posterior ventral cirri basally 

inflated; prostomial median sulcus invisible dorsally 

marovoi 

51(50). Eyes present; branchiae distinctly longer than notopodial cirri; ncuroaciculac 

emerging dorsal to parapodial midline; anterior notopodial cirri tapering 

52 

Eyes absent; branchiae about as long as notopodial cirri; neuroaciculae 

emerging at parapodial midline; anterior notopodial cirri basally 

inflated 

eugeniae, new species 

52(51). Ceratostyles tapering; anterior postsetal lobes form low transverse folds; 

narrow tips of ventral cirri tapering; eyes behind bases of A-I 

mucronata 

Ceratostyles digitiform; anterior postsetal lobes follow outline of acicular lobes 

closely; narrow tips of ventral cirri short and button-shaped; eyes between 

bases of A-I and A-II 

congesta 

53(41). Ceratostyle articulations long or short cylinders [Figure 75f] 54 

Ceratostyle articulations moniliform or drop-shaped distally [Figure 115j] 

57 

All ceratostyle articulations moniliform [Figure 83f] 59 


Subacicular hooks paired in most setigers 

aucklandica 

Three or more subacicular hooks in most setigers 56 

55(54). Peristomial cirri articulated; antennae with A-III isolated by a gap; peristomial 

cirri tapering; narrow tips of ventral cirri tapering 

multicylindri 

Peristomial cirri without articulations; antennae with A-I isolated by a gap; 

peristomial cirri digitiform; narrow tips of ventral cirri 

digiliform 

unifrons 

56(54). Pectinate setae tapering in both anterior and posterior setigers; prostomial 

median sulcus invisible dorsally 

vittata 

Pectinate setae flaring in both anterior and posterior setigers; prostomial median

NUMBER 523 33 

sulcus shallow 

tentaculata 

57(53). Peristomial cirri reach middle or anterior end of peristomium; peristomial cirri 

digitiform; narrow tips of ventral cirri tapering; prostomium at least as deep 

as V2 of peristomium 58 

Peristomial cirri reach beyond prostomium; peristomial cirri tapering; narrow 

tips of ventral cirri digitiform; prostomium less than V2 as deep as 

peristomium 

vittatopsis 

58(57). A-II and III similar in length with A-I shorter; peristomial cirri without 

articulations; ceratostyles digitiform; branchiae about as long as notopodial 

cirri 

kerguelensis 

Length of antennae increasing from A-I to A-III; peristomial cirri articulated; 

ceratostyles tapering; branchiae distinctly longer than notopodial cirri 

australis 

59(53). Ncuroaciculac distally pointed (sharp or blunt); anterior notopodial cirri 

tapering 

parva 

Ncuroaciculac distally expanded, knurled and knobbed; anterior notopodial 

cirri basally inflated 

papeetensis 

Ncuroaciculac distally hammer-headed; anterior notopodial cirri medially 

inflated 

pellucida 

60(2). Neuroaciculae distally pointed (sharp or blunt); appendages of compound 

falcigers distally bidentate; ceratophores ring-shaped in all antennae; 

peristomial cirri digitiform 

flaccida 

Neuroaciculae distally bifid (bidentate); appendages of compound falcigers 

distally tridentate; ceratophores long in A-I and ring-shaped in other three; 

peristomial cirri tapering 

elseyi 

61(1). Compound spinigcrs present [Figure 57c] 62 

Compound spinigers absent 64 

62(61). Branchiae present to near posterior end; neuroaciculae dark honey-colored to 

black; branchiae distinctly longer than notopodial cirri; neuroaciculae 

emerging dorsal to parapodial midline 63 

Branchiae terminating well before posterior end; neuroaciculae light yellow or 

translucent; branchiae about as long as notopodial cirri; neuroaciculae 

emerging at parapodial midline 

tubicola 

63(62). Ceratostyles articulated; branchiae palmate; neuroaciculae distally pointed 

(sharp or blunt); three or more subacicular hooks in most setigers 

impexa 

Ceratostyles without articulations; branchiae pectinate; neuroaciculae distally 

flattened with rounded tabs; subacicular hooks always single (except for 

replacements) 

tubifex 

64(61). Pseudocompound falcigers present pelamidis 

Pseudocompound falcigers absent 65 

65(64). Branchiae present 66 

Branchiae absent 145 

66(65). Subacicular hooks tapering to simple falcate tips [Figure 49a] 67 

Subacicular hooks bidentate 70 

Subacicular hooks tridentate with teeth in tandem Figure 96p] 144 



68(67). Branchiae pectinate; three or more subacicular hooks in most setigers; length of 

all antennae similar (short); antennae similar in thickness .... sebastiani 

Branchiae single filaments; subacicular hooks always single (except for 

replacements); A-II and III similar in length with A-I shorter; A-I thicker 

than other three antennae 

marenzelleri 

69(67). Ceratostyles articulated; peristomial cirri reach middle or anterior end of


peristomium; length of antennae increasing from A-I to A-III; neuroaciculae 


sonorae 

Ceratostyles without articulations; peristomial cirri reach middle or front of 

prostomium; length of A-II and III similar and A-I shorter, neuroaciculae 

emerging dorsal to parapodial midline 

fucata 


Branchiae present on less than 55% of total number of setigers 129 



72(71). Branchiae pectinate [Figure 44b] 73 

Branchiae palmate [Figure 92e] 11 4 

Branchiae single filaments 

pruvoti, new name 



74(73). Ceratostyle articulations long or short cylinders [Figure 44a) 75 

Ceratostyle articulations moniliform or drop-shaped dislally [Figure 63a] 

98 

All ceratostyle articulations moniliform [Figure 109a] 106 

75(74). Guards of compound falcigers distally mucronatc 76 

Guards of compound falcigers distally without mucros 78 

76(75). Peristomial cirri reach middle or anterior end of peristomium; without 

articulations; pectinate setae distally furled 

fauveli 

Peristomial cirri reach middle or front of prostomium; articulated; pectinate 

setae distally flat 77 

77(76). Peristomial cirri tapering; branchiae distinctly shorter than notopodial cirri; 

median acicular lobes triangular or conical; neuroaciculae emerging at 

parapodial midline 

franklini 

Peristomial cirri basally inflated; branchiae distinctly longer than notopodial 

cirri; median acicular lobes distally truncate; neuroaciculae emerging dorsal 

to parapodial midline 

nigricans 

78(75). Neuroaciculae distally pointed (sharp or blunt) 79 

Neuroaciculae distally expanded, knurled and knobbed 

bilobata 

79(78). Peristomial cirri reach middle or anterior end of peristomium 80 

Peristomial cirri reach middle or front of prostomium 93 


bipapillata 

80(79). Narrow tips of ventral cirri short and button-shaped 81 

Narrow tips of ventral cirri tapering 84 

Narrow tips of ventral cirri digitiform 89 

81(80). A-II and III similar in length with A-I shorter; pectinate setae flaring ... 82 

Length of all antennae similar (short); pectinate setae tapering 

petersi, new name 

82(81). Anterior postsetal lobes follow outline of acicular lobes closely 

purpurea 

Anterior postsetal lobes form low transverse folds 

ehlersi 

Anterior postsetal lobes project as free lobes 83 

83(82). Subacicular hooks paired in most setigers; median acicular lobes bilobed; 

neuroaciculae emerging at parapodial midline; pectinate setae tapering 

roussaei 

Three or more subacicular hooks in some setigers; median acicular lobes 

distally truncate; neuroaciculae emerging dorsal to parapodial midline; 

pectinate setae flaring borneensis 


Subacicular hooks paired in some setigers 88 

85(84). Peristomial cirri tapering g6

NUMBER 523 35 

Peristomial cirri digitiform 87 

Peristomial cirri basally inflated 

tribranchiata 

Peristomial cirri ovate 

magellanica 

86(85). A-II and III similar in length with A-I shorter; peristomial cirri articulated; eyes 

between bases of A-I and A-II; neuroaciculae emerging dorsal to parapodial 

midlinc 

violaceomaculata 

Length of antennae increasing from A-I to A-I 11; peristomial cirri without 

articulations; eyes lateral to the bases of A-I; neuroaciculae emerging at 

parapodial midline 

harassii 

87(85). A-II and III similar in length with A-I shorter; peristomial cirri without 

articulations; eyes present; median acicular lobes distally 

rounded 

afra 

Length of antennae increasing from A-I to A-I 11; peristomial cirri articulated; 

eyes absent; median acicular lobes distally truncate argentinensis 

88(84). Peristomial cirri tapering; branchiae about as long as notopodial cirri; median 

acicular lobes triangular or conical; anterior postsetal lobes follow outline of 

acicular lobes closely 

kinbergi 

Peristomial cirri digitiform; branchiae distinctly longer than notopodial cirri; 

median acicular lobes distally truncate; anterior postsetal lobes form low 

transverse folds 

laticeps 

89(80). Anterior notopodial cirri basally inflated 90 

Anterior notopodial cirri medially inflated 91 

Anterior notopodial cirri digitiform 92 

90(89). Neuroaciculae emerging at parapodial midline; ceratostyles tapering; pectinate 

setae tapering 

pauroneurata 

Neuroaciculae emerging dorsal to parapodial midline; ceratostyles digitiform; 

pectinate setae flaring 

bipapillata 

91(89). Length of antennae increasing from A-I to A-III; antennae with A-III isolated 

by a gap; inflated bases of ventral cirri thick, transverse welts; posterior 

ventral cirri without basal inflation 

pacifica 

Length of all antennae similar (short); antennae evenly spaced; inflated bases of 

ventral cirri ovate or spherical; posterior ventral cirri basally 

inflated 

eimeorum, new species 

92(89). Peristomial cirri articulated; neuroaciculae emerging dorsal to parapodial 

midline; ceratostyles digitiform; antennae in a horseshoe .... longisetis 

Peristomial cirri without articulations; neuroaciculae emerging at parapodial 

midline; ceratostyles tapering; antennae in a transverse row . . . mutilata 

93(79). A-II and III similar in length with A-I shorter 94 

Length of antennae increasing from A-I to A-III 96 

Length of all antennae similar (short) 

bipapillata 

94(93). Subacicular hooks always single (except for replacements); branchiae not 

reduced in mid-body region; antennae evenly spaced; neuroaciculae 

emerging dorsal to parapodial midline 95 

Three or more subacicular hooks in some setigers; branchiae reduced in 

mid-body region; antennae with A-III isolated by a gap; neuroaciculae 


microprion 

95(94). Branchiae distinctly longer than notopodial cirri; median acicular lobes distally 

truncate; anterior postsetal lobes follow outline of acicular lobes closely; 

anterior ventral cirri tapering from narrow bases .... violaceomaculata 

Branchiae distinctly shorter than notopodial cirri; median acicular lobes distally 

rounded; anterior postsetal lobes form low transverse folds; anterior ventral 

cirri digitiform 

pulvinopalpata 

96(93). Subacicular hooks always single (except for replacements); prostomial lobes 

dorsally flattened 97

36 


Subacicular hooks paired in some setigers; prostomial lobes dorsally excavate 

with thickened rim 

norvegica 

Three or more subacicular hooks in some setigers; prostomial lobes dorsally 

inflated 

philocorallia 

97(96). Anterior postsetal lobes follow outline of acicular lobes closely; notopodial 

cirri articulated in anterior setigers 

mindanavensis 

Anterior postsetal lobes form low transverse folds; notopodial cirri without 

articulations 

nesiotes 

Anterior postsetal lobes project as free lobes; notopodial cirri articulated 

throughout body 

jagori 


Subacicular hooks paired in some setigers 104 

Subacicular hooks paired in most setigers 

quinquefida 


Peristomial cirri reach middle or front of prostomium 

laurillardi 


longicirris 

100(99). Peristomial cirri articulated; neuroaciculae emerging dorsal to parapodial 

midlinc; anterior ventral cirri digitiform 101 

Peristomial cirri without articulations; ncuroaciculac emerging at parapodial 

midline; anterior ventral cirri tapering from narrow bases 103 

101(100). Antennae evenly spaced; branchiae distinctly longer than notopodial cirri; 

anterior postsetal lobes form low transverse folds; narrow tips of ventral cirri 

tapering 102 

Antennae with All I isolated by a gap; branchiae about as long as notopodial 

cirri; anterior postsetal lobes project as free lobes; narrow tips of ventral cirri 

digitiform 

pyenobranchiata 

102(101). Inflated bases of ventral cirri ovate or spherical; peristomial cirri tapering; 

notopodial cirri articulated throughout body; anterior pectinate setae distally 

flat 

biformicirrata 

Inflated bases of ventral cirri thick, transverse welts; peristomial cirri 

digitiform; notopodial cirri articulated in anterior setigers; anterior pectinate 

setae distally furled 

multipectinata 

103(100). Antennae evenly spaced; peristomial cirri digitiform; eyes behind bases of A-I; 

median acicular lobes distally rounded 

collaris 

Antennae with A-I isolated by a gap; peristomial cirri medially inflated; eyes 

behind bases of A-I I; median acicular lobes triangular or conical 

northioidea 

104(98). Peristomial cirri reach middle or anterior end of peristomium; A-II and III 

similar in length with A-I shorter; branchiae not reduced in mid-body 

region; ceratostyles tapering 105 

Peristomial cirri reach middle or front of prostomium; length of antennae 

increasing from A-I to A-III; branchiae reduced in mid-body region; 

ceratostyles digitiform 

floridana 

105(104). Antennae in a horseshoe; median acicular lobes distally truncate; anterior 

postsetal lobes follow outline of acicular lobes closely; anterior ventral cirri 

tapering from wide, triangular bases 

grubei 

Antennae in a transverse row; median acicular lobes triangular or conical; 

anterior postsetal lobes project as free lobes; anterior ventral cirri digitiform 

bowerbanki 

106(74). Peristomial cirri reach middle or anterior end of peristomium; subacicular 

hooks paired in some setigers; antennae in a horseshoe; peristomial cirri 

tapering 

torquata 

Peristomial cirri reach middle or front of prostomium; subacicular hooks 

always single (except for replacements); antennae in a transverse row;

NUMBER 523 37 

peristomial cirri digitiform 

annulicornis 


Subacicular hooks paired in some setigers 

guttata 

Subacicular hooks paired in most setigers 113 

Three or more subacicular hooks in most setigers 

scombrinis 


Peristomial cirri reach middle or front of prostomium 112 

109(108). Peristomial cirri digitiform 110 

Peristomial cirri basally inflated 

Ill 

Peristomial cirri ovate 

aciculata 

110(109). A-II and III similar in length with A-I shorter; anterior notopodial cirri basally 

inflated; eyes on ceratophores of A-II; prostomial lobes frontally rounded 

perrieri 

Length of antennae increasing from A-I to A-III; anterior notopodial cirri 

medially inflated; eyes behind bases of A-I; prostomial lobes frontally 

obliquely truncate 

dilatata 

Length of all antennae similar (short); anterior notopodial cirri digitiform; eyes 

between bases of A-I and A-II; prostomial lobes frontally 

truncate 

perimensis 

111(109). Inflated bases of ventral cirri ovate or spherical; anterior postsetal lobes form 

low transverse fold 

djiboutiensis 

Inflated bases of ventral cirri thick, transverse welts; anterior postsetal lobes 

project as free lobes 

mutabilis 

Inflated bases scoop-shaped; anterior postsetal lobes follow outline of acicular 

lobes closely 

aphroditois 

112(108). Antennae similar in thickness; peristomial cirri digitiform; ncuroaciculae 

emerging at parapodial midline; inflated bases of ventral cirri ovate or 

spherical 

polybranchia 

A-I thicker than other three antennae; peristomial cirri basally inflated; 

neuroaciculae emerging dorsal to parapodial midline; inflated bases of 

ventral cirri thick, transverse welts 

mutabilis 

113(107). Neuroaciculae emerging at parapodial midline; inflated bases of ventral cirri 

thick, transverse welts; narrow tips of ventral cirri short and button-shaped; 

anterior notopodial cirri basally inflated 

roussaei 

Neuroaciculae emerging dorsal to parapodial midline; inflated bases of ventral 

cirri ovate or spherical; narrow tips of ventral cirri tapering; anterior 

notopodial cirri medially inflated 

rullieri, new name 



115(114). Ceratostyle articulations long or short cylinders 116 

Ceratostyle articulations moniliform or drop-shaped distally 119 

116(115). A-II and III similar in length with A-I shorter; prostomium less than x li as deep 

as peristomium 117 

Length of all antennae similar (short); prostomium at least as deep as V2 of 

peristomium 118 

117(116). Antennae evenly spaced; branchiae distinctly longer than notopodial cirri; 

anterior postsetal lobes project as free lobes; prostomial lobes dorsally 

excavate with thickened rim plicata 

Antennae with A-I isolated by a gap; branchiae distinctly shorter than 

notopodial cirri; anterior postsetal lobes form low transverse folds; 

prostomial lobes dorsally inflated 

prognatha 

Antennae with A-I 11 isolated by a gap; branchiae about as long as notopodial 

cirri; anterior postsetal lobes follow outline of acicular lobes closely; 

prostomial lobes dorsally flattened 

excariboea

38 

SMITHSONIAN CONTRIBUTIONS TO ZOOIXJGY 

118(116). Antennae in a semicircle; anterior postsetal lobes project as free lobes; anterior 

ventral cirri digitiform; inflated bases of ventral cirri ovate or spherical 

guanica 

Antennae in a transverse row; anterior postsetal lobes form low transverse 

folds; anterior ventral cirri tapering from narrow bases; inflated bases of 

ventral cirri thick, transverse welts 

conglomerans 

119(115). Subacicular hooks always single (except for replacements); inflated bases of 

ventral cirri ovate or spherical; notopodial cirri without articulations .... 

120 

Subacicular hooks paired in some setigcrs; inflated bases of ventral cirri thick, 

transverse welts; notopodial cirri articulated in anterior 

setigers 

frauenfeldi 

120(119). A-II and III similar in length with A-I shorter; ceratostyles tapering; median 

acicular lobes triangular or conical; anterior notopodial cirri tapering 

goodsiri 

Length of all antennae similar (short); ceratostyles digitiform; median acicular 

lobes distally truncate; anterior notopodial cirri medially 

inflated 

rubella 

121(114). Ncuroaciculae distally pointed (sharp or blunt) 122 

Neuroaciculae distally hammer-headed 125 

122(121). Subacicular hooks always single (except for replacements); pcristomial cirri 

tapering; pectinate setae flaring 123 

Subacicular hooks paired in most setigcrs; peristomial cirri ovate; pectinate 


tenuis 

123(122). Length of antennae increasing from A-I to A-III; ceratostyles tapering 

crassitentaculata 

Length of all antennae similar (short); ceratostyles digitiform 

124 

124(123). Peristomial cirri articulated; antennae in a semicircle; ceratophores ring-shaped 

in all antennae; anterior postsetal lobes follow outline of acicular lobes 

closely 

parvibranchis 

Peristomial cirri without articulations; antennae in a horseshoe; ceratophores 

long in all antennae; anterior postsetal lobes project as free lobes 

bucciensis 

125(121). Guards of compound falcigers distally mucronate; A-II and III similar in length 

with A-I shorter; ceratostyles digitiform; anterior notopodial cirri basally 

inflated 

filamentosa 

Guards of compound falcigers distally without mucros; length of all antennae 

similar (short); ceratostyles tapering; anterior notopodial cirri digitiform 

126 

126(125). Antennae in a horseshoe; ceratophores ring-shaped in all antennae; peristomial 

cirri digitiform; neuroaciculae emerging at parapodial midline 

spongicola 

Antennae in a transverse row; ceratophores long in all antennae; peristomial 

cirri tapering; neuroaciculae emerging dorsal to parapodial 

midline 

denticulata 



guildingi 


tibiana 

128(127). Branchiae pectinate; anterior notopodial cirri digitiform . . . . flavapunctata 

Branchiae palmate; anterior notopodial cirri tapering 

goodsiri 

Branchiae single filaments; anterior notopodial cirri basally 

inflaled 

modesta 

129(70). Branchiae pectinate [Figure 61k] .130

NUMBER 523 39 

Branchiae palmate [Figure 39g] 135 

Branchiae single filaments 142 



elegans 


langi 

131(130). Ceratostyles articulated; prostomium at least as deep as ! /2 of peristomium; 

prostomium distinctly narrower than peristomium 132 

Ceratostyles without articulations; prostomium less than V2 as deep as 

peristomium; prostomium about as wide as peristomium 134 

132(131). A-II and III similar in length with A-I shorter reducta 

Length of antennae increasing from A-I to A-I 11 

coccinea 

Length of all antennae similar (short) 133 

133(132). Peristomial cirri articulated; ceratostyles digitiform; antennae in a transverse 

row; peristomial cirri digitiform 

fimbriata 

Peristomial cirri without articulations; ceratostyles medially inflated; antennae 

in a horseshoe; peristomial cirri medially inflated 

samoae 

134(131). Subacicular hooks always single (except for replacements); ceratostyles 

digitiform; peristomial cirri medially inflated; branchiae distinctly longer 

than notopodial cirri 

coccinioides 

Subacicular hooks paired in some setigers; ceratostyles tapering; peristomial 

cirri tapering; branchiae about as long as notopodial cirri. . . barvicensis 



136(135). Peristomial cirri reach middle or anterior end of peristomium; antennae similar 

in thickness 137 

Peristomial cirri reach middle or front of prostomium; antennae with A-I 

thicker than other three 

dubitata 

137(136). Subacicular hooks always single (except for replacements); branchiae 

distinctly longer than notopodial cirri; inflated bases of ventral cirri ovate or 

spherical; cross-section of aciculae round 138 

Subacicular hooks paired in some setigers; branchiae distinctly shorter than 

notopodial cirri; inflated bases of ventral cirri thick, transverse welts; 

cross-section of aciculae flattened or knife-edged 

thomasiana 

138(137). A-II and III similar in length with A-I shorter; anterior notopodial cirri tapering; 

notopodial cirri without articulations; ceratostyles tapering 

goodsiri 

Length of antennae increasing from A-I to A-III; anterior notopodial cirri 

basally inflated; notopodial cirri articulated throughout body; ceratostyles 

digitiform 

stanleyi, new species 

139(135). Peristomial cirri reach middle or anterior end of peristomium; antennae evenly 

spaced; antennae similar in thickness; branchiae distinctly shorter than 

notopodial cirri 140 

Peristomial cirri reach middle or front of prostomium; antennae with A-I 

isolated by a gap; antennae with A-I thicker than other three; branchiae 

distinctly longer than notopodial cirri 

lita 

140(139). Eyes present; ceratostyles digitiform; inflated bases of ventral cirri thick, 

transverse welts; anterior pectinate setae distally furled 141 

Eyes absent; ceratostyles tapering; inflated bases of ventral cirri ovate or 

spherical; anterior pectinate setae distally flat collini 

141(140). A-II and III similar in length with A-I shorter; antennae in a horseshoe; 

peristomial cirri tapering; eyes between bases of A-I and A-II 

levibranchia 

Length of all antennae similar (short); antennae in a transverse row; peristomial 

cirri ovate; eyes behind bases of A-I 

mutilatoides

40 


142(129). Branchiae present to near posterior end; ceratostyles without articulations; 

branchiae distinctly longer than notopodial cirri; inflated bases of ventral 

cirri thick, transverse welts 

gagzoi 

Branchiae terminating well before posterior end; ceratostyles articulated; 

branchiae distinctly shorter than notopodial cirri; inflated bases of ventral 

cirri ovate or spherical 143 

143(142). Peristomial cirri reach middle or anterior end of peristomium; peristomial cirri 

without articulations; ceratostyles medially inflated; peristomial cirri 

medially inflated 

rosaurae 

Peristomial cirri reach middle or front of prostomium; peristomial cirri 

articulated; ceratostyles tapering; peristomial cirri tapering 

nicidioformis 

144(66). Branchiae present to near posterior end; branchiae present on more than 65% of 

total number of setigers; ceratostyle articulations long or short cylinders; 

branchiae palmate 

cirrobranchiata 

Branchiae terminating well before posterior end; branchiae present on less than 

55% of total number of setigers; ceratostyle articulations moniliform or 

drop-shaped distally; branchiae single filaments 

rosaurae 

145(65). Ceratostyles articulated; antennae in a transverse row 146 

Ceratostyles without articulations; antennae in a horseshoe 148 

146(145). Ceratostyle articulations long or short cylinders; antennae evenly spaced; 

pectinate setae flaring 147 

Ceratostyle articulations moniliform; antennae with A-III isolated by a gap; 

pectinate setae tapering 

curricirris 

147(146). Ceratostyles medially inflated; inflated bases of ventral cirri ovate or spherical; 

anterior notopodial cirri tapering; prostomial median sulcus deep 

fuscafasciata 

Ceratostyles digitiform; inflated bases of ventral cirri thick, transverse welts; 

anterior notopodial cirri medially inflated; prostomial median sulcus 

shallow 

brevis 

148(145). Peristomial cirri tapering wasinensis, new name 

Peristomial cirri digitiform 

imogena 

Peristomial cirri ovate 149 

149(148). Inflated bases of ventral cirri ovate or spherical; narrow tips of ventral cirri 

short and button-shaped; anterior notopodial cirri basally inflated; pectinate 


goodei, new name 

Inflated bases of ventral cirri thick, transverse welts; narrow tips of ventral cirri 

digitiform; anterior notopodial cirri tapering; pectinate setae flaring 

cariboea 

Description of Species 

The species are arranged in alphabetic order. The tables 

comparing similar taxa are grouped at the end of the paper. 

1. Eunice aciculata (Treadwell, 1922) 

FIGURE 3; TABLES 33, 37-39 

Leodice aciculata Treadwell, 1922:143-144, figs. 24-25; pi. 3: figs. 7-13. 

Eunice afra.—Hartman, 1956:282 [in part, not Eunice afra Peters, 1854]. 

MATERIAL EXAMINED.—Holotype, AMNH 1528, Suva 

Harbor, Fiji, April 1920, in rocks outside the entrance light, in 

crevices made by boring echinoids, coll. Treadwell. 

DESCRIPTION.—Holotype complete, of unknown sex, with 

202 setigers; total length 129 mm; maximal width 4.5 mm at 

setiger 15; length through setiger 10, 11 mm; width at setiger 

10, 4.0 mm. Anterior body ventrally flattened with highly 

convex dorsum; posterior body ventrally flattened, tapering 

toward the posterior end. Anal cirri unknown. 

Prostomium (Figure 3a) less than half as long and distinctly 

narrower than peristomium; as deep as : /2 the depth of the 

peristomium. Prostomial lobes frontally obliquely truncate; 

dorsally flattened; median sulcus deep. Prostomium sloping 

towards front and laterally. Eyes present; behind bases of A-I. 

Antennae in a horseshoe, evenly spaced, similar in thickness. 

Ceratophores cylindrical in all antennae, confluent with

NUMBER 523 

41 

.1 

.05 

1 

5 

d 

b c 

FIGURE 3.—Eunice aciculata (holotype, AMNH 1528): a, anterior end, lateral view; b, compound falciger, 

parapodium 28; c, pectinate seta, parapodium 28; d, subacicular hook, parapodium 28; e, parapodium 28, anterior 

view. (Scale bars in mm.)


prostomium, without articulations. Ceratostyles short, thick, 

digitiform; all similar in thickness and shape; wrinkled, but 

without articulations. A-I to middle of first peristomial ring; 

A-II to front edge of second peristomial ring; A-III incomplete. 

Peristomium cylindrical. Separation between rings distinct 

dorsally and ventrally; anterior ring ~ 3 A of total peristomial 

length. Peristomial cirri to middle of first peristomial ring, 

nearly ovate, without distinct articulations. 

Jaws not available for examination. 

Branchiae present, pectinate, distinctly longer than notopodial 

cirri (Figure 3e), not reduced in mid-body region, erect. 

Branchiae from setiger 18 to setiger 190. Branchiae present to 

near posterior end, present on more than 65% of total number 

of setigers. First and last 10 branchiae single filaments. 

Maximum number of filaments 5, reached at setiger 32; number 

of filaments decreasing slowly over next 100 segments. 

Posterior to setiger 150,2 filaments in each branchia (except in 

last 10). Branchial stems tapering, longer than filaments. 

Filaments digitiform, about as long as notopodial cirri. 

Neuropodial acicular lobes distally rounded with aciculac 

emerging dorsal to midline. Presetal lobes low transverse folds. 

First 20 postsetal lobes free, triangular, projecting barely 

beyond acicular lobes. Median and posterior postsetal lobes 

folds following outline of acicular lobes. Ventral cirri thick and 

tapering from wide bases in first 15-20 setigers. Median 

ventral cirri basally inflated; basal inflated region indistinct, 

ovate, noticeable in setigers 20-50; digitiform free tips present. 

Posterior ventral cirri without basal inflation, digitiform. 

Notopodial cirri thick and digitiform in anterior setigers, slowly 

reduced in thickness, but retaining similar shape in all setigers, 

without articulations. 

Slender, marginally smooth limbate setae projecting beyond 

compound falcigers in all setigers. All pectinate setae (Figure 

3c) flat, distally flaring, with slender, cylindrical shafts; one 

marginal tooth distinctly longer and thicker than all other teeth; 

a total of about 15 teeth present. Shafts of compound falcigers 

(Figure 3b) slender, distally distinctly inflated with distinct 

distal beaks, marginally finely and indistinctly serrated, with 

internal oblique striations. Appendages tapering towards 

distinct head, bidentate. Proximal teeth triangular and directed 

laterally, as large as distal tooth; distal teeth tapering, directed 

laterally. Guards distally symmetrically rounded, cutting edge 

finely serrated, without mucro. Pseudocompound falcigers and 

compound spinigers absent. Neuropodial aciculae dark brown, 

distally blunt, geniculate. Separation of acicular cores and 

sheaths indistinct. Cross-section of aciculae round. Notopodial 

aciculae clear, slender. Subacicular hooks (Figure 3d) dark 

brown, bidentate. Separation of cores and sheaths indistinct. 

Hooks first present from setiger 22, present in all setigers 

thereafter, always single (except for replacements). Hooks 

distally obliquely tapering; both teeth directed distally; 

proximal teeth larger than distal teeth. 

UNKNOWN MORPHOLOGICAL FEATURES.—Pygidium and 

anal cirri; jaw structure. 

EXPECTED STATES OF SELECTED UNKNOWN FEATURES.— 

Mx III forming arc with left Mx IV; Mx VI absent. 

CHARACTERS USED IN PREPARATION OF KEY NOT 

SCORED.—Inappropriate Characters: 56, 58, 59. Unknown 

Characters: None. 

ASSUMED STATES FOR PURPOSE OF PREPARING KEY.— 

None. 

REMARKS.—Eunice aciculata was referred to E. afra by 

Hartman (1956). The species is a member of group B-4 as the 

species groups were defined by Fauchald (1970) and is listed 

with similar species in Tables 33 and 37-39. Among the 

species in this group, only five have 10 or more anterior 

branchiated setigers with single filaments only. These species 

include E. denticulata, E. guanica, E. lita, and E. tenuis, in 

addition to E. aciculata. Of these species, E. aciculata and E. 

denticulata have long, more or less cylindrical ceratophores; 

the other three have short, ring-shaped ceratophores. Branchiae 

are first present from setiger 22 in E. aciculata. from setigers 

18-19 in E. denticulata, from setigcr 36 in E. guanica, from 

setiger 23 in E. lita, and from setigcr 124 in E. tenuis. The 

posterior half of the body is abranchiate in E. lita, the other 

species are branchiated (as far as known) to the posterior end. 

Treadwell (1922:144) also reported that he had seen 

specimens from Samoa and Hawaii. 

Eunice adriatica Schmarda, 1861 

Eunice adriatica Schmarda, 1861:124-125, 12 figs., pi. 32: fig.257. 

REMARKS.—No material is available; the species was 

referred to Palola siciliensis by Grube (1864:79 and 1867:68, 

see also Ehlers, 1868:358). Nothing in Schmarda's description 

and illustrations argue against this conclusion and the 

synonymy is here accepted. 

2. Eunice aedificatrix (Monro, 1933) 

FIGURE 4; TABLES 1, 46,49 

Eunice antennata aedificalrix Monro, 1933:60-61. 

MATERIAL EXAMINED.—Six syntypes, BM(NH) ZK 1932 

12.4.537-548, Balboa, Panama, coll. Crossland, St. George 

Panama and Galapagos Expedition, 1924. 

COMMENTS ON MATERIAL EXAMINED.—Monro (1933:60) 

gave the following locality data: "Balboa, rocks and rock pools, 

low tide at Panama (3); and from scrapings off a buoy at canal 

entrance (3); and from piles of quays (5)." Based on the 

information on the labels it cannot be determined which of 

these samples were combined to give the six syntypes 

examined. 

DESCRIPTION.—Syntype complete, of unknown sex, with 

270 setigers; total length 128 mm; maximal width 4 mm at 

setiger 10; length through setiger 10, 7 mm. Anterior body 

cylindrical, becoming dorsoventrally flattened in midbody, 

tapering abruptly frontally and slowly towards posterior end,

NUMBER 523 

43 

.02 

.1 c, d 

FIGURE 4.—Eunice aedificatrix (syntype, BM(NH) ZK 1932:12.24.537-548): a, anterior end, lateral view; b, 

parapodium 15, anterior view; c, aciculae, parapodium 50; d, subacicular hook, parapodium 50; e, compound 

falciger, parapodium 15. (Scale bars in mm.) 

with short, abruptly tapering tail. 

Prostomium (Figure 4a) distinctly shorter and narrower than 

peristomium, less than x li as deep as peristomium. Prostomial 

lobes frontally rounded, dorsally inflated; median sulcus deep. 

Eyes posterior to bases of A-I, black. Antennae in a horseshoe, 

evenly spaced and similar in thickness. Ceratophores ringshaped 

in all antennae, without articulations. Ceratostyles 

tapering, with moniliform articulations to bases; maximum 12 

articulations in A-III. A-I to middle of anterior peristomial 

ring; A-I I to posterior edge of posterior peristomial ring; A-III


TABLE 1.—Variable and invariable characters in the type lot of Eunice aedificatrix (N = number of individuals 

examined; SD = standard deviation; measurements in mm; * = SD not calculated). 

VARIABLE FEATURES 

No. of setigers 

Total length 

Maximal width 

Length through 10 

No. of antennal articulations 

No. of peristomial cirral articulations 

Branchiae from setiger no. 

Max. no. branchial filaments 

Ventral cirri inflated through setiger no. 

No. of articulations in notopodial cirri 

Subacicular hooks from setiger no. 

N 

2 

2 

11 

11 

10 

8 

11 

10 

11 

9 

9 

Max. 

395 

263 

5 

9 

18 

12 

5 

13 

100 

5 

35 

Min. 

73 

24 

2 

4 

0 

0 

4 

6 

39 

2 

24 

Mean 

234 

143.5 

3.6 

627 

11.09 

5.82 

4.36 

8.80 

61.73 

4.45 

29.27 

SD 

• 

• 

1.07 

1.44 

4.61 

3.71 

0.50 

2.30 

19.35 

1.04 

3.00 

INVARIABLE FEATURES 

Shape of pectinate setae 

No. of teeth in pectinate setae 

Shafts of compound falcigers 

No. of teeth in appendages 

Color of aciculae 

Aciculartip 

Color of subacicular hooks 

Teeth in subacicular hooks 

Core-sheath construction 

N=ll 

flat 

12 

inflated 

2 

yellow 

hammer-headed to bifid 

yellow 

3 

distinct 

to middle of setiger 1. Peristomium cylindrical, twice as long as 

prostomium and at least x h wider. Peristomial fold with lateral 

notches; lower lip distinctly muscular and with a median notch. 

Separation between rings distinct dorsally and ventrally and 

visible, but less distinct laterally; anterior ring 2 /3 of total 

peristomial length. Peristomial cirri to middle of prostomium, 

slender, inflated near bases, sometimes with 4 or 5 indistinct 


Summary maxillary formula for 3 syntypes 1+1, 6+7-8, 

7-8+0, 6-7+9-11, and 1+1. Mx III long, straight, located 

behind left Mx II; Mx VI absent. 

Branchiae from setiger 4 to setiger 250. First and last 10 

branchiae single filaments. All other branchiae erect (Figure 

4b), pectinate with maximum 11 filaments at setiger 15. Most 

branchiae posterior to setiger 35 with 3-4 filaments. No 

increase in number or length of filaments towards posterior 

end. Branchiae longer than notopodial cirri in all but first 4 

branchial setigers. Filaments as long as notopodial cirri, slender 

and digitiform. Stems longer than filaments, tapering. 

Neuropodial acicular lobes distally rounded or truncate with 

aciculae emerging near the middle. Pre- and postsetal lobes low 

transverse folds. First 3 ventral cirri thick and tapering, 

thereafter basally inflated. Inflated base ovate, reduced from 

about setiger 45, with distinct tapering narrow tips. Posterior 

ventral cirri tapering and increasingly slender in far posterior 

setigers. Notopodial cirri basally inflated, becoming narrower 

and increasingly digitiform in far posterior setigers; articulations 

absent. 

Limbate setae longer than compound hooks and marginally 

finely serrated. All pectinate setae tapering, furled, with about 

10 coarse teeth. One marginal tooth somewhat longer and 

distinctly coarser than other teeth. All shafts of compound 

hooks (Figure 4e) distally inflated and marginally serrated, 

distinctly slimmer than aciculae and subacicular hooks. 

Appendages slender and tapering. Proximal tooth triangular 

and directed laterally; distal tooth gently curved, tapering, 

longer than proximal tooth. Guards distally symmetrically 

sharply pointed, but not mucronate; margin serrated. Aciculae 

yellow, paired, round in cross-section. Superior aciculae 

hammer-headed; inferior aciculae bent and tapering in anterior 

setigers, becoming hammer-headed posterior to setiger 25 

(Figure 4c). Subacicular hooks (Figure 4d) yellow and 

tridentate, without distinct separation of cores and sheaths. 

Hooks first present from setiger 35, thereafter present in all 

setigers, always single. Hooks curved with teeth decreasing 

evenly in size from curved main fang. 

Tube composed of pliable pergamentaceous flakes. 

UNKNOWN MORPHOLOGICAL FEATURES.—Pygidium 

anal cirri. 


Anal cirri long, tapering, with moniliform articulations. 


SCORED.—All characters scored. 


None. 

REMARKS.—Eunice aedificatrix was described as a subspecies 

of E. antennata, with which it agrees in the possession of 

tridentate yellow subacicular hooks, branchiae distributed 

along the length of the body, and strongly articulated antennae. 

Eunice aedificatrix is listed with similar species in Tables 46 

and

NUMBER 523 45 

and 49. In addition to E. aedificatrix, only E. aequabilis among 

the species listed in Table 49 has ornamented aciculae. In E. 

aedificatrix, both pointed aciculae and hammer-headed aciculae 

are present; in E. aequabilis the aciculae are either distally 

knobbed or nearly bidentate. In addition, the latter has 

tridentate appendages in the compound falcigers; the appendages 

arc bidentate in E. aedificatrix. 

Eunice aenea Blanchard, 1849 

Eunice aenea Blanchard, 1849:19-20. 

Marphysa aenea.—Augcner, 1922c:196; 1923:65.—Hartman. 1944:128. 

REMARKS.—Eunice aenea was referred to the genus Marphysa 

by Augener (1922c: 196, see also Augener, 1923:65), 

who redescribed the species; additional descriptive detail was 

added by Hartman (1944:128). There is nothing in Blanchard's 

original description to indicate that this synonymy is in error 

and it is here accepted. 

3. Eunice aequabilis Grube, 1878 

FIOURE 5; TABLES 46,49 

Eunice aequabilis Gmbc, 1878a:102.—Fauchald. 1986:242-243, figs. 1-8. 

MATERIAL EXAMINED.—Syntypes, MPW 273, Zool. Mus. 

Wroclaw, and ZMB 3993, Cape York, Australia, coll. Salmin 

(the label for the Berlin specimen gives Grube as collector). 

DESCRIPTION.—Wroclaw syntype complete with 158 setigers; 

total length 175 mm; maximum width 7 mm at setiger 15. 

Length through setiger 10, 16 mm; width at setiger 10, 6 mm. 

Berlin syntype complete with 151 setigers; 183 mm long. Both 

syntypes of unknown sex and maturity. Anal cirri long, slender, 

with 6 articulations. 


peristomium, as deep as x li of the peristomium. Prostomium 

about 2 /3 as wide as peristomium. Prostomial lobes frontally 

obliquely truncate, dorsally flattened; median sulcus shallow. 

Eyes absent. Antennae in a horseshoe; A-I separated from other 

antennae by distinct gap, similar in thickness. Ceratophores 

ring-shaped, without articulations. Ceratostyles slender, tapering, 

with up to 12 cylindrical articulations in A-I I and III. A-I 

to posterior end of anterior peristomial ring; A-I I and III to 

setiger 3. Peristomium cylindrical; peristomial fold entire; 

lower lip entire, not distinctly set off from the rest of the 

peristomium. Separation between peristomial rings distinct 

dorsally and ventrally; anterior ring s /6 of total peristomial 

length. Peristomial cirri to front edge of peristomium, slender 

and tapering, with 5 articulations. 

Maxillary formula of Berlin syntype 1+1,6+5,8+0, 5+4, and 

1+1. Mx III long, located behind left Mx II. Mx VI missing. 

Branchiae present from setiger 6 to end. All branchiae 

strongly pectinate (Figure 5b); first branchia with 13 filaments; 

maximum 17 filaments at setiger 20; number decreasing slowly 

posteriorly, never less than 12 filaments present. Median region 

with reduced branchiae absent. Stems tapering. Most filaments 

filiform, about 2 /3 as long as stems, shorter than notopodial 

cirri. Branchiae longer than notopodial cirri. 

Neuropodial acicular lobes distally obliquely truncate. 

Pre-and postsetal lobes low transverse to oblique folds. First 3 

ventral cirri tapering, thereafter basally inflated. Inflated bases 

ovate; long tapering tips present in all setigers, decreasing from 

about setiger 30. Posterior ventral cirri tapering. Notopodial 

cirri long, basally inflated, as long as branchial stem in most 

setigers, without articulations. 

Limbate setae longer than compound hooks, marginally 

frayed, brittle. Pectinate setae (Figure 5h) narrow, tapering, flat. 

Both marginal teeth longer than other teeth, with about 12 

teeth. Shafts of compound hooks (Figure 5e) inflated, with 

distal beaks, marginally smooth. All appendages short, tridentate. 

Primary fangs by far largest, triangular, directed laterally. 

Secondary fangs narrowly triangular, directed obliquely distally. 

Tertiary fangs very small rounded. Fangs decreasing 

evenly in size distally. Guards short, narrow, distally symmetrically 

rounded, marginally serrated, without mucros. Pseudocompound 

hooks and compound spinigers absent. Aciculae 

yellow, paired; superior aciculae flattened; inferior ones with 

round cross-section, without distinct segregation of core and 

sheath. Superior aciculae laterally distally expanded into 

rounded tabs; inferior aciculae (Figure 5f,g) bidentate; proximal 

teeth larger than distal teeth; both directed obliquely 

distally. Subacicular hooks (Figure 5c,d) yellow, tridentate. 

First present from setiger 26 in Wroclaw syntype and from 

setiger 32 in Berlin syntype, thereafter present in all setigers, 

always single. Hooks straight, tapering. Primary fangs separated 

from other fangs by a wide gap, tapering, directed 

laterally or slightly basally. Secondary fangs and tertiary fangs 

on joint bases. Secondary fangs tapering, directed laterally. 

Tertiary fangs slender, tapering, directed obliquely distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Structure of 

pygidium; relation between Mx III and IV. 


Left Mx IV about as long as right Mx IV and curved only 

distally. 


SCORED.—Inappropriate Characters: 14, 56, 58-59. Unknown 



None. 

REMARKS.—Eunice aequabilis belongs to the same group as 

E. antennata, from which appears to differ in that the branchiae 

retain the high number of branchial filaments in all setigers, and 

that the articulations of the long antennae are cylindrical, rather 

than moniliform. It is listed with similar species in Tables 46 

and 49. It was discussed in relation to E. aedificatrix above. 

The species apparently has remained unrecorded until 

Fauchald (1986) redescribed the types in a study of Australian 

eunicids.

46 


.02 

.1 

1 

5 

p., h 

c, d. t q 

h 

. , 

FIGURE 5.—Eunice aequabilis (syntype, MPW 273): a, anterior end, lateral view; b, parapodium 55, anterior 

view; c, subacicular hook, parapodium 55; d, subacicular hook, parapodium 134; e, compound falciger, 

parapodium 134; f, g, aciculae, parapodium 55; h, pectinate seta, parapodium 134. (Scale bars in mm.)

NUMBER 523 47 

4. Eunice afra Peters, 1854 

FIGURE 6; TABLES 33,35 

Eunice afra Peters. 1854:611.—Ehlers, 1897:166. 

Eunice collaris Grube, 1869:494-495 [in part, see below]. 

MATERIAL EXAMINED.—Holotype, ZMB 46, Ibo, Mozambique, 

coll. Peters; 2 syntypes of E. collaris, ZMB F2004 and 

2015, Tor, Red Sea, coll. Ehrenberg. One slide marked "Type, 

BM(NH) ZB 1984.69." 

COMMENTS ON MATERIAL EXAMINED.—The description is 

based on the holotype, with comments on the 2 syntypes of E. 

collaris where appropriate. Peters (1854:611) gave the type 

locality as "Qucrimba-Inscln (Ibo)." 

DESCRIPTION.—Holotype incomplete of unknown sex, with 

43 sctigcrs; total length 32 mm; maximal width 6 mm; length 

through sctigcr 10, 10 mm. Anterior body cylindrical; dorsal 

and ventral surfaces convex, appearing truncate frontally. Body 

tapering very slowly posteriorly, becoming slightly flattened 

dorsovcnt.ally, tapering abruptly near pygidium. Anal cirri 

unknown. 

Prostomium (Figure 6a, 0 distinctly shorter than peristomium, 

about as wide as peristomium, as deep as x ji of the 

pcristomium. Prostomial lobes frontally rounded, dorsally 

inflated; median sulcus deep. Eyes posterior to bases of A-I, 

black. Antennae in a horseshoe, evenly spaced, with A-I 11 

slimmer than other four. A-I pyriform; A-I I and A-I 11 

digitiform; A-I 11 distinctly narrower than other antennae. 

Ceratophores short, ring-shaped, without articulations. Ceratostyles 

with up to 5 indistinct cylindrical articulations mostly 

confined to distal half of A-I I and A-I 11. A-I basally inflated; 

A-I I and A-I 11 digitiform. A-I to front edge of posterior 

peristomial ring; A-I I and III to setiger 1. Peristomium slightly 

inflated, cylindrical; lower lip distinct and muscular. Separation 

between rings distinct dorsally and ventrally; anterior ring 

about 3 /

48 


FIGURE 6.—Eunice afra (holotype, ZMB 46): a, anterior end, lateral view; b, pectinate seta, parapodium 40; c, 

subacicular hook, parapodium 40; d, compound falciger, parapodium 40; e, parapodium 40. Eunice afra (syntype 

of E. collaris ZMB F2O15): f, anterior end, lateral view; g, pectinate seta, parapodium 39; h, subacicular hook, 

parapodium 39; i, compound falciger, parapodium 39; j, parapodium 39, anterior view. Eunice afra (type, 

BM(NH) ZB 1984.69, slide): k, subacicular hook. (Scale bars in mm.)

NUMBER 523 49 

median antennae similar in length and clearly outreaching the 

prostomium. The bases of the median inflated ventral cirri are 

scoop-shaped in E. afra and thick, transverse welts in E. 

ehlersi. 

5. Eunice afuerensis Hartman, 1944 


Eunice afuerensis Hartman, 1944:108-109. pi. 6: figs. 127-134. 

MATERIAL EXAMINED.—Holotype, AHF 391-35, Islas 

Lobos de Afuera, Peru, 6°55'40"S, 80°43'40"W, 17 Jan 1935, 

shore, rock and algae. 

DESCRIPTION.—Holotype complete female with 687 setigers; 

total length 720 mm; maximal width 12 mm at setiger 15; 

length through setiger 10, 19 mm; width at setiger 10, 10 mm. 

Anterior body cylindrical, truncate frontally, tapering slowly 

posteriorly, becoming dorsoventrally flattened. Anal cirri with 

moniliform articulations, as long as last 10 setigers; dorsal edge 

of pygidium crenulatcd. 




Eyes not observed. Antennae in a straight line, evenly spaced, 

similar in thickness. All ceratophores short, ring-shaped 

without articulations. Ceratostyles tapering, with -20 short 

articulations; articulations not moniliform, except for distalmost 

ones. Antennae to middle of anterior peristomial ring. 

Peristomium ~3 times as long as prostomium with a distinct 

muscular lower lip. Separation between rings very well marked 

dorsally and ventrally, indistinct only for a short distance 

laterally; anterior ring - 3 A of total peristomial length. Peristomial 

cirri to middle of anterior peristomial ring, medially 

inflated, without articulations. 

Maxillary formula 1+1, 4+5, 4+0, 6+7, and 1+1. Mx III 

short, located behind left Mx II, forming part of distal arc with 

left Mx IV. Mx VI absent. Maxillae strongly calcified, brittle. 

Branchiae (Figure 7d,e,l) pectinate, distinctly longer than 

notopodial cirri, not reduced in mid-body region, erect. 

Branchiae from setiger 3 to second from last setiger. All 

branchiae luxuriously pectinate, with up to 25 slender 

filaments. Stems flexible, slender, cylindrical. Filaments longer 

than stems in most setigers and longer than notopodial cirri in 

all but the first 2-3 setigers. 

Neuropodial acicular lobes rounded in anterior setigers, 

obliquely truncate in median and posterior setigers. Pre- and 

postsetal lobes low, transverse folds. First 3 ventral cirri thick, 

tapering. Ventral cirri basally inflated from setiger 4; free tips 

tapering in all setigers. Inflated bases retained in all but last 

setigers, forming broad, transverse welts with tips attached 

distally. Notopodial cirri medially inflated, with up to 7 

articulations in anterior setigers; number and distinctness of 

articulations lost in first 20 branchial setigers. 

Limbate setae longer than other setae, slender, narrow, 

marginally serrated. Large numbers of pectinate setae present. 

Anterior pectinate setae (Figure 7b) with thick, cylindrical 

shafts, furled, tapering; one marginal tooth distinctly thicker 

and longer than all other teeth; 12 teeth present. Median and 

posterior pectinate setae (Figure 7f) with wide, flattened shafts. 

Blades furled, flaring. One marginal tooth longer, but not 

thicker than other teeth, with 20 teeth. True compound hooks 

present only in first 25 setigers, replaced by pseudocompound 

hooks in all other setigers. Shafts of compound hooks (Figure 

7c) very coarse, tapering, without serrations. Appendages 

small, tapering. Both teeth similar in size, directed distally, 

tapering, slightly curved towards each other. Guards not seen. 

Pseudocompound hooks (Figure 7h,k) in one preacicular and 

one postacicular row, up to 5 hooks present in each row; setae 

oriented roughly in a fan in each row (not well illustrated). 

Hooks with distinct fused joints, represented by bends on one 

side and bulges on opposite side. Proximal teeth thicker and 

more distinctly tapering than distal teeth in most hooks. Distal 

teeth often distally hooked. All compound and pseudocompound 

hooks clear yellow. Compound spinigers absent. 

Aciculae paired, yellow, tapering; cross-sections round. Inferior 

aciculae coarser than superior aciculae, distinctly bent in 

anterior setigers. Superior aciculae coarser than inferior 

aciculae (Figure 7i) in far posterior setigers, but both similar in 

shape. Subacicular hooks (Figure 7g,j) clear yellow, bidentate. 

Hooks present from setiger 100, thereafter present singly in all 

setigers. Hooks slender, smoothly tapering. Proximal teeth 

much larger than distal teeth, triangular, directed laterally. 

Distal teeth slender, erect. Guards not seen. 

UNKNOWN MORPHOLOGICAL FEATURES.—Guards of compound 

and subacicular hooks. 

EXPECTED STATE OF SELECTED UNKNOWN FEATURES.— 

Guards of compound hooks asymmetrically bluntly pointed, 

following outline of appendages closely, marginally smooth. 


SCORED.—Inappropriate Characters: 56, 60. Unknown 

Characters: 13-14,70. 

ASSUMED STATES FOR PURPOSE OF PREPARING KEY.—70,2. 

REMARKS.—Eunice afuerensis is very similar to E. pelamidis 

from the same geographical region; both species have the 

same branchial distribution and similar closely articulated 

antennae, and the basic setal structures are the same. The two 

species can be separated on the development of the basal 

portion of the ventral cirri, which retains an inflated region in 

most of the body in E. afuerensis and is inflated only in anterior 

setigers in E. pelamidis and by the color of the setae, aciculae 

and subacicular hooks; these are dark amber in E. pelamidis 

and clear yellow in E. afuerensis. 

Hartman (1944:109) considered the pseudocompound hooks 

as series of subacicular hooks; the two kinds of hooks are 

clearly different, both in structure and origin. The pseudocompound 

hooks originate distal to the subacicular hooks and point 

distally, whereas the single subacicular hook has the origin and 

orientation of the subacicular hooks in other eunicids. Eunice

50 


I 

2 

1 

.1 

1 

. a 

d 

c, g, h, i, j. k 

FIGURE 7.—Eunice afuerensis (holotype, AHF 391-35): a, anterior end, lateral view; b, pectinate seta, 

parapodium 11; c, compound falciger, parapodium 11; d, parapodium 11, anterior view; e, parapodium 346, 

anterior view; f, pectinate seta, parapodium 346; g, subacicular hook, parapodium 346; h, pseudocompound 

falciger, parapodium 346; i, aciculae, parapodium 346; j, subacicular hook, parapodium 662; k, pseudocompound 

falciger, parapodium 662; 1, parapodium 662. (Scale bars in mm.)

NUMBER 523 51 

afuerensis and E. pelamidis are compared in Tables 52 and 53. 

In addition to the types, Hartman (1944:108) also reported 

the presence of fragments from station 844-38, another 

intertidal collection at Islas Lobos de Afuera. 

6. Eunice americana Hartman, 1944 


Eunice americana Hartman, 1944:118-121, pi. 8: figs. 164-174,189. 

MATERIAL EXAMINED.—Holotype, AHF Poly 0725, off 

Redondo Beach, California, 33°49'30"N, 118°26'15"W to 

33°49'15"N, 118°26'30"W, 315-249 m, mud and worms or 

33°49'15"N, 118°26'20"W to 33°48'45"N, 118°25'55"W, 256- 

90 m, mud. Velero sta 1133-40 (dredge hauls 1 and 2). 

COMMENTS ON MATERIAL EXAMINED.—The holotype is 

very soft. Hartman (1944:121) listed 1130-40 as the type 

locality; this station was off Newport Beach, rather than 

Redondo Beach. The labels with the specimens agree with the 

other information given, so station 1133-40 is here considered 

the type locality. 

DESCRIPTION.—Holotype complete, of unknown sex with 


sctiger 10; length through setiger 10, 9 mm. 


peristomium, as deep as x li of the peristomium. Prostomial 

lobes frontally truncate, dorsally slightly inflated; median 

sulcus shallow. Two pairs of black eyes; anterior pair behind 

bases of A-I; posterior pair near peristomial fold. Antennae in 

a straight line, evenly spaced, similar in thickness. Ceratophores 

ring-shaped in all antennae, without articulations. Ceratostyles 

distinctly tapering, with up to 20 vaguely indicated, 

cylindrical rings. A-I to setiger 1; A-I I and III to setiger 4. 

Peristomium more than twice as long as prostomium, cylindrical. 

Separation between rings distinct dorsally and ventrally; 

anterior ring 2 /3 of total peristomial length. Peristomial cirri 

nearly to Up of prostomium, basally inflated, tapering to fine 

tip, without articulations. 


long, located behind left Mx II; Mx VI absent. 


cirri, not reduced in mid-body region, erect. Branchiae 

from setiger 3 through setiger 35, terminating well before 

posterior end, present on less than 55% of total number of 

setigers. All branchiae, except last 2 pairs, with at least 2 

filaments. Maximum 20 filaments at setiger 10-15. Stems 

slender, erect, longer than filaments. Filaments slender, shorter 

than stems and notopodial cirri. 

Anterior neuropodial acicular lobes distally truncate, symmetrically 

rounded posteriorly; aciculae emerging at midline. 

Pre- and postsetal lobes low transverse folds. First 5 ventral 

cirri thick, digitiform, basally strongly inflated in branchial 

region. Inflated bases decreasing in postbranchial setigers. 

Inflated bases ovate; distal tips digitiform. Ventral cirri 

becoming slender and digitiform in last setigers. Anterior 

notopodial cirri medially inflated; medial and posterior 

notopodial cirri increasingly slender and digitiform, shorter 

than in anterior parapodia. All notopodial cirri articulated, with 

up to 3 distinct articulations. 

Limbate setae longer than compound hooks, slender, 

marginally serrated. Pectinate setae (Figure 8c) with slender, 

cylindrical shafts. Blades flaring, flat. Both marginal teeth 

longer than other teeth; -15 teeth. Shafts of compound hooks 

(Figure 8b) inflated, marginally serrated. Appendages very 

long, narrow, tapering distally; heads very small, bidentate. 

Proximal teeth small, triangular elevations; distal teeth slender, 

nearly erect, longer than proximal teeth. Guards very slender, 

tapering to very long, symmetrically placed mucros. Pseudocompound 

falcigers and compound spinigers absent. Aciculae 

(Figure 8d) paired, yellow, tapering, distally asymmetrically 

bent; tips slightly flattened. Neither aciculae nor 

subacicular hooks with distinct separation between sheath and 

core. Subacicular hooks (Figure 8e) yellow, tridentate. Hooks 

first present from setiger 25, present in all setigers thereafter, 

single. Main fangs large, pointed, tertiary fangs often very 

small. 

UNKNOWN MORPHOLOGICAL FEATURES.—Body shape not 

described (specimens very soft); details of pygidium and anal 

cirri not described. Shape of left Mx IV not described. 


Left Mx IV about as long as right Mx IV; curved distally. 





None. 

REMARKS.—Eunice americana is listed with similar species 

in Tables 41 and 42. It is the only species in Table 42 with as 

many as 20 branchial filaments; the next highest number is 12. 

Two pairs of eyes have also been reported from E. gravieri; 

however, the latter has dark brown or black, bidentate 

subacicular hooks. 

Hartman (1944:121) listed her new species from a total of 19 

stations off western Mexico and southern California in shelf 

depths. 

7. Eunice amoureuxi Rullier, 1974 

TABLES, 24,26 

Eunice amoureuxi Rullier, 1974:45-47, fig. 3a-k. 

MATERIAL EXAMINED.—The original material of this species 

is not present in the collections in Paris or Angers. The 

material was collected by Bacescu and Gomoiu in Cuba in 

1969. Requests for information about the material sent to 

Roumania, where the type was deposited (Rullier, 1974:47) 

went unanswered. 

This species was originally described from Bahia de Habana,

52 


.02 b, c, e 

e 

FIGURE 8.—Eunice americana (syntype, AHF Poly 0725): a, anterior end, lateral view; b, compound falciger, 

parapodium 46; c, pectinate seta, parapodium 46; d, aciculae, parapodium 46; e, subacicular hook, parapodium 

46. (Scale bars in mm.) 

Cuba, in 15 m depth. The following summary of the original 

description reports all that is known about this species. 

DESCRIPTION.—Complete specimen 13 mm long; number of 

setigers not mentioned. 

Prostomium frontally rounded. Eyes present. Antennae 

evenly spaced, similar in thickness. Ceratophores ring-shaped 

in all antennae, without articulations. Ceratostyles digitiform, 

without articulations. A-II and III similar and A-I shorter; none

NUMBER 523 53 

reaching beyond anterior peristomial ring. Peristomial cirri to 

posterior end of anterior peristomial ring, without articulations. 

The apparent maxillary formula 1+1,5+5, 5+0, and +4 and 

1+1. 

Branchiae present, single filaments, distinctly longer than 

notopodial cirri, not reduced in mid-body region. Branchiae 

from setiger 9 to sctiger 24. Branchiae terminating well before 


Pectinate setae with one marginal tooth heavier than other 

teeth, with -10 teeth. Shafts of compound hooks inflated, 

marginally serrated. Appendages short, tapering, with very 

large heads, bidentate. Both teeth of about same size, sharply 

pointed. Distal teeth distinctly curved. Guards symmetrically 

rounded, marginally serrated. Pseudocompound falcigers and 

compound spinigers absent. Aciculae dark-colored, tapering, 

straight. Subacicular hooks from setiger 17, dark-colored, 

bidentate. Hooks strongly curved. Proximal teeth slightly larger 

than distal teeth, directed laterally. Distal teeth slender, nearly 

erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—Many features 

unknown. 


Species too poorly known to make meaningful predictions. 


SCORED.—Inappropriate Characters: 22. Unknown Characters: 

1, 3-6, 7-12, 15, 24-26, 28, 34, 38-68, 74, 75, 78, 81, 

82. 


None. 

REMARKS.—This species is very poorly known; it has been 

listed with similar taxa in Tables 24 and 26, but has not been 

included in the key. In some features it resembles juveniles 

recorded from Belize, Central America, but without access to 

specimens from the type locality, such a relationship cannot be 

established. Eunice amoureuxi is the only species listed in 

Table 26 with single branchial filaments throughout. 

8. Eunice amphiheliae Marion in Filhol, 1885 

TABLE 27 

Eunice amphiheliae Marion in Filhol, 1885:199.—Roule, 1896:446-449, pi. 

19: figs. 1, 2, pi. 20: fig. 3. pi. 23: figs. 16-21, pi. 25: figs. 25, 26. 

MATERIAL EXAMINED.—No specimens are available. The 

following description is summarized, with updated terminology, 

from Roule's description. 

DESCRIPTION.—Complete specimens with 140-150 segments; 

length 180-200 mm; width 7-10 mm. 

Prostomium distinctly shorter and distinctly narrower than 

peristomium. Peristomial lobes frontally truncate, dorsally 

apparently flattened; median sulcus deep. Eyes absent. Antennae 

apparently in straight line, similar in thickness. Ceratophores 


tapering, irregularly articulated. All antennae similar in 

length; none reaching beyond peristomium. Peristomium 

massive, cylindrical. Separation between rings distinct on all 

sides or perhaps interrupted for short distance laterally; anterior 

ring ~*/6 of total peristomial length. Peristomial cirri to anterior 

edge of peristomium, tapering, slender, without articulations. 

Apparent maxillary formula 1+1. 5+5, 8+0, +5, 1+1. 

Illustration confused; number of teeth of Mx III and IV and 

shape of maxillae cannot be determined from description or 

illustration. 



from setiger 4 to setigers 140-150, decreasing drastically in 

size and complexity in posterior 2 /3 of body. Branchiae present 

to near posterior end, in more than 65% of total number of 

setigers. Maximally 7 filaments present. Stems tapering, longer 

than filaments. Filaments digitiform, shorter than notopodial 

cirri. 

Median acicular lobes distally rounded to triangular or 

conical with aciculae emerging at midline. Pre- and postsetal 

lobes low, transverse folds. Ventral cirri tapering, thick. 

Notopodial cirri long, slender, tapering, without articulations. 

Shafts of compound hooks inflated. Appendages short, 

tapering, with large heads, bidentate. Proximal teeth triangular, 

directed laterally. Distal teeth smaller than proximal teeth, 

nearly erect. Guards symmetrically sharply pointed, mucronate. 

Pseudocompound falcigers and compound spinigers absent. 

Aciculae paired, black. Subacicular hooks presumably black 

and bidentate. 


unknown. 


Species too poorly known to make meaningful predictions. 



4-6, 12, 16, 39, 40, 42, 53-60, 65-68, 74, 75, 78, 

80-82. 


None. 

REMARKS.—The species was originally named and briefly 

characterized by Marion in Filhol (1885:199) as having 

parchment-like tubes attached to stony corals from the Golfe de 

Gascogne, collected during the 1880 cruise of the Travailleur; 

also specifically mentioned were the eyeless condition of these 

specimens. Roule (1896:446) redescribed the species on 

material from the same area, taken during the Caudan cruises in 

1895. He quoted Marion in Filhol as the author of the species; 

nevertheless, he proceeded to describe it as new. Clearly, 

Marion in Filhol (1885) is the author of the species, with Roule 

(1896) as first revisor. 

The species is incompletely known, and without access to 

the types or material from the type locality, it is impossible to 

decide if it is valid. It has been considered a synonym of E. 

harassii and of E. pennata (Fauvel, 1923:399-401). Based on 

the available information it is placed in group B-2 and is listed


with similar species in Table 27; it has not been included in the 

key. 

Eunice anceps Pruvot, 1930 

Eunice anceps Pruvot, 1930:69. 

Eunice afra.—Fauvel in Pruvot, 1930:69 [in part, not Eunice afra Peters, 1854]. 

REMARKS.—Eunice anceps was named in a list of synonyms 

of E. afra in a paper written by Pruvot, a paper that Fauvel 

edited and issued after Pruvot's death. Pruvot was named as the 

author of the species. The epithet has never been used as a valid 

name for the taxon and is thus not available as a name. The 

species is described below as E. pruvoti; the new name honors 

the original author of the species, G. Pruvot. 

10. Eunice annulicornis Johnston, 1865 

FIGURE 9; TABLE 27 

Eunice annulicornis Johnston, 1865:131-132.—Baird in Johnston, 1865:340. 

MATERIAL EXAMINED.—Holotype, BM(NH) ZK 1962.6.1. 

Spinola N.Sy. 23 Fcb 1821 (see comment below). 


145 setigers, last 33 in regeneration; total length 135 mm; 

maximal width 4 mm; length through setiger 10, 12.5 mm. 


peristomium, as deep as V2 the peristomium. Prostomial lobes 

frontally rounded, dorsally inflated; median sulcus deep. Eyes 

not observed. Antennae in a transverse row, evenly spaced, 

similar in thickness. Ceratophores ring-shaped in all antennae, 

without articulations. Ceratostyles tapering, with up to 13 

moniliform articulations in A-III; articulations limited to distal 

half of styles. A-I to middle of peristomium; A-II to setiger 1; 

A-III to setiger 2. Peristomium cylindrical. Separation between 

rings visible only dorsally between bases of peristomial cirri; 

anterior ring 3 A of total peristomial length. Peristomial cirri to 

middle of prostomium, slender and digitiform, with 6 articulations. 

Jaws not observed. 

Branchiae present, pectinate (Figure 9e), distinctly shorter 

than notopodial cirri in most setigers, longer than the 

notopodial cirri in the few setigers around setiger 15, not 

reduced in mid-body region, erect. Branchiae from setiger 3 to 

setiger 134, missing in 12 last (regenerating) setigers. Branchiae 

present to near posterior end, present on more than 65% 

of total number of setigers. Last 2-3 pairs single filaments; all 

other branchiae with at least two filaments. Maximum 9 

filaments from about setiger 15. Length of branchiae decreasing 

rapidly from about setiger 25; numbers of filaments 

reduced to 3-4; this number retained for rest of branchial 

region. Stems tapering, about as long as filaments. Filaments 

distinctly shorter than notopodial cirri, digitiform. 

Median acicular lobes distally truncate with aciculae 

emerging dorsal to midline. Pre- and postsetal lobes low, 

transverse folds. Ventral cirri tapering from narrow bases 

through setiger 5. Median ventral cirri basally inflated. Bases 

ovate or spherical, narrow tips digitiform. Inflated bases 

reduced from about setiger 30. Ventral cirri digitiform in 

posterior x h of body. All notopodial cirri slightly inflated 

basally, articulated in all setigers with 3-4 distinct articulations 

in most segments. 

Limbate setae slender. Shafts of pectinate setae (Figure 9c) 

wide, flat. Blades flaring, flat. One marginal tooth distinctly 

longer than other teeth, with -15 teeth. Shafts of compound 

falcigers (Figure 9b) inflated, with distinct internal striations, 

marginally smooth; distal beaks present. Appendages short, 

with wide bases, bidentatc. Proximal teeth shorter than distal 

teeth, broadly triangular, directed laterally. Distal teeth gently 

curved, directed laterally. Guards symmetrically bluntly 

pointed, without mucros. Pscudocompound falcigers and 

compound spinigcrs absent. Aciculae paired, black, tapering, 

gently curved; cross-section round. Subacicular hooks (Figure 

9d) black, bidentatc. Hooks first present from setigcr 35, 

present in all setigers thereafter, always single. Hooks distally 

rapidly tapering. Both teeth of similar size, directed obliquely 

distally. 


anal cirri not described; jaws not described. 


Mx III long, located between Mx II and Mx IV, but not part of 

arc with left Mx IV. 



Features: 4, 6, 13, 14, 75, 78. 


REMARKS.—The locality information on the label reads as 

indicated above. Johnston did not know to what locality the 

name Spinola referred. Johnston's 1865 work was issued 

posthumously, and Baird wrote a set of "Addenda" and 

"Corrigenda" to the volume printed as part of the volume; his 

comment (Baird in Johnston, 1865:340) reads: 

This is most probably not a British species. It is the E annulicornis of 

Maximilian Spinola, but not described by him. It is very likely, from the label 

upon the bottle, to have been sent to the Museum by Spinola, and is perhaps a 

native of the Gulf of Genoa. 

Eunice annulicornis agrees in most respects with E. torquata 

Quatrefages; Fauvel (1923:401) suggested that they might be 

synonymous. These two species arc the only ones listed in 

Table 27 to have moniliform articulations in the ceratostyles. 

They differ from each other in some minor details of the setae, 

including the presence of marginal teeth along the upper end of 

the shaft in E. torquata; such dentition is absent in E. 

annulicornis. The peristomial cirri reach the anterior end of the 

peristomium in E. torquata and to the middle of the 

prostomium in E. annulicornis. The pectinate setae arc tapering

NUMBER 523 55 

.02 

.1 

1 

b. c 

d 

FIGURE 9.—Eunice annulicornis (holotype, BM(NH) ZK 1962.6.1): a, anterior end, lateral view; b, compound 

falciger, parapodium 77; c, pectinate seta, parapodium 77; d, subacicular hook, parapodium 77; e, parapodium 77, 

anterior view. (Scale bars in mm.) 

in E. torquata and mildly flaring in E. annulicornis. Finally, the 

subacicular hooks are occasionally paired in E. torquata and 

always single in E. annulicornis. All these differences are 

minor and may well fall within the range of variability of a 

single species. Johnston's name predates Quatrefages', because 

Quatrefages' book was not issued until 1866. However, if the 

two are found to be synonymous, Quatrefages' name is to be 

preferred for two reasons: It is well known and has been


frequently used and it has a known type locality. Johnston's 

name does not appear to have been used separately. 

11. Eunice antarctica Baird, 1869 

Eunice antarctica Baird, 1869:348. 

Eunice pennala.—Hattman 

1776]. 


1964:118 [in part, not Eunice pennata Miiller, 

MATERIAL EXAMINED.—Three syntypes, BM(NH) ZB 

1972.70, Antarctic Seas, Erebus and Terror Southern Seas 

Expedition. 

COMMENTS ON MATERIAL EXAMINED.—The type material 

consists of three syntypes and two posterior fragments. Two of 

the three syntypes are incomplete. The third syntype is 

complete and is described in detail below. 

DESCRIPTION.—Syntype complete, of unknown sex, with 

122 setigers; total length 102 mm; maximal width 3.5 mm; 

length through setiger 10, 11 mm. 

Prostomium (Figure 10a) distinctly shorter than peristomium, 

about as wide as peristomium, less than x li as deep as 

peristomium. Prostomial lobes frontally rounded, dorsally 

inflated; median sulcus shallow. Eyes not observed. Antennae 

in a shallow horseshoe, evenly spaced, similar in thickness. 

Ceratophores ring-shaped, without articulations. Ceratostylcs 

digitiform, with up to 10 irregular, distinct or very indistinct 

cylindrical articulations in A-III. A-I to end of peristomium; 

A-II to setiger 1 (setiger 3 in another syntype); A-III to setiger 

3 (setiger 4 in another syntype). Peristomium cylindrical. 


anterior ring 4 /s of total peristomial length. Peristomial cirri to 

posterior end of prostomium, slender and digitiform, without 


Jaws not examined. 

Branchiae (Figure 10b) present, pectinate, distinctly shorter 

than notopodial cirri, not reduced in mid-body region, erect. 

Branchiae from setiger 3 to setiger 39-44. Branchiae terminating 

well before posterior end, present on less than 55% of total 

number of setigers. First 6-7 and last 2 branchiae with single 

filaments. Maximum 5 filaments at about setiger 15. Stems 

slender, cylindrical, shorter than filaments. Filaments shorter 

than notopodial cirri, tapering. 


emerging at midline. Pre- and postsetal lobes follow outline of 

acicular lobes closely. First 3 ventral cirri digitiform, about half 

as long as notopodial cirri, becoming basally inflated from first 

branchial setigers. Inflated bases nearly spherical in most of 

branchial region, retaining digitiform, distally truncate tips. 

Inflated bases gradually lost in postbranchial region; last 

30-40 ventral cirri slender, digitiform, increasing in length 

posteriorly, remaining less than half as long as notopodial cirri 

in all segments. Anterior notopodial cirri slender, medially 

inflated, usually with 1-2 articulations. Median and posterior 

notopodial cirri (Figure lOg) increasingly slender, digitiform in 

postbranchial region; articulations increasingly poorly marked; 

finally represented only by steps in width of cirri. 

Limbate setae rather slender, marginally serrated. Shafts of 

pectinate setae (Figure lOd) wide, cylindrical. Blades tapering, 

flat. One marginal tooth much longer than other teeth, with -10 

rather coarse teeth. Shafts of compound falcigcrs (Figure 10c) 

barely inflated; marginally smooth; distal beaks distinct. 

Appendages thick, bidentatc, with very short teeth. Proximal 

teeth much smaller than distal teeth, triangular, directed 

laterally. Distal teeth gently curved, directed laterally. Guards 

symmetrically bluntly pointed, marginally serrated, without 

mucros. Pseudocompound falcigcrs and compound spinigcrs 

absent. Aciculae usually paired, yellow, slender, tapering, 

gently curved ventrally (Figure 10c); cross-section round. 

Subacicular hooks (Figure lOf) yellow, bidentatc. Hooks first 

present from setiger 31-44, present in all sctigcrs thereafter, 

always single (except for replacements). Hooks slender, 

tapering smoothly, with very large heads. Proximal teeth larger 

than distal teeth; both teeth directed distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Jaw structure; 

pygidium and anal cirri. 


Mx III long, but forming distal arc with left Mx IV. 



Characters: 4, 6, 13, 14. 


None. 

REMARKS.—Eunice antarctica has been considered a synonym 

of E. pennata (Hartman, 1964:118). The two species can 

be separated on several features. Eunice antarctica has a 

maximum of five branchial filaments in branchiae that are 

shorter than the notopodial cirri; E. pennata has up to 12 

filaments in branchiae that are distinctly longer than notopodial 

cirri. In E. antarctica the notopodial cirri have at least one 

distinct articulation throughout the body, but lack distinct 

cirrophores; in E. pennata the notopodial cirri are articulated in 

prebranchial segments; they lack all articulations in the 

branchial region and have a distinct basal cirrophore in 

postbranchial setigers. Eunice antarctica is listed with similar 

species in Tables 19 and 21. It is the only species in Table 21 

with branchiae shorter than notopodial cirri and with the 

notopodial cirri articulated throughout the body. 

Baird (1869) suggested that his new species resembled E. 

havaica, one of Kinberg's species; the latter has strongly 

moniliform antennal articulations and tridentate subacicular 

hooks. Eunice antarctica has cylindrical antennal articulations 

and bidentate subacicular hooks.

NUMBER 523 57 

FIGURE 10.—Eunice antarctica (syntype, BM(NH) ZB 1972.70): a, anterior end, lateral view; b, parapodium 21, 

anterior view; c, compound falciger, parapodium 21; d, pectinate seta, parapodium 21; e, acicula, parapodium 

106; f, subacicular hook, parapodium 106; g, parapodium 106, anterior view. (Scale bars in mm.) 

12. Eunice antennata (Lamarck, 1818) 

FIGURE 11; TABLES 2, 46. 47 

Leodice antennata Lamarck, 1818:322.—Savigny, 1820:49-50, pi. 5: fig. 

1.1-27. 

Eunice antennata.—Audouin and Milne Edwards, 1833:219. 

MATERIAL EXAMINED.—BM(NH) ZH 1869.7.8.6, Gulf of 

Suez, presented by R. McAndrew, Esq. 

COMMENTS ON MATERIAL EXAMINED.—The type material is 

lost; the largest specimen in ZH 1869.7.8.6 is here described in 

detail; variability is indicated in Table 2. These specimens were 

collected a few years after the opening of the Suez Canal. It 

appears unlikely that the fauna would have been extensively 

modified compared to the one present during Napoleon I's 

expedition to Egypt. The material studied by Savigny and 

Lamarck was collected during the latter expedition. 

DESCRIPTION.—Specimen complete, of unknown sex, with 

98 setigers; total length 54 mm; maximal width 3 mm; length 

through setiger 10, 7 mm. 

Prostomium (Figure lla) nearly as long as anterior peristomial 

ring, distinctly narrower than peristomium, about as deep 

as 72 the depth of peristomium. Prostomial lobes frontally

58 


.01 

.02 

.1 

1 

b, c 

(1 

e 

FIGURE 11.—Eunice antennata (BM(NH) ZH 1869.7.8.6, large specimen): a, anterior end, lateral view; b, 

compound falciger, parapodium 66; c, pectinate seta, parapodium 66; d, subacicular hook, parapodium 66; e, 

parapodium 66, anterior view. (Scale bars in mm.) 

Peristomium cylindrical; lower lip inflated and muscular, 

Separation between peristomial rings distinct dorsally and 

vcntrally; anterior ring 3 A of total peristomial length. Peristo- 

mial cirri to middle of anterior peristomial ring, tapering, with 

3 long articulations, 

Jaws missing. 

rounded, dorsally flattened; median sulcus deep. Eyes posterior 

to bases of A-I. Antennae in a shallow horseshoe, evenly 

spaced, similar in thickness. Ccratophores ring-shaped in all 

antennae, without articulations. Ceratostylcs tapering, with up 

to 17 moniliform articulations in A-III. A-I to posterior 

peristomial ring; A-II to setigcr 1; A-III to setiger 3.

NUMBER 523 59 

TABLE 2.—Variable and invariable characters in the material examined of Eunice antennata (X = number of 

individuals examined; SD = standard deviation; measurements in mm). 


Maximal width 



No. of perislomial cirral articulations 

Branchial start 

Max. no of branchial filaments 

Ventral cirri inflated through setiger 

No. of notopodial articulations 


Subacicular hooks first present from setiger 

N 

6 

6 

6 

6 

6 

6 

6 

6 

6 

6 

Max. 

3.5 

7.5 

29 

4 

7 

11 

33 

5 

15 

29 

Min. 

2 

4 

16 

3 

5 

4 

24 

2 

12 

21 

Mean 

2.58 

5.50 

21.17 

3.67 

6.20 

7.50 

30.33 

3.83 

12.50 

26.33 

SD 

0.66 

1.18 

4.49 

0.52 

0.84 

2.26 

3.27 

0.98 

1.22 

3.33 


Length of perislomial cirri 

Reduced branchial section 

Pectinate setae 

Shaft of compound falcigers 

Acicular color 

Aciculartips 

Subacicular color 


No. of subacicular teeth 

N=6 

to middle of peristomium 

present 

flared and flat 

inflated 

yellow 

blunt or pointec1 

yellow 

invisible 

3 


cirri, reduced in mid-body region, erect. Branchiae from 

setiger 7 to setiger 92. Branchiae present to near posterior end, 

present on more than 65% of total number of setigers. All 

branchiae except the last 3 with at least 2 filaments; first 

branchia with 4 filaments, maximum 7 relatively short 

filaments. Stems short, tapering. Filaments (Figure lie) no 

longer than notopodial cirri. Number of filaments reduced to 2 

or 3 in setigers 50 to 70, increasing to 4 in setigers 75-85, 

thereafter branchiae decreasing rapidly both in size and 

numbers of filaments, absent in last 6 setigers. 

Anterior neuropodial lobes truncate; posterior acicular lobes 

somewhat more conical, but retaining broadly transverse shape 

in all setigers; aciculae emerging near midline. Pre- and 


tapering, becoming basally inflated from about setiger 5. 

Inflated bases relatively modest, ovate, lost by setiger 35; 

narrow tips digitiform. Posterior ventral cirri increasingly 

slender, elongated towards the posterior end, digitiform, and 

longer than notopodial cirri in last 10-15 setigers. All 

notopodial cirri basally inflated, with 2-3 long, cylindrical 

articulations. Notopodial cirri increasing by one-third in length 

in early branchial setigers, decreasing in length towards 

posterior end, finally being as long as in first setigers. 

Limbate setae slender, marginally smooth. Shafts of pectinate 

setae (Figure lie) flat, wide. Blades flat, flared. One 

marginal tooth longer than other teeth, with -15 teeth. Shafts of 

compound falcigers (Figure 1 lb) strongly inflated, marginally 

smooth; distinct beaks small. Appendages thick, with distinct 

bosses on cutting edge near base, bidentate. Proximal teeth 

directed laterally, slightly narrower than distal teeth. Distal 

teeth thick, curved. Guards asymmetrically bluntly pointed, 

without mucros. Pseudocompound falcigers and compound 

spinigers absent. Aciculae paired, yellow, tapering, blunt, 

straight, cross-sections round. Separation between core and 

sheath indistinct in both aciculae and subacicular hooks. 

Subacicular hooks (Figure 1 Id) yellow, tridentate with teeth in 

a crest. Hooks first present from setiger 25, present in all 

setigers thereafter, always single (except for replacements). 

Fangs decreasing evenly in size from large main fang to small 

tertiary fang. 




Mx III long, straight and located behind left Mx II. Distal arc 

absent; Mx VI absent. Anal cirri long, with moniliform 




Characters: 4, 6,42 


None. 

REMARKS.—As noted in the introduction, Lamarck is the 

author of both the genus Leodice and this species. Savigny was 

clearly responsible for recognizing, describing, and illustrating 

the species, but did not publish his description until 1820; in the 

meantime, Lamarck and Cuvier both had taken advantage of 

their access to his manuscript. 

Eunice antennata was described from the Gulf of Suez and 

has been reported from worldwide areas. Restriction of the 

species implied by the definition given above excludes a large 

number of morphs reported from other areas. Eunice antennata


is listed with similar species in Tables 46 and 47. Four species 

in Table 47 have reduced branchiae in a median region. In 

addition to E. antennata, this group includes E. margariticacea, 

E. oliga, and E. lorresiensis. Eunice margariticacea has 

tridentate appendages in the compound hooks; the other species 

have bidentate appendages. Eunice oliga has very poorly 

developed branchiae with a maximum of three filaments; the 

two other species have seven filaments where the branchiae are 

best developed. The ceratophores are long in E. torresiensis 

and are short and ring-shaped in E. antennata. Other 

differences can be seen by comparing the illustrations of the 

two species. 

Eunice antennata aedificatrix Monro, 1933 

Eunice antennata aedificatrix Monro, 1933:60-61. 

REMARKS.—Originally described as a subspecies, this 

morph is here considered sufficiently different from E. 

antennata to be treated as a distinct species. 

Eunice antennata gracilis Grube, 1866 

Eunice antennata gracilis Grube, 1866a:65. 

REMARKS.—This "variety" is distinctly different from E. 

antennata and is here treated separately as E. gracilis. 

13. Eunice antillensis Ehlers, 1887 


Eunice antillensis Ehlers, 1887:84-85, pi. 24: figs. 5-7 [in part]. 

MATERIAL EXAMINED.—Lectotype, one syntype of E. 

antillensis, MCZ 810, Blake sta 45, 25°23'N, 84°21'W, 185 m. 

COMMENTS ON MATERIAL EXAMINED.—The original material 

of E. antillensis consists of three specimens. The one 

described here was illustrated by Ehlers (1887, pi. 24: fig. 6) 

and is here selected as lectotype for the species; one incomplete 

specimen consisting of -35 setigers and devoid of antennae, 

branchiae, and all but the broken stubs of the setae; and one 

complete specimen described below as a new species, E. 

stanleyi. 

DESCRIPTION.— Lectotype an incomplete female with 62 

setigers; total length 30 mm; maximal width 3 mm; length 

through setiger 10, 7 mm. Body thick, cylindrical, anteriorly 

truncate. 

Prostomium (Figure 12b) distinctly shorter and narrower 

than peristomium, as deep as l /2 of the peristomium. Prostomial 

lobes frontally truncate, dorsally flattened; median sulcus 

shallow. Eyes outside bases of A-II, black, overlapped by 

peristomial fold. Antennae in a deep horseshoe; A-I and II 

emerging close together, separated by a distinct gap from A-I 11, 


without articulations. Ceratostyles slender and tapering, with 

up to 24 cylindrical, somewhat irregular articulations. A-I to 

setiger 3; A-II to setigcr 10; A-I 11 to setiger 16. Peristomium 

cylindrical. Separation between rings distinct dorsally, visible 

ventrally; anterior ring ~ A /s of total peristomial length. 

Peristomial cirri reaching well beyond prostomium, as long as 

A-I, slender and digitiform, with 5 articulations. 

Maxillary formula 1 + 1, 4+4, 6+0, 4+9, and 1 + 1. 

Branchiae present, pectinate, distinctly shorter than notopodial 

cirri (Figure 12c), not reduced in mid-body region, erect. 

Branchiae from setiger 4 through setiger 36. Branchiae 

terminating well before posterior end. First and last 2 pairs 

single filaments, maximum 6 filaments. Stems tapering. 

Filaments digitiform. 


emerging dorsal to midline. Prc- and posLsetal lobes low, 

transverse folds. Ventral cirri without inflation in 3 anterior 

setigers. Anterior ventral cirri very thick, tapering from narrow 

bases. Median ventral cirri basally inflated. Bases ovale or 

spherical; narrow lips tapering. Postbranchial ventral cirri 

without basal inflations, tapering. Anterior noiopodial cirri 

digitiform, with 4 or 5 articulations. In branchial region 

notopodial cirri very long, tapering to long slender tips, with 3 

or 4 indistinct articulations. Postbranchial notopodial cirri 

(Figure 12a) considerably shorter than those in branchial 

region, with 2 or 3 articulations. 

Limbate setae slender, marginally serrated. Shafts of 

pectinate setae (Figure 12g) wide, cylindrical. Blades tapering, 

flat. Both marginal teeth longer than other teeth, with -12 teeth. 

Shafts of compound falcigers (Figure 120 slightly inflated, 

marginally indistinctly serrated, without distal beaks. Appendages 

short, bidentate; teeth prominent. Proximal teeth very 

much larger than distal teeth, tapering, directed basally. Distal 

teeth tapering, hooked, rather than smoothly curved. Guards 

symmetrically smoothly rounded, distinctly serrated basally, 


spinigers absent. Aciculae paired, yellow; cross-section of shaft 

round, distally flattened and expanded into rounded tabs 

(Figure 12d). Subacicular hooks (Figure 12e) yellow, bidentate. 


thereafter, paired in some setigers. Hooks tapering distally; 

heads large. Proximal teeth very much larger than distal teeth, 

slender, tapering, directed laterally. Distal teeth slender, 

tapering, nearly erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—All features associated 

with the posterior end, such as the shape of posterior 

parapodial features, pygidium, and anal cirri; the shape of Mx 

III and its relation to left Mx IV. 


None. 



Characters: 1, 2, 4, 6, 38, 42, 67, 68. 


None. 

REMARKS.—Eunice antillensis as here defined resembles E.

NUMBER 523 61 

.02 

.5 

.5 

.1 

2 

e. f, g 

r. 

a 

FIGURE 12.—Eunice antUlensis (MCZ, syntype 1): a, parapodium 55, anterior view; b, anterior end, lateral view; 

c, parapodium 11, anterior view; d, acicula, parapodium 11; e, subacicular hook, parapodium 55; f, compound 

falciger, parapodium 55; g, pectinate seta, parapodium 55. (Scale bars in mm.) 

websteri and related species. Most species in this group have 

branchiae from setiger 3; they are present from setiger 4 in E. 

antUlensis. Eunice antUlensis is listed with similar species in 

Tables 19 and 20. It is the only species in Table 20 with 

branchiae shorter than the notopodial cirri; and is one of four 

species in this table with fewer than 10 branchial filaments; the 

other three being E. articulata, E. biannulata, and E. kobiensis. 

These three species have articulated notopodial cirri limited to 

the anterior end; E. antUlensis has the notopodial cirri 

articulated throughout the body. 

Ehlers (1887:84) remarked that his type specimens differed 

in some respects; he believed the differences to be a case of


sexual dimorphism. One of the two specimens is of indeterminate 

sex; there is little evidence in the rest of the group that 

sexual dimorphism is present and the two specimens differ in 

most features used to characterize separate species. They are, as 

indicated above, considered as such here. 

14. Eunice aphroditois (Pallas, 1788) 

FIGURE 13a-d; TABLES 27,28 

Nereis aphroditois Pallas, 1788:229-230, pi. 5: figs. 1-7. 

Eunice aphroditois.—Cuvier, 1817:525.—Ehlers. 1868:306-310, pi. 15: figs. 

23-29.—Fauvel, 1917:215-220, fig. 18 [in part], pi. 7. 

Leodice gigantea Lamarck, 1818:322.—Savigny, 1820:49-50. 

MATERIAL EXAMINED.—MNHN, Paris, La Reunion, Indian 

Ocean, coll. M. Carriere, 1910. 

COMMENTS ON MATERIAL EXAMINED.—The type material is 

lost; it came from Sri Lanka; the specimen here described 

matches the original description closely. 

DESCRIPTION.—Specimen posteriorly incomplete, of unknown 

sex; with 529 setigers; total length 800 mm; maximal 

width 22 mm at setiger 25; length through setigcr 10, 55 mm. 

Body cylindrical anteriorly, becoming dorsoventrally flattened 

by setiger 200, but retaining about same width through setiger 

400, tapering from setiger 400 setiger to -10 mm in width at 

incomplete posterior end. 

Prostomium very short, distinctly narrower than peristomium, 

withdrawn inside nuchal fold, less than x /i as deep as 


inflated; median sulcus shallow. Eyes not observed. Antennae 

arranged in a horseshoe, evenly spaced, similar in thickness. 

Ceratophores ring-shaped in all antennae, without articulations. 

Ceratostyles thick, digitiform, tapering slightly, without articulations. 

Antennae to middle of peristomium; right A-II longest, 

to middle of posterior peristomial ring. Peristomium massive, 

cylindrical, about twice as wide as prostomium. Separation 

between rings distinct dorsally and ventrally; anterior ring 4 /5 of 

total peristomial length. Peristomial cirri to middle of first 

peristomial ring, thick, slightly inflated basally, without 



Branchiae (Figure 13c,d) present, pectinate, distinctly longer 


Branchiae from setiger 6 to end of fragment. All branchiae 

pectinate; maximum -30 filaments in setigers 30-100. Stems 

thick, erect, tapering, longer than notopodial cirri through 

setiger 250. Filaments slender, filiform, shorter than notopodial 

cirri. 

Anterior neuropodial acicular lobes truncate to rounded, 

becoming triangular or slightly conical in far posterior setigers; 

aciculae emerging at midline. Presetal lobes low, transverse 

folds. Postsetal lobes forming a collar around acicular lobes, 

about as high as acicular lobes in anterior and median setigers, 

projecting beyond acicular lobes in posterior setigers. Anterior 

ventral cirri thick, tapering from wide, triangular bases, not 

obviously inflated through setiger 250. Ventral cirri basally 

inflated in posterior half of fragment. Median and posterior 

bases first scoop-shaped, becoming triangular welts in last 

setigers present, retaining short, tapering tips in all setigers. 

Anterior notopodial cirri basally inflated, tapering to thick lips, 

becoming very large and very strongly inflated in posterior 

setigers (Figure 13c), by far dominant parapodial structures in 

far posterior setigers. All notopodial cirri without articulations. 

Limbatc setae longer than other setae, slender, marginally 

finely frayed. Pectinate setae (Figure 13a) in thick fascicles, 

slender; shafts flattened. Blades tapering, gently furled. 

Marginal teeth no longer than other teeth, with -15 teeth. Shafts 

of compound hooks about as thick as aciculac and subacicular 

hooks, tapering, marginally smooth; beak indistinct, brasscolored. 

Appendages not seen. Aciculac paired, with dark 

brown, rarely black cores and clear sheaths, slender, tapering, 

distally pointed, usually bent dorsally at tip; cross-section 

round. Subacicular hooks (Figure 13b) with medium to dark 

brown cores and clear sheaths, bidentatc. Hooks first present 

from setigcr 200, thereafter missing in many setigers, always 

single (except for replacements). Hooks tapering to very 

slender, indistinct heads. Proximal teeth larger than distal teeth; 

both directed distally; both teeth indistinct. 

UNKNOWN MORPHOLOGICAL FEATURES.—Features associated 

with far posterior setigers, pygidium, and anal cirri; jaw 

structure. 

EXPECTED STATE OF SELECTED UNKNOWN FEATURES.—Mx 

III and IV forming distal arc; Mx VI missing. 



Characters: 1, 2, 6, 13, 14, 36-38, 40, 57-59, 69, 70. 

ASSUMED STATES FOR PURPOSE OF PREPARING KEY.—37,1; 

38,1; 57,1; 58,2; 59,2; 69,1; 70,2. 

REMARKS.—The synonymy of this species was discussed in 

the introduction. Eunice aphroditois has been widely reported 

and a number of other named taxa has been synonymized with 

this species. The specimen described above agrees well with 

current definitions of the species; however, the description 

represents a restriction of the concept of this species compared 

to the definition given by Fauvel (1917:215); the species is 

listed with similar species in Tables 27 and 28. 

A series of species lacking articulations in the ceratostyles is 

listed in Table 28; several of these are known to have simple 

spine-like, relatively large subacicular hooks only; other 

species have relatively small, bidentate subacicular hooks. 

Included among the latter are, in addition to E. aphroditois. E. 

contingens, E. djiboutiensis, E. guttata, E. investigatoris, E. 

palauensis, E. polybranchia, E. roussaei, E. suviensis, and E. 

rullieri. Eunice sebastiani has been reported to have both 

simple falcate and bidentate subacicular hooks. Characteristic 

of E. aphroditois and related taxa are the large, medially 

inflated notopodial cirri; among the species listed above, such 

cirri are present in E. contingens, E. roussaei, and E. rullieri. 

Eunice djiboutiensis and E. sebastiani have basally inflated, 

basally distinctly pendulous, notopodial cirri. Eunice aphrodi-

NUMBER 523 63 

.02 

1 

.1 

f, y, h 

C 

FIGURE 13.—Eunice aphroditois (MNHN, Paris): a, pectinate seta, parapodium 150; b, subacicular hook, 

parapodium 260; c, parapodium 400; d, parapodium 150. Eunice arenosa (holotype, RM 426): e, anterior end, 

lateral view; f, compound falcigcr, parapodium 23; g, subacicular hook, parapodium 23; h, acicula, parapodium 


tois and E. rullieri have short antennae barely outreaching the 

prostomium and with A-II and A-III similar in length. Eunice 

contingens and E. roussaei have longer antennae; A-III is 

distinctly longer than A-II and reaches setiger 4. 

Eunice aphroditois has slender, tapering subacicular hooks 

with both teeth directed distally; E. rullieri has relatively larger 

subacicular hooks with distinct heads and the teeth directed 

laterally.


Eunice aphroditois djiboutiensis Gravier, 1900 

Eunice aphroditois djiboutiensis Gravier, 1900:224-229, figs. 73-77; pi. 13: 

figs. 63-67. 

REMARKS.—Eunice aphroditois djiboutiensis was separated 

from the stem species on the structure of the subacicular hooks. 

It differs from other members of group B-2 in the features that 

characterize distinct species and it is so considered here. 

Eunice aphroditois punctata Fishelson and Rullier, 1969 

Eunice aphroditois punctata Fishelson and Rullier, 1969:74-76, fig. 2a-e. 

REMARKS.—Originally described from the southern end of 

the Red Sea, the material currently present in the Zoological 

Museum, Tel-Aviv University, includes two specimens of a 

species related to, but different from E. aphroditois. The 

combination Eunice punctata is preoccupied; the species is 

here renamed as Eunice rullieri. 

15. Eunice arcturi (Treadwell, 1928) 

TABLES 19,20 

Leodice arcturi Treadwell, 1928:475-477, fig. 178:32-39. 

Eunice norvegica.—Hartman, 1956:283 [in part, not Eunice norvegica 

Linnaeus. 1767]. 

MATERIAL EXAMINED.—Holotype, AMNH 25-VII-1925- 

3530, Atlantic Ocean, 125° SE of New York City Hall, sta 

113:D-1. 

COMMENTS ON MATERIAL EXAMINED.—The holotype has 

been completely dried and no meaningful illustrations can be 

made. 

DESCRIPTION.—Holotype complete with 110 setigers, approximately 

80 mm long. 

Branchiae from setiger 6 to setiger 42. Branchiae terminating 


number of setigers. Maximum at least 10 filaments. 

Aciculae paired, yellow, tapering; tips conical, distinctly 

bent dorsally; cross-section round. Separation between core 

and sheath indistinct in both aciculae and subacicular hooks. 

Subacicular hooks yellow, bidentate. Hooks first present from 

setiger 36. 

UKNOWN MORPHOLOGICAL FEATURES.—All features associated 

with pro- and peristomium; jaws, parapodial features; 

limbate and pectinate setae and compound falcigers. 


The species is far too poorly known to make meaningful 

predictions. 


SCORED.—Inappropriate Characters: Too poorly known to 

determine which of the unknown characters might be inappropriate. 

Unknown Characters: 3-29, 31-34, 39, 40, 43-72, 

81, 82. 


72,2. 

REMARKS.—Eunice arcturi was considered a synonym of E. 

norvegica by Hartman (1956) after examination of the type of 

the former. This synonymy cannot be correct: the latter has 

dark subacicular hooks and branchiae continued to the posterior 

end. Eunice arcturi has branchiae terminating well before the 

posterior end and yellow subacicular hooks. The species is too 

poorly known to be adequately characterized; however, it 

differs from most members in group A-l by the late start of the 

branchiae; it is listed with other species in this group in Tables 

19 and 20. 

16. Eunice arenosa Kinberg, 1865 

FIGURE 13e-h; TABLES 19.20 

Eunice arenosa Kinberg, 1865:563.—Hartman, 1948:78. 

MATERIAL EXAMINED.—TWO syntypes, RM 426, Tahiti, 16 

m, Eugenie expedition. 

COMMENTS ON MATERIAL EXAMINED.—Kinberg 

(1865:563) gives the locality information as "Marc pacificum 

juxta urbcm Papicti insulae Tahiti, fundo arenosa 9 orgyiarum." 

One syntype is complete; the other is currently in two pieces. 

The latter specimen is here illustrated and described in detail. 

Both specimens are mature females with large eggs in the body 

cavity. 

DESCRIPTION.—Complete syntype with 79 setigers; other 

syntype incomplete, with 60 setigers, 16 mm long, maximal 

width 0.75 mm at setiger 10; length through setiger 10,2.5 mm. 

Body cylindrical anteriorly, dorsoventrally flattened posteriorly. 

Prostomium (Figure 13e) about as long as peristomium, 

about as wide as peristomium, as deep as x ji of the 

peristomium. Prostomial lobes elongated, frontally rounded, 

dorsally flattened; median sulcus shallow. Palpal region 

separated by distinct frontal, horizontal grooves. Eyes not 

observed. Antennae in a horseshoe, evenly spaced, similar in 

thickness. Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostyles slender and digitiform, with up to 8 

long, cylindrical articulations in A-III. A-I to setiger 1; A-II to 

setiger 3; A-III to setiger 5. Peristomium tapering towards 

anterior end. Separation between rings distinct on all sides; 

anterior ring x li of total peristomial length. Peristomial cirri to 

middle of prostomium, slender and tapering, with 4 articulations. 

Jaws unknown. 



from setiger 3 through setiger 25-26. Branchiae terminating 


number of setigers. First 5 and last 2-3 pairs single filaments; 

maximum 5 filaments first reached at about setiger 15. 

Filaments short, digitiform, about as long as notopodial cirri 

where best developed. 

Anterior neuropodial acicular lobes truncate, becoming 

distally rounded in posterior setigers; aciculae emerging at

NUMBER 523 65 

midline. Pre- and postsetal lobes low, transverse folds. Anterior 

ventral cirri thick and tapering. Median ventral cirri basally 

inflated from about setiger 5 through setiger 20. Bases ovate, 

narrow tips tapering. Posterior ventral cirri increasingly 

digitiform, increasing in length, nearly as long as notopodial 

cirri in far posterior setigers. Anterior notopodial cirri thick and 

tapering, decreasing in girth, but not in length posteriorly. 

Limbate setae longer than all other setae, marginally 

serrated. Pectinate setae unknown. Shafts of compound 

falcigers (Figure 130 slender, distally tapering without distinct 

marginal serrations; distinct distal beak absent. Appendages 

long, narrow; head large, bidentate. Anterior proximal teeth 

short, slender, becoming triangular in posterior setigers, 

directed slightly basally. Distal teeth longer than proximal 

teeth, curved, directed laterally. Guards narrow, symmetrical, 

slender, terminating in distinct mucros. Most guards distally 

rounded with mucro appended, but mucros appearing as 

continuations of guards proper in some setae. Pseudocompound 


single, yellow, tapering with straight, narrow tips; cross-section 

round. Some aciculae with narrow sharply pointed guards 

(Figure 13h). Separation between core and sheath indistinct in 

aciculae, distinct in subacicular hooks. Subacicular hooks 

(Figure 13g) yellow, tridentate with teeth in a crest. Hooks first 

present from setiger 16, present in all setigers thereafter, always 

single (except for replacements). Main fang curved, strongly 

tapering; secondary fang curved, rather slender; distal fang very 

poorly developed, blunt, indistinct in some posterior setigers. 

UKNOWN MORPHOLOGICAL FEATURES.—Jaw structure; 



III long, straight, located behind left Mx II. 



Characters: 1,2, 13, 14, 65-68. 


None. 

REMARKS.—A minor discrepancy with the original description 

can be noted. Kinberg found that the antennae stretched to 

segment 9 (corresponding to setiger 7); in both specimens the 

longest antenna, A-III, reaches setiger 5. One specimen had 

been dissected for the jaws, which are now missing. No 

attempts were made at examining the jaws in the other, intact 

specimen. 

Eunice arenosa is listed with similar species in Tables 19 

and 20. It can be characterized only by a combination of the 

characters used, including relatively long antennae and peristomial 

cirri, combined with a total lack of articulations of the 

notopodial cirri. 

17. Eunice argentinensis (Treadwell, 1929) 


Ltodice argentinensis Treadwell, 1929:3-5, figs. 7-12. 

MATERIAL EXAMINED.—Three syntypes, USNM 19280, 

Mar del Plata, Argentina, Felippone # 3008, coll. Florentino 

Felippone. 

COMMENTS ON MATERIAL EXAMINED.—One syntype is in 

good condition, but is in posterior regeneration; another has 

been frontally dissected for the jaw apparatus, the third 

syntype, which is currently in overall the best condition, is 

described in detail here. 

DESCRIPTION.—Specimen incomplete, of unknown sex, 

with 71 setigers; length 67 mm; maximal width 5 mm at setiger 

10; length through setiger 10, 12 mm. Body cylindrical, 

abruptly tapering frontally. 

Prostomium (Figure 14a) distinctly shorter and narrower 

than peristomium, as deep as 1 /2 of the peristomium. Prostomial 


Eyes absent. Antennae in transverse row, evenly spaced, 


without articulations. Ceratostyles digitiform, with up to 7 

short, slightly inflated articulations in A-III. A-I to posterior 

peristomial ring; A-II to setiger 1; A-III to setiger 4. 


ventrally, less so dorsally; anterior ring 3 /4 of total peristomial 


and digitiform, with 3 articulations. 

Maxillary formula 1+1,5+6,7+0,7+9, and 1+1 according to 

Treadwell (1929:5 and his fig.11). Mx III part of distal arc with 

left Mx IV; Mx VI missing. 

Branchiae (Figure 14b) present, pectinate, distinctly longer 

than notopodial cirri where best developed, not reduced in 

mid-body region, erecL Branchiae from setiger 3 to near end of 

body, present on more than 65% of total number of setigers. All 

branchiae pectinate except in regenerating posterior end of a 

complete syntype where branchiae are single filaments, 

missing in last 15 setigers in this syntype. First branchia with 5 

filaments, maximum 10 filaments, first reached at setiger 10. 

Stems thick, tapering, shorter than notopodial cirri. Filaments 

digitiform, shorter than notopodial cirri in all branchiae. 

Number and length of filaments decreasing posteriorly, from 

about setiger 30, 3-4 short filaments present. 

Neuropodial acicular lobes truncate with aciculae emerging 

dorsal to midline. Pre- and postsetal lobes low, transverse folds. 

First 3 ventral cirri thick, digitiform. Median ventral cirri 

basally inflated; inflated bases ovate; narrow tips tapering. 

Ventral cirri basally inflated through rest of fragments, without 

basal inflation only in regenerating posterior end in one 

syntype. Notopodial cirri medially inflated, with at least 1 long 

distal articulation; some anterior cirri with 2 or 3 articulations. 

Notopodial cirri large, becoming dominant parapodial feature 

in posterior segments. 

Limbate setae distinctly longer than other setae, slender, 

marginally serrated. Shafts of pectinate setae (Figure 14f) wide, 

cylindrical. Blades tapering, flat. One marginal tooth distinctly 


falcigers (Figure 14c) massive, smoothly tapering, with internal 

striations (not shown), but without marginal teeth; distal beak

66 


.02 

2 

.1 

.1 

c. f 

a 

FIGURE 14.—Eunice argeniinensis (paratype, USNM 19280): a, anterior end, lateral view; b, parapodium 52, 

anterior view; c, compound falciger, parapodium 52; d, acicula, parapodium 52; e, subacicularhook, parapodium 

52; f, pectinate seta, parapodium 52. (Scale bars in mm.) 

indistinct. Appendages small, slender; head very small, 

bidentate. Proximal teeth reduced triangular, directed laterally; 

distal teeth nearly erect; both teeth of similar size. Guards 

asymmetrical, bluntly pointed, without mucros. Pseudocompound 


paired, black; cross-section flattened, knife-shaped (Figure 

14d); thickened edge of aciculae ventral. Aciculae considerably 

heavier than subacicular hooks; tips bluntly pointed, curved.

NUMBER 523 67 

Subacicular hooks (Figure 14e) black, bidentate, slender and 

tapering. Hooks first present from setiger 19-22, present in all 

setigcrs thereafter, always single (except for replacements). 

Proximal teeth larger than distal teeth, directed laterally; distal 

teeth short, blunt, directed distally. 

UKNOWN MORPHOLOGICAL FEATURES.—Features associated 

with far posterior sctigers. 


None. 



Characters: 1, 2, 36, 37, 57-60, 63, 74, 78. 


REMARKS.—Eunice argentinensis is listed with similar 

species in Tables 27 and 29. Among the species in group B-2, 

E. argentinensis, E.floridana, and E. prognatha have at least in 

part knife-edged aciculae (£. argentinensis is the only one of 

these in Table 29). Eunice argentinensis has subacicular hooks 

first present from setigers 19-22, E. floridana has them from 

setigers 29-40, and E. prognatha from setiger 35. Eunice 

floridana has very long, slender antennae, with A-III distinctly 

longer than the other antennae; the other two species have short 

antennae where A-II and A-III are rather similar in size. In E. 

prognatha the first 8-10 branchiae are simple filaments; all 

anterior branchiae are pectinate with at least five filaments in E. 

argentinensis. 

18. Eunice armillata (Treadwell, 1922) 


Leodice armillata Treadwell, 1922:144-146, figs. 26-29, pi. 3: figs. 14-19. 

MATERIAL EXAMINED.—Holotype, AMNH V-1920-1542, 

Aua or Utile reef, Pago Pago Harbor, Samoa. 



setiger 15; length through setiger 10,6 mm; width at setiger 10, 

1.75 mm. Anterior body cylindrical, becoming dorsoventrally 

flattened medially and tapering slowly towards posterior end. 

Prostomium (Figure 15a) distinctly shorter than and narrower 

than peristomium, less than l /2 as deep as peristomium. 

Prostomial lobes frontally rounded, dorsally inflated; median 

sulcus deep. Eyes not observed. Antennae in a transverse line, 

evenly spaced, similar in thickness. Ceratophores erect cylinders 

in all antennae, without articulations. Ceratostyles tapering, 

with up to 14 moniliform articulations. A-I to posterior 


Peristomium cylindrical. Separation between rings distinct on 

all sides; anterior ring 4 /5 of total peristomial length. Peristomial 

cirri barely outreaching posterior peristomial ring, slender 

and digitiform, without articulations. 


Branchiae (Figure 15e,f) present, pectinate, distinctly longer 




of setigers. First 2 pairs single filaments, thereafter 2 filaments 

in most setigers to setiger 80; a third, short filament present 

scattered in a few setigers. From setiger 80, 1 or 2 filaments 

irregularly scattered in about equal numbers of setigers. Stems 

short, tapering. Filaments digitiform, longer than notopodial 

cirri. 

Neuropodial acicular lobes truncate with aciculae emerging 

in upper 73 of lobe. Presetal lobes transverse folds. Anterior 

postsetal lobes longer than acicular lobes, rounded, reduced to 

transverse folds from setiger 15. Ventral cirri without basal 

inflation in 5 anterior setigers; anterior ventral cirri digitiform. 

Median ventral cirri basally inflated. Bases ovate or nearly 

spherical, narrow tips tapering. Inflation rapidly reduced from 

setiger 45 and missing posterior to setiger 60. Posterior ventral 

cirri without basal inflation, becoming increasingly digitiform. 

Notopodial cirri digitiform, decreasing somewhat in length 

posteriorly, with up to 3 indistinct articulations in anterior and 

median setigers. Far posterior notopodial and ventral cirri 

similar in length and shape. 

Limbate setae longer than all other setae, coarsely limbate, 

marginally serrated. Pectinate setae (Figure 15b) very small; 

shafts cylindrical, thick. Blades distally narrowly flaring, flat. 

Both marginal teeth barely longer than other teeth, with about 

10 teeth. Shafts of compound falcigers in anterior setigers 

(Figure 15c) tapering, serrated, with distal beak. In posterior 

setigers tapering, without serrations and beaks (Figure 15d). 

Appendages very short, stocky, bidentate. Proximal teeth 

slightly shorter than distal teeth, narrowly triangular, directed 

obliquely distally. Distal teeth tapering, directed laterally, 

distinctly more curved in posterior than in anterior setigers. 

Guards asymmetrically bluntly pointed, marginally finely 

serrated, without mucros. Pseudocompound falcigers and 

compound spinigers absent. Aciculae paired, yellow, very 

heavy, distally tapering to sharp tips; inferior aciculae gently 

curved; cross-section of all aciculae round. Separation of core 


Subacicular hooks yellow, bidentate. Hooks present from 

setiger 34, present in all setigers thereafter, always single 

(except for replacements); first several slender, becoming 

increasingly heavy in posterior setigers, very coarse in far 

posterior setigers (Figure 150- Hooks slightly tapering subdistally; 

heads indistinct. Both teeth similar in size, directed 

laterally. 

UKNOWN MORPHOLOGICAL FEATURES.—Pygidium and anal 

cirri; jaw apparatus. 


III located immediately in front of left Mx IV; forming part of 

a distal arc. 



Characters: 13,14,40. 


None.

68 


b, c 

e, t 

FIGURE 15.—Eunice armillata (holotype, AMNH V-1920-1542): a, anterior end, lateral view; b, pectinate seta, 

parapodium 25; c, compound falciger, parapodium 25; d, compound falciger, parapodium 88; e, parapodium 25, 

anterior view; f, parapodium 88. (Scale bars in mm.)

NUMBER 523 69 

REMARKS.—Eunice armillata was compared to E. 

mindanavensis by Hartman (1956:282). It is listed with species 

from group A-2 (Fauchald, 1970) in Table 23, rather than with 

species in group B-2, to which E. mindanavensis belongs, 

because both subacicular hooks and aciculae are clear yellow, 

rather than brown or black. In addition to this species, £. 

leptocirris, E. schizobranchia, and E. stigmatura have branchiae 

continued to near the posterior end; the other species in 

Table 23 have the branchiae terminating posterior to setiger 

100, but well before the posterior end. The ceratostyles of E. 

leptocirris and E. schizobranchia lack articulations; the other 

two species have articulated ceratostyles. Eunice stigmatura 

has both bi- and tridentate subacicular hooks, and resembles 

species listed in Table 46 more closely than the species in Table 

22. Eunice armillata has only bidentate subacicular hooks and 

is furthermore characterized by a low number of branchial 

filaments compared to most other species in the group. 

19. Eunice articulate Ehlers, 1887 

FlOURE 16a-e; TABLES 19, 20 

Eunice articulala Ehlers, 1887:83, pi. 24: figs. 8-10. 

MATERIAL EXAMINED.—Two syntypes, MCZ 140 and 799, 

Blake, 3 Apr 1869, off French Reef, Cast No. 1, 15 fathoms. 

COMMENTS ON MATERIAL EXAMINED.—The two syntypes 

do not belong to the same taxon; the specimen from MCZ 140 

belongs to group B, probably to subgroup 2, the one from MCZ 

799 to group A-1. The species has been considered related to E. 

websteri (= E. longicirrata Webster) and Ehlers description 

matches the specimen from MCZ 799; thus the following 

description is based on this specimen. The specimen from MCZ 

140 is briefly characterized at the end of this description. The 

syntype from MCZ 799 previously has been dry. 

DESCRIPTION.—Syntype incomplete, of unknown sex, with 

132 setigers; length about 50 mm; width more than 3 mm; 

length through setiger 10 not determined. Body cylindrical, 

dorsoventrally flattened medially. 


than peristomium, less than V2 as deep as peristomium, 

bent dorsally. Prostomial lobes frontally truncate, dorsally 

inflated; median sulcus deep. Eyes lateral to bases of A-I. 

Antennae in transverse row, evenly spaced; A-I 11 distinctly 

heavier than other antennae. Ceratophores ring-shaped in all 

antennae, without articulations. Ceratostyles slender and 

tapering; articulations cylindrical, becoming increasingly dropshaped 

distally; maximum 17 articulations in A-III. A-I to 

setiger 5; A-II to setiger 7; A-III to setiger 11. Peristomium 

cylindrical. Separation between rings distinct only dorsally; 

anterior ring 3 A of total pcristomial length. Peristomial cirri 

reaching well beyond prostomium, about as long as A-I, 

extremely slender, nearly filiform, with 6 very long articulations. 

Maxillary formula 1+1, 5+6, 5+0, 4+7, and 1+1. 

Branchiae (Figure 16e) present, pectinate, distinctly longer 



terminating well before posterior end, present on less than 55% 

of total number of setigers. First and last 2 or 3 pairs single 

filaments; maximum 6 filaments at about setiger 15. Branchiae 

and filaments shorter than notopodial cirri in anterior branchial 

region; both longer than notopodial cirri in posterior branchial 

setigers. Stems thick, tapering, longer than filaments. Filaments 

digitiform, slender in anterior setigers. 

Shape of neuropodia indeterminable. Median ventral cirri 

basally inflated; postbranchial ventral cirri slender, digitiform. 

Anterior notopodial cirri long, slender, nearly filiform, with 3 

cylindrical articulations, much longer than branchiae. Median 

and postbranchial notopodial cirri short, tapering, without 


Limbate setae straight, narrowly limbate. Pectinate setae 

numerous. Shafts of pectinate setae (Figure 16d) cylindrical, 

slender. Blades tapering, flat. One marginal tooth longer than 

other teeth; 15 teeth present. Shafts of compound falcigers 

(Figure 16c) tapering, marginally smooth, without internal 

striations or distal beak. Appendages large, with parallel sides; 

basal boss distinct; head large, bidentate. Proximal teeth 

triangular, directed laterally. Distal teeth larger than proximal 

teeth, thick, directed obliquely distally. Guards symmetrically 

bluntly pointed, marginally serrated, without mucros. Pseudocompound 

falciger and compound spinigers absent. Aciculae 

yellow, 2-3 in a parapodium, distally pointed, straight; 

cross-section round. Separation of core and sheath indistinct in 

both aciculae and subacicular hooks. Subacicular hooks (Figure 

16b) yellow, bidentate. Hooks first present from setiger 40, 

present in all setigers thereafter, always single (except for 

replacements). Hooks tapering smoothly to tips; heads small. 

Proximal teeth larger than distal teeth, directed laterally. Distal 

teeth tapering, directed obliquely distally. 

Syntype from MCZ 140 frontally deeply dissected, with 80 

setigers; length 55 mm. Maxillary formula 1+1,4+4, 6+0,4+7, 

and 1+1. Branchiae from setiger 5 to end; all branchiae 

pectinate with erect stems. Shafts of pectinate setae slender, 

cylindrical; blades distinctly flaring, flat. One marginal tooth 

longer and thicker than the other teeth; -25 teeth. Aciculae 

paired, black, distally bent. Subacicular hooks from setiger 32, 

black, bidentate. 

UKNOWN MORPHOLOGICAL FEATURES.—Characters associated 

with posterior end; relationship between Mx III and left 

Mx IV; neuropodial structures. 


III long, straight, located behind left Mx II; distal arc absent; 

Mx VI absent. 



Characters: 1,2,4-6,43-49, 51, 52, 54, 55. 


None.

70 


.02 

.5 

.1 

2 

b. c. 0, t, g. i 

R 

h 

a 

FIGURE 16.—Eunice articulata (syntype, MCZ 799) a, anterior end, lateral view; b, subacicular hook, 

parapodium 40; c, compound falciger, setigcr 40; d, pectinate seta, parapodium 78; e, parapodium 78, anterior 

view. Eunice atlantica (holocype, RM 325) f, compound falciger, median parapodium; g, subacicular hook, 

median parapodium; h, aciculae, median parapodium; i, appendage of compound falciger, parapodium 9. (Scale 

bars in mm.)

NUMBER 523 71 

REMARKS.—Eunice articulata, as described above, belongs 

to group A-l and agree in most details with E. websteri; it is 

listed with similar species in Table 19. The maxillary formula 

given by Ehlers (1887:83) matches neither of the 2 syntypes. 

The other syntype clearly belongs to group B and, judging 

from the development of the branchiae in the last setigers 

present, probably to subgroup 2 (forms with branchiae beyond 

setiger 100). This specimen was seen by Ehlers: a note in his 

handwriting detailing some of the features of the specimen, 

including the branchial distribution and the maxillary formula, 

was present in the vial with the specimen. The specimen is not 

identifiable because details of the anterior end cannot be 

determined and is here considered an unidentifiable member of 

the genus. 

Eunice articulata may be separated from other species in the 

group by having branchiae from setiger 4 rather than from 

setiger 3; these branchiae are short, with thick, stubby 

filaments. In addition the antennae and peristomial cirri are 

unusually long, slender and strongly articulated, without being 

moniliform. It is listed with similar species in Tables 19 and 20. 

Eunice articulata (Hoagland, 1920) 

Leodice articulata Hoagland, 1920:615-616, pi. 50: figs. 9-12. 

Eunice articulata.—Hartman, 1959:316. 

REMARKS.—The specific name used by Hoagland (1920) is 

pre-occupied in the combination Eunice articulata Ehlers, 

1887. The species described by Hoagland, clearly different 

from Ehlers' species, is here re-named E. hirschi (see below). 

20. Eunice atlantica Kinberg, 1865 

FIGURE 16f-i; TABLES 41, 42 

Eunice atlantica Kinberg, 1865:563; 1910:42, pi. 16: fig. 18.—Hartman, 

1948:79. 

MATERIAL EXAMINED.—Holotype, RM 325, Praya Grande 

near Rio de Janeiro, Eugenie Expedition 305. 


consists of 4 fragments, one of which is an anterior end. The 3 

other fragments consists of 9 segments, 5 segments, and a 

single segment; these fragments must be from the median to 

posterior part of the body in that they lack branchiae and have 

subacicular hooks, assuming a common configuration of 

features. The shape of the compound hooks, aciculae, and 

limbate setae indicate that the median fragments came from a 

specimen of the same species as the anterior fragment, perhaps 

from the same specimen. 

DESCRIPTION.—Anterior fragment with 9 setigers, of unknown 

sex; length 4 mm; width 2 mm. 

Prostomium distinctly shorter than peristomium. Prostomial 

lobes frontally rounded, dorsally flattened; median sulcus 

shallow. Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostyles tapering, with up to 5 cylindrical 

articulations. A-I to middle of peristomium; A-II to setiger 1; 

A-III now broken (to segment 5 according to Kinberg, 1865). 


all sides; anterior ring 2 /3 of total peristomial length. Peristomial 

cirri to posterior third of prostomium, slender and 

tapering, with 3 cylindrical articulations. 


Branchiae from setiger 3; first branchiae single; maximum 5 

digitiform filaments. 

Parapodial shapes indeterminable. 

Limbate setae longer than other setae, slender, marginally 

serrated. Pectinate setae not seen. Shafts of compound falcigers 

(Figure 160 inflated, marginally coarsely serrated; distal beak 

present. Appendages long, narrow, tapering, bidentate. Proximal 

teeth short, reduced triangular, directed basally. Distal 

teeth slender, distally hooked, tapering, directed laterally. 

Guards distally symmetrically rounded, with distinct, slender 

mucros (Figure 16f,i), marginally smooth. Pseudocompound 

falcigers and compound spinigers absent. Aciculae paired, 

yellow, tapering, straight or gently curved (Figure 16h). 

Subacicular hooks (Figure 16g) yellow, tridentate with teeth in 

a crest. Primary fangs large, curved basally; 2 distal fangs with 

common stem; tertiary fangs very small. Distribution of 

subacicular hooks unknown. 


with mid-body and posterior parts of the body; jaw 

structure. 


The species is too poorly known to yield meaningful 

predictions. 


SCORED.—Inappropriate Characters: None known. Unknown 

Characters: 1, 2, 4-6, 11-17, 32, 34, 36-38, 40, 

42-59, 61-68, 80-82. 


37,2; 38,2; 53,1; 57,2. 

REMARKS.—The anterior end is poorly preserved. Kinberg 

stated that up to 12 branchial filaments should be present, but 

this cannot presently be confirmed. The description of the 

subacicular hooks given above assumes that the median 

fragments present belong to the same species as the anterior 

end. 

The label in the vial is marked Eunice atlantica (var). 

Kinberg (1865:563) described a variety of his new species, 

which should differ from the main form basically in the reduced 

number of branchial filaments. This variety should be present 

in sandy and rocky shores near the mouth of La Plata. The 

locality information given for the specimen examined here 

matches that given for the main form, and the specimen is here 

considered the type of the species. The difference between the 

two forms is minor and it is here assumed that both (lots of) 

specimens belonged to the same species.


Eunice atlantica is listed with similar species in Tables 41 

and 42; it is too poorly known to be included in the key. 

Eunice attenuata Grube, 1866 

Eunice attenuata Grube, 1866b:68; 1878a:99. 

REMARKS.—No material is available. It was originally 

characterized as having branchiae from setiger 6 through 

setiger 83 (of a total of more than 350 setigers) and with a 

maximum of 11 filaments. A-III was described as reaching 

setiger 2 (segment 4). The species was also characterized as 

resembling E. prayensis. Grube (1878a) grouped the species 

with those that had branchiae terminating in midbody, without 

articulations in the ceratostyles, and without mucronate hoods 

on the compound hooks. 

The available information is clearly inadequate to characterize 

the species and it is here considered indeterminable. 

21. Eunice aucklandica Averincev, 1974 

FIGURE 17a-f; TABLES 41,42 

Eunice aucklandica Averincev. 1974:172-173, pi. 35: figs. 9-12. 

MATERIAL EXAMINED.—Holotype, ZML 1/15807, Antarctic 

Ocean, 51°13'1"S, 165°37'7"E. OB sta 74, 6 Apr 1956, 183 m 

depth, coll. Averincev, Ushakov, Belijaev. 


79 setigers; total length 18 mm; maximal width 1 mm at setiger 

10; length through setiger 10, 3.2 mm. Body cylindrical 

throughout, anteriorly truncate, tapering posteriorly. Anal cirri 

articulated, as long as last 10 setigers. 


than peristomium, as deep as l li of the peristomium. Prostomial 

lobes large, frontally rounded, slightly dorsally flattened; 

median sulcus deep. Eyes lateral to bases of A-II, large. 

Antennae in horseshoe, evenly spaced, similar in thickness. 


Ceratostyles digitiform, with up to 8 large, short cylindrical 

articulations in A-III. A-I to second peristomial ring; A-II to 

setiger 2; A-III to setiger 3. Peristomium cylindrical. Separation 

between rings distinct on all sides, especially well marked 

dorsally and ventrally; anterior ring 3 /4 of total peristomial 

length. Peristomial cirri to middle of first peristomial ring, 

slender and tapering, without articulations. 


Branchiae present, palmate, distinctly longer than notopodial 

cirri, not reduced in mid-body region. Branchiae from setiger 8 

through setiger 21. Branchiae terminating well before posterior 

end, present on less than 55% of total number of setigers. Most 

branchiae single, digitiform filaments; some setigers with 

paired filaments. Filaments longer than notopodial cirri in 

middle of branchiated region. 

Anterior neuropodial acicular lobes broadly rounded, becoming 

slightly triangular in far posterior setigers; aciculae 

emerging at midline (Figure 17f). Pre- and postsetal lobes low, 

transverse folds. Anterior ventral cirri thick, tapering, becoming 

basally inflated from about setiger 6. Inflated bases ovate, 

gradually reduced from setiger 25, narrow lips tapering. 

Posterior ventral cirri thick, abruptly tapering. Notopodial cirri 

basally slightly inflated, tapering to digitiform tips, without 


Limbate setae marginally smooth. Shafts of pectinate setae 

(Figure 17e) narrow, cylindrical. Blades tapering, flat. Both 

marginal teeth thicker and slightly longer than other teeth, with 

about 7 teeth. Shafts of compound falcigers (Figure 17b,d) 

distally inflated, marginally smooth; distal beaks absent. 

Appendages short, slightly tapering with large heads, bidentate 

or tridentate (Figure 17b,d). Proximal teeth narrow, tapering, 

directed laterally. Distal teeth about as long as proximal tooth, 

curved; in some setae third, small teeth present, making distal 

end of appendage appear nearly furcate. Guards symmetrically 

bluntly pointed, marginally smooth, without mucros. Pseudocompound 

falcigers and compound spinigcrs absent. Aciculae 

at least paired in all sctigcrs, yellow, distally tapering, 

straight, bluntly pointed. Separation between cores and sheaths 

indistinct in both aciculae and subacicular hooks. Subacicular 

hooks (Figure 17c) yellow, tridentate with teeth in a crest. 


thereafter, paired in most setigers. Primary fangs large, curved. 

Both distal fangs emerging from joint, thick bases, short, 

triangular. 




III long and located behind left Mx II. 


SCORED.—Inappropriate Characters: 34, 56, 58, 59. Unknown 

Characters: 39, 40,42. 


None. 

REMARKS.—Eunice aucklandica is listed with similar 

species in Tables 41 and 42. A characteristic feature is the 

presence of paired subacicular hooks in most setigers. The 

maximum number of branchial filaments is only two; all other 

species in Table 42 have at least twice as many and have 

distinct branchial shafts; a branchial shaft is missing in E. 

aucklandica. 

Eunice auriculata Treadwell, 1900 

Eunice auriculata Treadwell, 1900:342-343; 1901:196-197, figs. 33-36b. 

Eunice ariculata Hartman, 1959:308 [lapsus]. 

REMARKS.—The species was first named and recognizably 

described in a short paper in which the characteristic rounded 

smooth glandular organ attached ventrally on the notopodia 

was mentioned. A more complete description was published 

the following year and has been considered the original

NUMBER 523 73 

FIGURE 17.—Eunice aucklandica (holotype, ZML 1/15807): a, anterior end, lateral view; b, compound falciger, 

parapodium 40; c, subacicular hook, parapodium 40; d, compound falciger, parapodium 50; e, pectinate seta, 

parapodium 50; f, parapodium 40, anterior view. Eunice australis (holotype, MNHN, A.l(R.)-1868-no. 56a): g, 

anterior end, lateral view; h, parapodium 21, anterior view; i, compound falciger, parapodium 21; j, parapodium 

60, anterior view; k, aciculae, parapodium 60; 1, subacicular hook, parapodium 60. (Scale bars in mm.) 

description by most authors. Hartman (1959:308) indicated that 

Treadwell misspelled the specific name in the first publication; 

this is incorrect. 

Eunice auriculata belongs to the genus Euniphysa as this 

genus is currently defined and is here newly referred to that 

genus.


22. Eunice australis Quatrefages, 1866 

FIGURE 17g-l; TABLES 41,43 

Eunice australis Quatrefages, 1866:321-322.—Fauchald. 1986:244-245, figs. 

9-14. 

Eunice leuconuchalls Bcnham, 1900:21-22. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, A.1(R.)- 

1868-no. 56a, New Zealand, coll. Quoy and Gaimard. 


109 setigers, last 10 in regenerating posterior end; total length 

70 mm; maximal width 4 mm at setiger 15; length through 

setiger 10,9 mm; width at setiger 10,8.5 mm. Body cylindrical 

anteriorly, dorsoventrally flattened posteriorly, truncate anteriorly, 

tapering slowly posteriorly. 

Prostomium (Figure 17g) distinctly shorter than peristomium, 

about as wide as peristomium, as deep as x h of the 


slightly flattened; median sulcus deep. Eyes not observed. 



Ceratostyles slender and tapering; articulations cylindrical 

basally, increasingly drop-shaped or moniliform distally, 

especially in A-II and A-III; maximum 15 articulations in A-III. 

A-I to middle of peristomium; A-II to setiger 1; A-III to setiger 

3. Peristomium cylindrical. Separation between rings distinct 

dorsally and ventrally; anterior ring 3 A of total peristomial 

length. Peristomial cirri to front x h of peristomium, slender and 

digitiform, with 7 articulations. 


Branchiae (Figure 17h) present, pectinate, distinctly longer 



terminating well before posterior end, on less than 55% of total 

number of setigers. Last 2 pairs single filaments; all other 

branchiae with at least 2 filaments. Maximum 8 filaments by 

setiger 10. Stems erect, slender, tapering, longer than filaments. 

Filaments slender, digitiform, about as long as notopodial cirri. 

Anterior neuropodial acicular lobes symmetrically truncate, 

becoming increasingly obliquely rounded posteriorly with 

aciculae emerging superior to midline. Pre- and postsetal lobes 

low, transverse folds. Ventral cirri digitiform in pre- and 

post-branchial setigers, basally inflated with short, strongly 

tapering tips in branchial region (Figure 17h,j). Inflated bases 

ovate. Prebranchial notopodial cirri digitiform, basally somewhat 

inflated, with 2 long articulations. Notopodial cirri of 

branchial region more distinctly basally inflated without 

articulations. Postbranchial notopodial cirri increasingly slender, 

retaining similar lengths in all setigers. 

Limbate setae longer than all other setae, slender, marginally 

serrated. Shafts of pectinate setae cylindrical, slender. Blades 

flat, flaring. Both marginal teeth longer than other teeth, with 

-12 teeth. Shafts of compound falcigers (Figure 17i) distally 

inflated, marginally very finely serrated; distal beak distinct. 

Appendages short, relatively thick, tapering; head small, 

bidentate. Proximal teeth reduced triangular, directed laterally 

or slightly basally. Distal teeth longer and thicker than 

proximal teeth, abruptly tapering distally, directed laterally. 

Guards asymmetrically bluntly pointed, marginally serrated, 

without mucros. Pscudocompound falcigers and compound 

spinigers absent. Aciculae (Figure 17k) paired, yellow, similar 

in size and shape, tapering, blunt-tipped, bent dorsally; 

cross-section round. Separation between core and sheath 


hooks (Figure 171) yellow, tridentate with teeth in a cresL 


thereafter, always single (except for replacements). Primary 

fangs very large, triangular, curved basally. Distal fangs 

emerging from common base; tertiary fangs very small, closely 

appresscd to secondary fangs. 

UKNOWN MORPHOLOGICAL FEATURIS.—Pygidium and anal 

cirri; relationship between Mx III and left Mx IV. 


Mx III long; located behind left Mx II. 



Characters: 13, 14. 


None. 

REMARKS.—Eunice australis is listed with similar species in 

Tables 41 and 43. It has notopodial articulations limited to the 

anterior end; other species in Table 43 either have all 

notopodial cirri articulated or they lack articulations completely. 

Justification for accepting the synonymy of E. leuconuchalis 

with E. australis as originally suggested by Ehlers (1907) has 

been given elsewhere (Fauchald, 1986:252-253). 

Eunice badia Grube, 1878 

Eunice badia Grube. 1878b:148-149. pi. 9: fig. 4. 

REMARKS.—Grube had only a single specimen, from the 

Philippines, which now is lost. Grube's description is summarized, 

with updated terminology, below. 

Prostomial antennae with distinct, but not moniliform 

articulations. A-III longest, apparently reaching setiger 2. 

Peristomial cirri short, vaguely articulated. Maxillary formula 

apparently 1+1, 4+4, 6+0, 3+8, and 1+1. Branchiae from 

setiger 8, with up to 19 filaments, illustrated as having a strong 

branchial stem and slender filaments. Ventral cirri pad-shaped 

posterior to setiger 5; notopodial cirri short. Aciculae black; 

subacicular hooks black and bidentate. 

Eunice badia clearly belongs to group B-2 and resembles E. 

aphroditois and related species. Without access to type material 

the species is however indeterminable. 

Eunice balfouriana (Mclntosh, 1885) 

FIGURE 18a 

Nicidion balfouriana Mclntosh, 1885:301-303, figs. 58, 59, pi. 39: figs. 4-6; 

pi. 20A: figs. 17, 18.

NUMBER 523 75 

• 02 

, d 

a 

FIGURE 18.—Eunice balfouriana (holotype, BM(NH) ZK 1885, 12.1.209): a. anterior end, lateral view. Eunice 

barvicensis (BM(NH) ZK 1885.12.1.203a): b, anterior end, lateral view; c, subacicular hook, parapodium 75; d, 

compound falciger, parapodium 75. (Scale bars in mm.) 

MATERIAL EXAMINED.—Holotype, BM(NH) ZK 

1885.12.1.209, off Kermadec Islands, 29°55'S, 178°14'W, 978 

m, volcanic mud; Challenger sta 170, 14 Jul 1874. 

COMMENTS ON MATERIAL EXAMINED.—A. Muir (in litt.) 

pointed out that the official station list for the Challenger states 

that the substrate should be rock, rather than volcanic mud; in 

addition, two positions were given, one as quoted above, the 

other 29°45'S, \1%°WV1, 1184 m, rock. 

The holotype is in two pieces, one anterior end, which has 

been dissected frontally, and a median piece of about 33 

setigers. The two pieces do not match each other. The jaws are 

missing. 

DESCRIPTION.—Anterior end with 17 setigers; length 7.2 

mm; maximum width 2.0 mm. Body cylindrical, truncate 

anteriorly. 

Prostomium (Figure 18a) slightly narrower than peristo-

76 


mium, short, posteriorly overlapped by nuchal fold. Prostomial 

lobes frontally rounded, dorsally inflated; median groove deep. 

Antennae slender, tapering. A-I and II articulated with long 

basal articulation and shorter distal articulations; A-III without 

articulations. A-I with 5 articulations; A-I I with 4. A-I to setiger 

1; A-II and III to setiger 5. Peristomium cylindrical; anterior 

ring 4 /5 of total peristomial length. Separation between rings 

distinct dorsally and ventrally. Peristomial cirri to front edge of 

peristomium, digitiform, without articulations. 

Maxillary formula (rewritten from Mclntosh) 1+1,7+6,8+0, 

2+3, and 10+11. Mx-III part of distal arc with left Mx-IV; 

Mx-VI missing. 

Branchiae absent. 

Neuropodial acicular lobes symmetrically triangular. All 

presetal lobes low folds; postsetal lobes about as long as 

acicular lobes, rounded. Ventral cirri tapering through setiger 8, 

thereafter basally swollen in remaining setigers. Notopodial 

cirri basally inflated, without articulations, less than half as 

long as width of body. 

Limbate setae present. Pectinate setae with -15 teeth. 

Aciculae black, single. Subacicular hooks not present in 

anterior fragment. 

Median fragment not matching anterior end; at least 20 

setigers missing. Fragment abranchiate, with relatively short, 

tapering notopodial cirri; ventral cirri basally inflated. All 

segments with black, bidentate subacicular hooks in addition to 

tapering black aciculae and limbate and pectinate setae. 

REMARKS.—The anterior fragment may belong to the same 

specimen as the median fragment, but too many segments are 

missing to justify describing the median fragment as belonging 

to the same species. The distribution of the subacicular hooks 

remains unknown. Eunice balfouriana differs clearly from all 

other abranchiate species in having short second ring on the 

peristomium and in the development of the antennae. In 

abranchiate species (often described as Nicidion) the two 

peristomial rings are similar in length and antennae are short, 

barely projecting beyond the tip of the prostomium. 

The jaw apparatus is currently missing. The very high 

number of teeth reported by Mclntosh for Maxilla V is, as far 

as known, completely unique in the family. Eunice balfouriana 

is here considered indeterminable. 

23. Eunice barvicensis Mclntosh, 1885 

FIGURE 18b-d; TABLES 24,26 

Eunice barvicensis Mclntosh, 1885:292-294, figs. 52, 53. pi. 39: fig. 12, pi. 

21A: figs. 1-3. 

MATERIAL EXAMINED.—One syntype, BM(NH) ZK 

1885.12.1.203a, Bermuda, between tidemarks, June 1873. 

DESCRIPTION.—Syntype incomplete with 112 setigers; 

length 41 mm; maximal width 3.5 mm at setiger 10. Length 

through setiger 10, 6 mm. Syntype frontally dissected, at one 

time desiccated. Body apparently cylindrical, anteriorly truncate. 

Prostomium (Figure 18b) distinctly shorter than peristomium, 

about as wide as peristomium, less than x h as deep as 

peristomium. Prostomial lobes large, frontally rounded, dorsally 

flattened; median groove deep. Eyes large, lensed, 

between bases of A-I and A-II. Antennae poorly preserved, in 

a horseshoe, evenly spaced, similar in thickness. Ceratophores 

long in all antennae, without articulations. Ceratostyles 

tapering, apparently not articulated (but see Mclntosh, 

1885:292-293). A-I to posterior margin of peristomium; A-II 

to setiger 1; A-III to setiger 3. Peristomium short, tapering 

towards anterior end (artefact); anterior ring ~ 3 A of total 

peristomial length. Separation between rings distinct dorsally 

and ventrally. Peristomial cirri to front edge of pcrisiomium, 

tapering, without articulations. 

Jaws present in vial. Maxillary formula 1+1, 5+5, 5+0, 5+8, 

1+1. Mx III very short and forming distal arc with left Mx IV 

and V. Only 3 teeth on left Mx IV well developed; others low 

tubercles. 

Branchiae present, pectinate, about as long as notopodial 

cirri, not reduced in mid-body region, erect. Branchiae from 

setiger 6 to setiger 65, terminating well before the posterior 

end. First 3 and last 10 pairs single filaments; maximum 4 

filaments at setigers 25-30. Branchial stems short, tapering. 

Filaments shorter than notopodial cirri, short, flattened. 

Anterior neuropodial acicular lobes symmetrically rounded, 

becoming distinctly conical by setiger 60; aciculae emerging at 

midline. Pre- and postsetal lobes low folds. Ventral cirri 

digitiform through setiger 4. Ventral cirri basally inflated from 

setiger 5 through about setiger 25. Inflated bases ovate; narrow 

tips digitiform. Posterior ventral cirri without basal inflation, 

digitiform. All notopodial cirri digitiform, without articulations. 

Far posterior notopodial and ventral cirri similar in shape 

and size. 

Limbate setae present. Up to 10 pectinate setae in a 

parapodium in posterior setigers. Shafts of pectinate setae 

slender, cylindrical; blades flat, flared. Both marginal teeth 

longer than other teeth; number of teeth 15. Shafts of 

compound falcigers (Figure 18d) distally inflated, without 

serrations; distal beak distinct. Appendages short, tapering; 

head large, bidentate. Proximal teeth triangular with narrow 

base, directed laterally; distal teeth as large as proximal teeth, 

tapering, directed laterally. Guards symmetrically bluntly 

pointed, marginally serrated; mucros absent Pseudocompound 

falcigers and compound spinigers absent. Aciculae single in 

anterior and posterior parapodia, paired in median parapodia, 

light brown in anterior and posterior setigers, dark brown in 

median setigers, pointed, straight; cross-sections round. Subacicular 

hooks (Figure 18c) with dark brown cores and clear 

sheaths, bidentate. Hooks first present from setiger 19, present 

in all setigers thereafter, paired in some setigers. Hooks 

distinctly bent distally, with narrow necks and large heads.

NUMBER 523 77 

Proximal teeth larger than distal teeth, directed laterally, 

distally slightly upturned; distal teeth sharply pointed, directed 

distally. 

UKNOWN MORPHOLOGICAL FEATURES.—Features associated 

with pygidium, anal cirri, and far posterior setigers. 

EXPECTED STATES OF UNKNOWN MORPHOLOGICAL FEA- 

TURES.—None. 




ASSUMED STATES FOR PURPOSE OF PREPARING KEY — 

None. 

REMARKS.—Eunice barvicensis was considered a synonym 

of E. mutilata by Hartman (1944:113); however, it has 

branchiae present only through setiger 65, leaving nearly half 

of the body abranchiate; branchiae are missing only in the last 

15-20 setigers in E. mutilata; this number represents less than 

15% of the total body length. Eunice barvicensis is listed with 

similar species in Tables 24 and 26. Other species in this group 

with branchiae from setiger 6 include E. samoae and E. 

coccinea. Both these species have articulated ceratostyles; E. 

barvicensis lacks such articulations. The shape of the subacicular 

hooks resembles the shape present in E. cariboea and 

similar abranchiate species listed in Tables 33 and 40, in that 

the proximal tooth is tilted up, rather than strongly beaked as is 

characteristic of E. mutilata and related species. 

24. Eunice bassensis Mclntosh, 1885 

FIGURE 19; TABLES 46, 47 

Eunice bassensis Mclntosh, 1885:298, pi. 39: fig. 16, pi. 21A: figs. 8, 

9.—Fauchald, 1986:245, figs. 15, 16. 

Eunice anlennata.—Hartman, 1959:309 [in part, not Eunice antennata 

Lamarck, 1818]. 


1885.12.1.207, off East Montcoeur Island, Bass Strait, South 

Australia, 39°10'S, 146°37'E, 2 Apr 1874, 69-73 m, dredged, 

sand, Challenger sta 162. 

COMMENTS ON MATERIAL EXAMINED.—The holotype was 

reported as anteriorly incomplete when first described. The 

type locality was corrected by Mr. A. Muir (in litt.) 

DESCRIPTION.—Holotype posterior fragment of 55 setigers 

of which last 22 in regeneration; 50 mm long; 3 mm wide; 

apparently middle and last third of very large specimen. The 

integument is light colored and contains a number of whitish 

granules. 

Branchiae (Figure 19b) present except in last 3 setigers, 

longer than notopodial cirri, erect. All branchiae branching 

except in regenerating portion where either 1 or 2 filaments 

present. Maximal number of filaments 9; most branchiae with 

6-8 filaments through setiger 33 of fragment. Stems erect, 

longer than filaments, tapering. Filaments about as long as 

notopodial cirri, digitiform. 

Posterior neuropodial acicular lobes truncate with aciculae 

emerging in midline. Pre- and postsetal lobes low folds. Ventral 

cirri moderately basally inflated, becoming digitiform in 

posterior setigers. Notopodial cirri tapering, with up to 5 

distinct articulations. 

Compound falcigers with bidentate appendages; guards 

symmetrically bluntly pointed, without mucros. Pseudocompound 


paired, yellow, tapering; tips pointed; cross-section round. 

Separation between core and sheath indistinct in both aciculae 

and subacicular hooks. Subacicular hooks (Figure 19a) yellow, 

tridentate with teeth in a crest. Hooks tapering towards distinct 

head; teeth decreasing evenly in size distally, proximal teeth 

directed laterally. 


with prostomium and peristomium; jaw structure; 

distribution of branchiae and subacicular hooks; structure of 

anterior and median parapodia. 

EXPECTED STATE OF UNKNOWN MORPHOLOGICAL FEA- 

TURES.—A-I through A-III with moniliform articulations. Mx 

III long, located behind left Mx II; distal arc absent. Branchiae 

from approximately setiger 6; with reduced number of 

filaments in mid-body region. Subacicular hooks present from 

about setiger 25. 



Characters: 1-29, 33, 35, 36, 39, 40, 42, 45, 46, 48, 49, 

51-56,61,62,65-68,80-82. 


REMARKS.—Eunice bassensis was considered a junior 

synonym of E. antennata by Hartman (1959:309); it resembles 

that species in having tridentate subacicular hooks and 

branchiae continued to near the posterior end; however, the 

very strong articulation of the posterior notopodial cirri, the 

shape of the branchiae, and the subacicular hooks differ as do 

the shape of the aciculae. 

Eunice bassensis is listed with similar species in Tables 46 

and 47; it is too poorly known to be included in the key. 

Eunice bellii Audouin and Milne Edwards, 1833 

Eunice bellii Audouin and Milne Edwards, 1833:223-224, pi. 10: figs. 1-4, 

8-9, table 27. 

Marphysa bellii.—Quatrefages, 1866:333-334. 

REMARKS.—Eunice bellii was referred to the genus Marphysa 

by Quatrefages (1866). As far as known, there has never 

been any doubt that it belongs to that genus as currently is 

defined. The earlier use of the name Eunice bellii in Cuvier 

(1830:200) is a nomen nudum, thus the name was available for 

use by Audouin and Milne Edwards (1833).

78 


FIGURE 19.—Eunice bassensis (holotype, BM(NH) ZK 1885.12.1.207): a, subacicular hook, posterior 

parapodium; b, posterior parapodium, anterior view. (Scale bars in mm.) 

25. Eunice benedicti (Verrill, 1885) 


Leodice benedicti Verrill. 1885:427-428. 

Eunice benedicti.—Hartman, 1942:52, figs. 88-90. 

MATERIAL EXAMINED.—Lectotype, YPM 2729, south of 

Cape Cod, Massachusetts, 40°03'48"N, 70°45'54"W, 130 m, 

Fish-Hawk sta 922, 16 Jul 1881 (1 specimen). 

COMMENTS ON MATERIAL EXAMINED.—This specimen was 

listed as catalog 60-3 in Hartman (1942). It must have been 

exposed to strong alcohol at one time, because it now appears 

slightly dehydrated. The two localities listed by Verrill 

(1885:428) are Fish Hawk stations 865 and 943. Hartman 

(1942:52) found material from stations 922 and 1032 in the 

Pcabody Collections; all specimens are from the same area and 

similar depths. The specimen from station 922 is not, strictly 

speaking, part of the type series, because neither station 922 nor 

station 1032 were mentioned in the original description. 

However, one label in YPM 2729 is in Verrill's handwriting 

and names the species as benedicti; thus this specimen was 

identified by Verrill. It came from a set of stations studied by 

Verrill in preparation for the publication of his new species; for 

that reason, it is here considered part of the original type series 

and eligible to be considered a lectotype. 

The specimens from Fish Hawk station 1032 seen by 

Hartman (1942) are cataloged in our collections as USNM 

14362. USNM 14361, quoted by Hartman for the specimen 

from station 922, must at one time have been transferred to the 

Peabody Museum; the number is correctly present in the 

ledgers, but the corresponding card and the specimen are no 

longer part of our collections. 

DESCRIPTION.—Complete mature female, with 105 setigers; 

total length 38 mm; maximum width 2.5 mm at setiger 15. 

Length through setiger 10, 5 mm; width at setiger 10, 2 mm. 

Body cylindrical throughout, tapering from anterior to poste-

NUMBER 523 79 

FIGURE 20.—Eunice benedicti (holotype, YPM 2729) a, anterior end, lateral view; b, parapodium 76, anterior 

view; c, compound falciger, parapodium 4; d, pectinate seta, parapodium 76; e, compound falciger, parapodium 

76; f, subacicular hook, parapodium 32; g, subacicular hook, parapodium 76; h, acicula, parapodium 76; i, 

parapodium 32. (Scale bars in mm.)

SO 

rior end. Prostomium (Figure 20a) distinctly shorter than and 

narrower than peristomium, less than V2 as deep as peristomium. 

Prostomial lobes frontally rounded, inflated dorsally; 

median sulcus shallow. Eyes faded, posterior to the bases of 

A-I. Antennae in a horseshoe, evenly spaced, similar in 


articulations. Ceratostyles tapering, without articulations. A-I 

to second peristomial ring; A-II to setiger 3; A-I 11 to setiger 4. 


all sides; anterior ring roughly 2 /3 of total peristomial length. 

Peristomial cirri to middle of prostomium, tapering, without 


Jaws not examined. Hartman (1942:52) reported maxillary 

formula as 1+1, 6+7, 9+0, 5+11, and presumably 1+1. 

Branchiae (Figure 20i) present, pectinate, distinctly longer 

than notopodial cirri, not reduced in mid-body region, flexible. 



of total number of setigers. Three first pairs single filaments; all 

other branchiae with 2 or more filaments; maximum number 

12, at or near setiger 15. Number of filaments maintained near 

maximum number to 3 segments before end of branchiated 

region. Branchial stems longer than filaments, flexible, 

tapering. Filaments longer than notopodial cirri, tapering 

distally to slender, nearly translucent tips. 

Anterior neuropodial acicular lobes symmetrically truncate; 

median acicular lobes asymmetrically triangular, with aciculae 

emerging superior to midline; posterior acicular lobes nearly 

trapezoidal with aciculae and the subacicular hooks supporting 

distal corners (Figure 20b). All pre- and postsetal lobes low 

transverse folds. First 4 ventral cirri thick, tapering from narrow 

bases. Ventral cirri basally inflated from about setiger 5; 

inflated bases thick transverse welts in all remaining setigers. 

Narrow tip of ventral cirri digitiform in anterior and median 

setigers, becoming reduced from about setiger 40 and 

completely missing in last 45 setigers. Remnant thick pad-like 

structure moved to posterior face of parapodia visible only as a 

modest pad-like swelling on postero ventral edge of neuropodia 

in far posterior setigers. Anterior notopodial cirri medially 

inflated, increasing in length through setiger 15, thereafter 

decreasing through remainder of branchial region, retaining 

approximately same shape. Far posterior notopodial cirri short 

and digitiform. Notopodial cirri without articulations. 

Limbate setae more than twice as long as other setae in 

anterior and median setigers, clearly longer than all other setae 

in all setigers, slender, marginally smooth, very narrowly 

limbate, appearing nearly capillary. Pectinate setae (Figure 

20d) very long, narrow; shafts cylindrical, slender; blades 

tapering, flat. One marginal tooth distinctly longer than other 

teeth; total number about 10. Shafts of compound falcigers 

(Figure 20c ,e) extremely long compared to length of appendages, 

distally inflated with serrated margins; distal beak 

present. Appendages short, tapering, bidentate. Proximal teeth 

triangular, directed laterally, larger in posterior than in anterior 


setigers; distal teeth larger than proximal teeth, tapering, 

slightly less curved in posterior than in anterior setigers. 

Guards symmetrically sharply pointed with distinct short 

mucros, marginally serrated. Pseudocompound falcigers and 

compound spinigcrs absent. Aciculae paired, yellow, tapering; 

cross-section round. Superior aciculae (Figure 20h) considerably 

heavier than inferior aciculae, gently curved, sharply 

tapering. Inferior aciculae similar in shape, but distally straight. 


and subacicular hooks. Subacicular hooks (Figure 20f,g) 

yellow, bidentate. Hooks first present from setiger 21, present 


tapering, with distinct head. Proximal teeth large, slender, 

directed laterally. Distal teeth, small, nearly erect. Guards 

rounded truncate. 


cirri; relation between Mx III and left Mx IV. 

EXPECTED STATES OF UNKNOWN MORPHOLOGICAL FHA- 

TURES.—The number of teeth in Mx III and left Mx IV 

indicates that Mx III is long and located behind left Mx II rather 

than as part of a distal arc. 

CHARACTERS USED IN PREPARATION OF- KI:Y NOT 



ASSUMED STATES TOR PURPOSE OF PREPARING KKY.— 

None. 

REMARKS.—Eunice benedicti is listed with similar species 

in Tables 19 and 21. Species in this group combining branchiae 

starting from setiger 3 with a lack of articulations of the 

ceratostyles include E. hawaiensis, E. leplocirris, E. megabranchia, 

E. rubrocincta, and E. validobranchiata. All of these 

species except E. benedicti have at least two anterior pairs of 

branchiae as single filaments; all anterior branchiae are 

pectinate in E. benedicti. The branchiae are folded over, nearly 

coiled in E. benedicti, and erect in all the other species listed. In 

addition, the bases of the ventral cirri are inflated through the 

posterior part of the body in E. benedicti; the inflated bases are 

reduced in posterior setigers in all the other species. Eunice 

benedicti has mucronate compound falcigers; the compound 

falcigers are at best pointed in the other species listed. 

Eunice bertolonii (Chiaje, 1828) 

Nereis Bertoloni Chiaje, 1828:174, see also p. 163.—Grube, 1850:291.— 

Quatrefages, 1866: 354. 

Eunice Bertoloni.—Audouin and Milne Edwards, 1833:220. 

Eunice Bertoloniana.—Fauvel, 1923:451. 

REMARKS.—The species has been quoted in the genus 

Eunice due to Chiaje's placement of the species under the 

genus Leodice in his formal diagnosis (1828:174). Chiaje 

specifically mentions, both in his Latin diagnosis and in the 

Italian description, the presence of spiralled branchiae. However, 

Audouin and Milne Edwards (1833:220) stated that 

although the species "had the appearance of a Eunice" they had

NUMBER 523 81 

been unable to see that the branchiae were pectinate. Grube 

(1850:291) indicated that this species must belong in the 

vicinity of the onuphid genus Diopatra; similarly Quatrefages 

(1866:354) indicated that he was inclined to refer it to 

Diopatra. Fauvel (1923:451) in a table listed the species as 

Eunice Bertoloniana [sic] and referred it under doubt to 

Diopatra. As far as can be ascertained, no material of this 

species exist and it is here considered indeterminable. 

26. Eunice biannulata Moore, 1904 


Eunice biannulata Moore, 1904:487-490, pi. 37: figs. 10-18, pi. 38: fig. 

42.—Fauchald, 1969:2-4, fig. la-e. 

MATERIAL EXAMINED.—Holotype, CAS Type 35 (Inv. cat. 

no. 019712), San Diego, California, coll. E.C. Starks. 

DESCRIPTION.—Holotype complete, with 139 setigers; total 

length 105 mm; maximum width 3 mm at setiger 15. Length 

through setiger 10, 12 mm; width at setiger 10, 2.75 mm. Body 

slender, cylindrical throughout. 

Prostomium (Figure 21b) distinctly shorter than peristomium, 

about as wide as peristomium, as deep as x li of the 

pcristomium. Prostomial lobes frontally truncate, dorsally 

flattened; median sulcus deep. Eyes dark, posterior to and 

between A-I and A-II. Antennae in a horseshoe, evenly spaced, 


without articulations. Ceratostyles slender and tapering; basal 

V3-V2 of ceratostyle without articulations; distal portions 

increasingly articulated; final 5-6 articulations rounded, 

drop-shaped. A-I to second peristomial ring; A-II to setiger 2; 

A-III to setiger 4. Peristomium slightly flaring ventrally, with 

distinct, muscular lower lip. Separation between rings distinct 

on all sides; anterior ring 3 A of total peristomial length. 

Peristomial cirri to front edge of peristomium, basally slightly 

inflated, with 4 cylindrical articulations. 

Jaws not examined. Mandibles strongly calcified. 



from setiger 3 through setiger 50. Branchiae terminating well 

before posterior end, present on less than 55% of total number 

of setigers. First and the last 2-3 pairs single filaments, 

maximum 8 filaments at about setiger 10. Branchial stem erect, 

tapering, longer than filaments. Filaments slender and digitiform, 

about as long as notopodial cirri. 


becoming asymmetrically rounded with aciculae emerging 

superior to midline in far posterior setigers (Figure 21a). 

Pre-and postsetal lobes low, transverse folds. Anterior ventral 

cirri thick, tapering, becoming distinctly inflated basally from 

about setiger 6. Inflated bases spherical; narrow tip rather large, 

digitiform. Inflated bases decreasing posterior to setiger 90, 

indistinct in last 10-15 setigers. Far posterior ventral cirri 

digitiform. All notopodial cirri medially inflated; anterior and 

median ones with 3-4 cylindrical articulations; posterior ones 


Limbate setae about as long as compound hooks, smoothedged. 

Shafts of pectinate setae (Figure 21c) wide, cylindrical; 

blades furled, tapering. Both marginal teeth distinctly longer 

than other teeth, -10 very large teeth present. Shafts of 

compound falcigers (Figure 2If) tapering in most setigers, 

slightly inflated distally in a few setigers, marginally smooth; 

distal beak absent. Appendages thick, barely tapering; heads 

large, bidentate. Proximal teeth triangular, directed laterally; 

distal teeth shorter than proximal teeth, bent, directed laterally. 

Guards distally symmetrically rounded, marginally serrated; 

mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae (Figure 2Id) paired, yellow, tapering 

to blunt, distinctly curved tips; cross-sections round. Separation 

of core and sheath indistinct in both aciculae and subacicular 

hooks. Subacicular hooks (Figure 21e) yellow, bidentate. 


thereafter, paired in some setigers. Hooks barely tapering; head 

indistinct. Proximal teeth large, directed laterally; distal teeth 

short, narrow, directed obliquely distally. 




TURES.—Mx III long and located behind left Mx II. 





None. 

REMARKS.—Eunice biannulata was reviewed in Fauchald 

(1969); it is listed with similar species in Tables 19 and 20. In 

common with several other species in Table 20, it has 

articulated ceratostyles in which the articulations become 

bead-shaped or moniliform distally; this feature is uniquely 

exaggerated in this species compared to the other species listed 

in the table. Eunice biannulata can also be separated from other 

species in the group by the distribution and development of the 

branchiae as suggested in Fauchald (1969). 

Eunice biannulata mexicana Fauchald, 1970 

Eunice biannulata mexicana Fauchald, 1970:27-28, pi. 1: figs. f-g. 

REMARKS.—Originally described as a subspecies, this form 

is here considered a distinct species and is discussed below as 

E. mexicana. 

27. Eunice bicirrata Rullier, 1964 

TABLES 41, 42, 46-48 

Eunice bicirrala Rullier, 1964:178-180, fig. 15a-h. 

COMMENTS ON MATERIAL EXAMINED.—This species was 

described from a single specimen from the Cape Verde Islands

82 


FIGURE 21.—Eunice biannulata (hololype, CAS Type scries 35 (CAL 019712)): a, parapodium 79, anterior view; 

b, anterior end, lateral view; c, pectinate seta, parapodium 79; d, aciculae, parapodium 79; e, subacicular hook, 

parapodium 79; f, compound falciger, parapodium 79. Eunice biformicirrata (AMNH 1527): g, anterior end, 

lateral view; h, parapodium 111, anterior vie w; i, parapodium 11, anterior view; j, compound falciger, parapodium 

111; k, acicula, parapodium 111; I, pectinate seta, parapodium 111; m, subacicular hook, parapodium 111. (Scale 

bars in mm.)

NUMBER 523 83 

in 120-150 m depth. The specimen was not available; the 

description below is summarized from the original description. 

DESCRIPTION.—Holotype (and only specimen) incomplete, 

with 50 setigers; total length 12 mm; maximal width 1 mm. 

Ceratostyles slender and digitiform, articulated, especially 

distally, with long, cylindrical articulations, becoming apparently 

drop-shaped distally. A-III to setiger 10. Two pairs of 

peristomial cirri present; outer pair to posterior end of 

prostomium; medial pair somewhat shorter. 

Branchiae present, palmate. Branchiae from setiger 3 

through setiger 48. Maximum number of filaments 2. 

Limbate setae illustrated as marginally smooth. Pectinate 

setae absent. Shafts of compound falcigers inflated. Appendages 

small, linear, bidentate. Proximal teeth triangular, distal 

teeth slender, curved. Guards illustrated as symmetrically 

sharply pointed, but not mucronate. Pseudocompound falcigers 

and compound spinigers absent. Aciculae yellow, tapering to 

blunt tips and distally curved. Subacicular hooks yellow, 

trideniate with teeth in a crest. Hooks first present from setiger 

45. Two distal teeth emerging from common base. 

UKNOWN MORPHOLOGICAL FEATURES.—Shape features 

associated with pro- and peristomium; jaw structure (but see 

comments below); all parapodial features; setal distribution and 

shapes must be confirmed; pygidium and anal cirri. 


The species is too poorly known to make meaningful 

predictions. 



Characters: 1, 2, 4-19, 24-26, 28, 29, 32-34, 

37-68,74,75,78,81,82. 


None. 

REMARKS.—Rullier considered the peristomium as two 

asetigerous segments; the distribution of branchiae and 

subacicular hooks has been recalculated to the conventions 

used in this paper; thus the numbers will not agree with those 

originally stated. 

The jaws appear to be highly unusual. Rullier specifically 

stated that maxillae were absent and that the whole apparatus 

consisted of two pairs of mandibles. The illustration, seen from 

the ventral side, shows a pair of very slender forceps and a pair 

of curved, medially excavated lateral pieces fused to a narrow 

carrier in the midline posteriorly; the dorsal and distal ends of 

each half are strongly denticulated with very small, even teeth. 

These jaws resemble those of juvenile eunicids and onuphids. 

The structure of the jaws as illustrated and the other features 

specified indicate that Rullier had a juvenile specimen, perhaps 

anteriorly damaged, suggested by the doubled peristomial cirri, 

of group C-l orC-2. 

Eunice bicirrata is here considered a juvenile, possibly 

indeterminable. It is listed with similar species in Tables 41 and 

42 and in Tables 46-48. It is too poorly known to be included 

in the key. The description of a new species based on a juvenile 

does not, per se, make the taxon invalid. 

28. Eunice biformicirrata (Treadwell, 1922) 

FIGURE 21g-m; TABLES 27,32 

Leodice biformi-cirrata Treadwell. 1922:148-149, figs. 34, 35, pi. 4: figs. 

6-11. 

Eunice grubei Hartman, 1956:282 [not Eunice grubei Gravier, 1900]. 

MATERIAL EXAMINED.—Holotype, AMNH 1527, Suva, Fiji, 

Apr 1920, coll. A.L. Treadwell. 

DESCRIPTION.—Holotype complete male with 137 setigers; 

total length 217 mm; maximum width 7 mm at setiger 15. 

Length through setiger 10,17 mm; width at setiger 10,6.5 mm. 

Anterior body cylindrical with strongly convex dorsum, 

becoming ventrally flattened in median setigers and dorsoventrally 

flattened in posterior setigers. Anterior end truncate, 

tapering slowly posteriorly. 

Prostomium (Figure 21g) distinctly shorter than and narrower 

than peristomium, less than x ji as deep as peristomium. 


sulcus deep. Eyes not seen. Antennae in a horseshoe, evenly 

spaced, similar in thickness. Ceratophores ring-shaped, without 

articulations. Ceratostyles tapering; basal 73 without articulations; 

styles increasingly articulated in median l /i and 

articulations distinctly moniliform in distal x h of each antenna. 

A-I to middle of peristomium; A-II and A-III to setiger 1; 

maximally -10 articulations. Peristomium cylindrical. Separation 

between rings distinct ventrally and at bases of peristomial 

cirri; anterior ring 3 A of total peristomial length. Peristomial 

cirri to middle of first peristomial ring, thick and tapering, with 

about 7 short, moniliform articulations. 

Jaws missing; according to Treadwell, maxillary formula 

1+1,5+5, 6+0, +8, and 1+1. 

Branchiae (Figure 21h,i) present, pectinate, distinctly longer 




of setigers. First and last 5 branchiae single filaments; 

maximum number of filaments 7 reached at setiger 10, 

thereafter number of filaments decreasing slowly; by setiger 

100 (Figure 21h) only 2 filaments left. Branchial stems thick, 

tapering, shorter than filaments. Filaments digitiform. 

Anterior neuropodial acicular lobes short, asymmetrically 

truncate or rounded with aciculae emerging well above 

midline; median and posterior acicular lobes broadly asymmetrically 

truncate. All pre- and postsetal lobes low, continuous 

folds. Anterior ventral cirri digitiform in 3 first setigers. Ventral 

cirri strongly inflated from about setiger 4. Inflated bases 

spherical; narrow tips digitiform. Inflation decreasing posterior 

to setiger 60; far posterior ventral cirri digitiform. All 

notopodia tapering with 2 or 3 cylindrical articulations. 

Limbate setae longer than other setae, marginally serrated. 

Pectinate setae (Figure 211) very small; shaft cylindrical, 

narrow; blades flat, flared. One marginal tooth longer than 

other teeth; -20 very fine teeth. Shafts of compound falcigers 

(Figure 21j) tapering, internally striated, marginally smooth,


with distinct distal beak. Appendages short, tapering with 

distinct heads, bidentate. Proximal teeth curved, triangular, 

directed laterally; distal teeth longer than proximal teeth, 

tapering, curved, directed laterally. Guards asymmetrically 

bluntly pointed, marginally smooth, without mucros. Pscudocompound 


(Figure 21k) paired, black, blunt, gently curved in most 

setigers; cross-section round. Subacicular hooks (Figure 21m) 

black, bidentate. Hooks first present from setiger 27, present in 

all setigers thereafter, always single (except for replacements). 

Hooks with large heads and narrow necks. Proximal teeth 

slender, tapering, directed obliquely distally; distal teeth 

shorter than proximal teeth, but distinctly thicker, directed 

obliquely laterally. 


cirri; relation between Mx III and left Mx IV. 


TURES.—Mx III part of distal arc with left MX IV; short, 

curved. Left Mx IV distinctly shorter than right Mx IV, curved. 



Characters: 13, 14, 74, 78. 


None. 

REMARKS.—The specimen is very well preserved; it is a 

nearly mature male with sperm discs in the coelom. 

Eunice biformicirrata is listed with similar species in Tables 

27 and 32. In addition to E. biformicirrata, two species in Table 

32, E. laurillardi and E. longicirris, have the notopodial cirri 

articulated throughout the body; the other species either lack 

such articulations altogether or the notopodial cirri are 

articulated only in the anterior, usually pre-branchial and early 

branchial setigers. In E. biformicirrata, the peristomial cirri 

reach the middle of the peristomium; in both the other species 

the peristomial cirri are longer, reaching at least the middle of 

the prostomium. 

29. Eunice bilobata Treadwell, 1906 


Eunice bilobata Treadwell, 1906:1168-1169, figs.47, 48. 

MATERIAL EXAMINED.—Holotype, USNM 5212, Albatross 

sta 3871, Auau Channel, between Maui and Lanai islands, 

20°52'20"N, 156°41'35"W, 24-79 m, fine white sand, 12 Apr 

1902. 

COMMENTS ON MATERIAL EXAMINED.—The holotype consists 

of a well-preserved anterior end and three median 

fragments, which probably all derive from the same specimen; 

size and general constitution are similar, but several intermediate 

sections must be missing, because the fragments do not 

match well. The description is based on the anterior end only. 

DESCRIPTION.—Anterior fragment with 35 setigers; length 

18 mm; maximal width 6 mm; length through setiger 10,7 mm. 

Head and anterior 5-6 segments cylindrical; remainder of 

fragment dorsoventrally flattened. 


than peristomium, as deep as l /i of the peristomium, nearly 

hidden under large antennae. Prostomial lobes frontally 

obliquely truncate, dorsally flattened; median sulcus deep. 

Eyes absent. Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostylcs digitiform, with up to 12 irregular, 

indistinct cylindrical articulations in A-II. A-I to posterior 

margin of peristomium; A-II reach setiger 7; A-III incomplete; 

at least as long as A-II. Peristomium strongly flaring anteriorly; 

with distinct, muscular lower lip. Separation between rings 

distinct dorsally, possibly also ventrally, but distorted by 

overlapping fold; anterior ring 9 /io of total peristomial length. 

Peristomial cirri to middle of prostomium, digitiform, without 



Branchiae (Figure 22e) present, pectinate, distinctly shorter 


Branchiae from setiger 7, present on all fragments. All 

branchiae pectinate with up to 18 filaments. Branchial stems 

curved dorsally and twisted posteriorly in most segments. 

Filaments short, slender. 

Neuropodial acicular lobes broadly rounded; aciculae 

emerging dorsal to midline. Pre- and postsetal lobes low, 

transverse folds. First 3 ventral cirri thick, tapering, becoming 

basally inflated from setiger 4. Inflated bases thick, ovate; 

narrow tips tapering. Anterior notopodial cirri long, basally 

inflated, tapering to thick, digitiform tips; in branchial region 

notopodial cirri shorter, tapering from bases. Notopodial cirri 

with up to 7 articulations anteriorly; number of articulations 

decreasing posteriorly; median and posterior notopodial cirri 

with 3 or 4 articulations. 

Limbate setae long, narrow. Pectinate setae (Figure 22c) 

large, tapering, flat. Marginal teeth no longer than other teeth, 

about 10 long, distally tapering teeth present. Shafts of 

pectinate setae distinctly punctate. Shafts of compound 

falcigers (Figure 22b) tapering. Appendages short, bidentate. 

Both teeth similar in size. Proximal teeth directed laterally. 

Distal teeth directed obliquely distally. Guards symmetrically 

rounded, without mucros. Pseudocompound falcigers and 

compound spinigers absent. Aciculae paired, dark, superior 

aciculae bent with irregular knobs (Figure 22d); other aciculae 

tapering to straight, blunt points; cross-section of aciculae 

round. Subacicular hooks black, bidentate. Hooks first present 

from setiger 25, present in all setigers thereafter, always single 

(except for replacements). 

UNKNOWN MORPHOLOGICAL FEATURES.—Relation between 

Mx III and left Mx IV; features associated with posterior 

setigers; pygidium and anal cirri. 


TURES.—Branchiae continued to the far posterior end. 


SCORED.—Inappropriate Characters: 14, 56. Unknown

NUMBER 523 85 

FIGURE 22.—Eunice bilobata (holotype, USNM 5212): a, anterior end, lateral view; b, compound falciger, 

parapodium 32; c, pectinate seta, parapodium 32; d, acicula, parapodium 32; e, parapodium 32, anterior view. 

(Scale bars in mm.) 

Characters: 1, 2, 4, 6, 15-17, 23, 36-38, 40, 42, 57-60, 74, 

78. 


38,1. 

REMARKS.—Eunice bilobata is listed in Tables 27 and 29. 

Enough fragments are present to indicate that the branchiae 

continue beyond setiger 100, but whether the branchiae 

continue to the end of the body cannot be determined based on 

the fragmentary type. Eunice bilobata is the only species listed 

in Table 29 with knobbed, distally expanded aciculae. 

Eunice binominata Quatrefages, 1866 

Eunice binominata Quatrefages, 1866:327.—Ehlers, 1887:85. 

Eunice punctata Grube, 1856:59-60 [not Eunice punctata Risso, 1826]. 

REMARKS.—Quatrefages (1866) renamed Grube's species 

because Grube's name was preoccupied and did not see any 

material. Grube had a single specimen collected at St. Croix in 

the West Indies at his disposal. The specimen is now missing. 

Ehlers did not examine types in his review of the species. The 

following is a summary of Grube's description:


Type yellow with white punctations, with 96 setigers. 

Antennae with up to 12 long articulations in A-III. A-I to 

posterior peristomial ring; A-II to setiger 3; A-III to setiger 5. 

Branchiae from setiger 3 through setiger 27; all pectinate 

with up to 9 filaments. 

Notopodial cirri about as long as branchiae, vaguely 

articulated. 

Aciculae yellow. 

The branchial distribution combined with the information 

about the antennae and the color of the aciculae indicate that 

this species might be a member of C-l, but the information is 

clearly insufficient to identify the species and it is here 

considered indeterminable. 

30. Eunice bipapillata Grube, 1866 

Eunice bipapillata Grube, 1866a:64. 

FIGURE 23; TABLES 27,30, 31 

MATERIAL EXAMINED.—Part of holotype, ZMB F 1993b 

(marked Eunice magnified, see comment below), Samoa, 3 

parapodia. 

COMMENTS ON MATERIAL EXAMINED.—G. Hartwich (in 

litt.) informed me that parapodial preparations of E. bipapillata 

and E. magnifica originally were stored together in the same 

vial; at one time they were split apart into two vials. The three 

parapodia referred to above agree with E. bipapillata and 

cannot agree with E. magnifica due to differences in the 

distribution of branchiae and the different maximal numbers of 

branchial filaments reported by Grube for the two species. 

Therefore, it is assumed that the labels for the two vials were 

confused. The description given below is a summary of 

observations on the parapodial preparations and the original 

description. 

DESCRIPTION.—Type copper-colored with white band across 

setiger 4. 

Ceratostyles articulated. 

Branchiae (Figure 23d,h,i) present, pectinate, distinctly 

shorter than notopodial cirri, not reduced in mid-body region. 

Branchiae from setiger 4 to end of body. Branchiae present to 


of setigers. Up to 4 filaments from setiger 8 through setiger 18; 

paired filaments in setiger 77. Branchial stems short, tapering. 

Filaments digiuform. 

Anterior acicular lobes (Figure 23d,e) asymmetrically 

truncate, with aciculae emerging above midline, becoming 

slightly triangular in posterior setigers (Figure 23i). All 

pre- and postsetal lobes low folds. Ventral cirri distinctly 

inflated basally in setigers 15 and 18; inflated base lost by 

setiger 77; narrow tip digitiform in all setigers. Ventral cirrus in 

setiger 77 digitiform. All notopodial cirri similar in size and 

shape; bases slightly inflated, tapering to blunt, digiuform tips, 


Limbate setae longer than other setae, slender, narrowly 

limbate, marginally serrated. Shafts of pectinate setae (Figure 

23b,h) cylindrical, slender; blades slightly flared, flat. One 

marginal tooth longer than other teeth; up to 15 teeth. Shafts of 

compound falcigers (Figure 23a,c) tapering, marginally 

smooth; distal beak indistinct. Appendages short, relatively 

thickset with large heads, bidentate. Proximal teeth triangular, 

directed laterally or slightly distally, with distinct distal 

upturned tip; distal teeth longer than proximal teeth, tapering or 

blunt, directed obliquely distally. Guards asymmetrically 

bluntly pointed, without mucros. Pseudocompound falcigers 

and compound spinigcrs absent. Aciculae paired, black, 

tapering, straight, blunt-tipped; cross-section round. Subacicular 

hooks (Figure 230 black, bidentate. Hooks absent in 

parapodia 15 and 17, present in parapodium 77. Proximal tooth 

thicker than distal tooth, but no longer, curved, directed 

laterally. Distal tooth nearly erect, tapering. Guard short, 

truncate. 


with prostomium and pcrisiomium; jaw structure; 

distribution of branchiae and subacicular hooks. 


TURES.—Antennae not reaching beyond pcristomium; with 

cylindrical articulations or without articulations; digitiform. 

Mx III short; located in front of left Mx IV, forming part of a 

distal arc. Branchiae continued to far posterior setigers. 



Characters: 1-20, 22-29, 33, 34, 36-40, 51, 52, 74, 78, 

80-82. 


22,2; 23,4; 33,2; 37,1; 38,1. 

REMARKS.—Eunice bipapillata is listed with similar species 

in Tables 27, 30, and 31; it resembles in certain respects E. 

aphroditois, but cannot be completely identified without access 

to more material. 

Eunice bitorquata Grube, 1870 

Eunice bitorquata Grube, 1870b:56. 

Eunice siciliensis.—Grube, 1878a: 101. 

REMARKS.—The original description contains the following 

statements (K. Fauchald translation): 

The species is brown with golden to brown cross bars on the peristomium. 

Branchiae are present from setiger 104 and are present only as single filaments, 

far outreaching the notopodial cirri. Segments are remarkably short and the 

median antenna is as long as four segments. The maxillae have coarse teeth; Mx 

II has 4 teeth. 

Grube (1878a: 101) referred this species questionably to his 

own Eunice siciliensis, but gave no comment or any supportive 

statements. 

The original description is wholly inadequate to characterize 

the species; the information given is consistent with the 

synonymy suggested by Grube (1878a), but is not sufficient to 

be conclusive. Eunice bitorquata is here considered indeterminable.

NUMBER 523 87 

FIGURE 23.—Eunice bipapillata (type, ZMB F1993b): a, compound falciger, parapodium 15; b, pectinate seta, 

parapodium 15; c. compound falciger, parapodium 18; d, parapodium 15; e, parapodium 18; f, subacicular hook, 

parapodium 77; g, compound falciger, parapodium 77; h, pectinate seta, parapodium 77; i, parapodium 77. The 

parapodium numbers indicated do not appear to be entirely accurate (see text). (Scale bars in mm.)


31. Eunice borneensis Grube, 1878 


Eunice (Eriphyle) borneensis Grube, 1878a: 102. 

MATERIAL EXAMINED.—Three syntypes, ZMB F2033, 

North Borneo. 

COMMENTS ON MATERIAL EXAMINED.—All three syntypes 

are complete, one is in two pieces; they vary from 103 to 129 

setigers and from 40 to 64 mm in length. The syntype described 

is the largest of the complete syntypes. 

DESCRIPTION.—Syntype with 120 setigers; total length 42 

mm; maximum width 6 mm at about setiger 50. Length through 

setiger 10,7 mm; width at setiger 10,4.5 mm. Anterior body up 

to about setiger 30 cylindrical, becoming dorsoventrally 

flattened, strongly flattened near posterior end. 


than peristomium, less than x li as deep as peristomium. 

Prostomial lobes frontally truncate, dorsally flattened; median 

sulcus deep. Eyes not observed. Antennae in a shallow 

horseshoe, evenly spaced, similar in thickness. Ceratophorcs 


slender and tapering; maximum 6 cylindrical articulations in 

A-II and A-111. A-I to middle of peristomium; A-II and A-111 to 

setiger 2. Peristomium massive and muscular; cylindrical. 

Separation between rings distinct dorsally and ventrally, 

indistinct only in narrow section laterally; anterior ring 6 /i of 

total peristomial length. Lower lip scalloped. Peristomial cirri 

to middle of anterior peristomial ring, slender and tapering, 

with 3 indistinct, cylindrical articulations (not shown in 

illustration). 

Maxillary formula 1+1, 4+4, 7+0, 5+11, and 1+1. Teeth of 

Mx II very large and coarse compared to other teeth. 





of setigers. First branchia with 3 filaments; maximal number of 

filaments 9 by setiger 15; last 10 setigers with single filaments. 

Branchial stem as thick as filaments, short, flexible, tapering. 

Filaments digitiform, longer than notopodial cirri. 

Anterior acicular lobes (Figure 24c) broadly, asymmetrically 

truncate with aciculae emerging above midline. Median and 

posterior neuropodial acicular lobes narrowly, asymmetrically 

rounded with aciculae emerging above midline. All presetal 

lobes low truncate folds. Anterior postsetal lobes free, truncate, 

longer than acicular lobe, becoming rounded in posterior 

setigers, but remaining longer than acicular lobes in all setigers. 

Four first ventral cirri thick, tapering. Ventral cirri basally 

inflated from about setiger 5. Inflated bases thick, transverse 

welts in all remaining setigers; narrow tips of ventral cirri 

small, nearly button-shaped. All notopodial cirri basally 

inflated with distinct rounded basal tubercular protuberance, 


Limbate setae longer than other setae, slender, nearly 

straight, limbations narrow, marginally smooth. Pectinate setae 

(Figure 24g) short; shafts slender, cylindrical; blades flaring, 

fiat. One marginal tooth longer than other teeth; total of -15 

teeth. Shafts of compound falcigcrs (Figure 24e) distally 

inflated, marginally smooth, with distinct distal beak. Appendages 

short, thick, barely tapering; heads large, bidentate. 

Proximal teeth tapering, directed laterally; distal teeth about as 

long as proximal teeth, but thicker, tapering, directed laterally. 

Guards asymmetrically bluntly pointed, marginally smooth, 

without mucros. Pscudocompound falcigcrs and compound 

spinigers absent. Aciculae paired anteriorly; 3 aciculae from 

about setiger 50, black, tapering, with narrowly blunt tips 

(Figure 24d), gently curved or straight; cross-section round. 

Subacicular hooks black (Figure 241), bidentate. Hooks first 

present from sctigcr 30-35, present in all setigers thereafter, 

single in most setigers, up to 3 in a few sctigcrs. Both teeth 

directed distally; proximal teeth slightly larger than distal teeth. 

UKNOWN MORPHOLOGICAL FEATURES.—Relation between 

Mx III and left Mx IV; pygidium and anal cirri. 


TURES.—Mx III short, located in front of left Mx IV; forming 

part of distal arc. 



Characters: 13,14,74,78. 


None. 

REMARKS.—Eunice borneensis is listed with similar species 

in Tables 27 and 31. It appears to differ from other species 

listed in Table 31 especially in the presence of well-developed 

postsetal lobes, in the shape of the notopodial cirri, and in 

having the ventral cirri basally inflated to the posterior end. 

32. Eunice bottae Quatrefages, 1866 


Eunice bottae Quatrefages, 1866:320.—Grube, 1870a:292. 


1868-no. 55b, Red Sea, coll. Botta. 

DESCRIPTION.—Holotype incomplete with 60 setigers; 

length 28 mm; maximum width 2 mm at about setiger 40. 

Length through setiger 10,7 mm; width at setiger 10,1.75 mm. 

Body slender, cylindrical, becoming somewhat dorsally flattened 

towards posterior end. 


than peristomium, as deep as '/2 of the peristomium. Prostomial 

lobes irregularly rounded, dorsally flattened; median sulcus 

shallow; frontal groove marking off palpal region present. Eyes 

not seen. All antennae incomplete, except left A-I, in a shallow 

horseshoe, evenly spaced, similar in thickness. Ceratophores 


digitiform with maximum 15 moniliform articulations in A-I 11.

NUMBER 523 89 

FlOURE 24.—Eunice borneensis (ZMB F2033): a, anterior end, lateral view; b, parapodium 42, anterior view; c, 

parapodium 3, anterior view; d, acicula, parapodium 42; e, compound falciger, parapodium 42; f, subacicular 

hook, parapodium 42; pectinate seta, parapodium 42. (Scale bars in mm.)

90 


FIGURE 25.—Eunice bottae (type, MNHN, Paris, A.l(R.)-1868-no. 55b): a, anterior end, lateral view; b, 

compound falciger, parapodium 28; c, subacicular hook, parapodium 28; d, aciculae, parapodium 28; e, 

parapodium 28, anterior view. (Scale bars in mm.) 

A-I to second peristomial ring; A-II to setiger 4 and A-III to 

setiger 3. Peristomium cylindrical with flaring lower lip. 


anterior ring 5 /6 of total peristomial length. Peristomial cirri to 

middle of first peristomial ring, tapering, with 4 long, 

cylindrical articulations. 



than notopodial cirri, erect. Branchiae from setiger 6 to end of 

fragment. First and last branchiae with 3 filaments; maximum 

number of filaments 8 by setiger 15. Filaments shorter than 

notopodial cirri. Branchial stems slender, erect, tapering, longer 

than filaments. Filaments digitiform. 

All neuropodial acicular lobes distally asymmetrically 

truncate or rounded with aciculae emerging above midline. All 

pre- and postsetal lobes low folds. Anterior and posterior 

ventral cirri thick, tapering. Median ventral cirri distinctly, 

nearly spherically basally inflated, with long, distally truncate 

narrow tips. Notopodial cirri distinctly basally inflated in 

anterior and early median setigcrs, becoming tapering and 

slender towards the posterior end. All notopodial cirri with 4 or 

5 long, cylindrical articulations. 


Pectinate setae present, but not observed in detail. Shafts of

NUMBER 523 91 

compound falcigcrs (Figure 24b) slightly inflated, marginally 

serrated, with distinct distal beak. Appendages short, thick with 

short, nearly truncate heads, bidentate. Proximal teeth triangular, 

directed laterally; distal teeth larger than proximal teeth, 

bent, blunt, directed laterally. Guards asymmetrically bluntly 

pointed, marginally serrated, without mucros. Pseudocompound 


(Figure 24d) paired, yellow, tapering, with narrow, straight or 

gently curved tips; cross-section round. Separation between 

core and sheath indistinct in both aciculae and subacicular 

hooks. Subacicular hooks (Figure 24c) yellow, tridentate with 

teeth in a crest. Hooks first present from setiger 25, present in 


Teeth decreasing rather evenly in size from proximal to distal. 

All 3 teeth distinctly curved. 

UKNOWN MORPHOLOGICAL FEATURES.—Presence and location 

of eyes; jaw structure; various features associated with the 

incomplete nature of the type such as the detailed distribution 

of branchiae. 


TURES.—Mx III long, located behind left Mx II; not part of 

distal arc. 



Characters: 1,2, 13,14,16,33,36-38,40,47,50,51,64-68. 


38,1. 

REMARKS.—Eunice bottae has not been reported since the 

original description and the review of the original material 

given by Grube (1870a). It is listed with similar species in 

Tables 46 and 47. The anterior and posterior rings of the 

peristomium are completely distinct in E. bottae and E. lucei; 

in all other species in Table 47, the separation is indistinct 

laterally. The inflated bases of median ventral cirri are ovate in 

E. bottae and form thick, transverse welts in E. lucei. 

33. Eunice bowerbanki Baird, 1869 


Eunice bowerbanki Baird, 1869:349-350.—Fauchald, 1986:245-246, figs. 

17-23. 

MATERIAL EXAMINED.—Holotype, BM(NH) ZH 

1863.923.41, Australia, coll. Dr. Bowerbank. 


length 95 mm; maximal width 7 mm at about setiger 20. Length 

through setiger 10, 11 mm; width at setiger 10, 6.5 mm. Body 

anteriorly cylindrical, becoming dorsoventrally flattened. 




sulcus deep. Antennae in a transverse line, evenly spaced, 


without articulations. Ceratostyles tapering, distinctly articulated; 

inner articulations cylindrical; outermost articulations 

moniliform; maximal number of articulations 8 in A-III. A-I to 

posterior peristomial ring; A-I I and A-III to setiger 1. 

Peristomium massive, cylindrical, with distinct muscular lower 

lip. Separation between rings distinct dorsally and ventrally; 


middle of anterior peristomial ring, digitiform, with 4 



curved, forming part of distal arc with left Mx IV. Mx VI 

absent. 


than notopodial cirri, reduced in mid-body region, flexible. 



of setigers. Posterior half of branchiated setigers with single 

filaments only; first branchia with 3 filaments; maximum 

number of filaments 15 by setiger 12; number and length of 

filaments rapidly decreasing posteriorly; by setiger 25, 5-6 

short, digitiform filaments present. Branchial stems erect, 

tapering, longer than filaments. Filaments thick, digitiform, 

longer than notopodial cirri. 

Neuropodial acicular lobes triangular, nearly symmetrical. 

All presetal lobes low folds. Anterior postsetal lobe free, as 

long as acicular lobes, rounded; postsetal lobes reduced to low 

folds by setiger 20. Ventral cirri thick, digitiform in all setigers; 

basal inflation indistinct. Notopodial cirri supported by paired 

aciculae, increasing in length through setiger 3, decreasing in 

length through next several setigers. Prebranchial and early 

branchial notopodial cirri digitiform, with 2 to 3 articulations, 

tapering, becoming more pyriform; basal inflation decreasing 

posteriorly; by setiger 20 notopodial cirri, thick, digitiform, 



Pectinate setae (Figure 26e) short. Shafts coarse, flattened; 

blades tapering, flat. Both marginal teeth somewhat thicker 

than the other teeth, but no longer than other the teeth; -15 teeth 

present. Shafts of compound falcigers (Figure 260 tapering, 

with internal striations, marginally smooth; distal beak indistinct. 

Appendages short, tapering, with large head, bidentate. 

Proximal teeth triangular, directed laterally; distal teeth longer 

than proximal teeth, tapering, nearly erect. Guards asymmetrically 

bluntly pointed, without mucros. Pseudocompound 

falcigers and compound spinigers absent. Aciculae usually 

paired, black; superior aciculae slender, tapering to straight 

tips; inferior aciculae (Figure 26c) thicker than superior 

aciculae, distinctly bent ventrally; cross-section round. Subacicular 

hooks (Figure 26d) black, bidentate. Hooks first present 

from setiger 31, present in all setigers thereafter, paired in some 

setigers. Hooks tapering smoothly to tips; teeth similar in 

length; proximal teeth thicker than distal teeth; both directed 

distally; guards not observed. 

UKNOWN MORPHOLOGICAL FEATURES.—Presence and position 

of eyes; pygidium and anal cirri.


.02 

.1 

1 

5 

e 

c. d, f 

h 

^ . 7 f f c e bowerbanki (holotype, BM(NH) ZH 1863.9.23.41): a, anterior end lateral view b 

parapodium 34. antenor view; c, acicula. parapodium 34; d. subacicular hook, parapodium 34; e pectinate seta 

parapodium 34; f. compound falciger, parapodium 34. (Scale bars in mm.)

NUMBER 523 93 




SCORED.—Inappropriate Characters: 51, 54, 55, 58, 59. 

Unknown Characters: 13, 14, 74, 78. 


None. 

REMARKS—Eunice bowerbanki is listed with similar species 

in Tables 27 and 32. Of the species listed in Table 32, E. 

bowerbanki, E. laurillardi, and E. multipeclinata have 10 or 

more branchial filaments where the branchiae are best 

developed. In E. laurillardi the notopodial cirri are articulated 

throughout the body; in the other two species, only anterior 

notopodial cirri are articulated. In E. bowerbanki the appendages 

of the compound falcigcrs are tapering; the proximal teeth 

are reduced and the distal teeth nearly erect. In E. multipectinata 

the appendages do not taper appreciably; the proximal 

teeth are much larger than the distal teeth and the distal teeth are 

distinctly bent. 

Eunice brasiliensis Kinberg, 1865 

Eunice brasiliensis Kinberg, 1865:563; 1910:42. pi. 16: fig.19. 

REMARKS.—Eunice brasiliensis was not reviewed by Hartman 

(1948) nor does any material exist in Riksmuseet, 

Stockholm, where the rest of Kinberg's types were deposited 

(R. OlerOd, in litt.). 

Kinberg's description and illustrations are of a species with 

strongly moniliform, slender antennae and peristomial cirri. 

Branchiae start between setigers 4 and 10; at the latter 

segment Kinberg's illustrations shows that nine filaments are 

present. The notopodia are articulated at least through the first 

30 segments. 

The species, originally reported from Pernambuco, Brazil, in 

30-36 m depth, is not identifiable based on the descriptions 

and illustrations alone and is here considered indeterminable. 

34. Eunice brevis (Ehlers, 1887) 


Nicidion brevis Ehlers, 1887:98-99, pi. 28; figs. 9-14, pi. 29: figs. 1,2. 

MATERIAL EXAMINED.—Holotype, MCZ 833, Key West, 

Florida, 2-3 m, Blake cruises. 


been dried out at one time and is somewhat distorted. 

DESCRIPTION.—Holotype complete with 130 setigers; total 

length 18 mm; maximal width 1.5 mm wide at about setiger 10. 

Length through setiger 10, 2.5 mm. Anterior part of body 

inflated with highly convex dorsum, becoming flattened in 

posterior setigers; ventrum flattened in whole specimen. 


than peristomium, as deep as x li of the peristomium. 

Prostomial lobes irregularly truncate, dorsally somewhat 

flattened; median sulcus shallow. Eyes not observed. Antennae 

in nearly straight line, evenly spaced, similar in thickness. 


Ceratostyles digitiform, with 2 to 3 irregular, cylindrical 

articulations. All antennae to posterior peristomial ring. 

Peristomium ventrally flaring, with distinct muscular lower lip. 

Separation between rings distinct dorsally only; anterior ring 

less than 2 /3 of total peristomial length. A transverse crease 

along ventral surface not matching dorsal separation into rings. 

Peristomial cirri barely to first peristomial ring, slender and 

digitiform, without articulations. 

Jaws now missing. Maxillary formula 1+1, 5+5, 4+0, 3+4, 

and 1+1. Ehlers (1887:98, pi. 29: fig. 1) illustrated the jaws as 

symmetrical; probably Mx III form distal arc with left Mx IV. 


Anterior neuropodial acicular lobes symmetrically rounded 

or truncate, becoming symmetrically triangular in posterior 

setigers; aciculae emerging at midline. All pre- and postsetal 

lobes low folds. Ventral cirri thick, tapering. Median ventral 

cirri basally strongly inflated in setigers 8 and 28 according to 

Ehlers' illustrations; inflated bases thick, transverse welts. 

Posterior ventral cirri without basal inflation, tapering. All 

notopodial cirri short, medially inflated, without articulations. 

Limbate setae (Figure 27c) longer than other setae, narrowly 

limbate, marginally smooth, geniculate. Pectinate setae (Figure 

27e) large; shafts cylindrical, slender, blades strongly flared, 

flat. One marginal tooth longer than other teeth; about 15 teeth 

present. Shafts of compound falcigers (Figure 27b) distally 

inflated, marginally smooth, without distal beaks. Appendages 

short, slightly tapering, with large heads, bidentate. Proximal 

teeth slender, tapering, directed laterally, but curved distally. 

Distal teeth about the same size as the proximal teeth, slender, 

tapering, directed obliquely distally. Guards symmetrically 

rounded, marginally serrated, without mucros. Pseudocompound 


(Figure 27f) single, black, tapering, sharply pointed, straight; 

posterior aciculae thick; cross-section round. Subacicular 

hooks (Figure 27d) black, bidentate. Hooks first present from 


(except for replacements). Hooks with narrow neck, distinct 

head; proximal teeth larger than distal teeth, curved; distal teeth 

slender, tapering; both teeth directed distally. 

UKNOWN MORPHOLOGICAL FEATURES.—Relationship between 



TURES.—Mx III short, curved, and forming part of distal arc 

with left Mx IV. 


SCORED.—Inappropriate Characters: 31-42, 56, 58, 59. 



None. 

REMARKS.—Eunice brevis is listed with similar species in 

Tables 33 and 40. Among the abranchiate species, three have

94 


FIGURE 27.—Eunice brevis (holotype of Nicidion brevis, MCZ 833): a, anterior end, lateral view; b, compound 

falciger, parapodium 100; c, limbate seta, parapodium 100; d, subacicular hook, parapodium 100; e, pectinate 

seta, parapodium 100; f, acicula, parapodium 100. (Scale bars in mm.) 

articulated ceratostyles; in addition to E. brevis, these species 

are E. curticirris and E. fuscafasciata. Eunice curticirris has 

moniliform articulations; the other two species have cylindrical 

articulations. In E. brevis the inflated bases of median ventral 

cirri forms thick, transverse welts; in E. fuscafasciata, they are 

ovate. 

35. Eunice bucciensis (Treadwell, 1921) 


Leodice bucciensis Treadwell, 1921:54-56, figs. 174-183. 


MATERIAL EXAMINED.—Holotype, AMNH IV-1918-1286, 

Buccoo Bay, Tobago Island, Mar 1918. 

COMMENTS ON MATERIAL EXAMINED.—The anterior end 

had been deeply dissected and the illustration is somewhat 

distorted; especially, the lower lip may be expanded over the 

normal condition. 

DESCRIPTION.—Holotype incomplete, currently in 3 pieces, 

with a total of 226 setigers; anterior fragment with 66 setigers; 

length 41 mm; maximal width 3.5 mm at about setiger 15. 

Length through setiger 10, 7.5 mm; width at setiger 10, 3.25 

mm. Body anteriorly cylindrical, becoming dorsoventrally 

flattened, truncate anteriorly, tapering posteriorly. 

Prostomium (Figure 28a) distinctly shorter and narrower

NUMBER 523 95 

FIGURE 28.—Eunice bucciensis (holotype, AMNH IV-1918-1286): a, anterior end, lateral view; b, acicula, 

parapodium 58; c, subacicular hook, parapodium 58; d, parapodium 58, anterior view; e, compound falciger, 

parapodium 58; f, pectinate seta, parapodium 58. (Scale bars in mm.)


than peristomium, less than V2 as deep as peristomium. 

Prostomial lobes frontally rounded, dorsally excavate with 

thickened rim; median sulcus deep, directed ventrally. Eyes not 

observed. Antennae in a shallow horseshoe, evenly spaced; 

A-III slimmer than other antennae. Ccratophorcs long in all 

antennae, without articulations. Ccratostyles digiliform, without 

articulations. No antennae reaching beyond first peristomial 

ring; A-I shortest; A-III longest, but difference in length less 

than one-third of length of longest antenna. Peristomium flared 

anteriorly, with distinct, muscular lower lip. Separation 

between rings distinct on all sides; anterior ring ~ 2 /3 of total 

peristomial length. Peristomial cirri to posterior ! /3 of first 

peristomial ring, without articulations, tapering, less than half 

as long as A-I. 


and 1+1 according to Trcadwell (1921:56). 

Branchiae (Figure 28d) present, palmate, distinctly longer 


Branchiae from setiger 18 to end of fragments. Branchiae 

present to near posterior end, present on more than 65% of total 

number of setigers. First 3 branchiae single filaments; most 

branchiae with paired filaments; in a posterior fragment up to 3 

filaments present. Branchial stems very short and truncate. 

Filaments long, strap-like. 

All neuropodial acicular lobes asymmetrically truncately 

rounded; aciculae emerging above midline. All presetal lobes 

low folds. All postsetal lobes free, rounded to truncate 

projecting lobes, visible behind acicular lobes. First 20 ventral 

cirri thick, tapering. Median ventral cirri moderately inflated 

basally; narrow tips digitiform. Posterior ventral cirri tapering, 

digitiform. Notopodial cirri of first setiger short, digitiform, 

without articulations. All other notopodial cirri basally inflated, 


Limbate setae longer than all other setae, coarse, gently 

curved, marginally serrated. Shafts of pectinate setae (Figure 

28f) cylindrical, slender; blades flared, flat. Both marginal teeth 

slightly longer than other teeth; total of -15 teeth present. 

Shafts of compound falcigers (Figure 28e) distinctly inflated, 

internally striated, marginally nearly smooth, with distinct 

distal beak. Appendages long, tapering; head large, bidentate. 

Proximal teeth triangular, directed laterally; distal teeth slightly 

shorter than proximal teeth, sharply tapering, directed laterally. 

Guards symmetrically rounded, marginally smooth, without 

mucros. Pseudocompound falcigers and compound spinigers 

absent. Aciculae (Figure 28b) single, with dark cores and clear 

sheaths, tapering towards blunt, slightly curved tips. Subacicular 

hooks (Figure 28c) with dark cores and clear sheaths, 

bidentate (nearly bifid). Hooks first present from setiger 34, 


replacements). Hooks tapering abruptly just below head. 

Proximal teeth narrower than distal teeth; both teeth directed 

distally. 

UKNOWN MORPHOLOGICAL FEATURES.—Presence and position 

of eyes; relationship between Mx III and left Mx IV; 



TURES.—Mx III short, forming part of distal arc with left Mx 

IV. 



Characters: 1,2, 13, 14, 36,40, 42. 


None. 

REMARKS.—Eunice bucciensis was considered a synonym 

of E. afra by Hartman (1956). In E. bucciensis branchiae arc 

palmate with very long filamcnLs; in E. afra branchiae arc 

distinctly pectinate. The antennae lack articulations in E. 

bucciensis and have distinct, cylindrical articulations in E. afra. 

Eunice bucciensis is listed with similar species in Tables 33 and 

39. Among the species in Table 39, E. bucciensis and E. 

denticulata stand out for having elevated, long ccratophorcs. 

Eunice bucciensis has A-III slimmer than the other antennae 

and branchiae first present from sctiger 18; in E. denticulata, all 

antennae arc similar and branchiae are not present until sctigcr 

27. 

Eunice burmeisteri Grubc, 1878 

Eunice (Marphysa) Burmeisteri Grubc. 1878a: 101. 

REMARKS.—Grubc listed this species as a possible synonym 

of Marphysa sanguinea and indicated that it was named by 

Fritz Miillcr in a letter to him. As far as can be ascertained, no 

description of the species was ever issued, nor does any 

material exist. Due to the reference by Grube (1878a) of the 

possible synonymy with Marphysa sanguinea, the species 

must be considered indeterminable rather than a nomen nudum; 

the issue will be considered further in a planned review of the 

genus Marphysa. 

36. Eunice caeca Shisko, 1981 

FIGURE 29a-f; TABLES 19, 20 

Eunice caeca Shisko, 1981:968-971, fig. la-f, table 1. 

MATERIAL EXAMINED.—Holotype, USNM 63065, Bureau 

of Land Management sta 24774BFI, off Tanner Bank, 

California, 32°49.8'N, 119°29.86'W, 1357 m, trawl. 

DESCRIPTION.—Holotype complete with 90 setigers; last 7 

in regeneration; total length 103 mm; maximal width 6 mm at 

setiger 10. Length through setiger 10, 14 mm. Body ventrally 

flattened, dorsally moderately inflated throughout, slightly 

wider in branchial region than in postbranchial region, abruptly 

tapering anteriorly, slowly tapering posteriorly; regenerating 

end abruptly narrower than remainder of body. Anal cirri 

slender, digitiform, without articulations. 



Prostomial lobes frontally rounded, dorsally flattened; median

NUMBER 523 97 

FIGURE 29.—Eunice caeca (holotype, USNM 63065): a, anterior end, lateral view; b, parapodium 9, lateral view; 

c, compound falciger, parapodium 9; d, pectinate seta, parapodium 9; e, pectinate seta, parapodium 75; f, 

subacicular hook, parapodium 75. Eunice cariboea (ZMB 4004, syntype): g, anterior end, lateral view; h, 

pectinate seta, parapodium 16; i.acicula, parapodium 16; j, compound falciger,parapodium 16; k, parapodium 16, 

anterior view; 1, parapodium 75, anterior view; m, subacicularhook, parapodium 75; n, pectinate seta, parapodium 

75. Eunice cariboea (holotype of Nic'uiion incerta): o, subacicularhook, parapodium 127; p, compound falciger, 

parapodium 127; q, anterior end, lateral view. (Scale bars in mm.)


sulcus deep. Diffuse eye spots present near bases of A-I. 

Antennae in a horseshoe; median 3 in a group, well separated 

from A-I, similar in thickness. Ceratophores ring-shaped in all 


tapering; maximum 10 cylindrical articulations in left A-I I. A-I 

to setiger 1; A-II to setiger 3; A-III incomplete. Pcristomium 

inflated cylindrical with distinct muscular lower lip. Separation 

between rings distinct on all sides, especially deeply marked 


length. Peristomial cirri to posterior x h of prostomium, slender 

and tapering, with 7 cylindrical articulations. 

Maxillary formula 1+1,6+8,9+0,7+8, and 1+1. Mx III long 

straight, located behind left Mx II. Mx VI absent 





of total number of setigers. First branchiae single filaments; all 

other branchiae very strongly pectinate; maximum 24 filaments 

by setiger 15. Branchial stem slim, tapering, erect, longer than 

filaments; in last branchiated setigers branchial stems shortening 

rapidly. Filaments about as long as notopodial cirri where 

branchiae best developed, slender, nearly filiform; branchial 

filaments remaining long and slender throughout the branchial 

region. 

Anterior neuropodial acicular lobes asymmetrically rounded 

with aciculae emerging above the midline. Median and 

posterior neuropodial acicular lobes symmetrically triangular 

with superior aciculae emerging from apex. All presetal lobes 

low transverse folds. Anterior postsetal lobes free, evenly 

rounded, as long as acicular lobe, becoming reduced by 

midbranchial setigers to low folds. Ventral cirri tapering; thick 

in first 3 setigers. Ventral cirri in branchial region basally 

inflated. Inflated bases ovate; narrow tips tapering. Postbranchial 

ventral cirri gradually becoming short, tapering. Anterior 

notopodial cirri long, slender, with 4 to 5 irregular, indistinct 

cylindrical articulations. Articulations lost in first branchiated 

setigers. Posterior notopodial cirri short; slender. 

Limbate setae barely longer than other setae in most setigers, 

slender, marginally smooth. Pectinate setae (Figure 29d,e) 

nearly as long as limbate setae, very narrow; shafts relatively 

wide, flattened; blades tapering, flat. One marginal tooth longer 

and thicker than other teeth; up to 8 teeth present; anterior and 

posterior pectinate setae similar. Shafts of compound falcigers 

(Figure 29c) distally inflated, marginally smooth, with distinct 

distal beaks. Appendages narrow, with parallel sides; head 

indistinct, bidentate. Proximal teeth low triangular protuberance, 

directed laterally; distal teeth longer, blunt, erect. Guards 

symmetrically bluntly pointed, marginally serrated, without 

mucros. Pseudocompound falcigers and compound spinigers 

absent. Aciculae paired in anterior setigers; up to 4 in a 

parapodium in posterior setigers, yellow, tapering to blunt, 

nearly straight tips; cross-sections round. In posterior setigers 2 

aciculae emerge distally; 2 aciculae emerge along ventral edge 

of parapodia. Separation between core and sheath indistinct in 

both aciculae and subacicular hooks. Subacicular hooks (Figure 

290 yellow, bidentate. Hooks first present from setiger 42, 


replacements). Shafts of hooks thick, tapering distally to very 

small heads. Proximal teeth very much larger than distal teeth, 

curved, directed laterally; distal teeth small, tapering, erect. 

UKNOWN MORPHOLOGICAL FEATURES.—None. 

EXPECTED STATE OF UNKNOWN 


MORPHOLOGICAL FEA- 



Characters: 23. 


None. 

REMARKS.—Eunice caeca is listed with similar species in 

Tables 19 and 20. An unusual feature of this species is the 

emergence of two of the four aciculae along the ventral edge of 

the ncuropodia rather than distally. Eunice caeca is the only 

species listed in Table 20 with more than 20 branchial filaments 

in each branchia where the branchia are best developed. 

Eunice capensis Schmarda, 1861 

Eunice capensis Schmarda, 1861:126, 8 figs. 

Marphysa capensis.—Willey. 1904:263 — Day. 1967:400. 

REMARKS.—The original description, while vague, fits 

species of Marphysa. 

Eunice capensis (Kinberg, 1865) 

Eriphyle capensis Kinberg, 1865:561; 1910:41, pi. 15: fig. 16. 

Eunice kinberg i; Ehlers, 1868:306. 

REMARKS.—As pointed out by Ehlers (1868:306), the 

species name proposed by Kinberg is preoccupied in the 

combination Eunice capensis Schmarda (1861), because the 

generic name originally used by Kinberg is a junior synonym of 

Eunice; the species is further discussed as E. kinbergi. 

37. Eunice cariboea Grube, 1856 

FIGURE 29g-q; TABLES 3,33,40 

Eunice cariboea Grube, 1856:57. 

Eunice gagzoi Augener, 1922b:45 [in pan, see also E. gagzoi below]. 

Nicidion incerla Hansen, 1882:8, pi. 2: figs. 19-21. 

Eunice (Nicidion) incerla.—Hartman, 1959:313. 

MATERIAL EXAMINED.—Lectotype and 3 paralectotypes, 

ZMC, and 5 paralectotypes, ZMB 4004, Christiansted, St. 

Croix, Virgin Islands, coll. A.S. Orstcd and H. Kroyer. 

Holotype of Nicidion incerta, Leiden no. 1510, Rio de 

Janeiro, coll. E. v. Beneden. 

COMMENTS ON MATERIAL EXAMINED.—Six of the types of 

E. cariboea are complete. The lectotype consists of 157 setigers

NUMBER 523 99 

TABLE 3.—Variable and invariable features of the type lot of Eunice cariboea (N = number of individuals 

examined; SD = standard deviation; measurements in mm). 


N 

Max. 

Min. 

Mean 

SD 


Total length 

Maximum width 


Subacicular hooks first present from setiger no. 

4 157 

4 30 

4 1.5 

4 3 

4 32 

113 

18 

1 

2 

26 

125.75 

22.25 

1.13 

2.38 

28.5 

21.00 

5.32 

0.25 

0.48 

2.52 


A-I reach 

A-II reach 

A III reaches 

Anlennal articulations 

Peristomial cirri reach 

Peristomial cirral articulations 

Branchiae 


Acicular lip 


Number of subacicular teeth 

Tcdh in pectinate setae 

Compound shafts 

Teeth of compound setae 

N=4 

middle of peristomium 

front of peristomium 

middle of prostomium 

absent 

middle of peristomium 

absent 

absent 

black 

blunt or pointed 

black 

2 

12 

tapering 

2 

and is 30 mm long and 1 mm wide, and the length through 

setiger 10 is 2 mm. The larger of the two complete 

paralcctotypes in ZMB is here illustrated and described in 

detail. The holotype of Nicidion incerta is presently in two 

parts; it is, however, complete and in good condition. In order 

to account for minor differences between the two described 

taxa, the descriptions are presented separately. 

DESCRIPTION.—Paralectotype of E. cariboea complete with 

127 setigers; total length 24 mm; maximal width 1.2 mm wide 

at setiger 10. Length through setiger 10, 2.5 mm. Body 

anteriorly inflated, otherwise cylindrical throughout. 

Prostomium (Figure 29g) distinctly shorter and narrower 

than peristomium, as deep as x li of the peristomium. Prostomial 

lobes frontally obliquely truncate; lobes triangular, dorsally 

flattened; median sulcus shallow. Eyes small, indistinct, 

posterior to bases of A-I. Antennae in a deep horseshoe; A-I 

and A-II close together, separated from A-II I, similar in 


articulations. Ceratostyles digitiform, stout, nearly sausageshaped, 

without articulations. A-I to middle of first peristomial 

ring; A-II and A-III to middle of second peristomial ring. 

Peristomium slightly flaring anteriorly; lower lip distinct, 

muscular. Separation between rings distinct dorsally and 

ventrally, indistinct only for a short distance laterally; anterior 

ring 3 A of total peristomial length. Peristomial cirri to posterior 

end of first peristomial ring, nearly ovate, without articulations. 

Jaws examined in a ZMC paralectotype. Maxillary formula 

1+1, 5+5, 6+0, 4+9, and 1+1. Mx III long; located behind Mx 

II; Mx VI absent. 


Anterior neuropodial lobes (Figure 29k) symmetrically 

rounded, becoming symmetrically sharply pointed in posterior 

setigers (Figure 291); aciculae emerging at midline. Presetal 

lobes low, transverse folds; postsetal lobes following outline of 

acicular lobes closely. Four or 5 first ventral cirri thick, 

tapering. Ventral cirri basally inflated from about setiger 5. 

Inflated bases transversely elongated, welt-like; narrow tips 

digitiform. Inflated base decreasing in size from about setiger 

50. Last 40 ventral cirri without inflated bases, short, nearly 

tubercular. All notopodial cirri tapering, decreasing in length 

from anterior to posterior end, without articulations. Anterior 

notopodial cirri emerging close to parapodial bases; posterior 

notopodial cirri emerging from dorsum well above parapodial 

bases. 

Limbate setae short, stout, geniculate, marginally serrated. 

Two to 3 short pectinate setae (Figure 29h) in fascicle 

anteriorly; shafts coarse, cylindrical; blades tapering; flat. One 

marginal tooth longer than other teeth; -10 teeth present in a 

seta. In far posterior setigers up to 10 long pectinate setae 

(Figure 29n) in a fascicle; shafts slender, cylindrical; blades 

flaring, flat. One marginal tooth longer than other teeth; up to 

15 teeth in a seta. Shafts of compound falcigers (Figure 29j) 

distally inflated, marginally serrated; beak indistinct. Appendages 

very small, delicate; head small, bidentate. Proximal teeth 

curved, tapering, directed obliquely distally; distal teeth longer 

than proximal teeth, blunt, erect. Guards symmetrically bluntly 

pointed, marginally serrated, without mucros. Pseudocompound 


single; cross-sections round. Anterior aciculae tapering to sharp 

tips, light brown to dark yellow. Aciculae mucronate (Figure 

29i), bluntly pointed, dark brown or black from about setiger 

30. Subacicular hooks (Figure 29m) dark brown or black,

100 

bidentate. Hooks first present from setiger 24-31, present in all 


Hooks subdistally curved; head large; both teeth strongly 

curved, directed laterally. Proximal teeth about twice as large as 

distal teeth. Guards truncate. 

DESCRIPTION OF Nicidion incerta.—Holotype complete, of 

unknown sex, with 158 seugers; total length 38 mm; maximal 

width 1.5 mm at setiger 20; length through setiger 10,2.75 mm; 

width at setiger 10, 1.25 mm. Body ventrally flattened 

anteriorly, becoming cylindrical medially and retaining that 

shape for the rest of body. Truncate anteriorly, slowly tapering 

posteriorly. Pygidium is dorsally smooth; only the short anal 

cirri are present. 

Prostomial lobes (Figure 29q) frontally rounded, dorsally 

slightly inflated; median sulcus shallow. Eyes not observed 

(illustrated as being behind and between A-I and A-II in 

Hansen, 1882). Antennae in shallow horseshoe, evenly spaced 

and similar in thickness. Ceratophorcs short, ring-shaped. 

Ceratostyles thick, digitiform, without articulations. A-I to 

middle of anterior peristomial ring; A-II to middle of posterior 

peristomial ring; A-III to middle of posterior peristomial ring, 

slightly outreaching A-II. Nuchal fold with lateral notch; lower 

lip entire. Peristomium tapering anteriorly. Separation between 

rings distinctly dorsally and ventrally; anterior ring ~ 3 A» of total 

peristomial length. Peristomial cirri barely outreaching posterior 

peristomial ring, thick, ovate, without articulations. 



Neuropodial acicular lobes symmetrically rounded. Pre- and 

postsetal lobes low, transverse folds. Ventral cirri tapering 

through first 9 setigers, distinctly inflated from about setiger 

10. Inflated bases large, spherical; distal tips digitiform. 

Inflated bases decreasing in size from about setiger 60 and 

totally missing posterior to setiger 100. Posterior ventral cirri 

short, thick, tapering; distinctly shorter than notopodial cirri. 

Notopodial cirri well separated from neuropodial acicular 

lobes, thick, without articulations. Anterior neuropodial cirri 

digitiform. Median and posterior neuropodial cirri tapering. 

Limbate setae medially thick, marginally serrated, slightly 

longer than other setae. Pectinate setae not seen. Shafts of 

posterior compound falcigers (Figure 29p) distally inflated, 

without distinct beaks, marginally serrated, without internal 

striations. Appendages short, tapering, with distinct heads. 

Proximal teeth about as large as distal teeth, narrowly tapering, 

directed slightly to distinctly distally. Distal teeth tapering, 

directed obliquely distally, slender. Guards of posterior 

compound hooks symmetrically rounded, without mucros or 

marginal serrations. Posterior aciculae single, about as thick as 

subacicular hooks, brown, without distinct separation between 

sheath and core, tapering to blunt tips, straight. Subacicular 

hooks (Figure 29o) single, brown, without separation into 

sheath and core, bidentate. Hooks abruptly tapering to distinct, 

bent necks. Proximal teeth longer than distal teeth, erect, 

straight or slightly tilted distally, erect. Distal teeth short, thick, 

blunt, erect. 



cirri. 




SCORED.—Inappropriate Characters: 22, 31-42, 56, 58, 59. 

Unknown Characters: 74, 78. 


None. 

REMARKS.—According to G. Hartwich (in litt.), a label in 

the vial indicated that three of these specimens should be the 

types of E. gagzoi Augcner (1922b). This label reads (in 

translation from German): "Eunice Gagzoi n.sp. (cariboea Gr. 

partim). 2 complete specimens without, one complete specimen 

with a few scattered branchiae." The original description of E. 

gagzoi is very brief and contains no discussion of the 

relationship between E. cariboea and E. gagzoi. 

One clue to the identity of the specimens that he considered 

types of his new species is available. Augcner indicated that 

one of the types should consist of 161 segments and be 32 mm 

long. No such specimen is present in the Berlin material. Two 

posteriorly branchiaicd specimens in the Hamburg collections 

arc labeled as original material of E. gagzoi. and one of these 

matches the size and length mentioned by Augcner closely. 

These two specimens were selected as lectotypes of E. gagzoi 

and are described below. No specimens from ZMB belong to £. 

gagzoi; they are E. cariboea as here defined or go to E. 

excariboea (see below). 

Specimens of E. cariboea may be confused with small 

specimens oiE.filamentosa from which they clearly differ in 

the shape of the subacicular hooks, by the total absence of 

branchiae, and by the tapering, rather than hammer-headed 

acicula. Eunice cariboea can be separated from E. gagzoi on a 

similar set of characters, except that in the latter the aciculae are 

abruptly tapering with a short, bent tip, rather than hammerheaded 

as in E. filamentosa or smoothly tapering and gently 

curved as in E. cariboea. As indicated, E. cariboea is here 

restricted to include only abranchiate forms; the two other 

species confounded with E. cariboea in the type lot are both 

branchiated, but are otherwise dissimilar. One of these species, 

E. gagzoi, was discussed above; the other one is herein newly 

recognized as E. excariboea (see below). 

Eunice cariboea is listed with similar species in Tables 33 

and 40. Five abranchiate species lack antcnnal articulations, in 

addition to E. cariboea; these include E. cincta, E. goodei, E. 

imogena, and E. wasinensis. Of these, E. imogena has 

subacicular hooks first present from about setiger 50; the other 

four species have hooks present from about setiger 15 through 

setiger 31. Eunice cincta lacks inflated bases to the median 

cirri; the other species listed all have inflated bases. These bases 

are thick, transverse welts in E. cariboea and E. wasinensis and 

ovate in the other two species. Eunice cariboea and E. 

wasinensis can be separated on various differences in soft-body

NUMBER 523 101 

characters, best explored by comparing the illustrations of the 

two species. 

Hansen's complete description ofNicidion incerta consisted 

of the following statement (1882:8): 

The animal resembles N. gallopagensis Kinberg closely; the dorsal cirri are 

however slighily longer and the setae appear different from those illustrated by 

Kinberg. 

[Original in French, translation by K. Fauchald.] 

Hansen's illustration of the anterior end shows the prostomium 

as shallowly incised; the antennae are short; none 

reaching beyond the peristomium and without articulations. 

His illustration of the parapodium shows the ventral cirrus to be 

basally spherically inflated with a digitiform tip. 

Augcncr reviewed the type at one time and left a note in the 

vial: "Presumably the abranchiate form of Eunice cariboea Gr. 

(Kr.)." 

Hansen's species is here considered a synonym of E. 

cariboea. 

38. Eunice cedroensis Fauchald, 1970 

FIGURE 30a-e; TABLES 41, 42 

Eunice cedroensis Fauchald, 1970:29-31, pi. 2: figs. a-e. 

MATERIAL EXAMINED.—Holotype, AHF Poly 0336, Baja 

California, Mexico, 4 miles N of Isla Todos Santos, 

31°53'2O"N, 116°48'15"W,75 m, shell, mud, gray sand, 24 Feb 

1941, Velero III, sta 1245-41. 


length 10 mm; maximum width 2 mm at setiger 10. Length 

through setiger 10, 3 mm. Body cylindrical, becoming slightly 

ventrally flattened posteriorly. Anterior end abruptly tapered; 

body tapering slowly posteriorly from about setiger 10. 

Prostomium (Figure 30a) about as long and wide as 

peristomium, as deep as l /2 of the peristomium. Prostomial 

lobes frontally truncate, dorsally slightly inflated; median 

sulcus shallow. Eyes black, posterior to bases of A-I. Palpal 

region marked by 2 shallow circular grooves frontally on each 

prostomial lobe. Antennae in a horseshoe, evenly spaced, 



maximum 5 long, cylindrical articulations in A-III. A-I to 


tapering towards anterior end, longer ventrally than dorsally. 

Separation between rings distinct on all sides; anterior ring 2 h 

of total peristomial length. Peristomial cirri to middle of 

prostomium, slender and tapering, without articulations. 

Maxillary formula 1+1, 7+8, 9+0, 8+10, and 1+1. Mx VI 

absent; Mx III long, located behind left Mx II. 



Branchiae from setiger 3 through setiger 27. First 2 and last 3 

pairs single filaments; all others with 3 or more filaments; 

maximum 10 filaments by setiger 10. Branchial stems tapering. 

Filaments longer than notopodial cirri except in first 2 and last 

3 branchiated setigers; filaments digitiform, relatively slender. 

Anterior neuropodial acicular lobes obliquely truncate, with 

aciculae emerging at midline but ventral to high point; acicular 

lobes becoming symmetrically rounded in first branchial 

setigers, retaining that shape in rest of body. Anterior and 

median pre- and postsetal lobes low, transverse folds. Ventral 

cirri tapering in first 5 setigers. Ventral cirri basally inflated in 

branchial region, retaining modestly inflated, ovate bases in 

remainder of specimen; narrow tips tapering. Anterior and 

median notopodial cirri tapering, with 3 or 4 articulations. 

Limbate setae longer than other setae, slender, marginally 

smooth. Pectinate setae (Figure 30d) small; shafts coarse, 

cylindrical; blades tapering, flat. One marginal tooth longer and 

thicker than other teeth; a total of 10 teeth in a seta. Shafts of 

compound falcigers (Figure 30c) distally inflated, marginally 

serrated, with distinct distal beak. Appendages long, narrow, 

tapering; head very small, but distinct, bidentate. Proximal 

teeth triangular, directed laterally, about twice as large as distal 

teeth. Distal teeth tapering, erect, slender. Guards symmetrically 

tapering, with long, narrow mucros. Pseudocompound 

falcigers and compound spinigers absent. Aciculae single, 

yellow, tapering to blunt, straight tips; cross-section round. 

Separation of core and sheath indistinct in both aciculae and 

subacicular hooks. Subacicular hooks (Figure 30e) yellow, 

tridentate with teeth in a crest. Hooks first present from setiger 

17, present in all setigers thereafter, always single (except for 

replacements). Hooks slightly tapering. All teeth curved; teeth 

decreasing evenly in size distally. 


with the posterior setigers. 





1,2,37,38,47,50,57-60,63. 


38,2. 

REMARKS.—Eunice cedroensis is listed with similar species 

in Tables 41 and 42. Of the species in Table 42, five have the 

three median antennae similar in length. In addition to E. 

cedroensis, these species are E. americana, E. multicylindri, E. 

rubrivittata, and E. unifrons. Of these five, two species have 10 

or more branchial filaments, E. cedroensis and E. americana. 

The latter has up to 20 filaments; the former 10 filaments. 

Eunice challenged Mclntosh, 1885 

Eunice challenged Mclntosh, 1885:293. 

REMARKS.—This species is known only from the following 

quote in Mclntosh's discussion of E. barvicensis (1885:293): 

"The hook somewhat approaches that of Eunice challengeri, 

but the other differences between the species are sufficiendy


FIGURE 30.—Eunice cedroensis (hololype, AHF Poly 0336): a, anterior end, lateral view; b, parapodium 20, 

anterior view; c, compound falciger, parapodium 20; d, pectinate seta, parapodium 20; e, subacicular hook, 

parapodium, 20. Eunice cincta (holotype, RM 418): f, pectinate seta, parapodium 30; g, compound falciger, 

parapodium 40; h, subacicular hook, parapodium 50. (Scale bars in mm.)


diagnostic." 

As indicated elsewhere, E. barvicensis is similar to E. 

filamentosa, so presumably the name E. challenged was used 

by Mclntosh as a manuscript name for another, similar species 

from the Challenger expedition, which species cannot now be 

identified and no material is available under this name in 

BM(NH) (A. Muir, in litt.). 

39. Eunice cincta (Kinberg, 1865) 

FIGURE 30f-h; TABLES 33,40 

Nicidion cincta Kinberg, 1865:564; 1910:43, pi. 16: fig. 21. 

Eunice (Nicidion) cincta.—1 [artman, 1948:80-81. 

MATERIAL EXAMINED.—Holotype, RM 418, Eimeo, Society 

Islands, 0.3-1 m, Eugenie Expedition 1159. Kinberg 

(1865:564) gave the locality information as "Mare pacificum 

inter corallia mortua tacniarum exteriorum insulac Eimeo, 

summa aqua vel fundo 1-4 pedum." 


has been deeply dissected and the shape of the prostomium 

cannot be determined accurately. The last 15 setigers are 

extremely poorly preserved. 

DESCRIPTION.—Holotype incomplete with 65 setigers (last 

15 extremely poorly preserved); length through setiger 50 15 

mm; maximal width 1.1 mm; length through setiger 10, 3 mm. 


Ceratostyles thick and digitiform, wrinkled, but without 

articulations. All antennae to posterior peristomial ring. 

Separation between peristomial rings distinct on all sides; 

anterior ring ~ l /i of total peristomial length. Peristomial cirri to 

posterior part of anterior peristomial ring, ovate, without 


Jaws missing. 


Neuropodial acicular lobes rounded; aciculae emerging at 

midline. Presetal lobes low, transverse folds. Anterior postsetal 

lobes following outline of acicular lobes closely, becoming 

low, transverse folds from about setiger 25. Anterior and 

median ventral cirri thick, tapering. Anterior notopodial cirri 

medially inflated, becoming digitiform in last distinct segments 

in fragment, without articulations. 


pectinate setae (Figure 300 very slender. Pectinate setae 

flaring, flaL Both marginal teeth slightly longer than other 

teeth; -15 teeth present. Shafts of compound falcigers (Figure 

30g) distinctly inflated, marginally coarsely serrated. Appendages 

slender with nearly parallel sides, bidentate. Proximal 

teeth shorter than distal teeth, sharply pointed, directed basally. 

Distal teeth slender, directed laterally. Guards asymmetrically 

bluntly pointed, marginally serrated, without mucros. Pseudocompound 


brown, bluntly conical, straight; cross-section round. 

Subacicular hooks (Figure 30h) brown, bidentate. Hooks first 

present from setiger 21, present in all setigers thereafter, single 

at least through median setigers. Hooks with distinctly bent 

necks. Proximal teeth slightly larger than distal teeth, directed 

laterally. Distal teeth curved, directed laterally. 


with the posterior setigers and with the poor state of 

preservation. 




SCORED.—Inappropriate Characters: 22, 31-42, 54, 55. 

Unknown Characters: 1, 2, 4, 6-17, 24, 47, 50, 51, 57-60, 

63, 74, 78. 


None. 

REMARKS.—Eunice cincta is listed with similar species in 

Tables 33 and 40. The shape of the subacicular hook resembles 

the condition in E. mutilata; however, the latter is always 

branchiated, even as a juvenile. Among the abranchiate species, 

E. cincta is unusual in that it wholly lacks inflated bases in 

median ventral cirri. 

Eunice cingulata Claparede, 1868 

Eunice cingulata Claparede, 1868:444-445, pi. 7: fig. 1. 

Eunice purpurea.—Grube, 1878a:100. 

REMARKS.—No type material exists of this or any other 

species of Eunice described by Claparede. The following 

summary is based on his description. 

Prostomium deeply bilobed. Antennae short, with moniliform 

articulations. Anterior peristomial ring very nearly as long 

as posterior ring. Peristomial cirri to front edge of peristomium, 

with moniliform articulations. 

Branchiae present from setiger 8. First few single filaments, 

all others strongly pectinate; maximum number of filaments 

unknown. Presumably, branchiae continue to end of body. 

Limbate setae marginally serrated. Pectinate setae with 

marginal tooth very much longer and thicker than all other 

teeth. Shafts of compound hooks inflated. Appendages short, 

tapering. Proximal teeth triangular. Distal teeth curved. Guards 

distally asymmetrically bluntly pointed. Aciculae and subacicular 

hooks with dark cores and clear sheaths; aciculae 

tapering distally to blunt, gently curved tips. Subacicular hooks 

tapering to distinct heads. Proximal teeth directed laterally. 

Eunice cingulata was well characterized by Claparede, who, 

in addition to the description summarized above, emphasized 

the dark violet color and the violet-and-white banding of the 

anterior end and appendages. The species clearly is a member 

of group B-2, but, based on the available information, cannot be 

separated from a series of other species. It is here considered 

indeterminable.


Eunice cirribranchis Grube, 1870 

Eunice cirribranchis Grube, 1870b:55; 1878a:99. 

REMARKS.—Grube characterized this species briefly as 

having strongly articulated, moniliform antennae; branchiae 

present from setiger 6 through 72 (of 123 segments present) 

and with only a single branchial filament. Peristomial cirri and 

notopodial cirri lacked articulations. Branchiae and notopodial 

cirri were similar in length. 

Grube (1878a:99) listed the species among those that had 

branchiae limited to some anteromedian setigers, and distinctly 

articulated antennae, in his subgenus Leodice; most other 

members in this grouping are now considered members of 

group C-l. 

Eunice cirribranchis is, however, so poorly characterized 

that without access to the types and additional material, it must 

be considered indeterminable. It was originally described from 

Fiji. 

40. Eunice cirrobranchiata Mclntosh, 1885 

FIGURE 31a-h; TABLES 33,36,52, 53 

Eunice cirrobranchiata Mclntosh, 1885:277-278, pi. 38: figs. 9-11, pi. 19A: 

figs. 21-24. 

MATERIAL EXAMINED.—Two syntypes, BM(NH) ZK. 

1885.12.1.196a. Bermuda, between tidemarks, Jun 1873. 

COMMENTS.—Both syntypes are incomplete; the larger one 

has been frontally dissected and most setae are broken. 

DESCRIPTION.—Large syntype with 123 setigers; length 30 

mm; maximal width 3 mm; length through setiger 10, 6 mm. 

Small syntype with 45 setigers; length 18 mm; maximal width 

3 mm; length through setiger 10, 5.5 mm. All segments very 

short and crowded; parapodia supported on high triangular 

lateral ridges in posterior ends of both fragments. 

Prostomium (Figure 31a, illustration of small syntype) 

distinctly shorter and narrower than peristomium, as deep as x li 

of the peristomium. Prostomial lobes frontally rounded, 

dorsally excavate with a thickened rim; median sulcus shallow. 

Eyes posterior to bases of A-I, black. Antennae in horseshoe, 


in all antennae, without articulations. Ceratostyles thick and 

digitiform, with up to 3 irregular, cylindrical articulations each, 

in addition to wrinkles. Antennae to middle or posterior end of 

peristomium; A-I shortest; A-III longest. Peristomium cylindrical 

with distinct muscular lower lip. Separation between rings 

distinct dorsally; anterior ring 9 /io of total peristomial length; a 

shallow crease dividing ventral side with anterior part 5 /6 of 

total length of ventral side. Dorsal and ventral grooves not 

matching. Peristomial cirri to middle of peristomium, without 


Maxillary formula 1+1, 5+5, 6+0, 1+6, and 1+1. Mx VI 

absent. Mx III short, part of distal arc with Mx IV and V. 


cirri, not reduced in mid-body region. Branchiae from setigcr 

22 in small and from setiger 30 in large syntypc, continued to 

ends of fragments. Maximally 2 long, narrow branchial 

filaments present; most segments with single filaments. 

Anterior ncuropodial acicular lobes truncate, from about 

setigcr 25, acicular lobes reduced, represented by single, large 

aciculae emerging directly from body wall (Figure 310; 

aciculae emerging at midline. Body wall forming high 

transverse ridges on both sides of the body. Pre- and postsetal 

lobes follow outline of acicular lobes closely. First 4 ventral 

cirri thick and digitiform. Bases of ventral cirri increasingly 

elongated from setiger 5, with short digitiform tips in all 

setigers. From about setiger 20 ventral cirri elongated, 

glandular ridges making up lower edge of fleshy ridges 

supporting setae. All noiopodial cirri medially inflated, without 

articulations, longer in posterior than in anterior sciigcrs. 

Shafts of pectinate setae (Figure 3Id) slender, cylindrical. 

Blades flared, furled, forming an open scoop. Both marginal 

teeth slightly longer than other teeth, with -15 teeth. Shafts of 

compound falcigcrs (Figure 31c,h) inflated, marginally 

smooth; distal beaks and internal striations absent. Appendages 

short; heads relatively large, bidcntaic. Proximal teeth larger 

than distal teeth, tapering, directed laterally. Distal teeth short, 

tapering, directed obliquely distally or laterally. Guards 

symmetrically rounded, marginally frayed in most hooks, 


spinigers absent. Aciculae single (Figure 31b,g), very heavy, 

with light to dark brown cores and clear sheaths, with distinct 

narrow neck, asymmetrically hammer-headed distally; distal 

ends of acicular shafts curved in some parapodia; crosssections 

round. Subacicular hooks (Figure 31e) with dark 

brown cores and clear sheaths, bidentate or tridentate with teeth 

in tandem. Hooks first present from setiger 17 or 24, present in 


Hooks strongly curved, strongly beaked in appearance. Main 

fangs very large and curved, with paired lateral denticles in 

some hooks; distal fangs smaller but similar in shape. 


with far posterior setigers; pygidium and anal cirri. 


None. 



Characters: 1, 2,4, 6, 36-40,63. 


38,1. 

REMARKS.—Eunice cirrobranchiata is usually considered a 

synonym of E.fdamentosa (see Hartman, 1959) and little in the 

description of the two species differs. Mclntosh stated that he 

had only a single 30 mm anterior fragment at his disposal and 

that specimen had been dried out. The larger of the two 

specimens is certainly the one described by Mclntosh; all 

information given matches the specimen closely. It does not 

appear to have been seriously dried out. The provenance of the


FIGURE 31.—Eunice cirrobranchiata (holotype, BM(NH) ZK 1885 12.1.196a): a, anterior end, lateral view; b, 

acicula, parapodium 30; c, compound falciger, parapodium 30; d, pectinate seta, parapodium 30; e, subacicular 

hook, parapodium 30; f, parapodium 30, anterior view; g, acicula, parapodium 40; h, compound falciger, 

parapodium 40. Eunice coccinea (syntype, ZMB 866): i, pectinate seta, parapodium 10; j, compound falciger, 

parapodium 10; k, acicula, parapodium 10; 1, pectinate seta, parapodium 55; m, subacicular hook, parapodium 55; 

n, parapodium 10, anterior view; o, anterior end, lateral view; p, parapodium 55, anterior view. (Scale bars in 

mm.)


smaller syntype is unclear; it matches the other specimen 

closely and is here considered a second syntype. 

Eunice cirrobranchiata is listed with similar species in 

Tables 33 and 36. Eunice cirrobranchiata and E. conglomerans 

have modified acicular tips; however, E. cirrobranchiata is the 

only species listed in Table 36 to have subacicular hooks with 

the teeth in tandem. 

Eunice rosaurae also has the teeth in tandem in the 

subacicular hooks. The species with this characteristic (compared 

in Tables 52 and 53) can be separated as suggested in the 

discussion of E. rosaurae. 

Eunice claparedii Quatrefages, 1866 

Eunice claparedii Quatrefages, 1866:652-653. 

Eunice harassii.—Claparede, 1864:578-580, pi. 2: fig. 5 [not Eunice harassii 

Audouin and Milne Edwards, 1833]. 

Eunice torquala.—Gmbe, 1878a:99. 

REMARKS.—Quatrefages based his species on specimens 

from Port-Vendres, Mediterranean Sea, identified by Claparede 

as E. harassii. As far as I can see, Quatrefages based his 

description solely on Claparede's written report and never saw 

any specimens. Quatrefages differential diagnosis emphasized 

that the antennae in Claparede's form were placed in a 

semicircle and were short and had relatively few articulations. 

The notopodial cirri should be articulated. 

Claparede's description and illustration indicates that his 

specimens belong to group B-2. The prostomial antennae are 

short and are divided into seven articulations, reaching the 

posterior end of the peristomium. The peristomial cirri do not 

outreach the peristomium and are divided into five articulations. 

The notopodial cirri are also articulated with about five 

articulations. Branchiae are present from setiger 3 according to 

the illustration (Claparede, 1864, pi. 2: fig. 5) and may have 

seven filaments where best developed. Aciculae and subacicular 

hooks have black cores and clear sheaths; the subacicular 

hooks are illustrated as being very strongly curved, with a large, 

laterally directed proximal tooth and a small, erect distal tooth. 

The aciculae are tapering and apparently gently curved. Details 

of the other setae cannot be determined, nor can the distribution 

of the subacicular hook be determined. 

Claparede left no material identified by him, so the validity 

of this species depends solely on the description furnished. 

Grube (1878a:99) synonymized Quatrefages species with E. 

torquata, also described by Quatrefages. This synonymy 

appears unproblematic and is here accepted. 

41. Eunice coccinea Grube, 1878 

FIGURE 31i-p; TABLES 24,25,27,30 

Eunice coccinea Grube, 1878b:153-155, pi. 9: fig. 1. 

MATERIAL EXAMINED.—TWO syntypes, ZMB 866, Singapore, 

coll. v. Martens. 

COMMENTSON MATERIAL EXAMINED.—Crube (1878b:153) 

gave the Philippines (Bohol) as type locality; the specimens 

described here are marked types in the collections of the Berlin 

Museum. The large syntype is very soft and flabby; the small 

syntype is in excellent condition and forms the base for the 

re-description. 

DESCRIPTION.—Large syntype incomplete, of unknown sex, 

with 241 setigers; length 242 mm long. Small syntype 

incomplete; with 80 setigers; length 55 mm; maximal width 5 

mm; length through setiger 10, 10 mm. 

Prostomium (Figure 31o) distinctly shorter than peristomium, 

nearly as wide as peristomium, as deep as x li of 


inflated; median sulcus deep. Eyes not observed. Antennae in 

transverse row, evenly spaced, similar in thickness. Ccratophores 

ring-shaped in all antennae, without articulations. Ccratostyles 

digitiform, with up to 5 long, indistinctly marked, 

cylindrical articulations in A-III. A-I to middle of peristomium; 

A-II to setiger 1; A-I 11 to setiger 2. Peristomium flaring 

anteriorly; lower lip expanded into thick bolster-like structure, 

wider than rest of peristomium. Separation between rings 

distinctly dorsally and vcnirally; anterior ring 5 A of total 

peristomial length. Peristomial cirri to middle of anterior 

peristomial ring, digitiform, with 2 articulations. 

Maxillary formula of large syntype 1+1, 4+4, 7+0, 5+8, and 

1+1. 

Branchiae (Figure 3In) present, pectinate, distinctly longer 


Branchiae from setiger 6 to end of small syntype, to setiger 138 

in large syntype, leaving about half of incomplete specimen 

abranchiate. Branchiae terminating well before posterior end. 

First branchia with 3 filaments; maximum 10 filaments at about 

setiger 15. Number of filaments reduced from about setiger 30; 

from about setiger 45 rest of branchial region with 1 or 2 

filaments present. Stems longer than notopodial cirri, tapering, 

erect Filaments longer than notopodial cirri, digitiform, 

relatively slender. 

Anterior neuropodial acicular lobes truncate with aciculae 

emerging at superior corner; median and posterior acicular 

lobes (Figure 31p) short, rounded. Pre- and postsetal lobes low, 

transverse folds; postsetal lobes unusually low in relation to 

acicular lobes. First 4-5 ventral cirri thick, tapering. Median 

ventral cirri basally inflated from early branchial setigers; 

narrow tips absent. Inflated bases nearly spherical in setigers 

15-25, decreasing in size slowly posteriorly, developing a 

broad transverse shape and eventually a dorsal depression. Far 

posterior ventral cirri scoop-shaped, curving around emerging 

subacicular hooks, relatively narrow, nearly digitiform in 

anterior and posterior outline; narrow tips tapering. Anterior 

notopodial cirri tapering, with 2 or 3 indistinct, cylindrical 

articulations. Notopodial cirri loosing articulations, becoming 

shorter and more distinctly tapering from branchial region, 

retaining that shape to last segments. 

Limbate setae slender. Anterior pectinate setae (Figure 31i)


very short. Shafts slender and cylindrical. Blades flat, flaring. 

One marginal tooth longer and thicker than other teeth; with 

-15 teeth. In median and posterior setigers number and size of 

pectinate setae increasing, by setiger 50 dominant kind of setae 

in dorsal fascicles. Shafts of median and posterior pectinate 

setae (Figure 311) slender, cylindrical. Blades flaring, flat. One 

marginal tooth longer, but not thicker than other teeth; about 25 

teeth present. Shafts of compound falcigers (Figure 31j) 

tapering, marginally smooth, without internal striations or 

distal beaks. Appendages wide-based, triangular, bidentate. 

Proximal teeth a slight elevation on side of appendage. Distal 

teeth distinctly larger than proximal teeth, nearly erect, thick. 

Guards fit appendages closely, visible only on cutting edge, 


spinigers absent. Aciculae paired, amber to dark brown; 

anterior superior aciculae (Figure 31k) tapering distally to fine 

slender tips; other aciculae taper distally to blunt, truncate tip; 

cross-section round. Superior aciculae gently curved dorsally; 

inferior aciculae straight. Fine amber-colored notopodial 

aciculae present. Subacicular hooks (Figure 31m) ambercolored 

to dark brown, bidentate. Hooks first present from 

setiger 38-39, present in all setigers thereafter, always single 

(except for replacements). Hooks slender, tapering to distinct 

heads. Proximal teeth shorter, but wider than distal teeth, blunt, 

erect. Distal teeth tapering, erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—Relationship between 

Mx III and left Mx IV; distribution of branchiae on 

posterior end; pygidium and anal cirri. 


Mx III short, forming a distal arc with left Mx IV. Branchiae 

absent on posterior half of body. 



Characters: 1, 2,4, 6, 13, 14, 36, 38, 40, 74, 78. 


REMARKS.—Eunice coccinea is listed with similar species in 

Tables 24 and 25. It is also listed in Tables 27 and 30, because 

the branchiae continue beyond setiger 100 (one of the defining 

characters of group B-2 in Fauchald, 1970). 

Among the species in Table 25, E. rosaurae and E. coccinea 

have basally inflated ventral cirri continued to the posterior 

end. Eunice rosaurae has both bi- and tridentate subacicular 

hooks of which the latter have the teeth in tandem; E. coccinea 

has only bidentate subacicular hooks. In addition, E. rosaurae 

has only single branchial filaments; E. coccinea has pectinate 

branchiae with up to 10 filaments. 

Of species listed in Table 30, E. coccinea, E.fimbriata, and 

E. reducta have the branchiae terminating well before the 

posterior end. Of these three species, only E. coccinea has 

basally inflated ventral cirri in the posterior end; in both the 

other species, the inflated region is missing in the posterior end. 

42. Eunice coccinioides Augener, 1922 

FIGURE 32; TABLJES 24,26 

Eunice coccinioides Augener, 1922b:45. 

Eunice fucata.—Augener, 1918:316-319, fig. 31 [not Eunice fucata Ehlers, 

1887]. 

MATERIAL EXAMINED.—TWO syntypes, ZMH V-5693, Isla 

Annobon, Angola, 7 Oct 1911, coll. Arn. Schultze 

COMMENTS ON MATERIAL EXAMINED.—Augener (1918) 

gives a detailed description of the specimens from Isla 

Annobon; the description fits rather well with the specimens 

listed as types and Isla Annobon is here considered the type 

locality. 

The sample contains two anterior ends and a series of median 

and posterior ends; of these one median piece and three 

posterior ends fit the anterior ends. One posterior end belongs 

to a species in group B-2; it is strongly branchiated even in the 

last setigers present and has dark brown or black subacicular 

hooks. Five median and posterior fragments lack subacicular 

hooks entirely and may belong to a species of Palola. The 

longer of the syntypes is described in detail. 

DESCRIPTION.—Longer syntype incomplete, of unknown 

sex, with 133 setigers; length 80 mm long; maximal width 5 

mm at setiger 20; length through setiger 10, 8 mm. Short 

syntype incomplete, of unknown sex, with 56 setigers; total 

length 32 mm; maximal width 4.5 mm at setiger 15; length 





inflated; median sulcus deep. Eyes on ceratophores of A-I, 

black. Antennae in shallow horseshoe, evenly spaced, similar 

in thickness. Ceratophores of A-I short, truncate cones; A-I I 

and A-III with ring-shaped ceratophores; all ceratophores 

without articulations. Ceratostyles tapering, without articulations. 

A-I to middle of anterior peristomial ring; A-II to setiger 

2; A-III to setiger 3. Peristomium massive, slightly flaring 

anteriorly. Separation between rings distinct dorsally and 

ventrally; anterior ring 3 /4 of total peristomial length. Peristomial 

cirri to middle of anterior peristomial ring, medially 

inflated, without articulations. 

Maxillary formula of large syntype 1+1, 6+6, 10+0, 6+12, 

and 1+1; maxillary formula of small syntype 1+1, 5+6, 10+0, 

5+10, and 1+1. Mx III part of distal arc with left Mx IV. Mx VI 

absent. 

Branchiae (Figure 32g) present, pectinate, distinctly longer 


Branchiae from setigers 7 or 8, continued to setiger 78 on large 

syntype, leaving roughly half of body abranchiate. Maximum 9 

filaments. Stems much longer than notopodial cirri, tapering, 

slender, erect. Filaments longer than notopodial cirri, digitiform, 

slender. 

Anterior neuropodial acicular lobes rounded; posterior 

acicular lobes truncate; aciculae emerging dorsal to midline.

108 


FIGURE 32.—Eunice coccinoides (holotype, ZMH V-5693): a, anterior end, lateral view; b, compound falciger, 

parapodium 129; c, pectinate seta, parapodium 129; d, subacicular hook, parapodium 129; e, pectinate seta, 

parapodium 19; f, compound falciger, parapodium 19; g, parapodium 19, anterior view. (Scale bars in mm.)


Presetal lobes low, transverse folds. Anterior postsetal lobes as 

high as acicular lobes, projecting as free lobes on ventral sides 

of acicular lobes, becoming reduced to low, transverse folds 

from about setiger 25. First 4 ventral cirri thick, tapering. 

Ventral cirri basally inflated from setiger 5; inflated portion 

forming thick transverse ridges; narrow tips tapering. Inflated 

bases becoming reduced from about setiger 55; ventral cirri 

from about setiger 70 slender, tapering, about as thick as 

notopodial cirri. Notopodial cirri supported by internal aciculae 

in all setigers. Anterior notopodial cirri basally inflated, short, 

tapering, becoming slightly longer and distinctly narrower in 

the last 50 setigers present. All notopodial cirri without 

articulations, relatively short. 

Limbate setae marginally smooth. Shafts of pectinate setae 

slender, cylindrical. Blades of anterior pectinate setae (Figure 

32c) slightly flaring, becoming more distinctly flaring in 

posterior setigers (Figure 32c). One marginal tooth slightly 

longer than other teeth; -12 teeth present. Shafts of compound 

falcigers (Figure 32b, 0 slightly inflated, marginally smooth, 

without internal striations or distal beaks. Appendages short, 

tapering, with large heads, bidentate. Proximal teeth larger than 

distal teeth, narrow-based triangular, directed laterally, but 

tilted distally in anterior setigers (Figure 32f), directed distally 

(Figure 32b) in posterior setigers. Distal teeth tapering, 

distinctly bent in anterior setigers and gently curved in 

posterior setigers. Guards symmetrically bluntly pointed in and 

marginally serrated anterior setigers, rounded and marginally 

smooth in posterior setigers, without mucros. Pseudocompound 


single, light copper-colored in anterior setigers, becoming 

distinctly darker brown in posterior setigers, distinctly projecting, 

thick, tapering, straight, cross-section round. Subacicular 

hooks (Figure 32d) copper- to chestnut-colored, bidentate. 

Hooks first present from setiger 28 or 30, present in all setigers 


tapering to narrow, bent necks and distinct head. Proximal teeth 

larger than distal teeth, directed laterally, but tilted distally. 

Distal teeth shorter than proximal teeth, blunt, directed 

obliquely laterally. 




Branchiae missing on posterior half of body. 



Characters: 1,2, 38-40,42, 74, 78. 


REMARKS.—Eunice coccinioides is listed with similar 

species in Tables 24 and 26. It is characterized by the 

copper-chestnut colored aciculae and subacicular hooks, which 

sets it apart from other species in the group. The subacicular 

hooks have a distinctly bent neck and the proximal teeth of the 

subacicular hooks are distally tilted up, resembling the 

condition in some of the abranchiate species in group B-4. 

43. Eunice collaris Grube, 1869 


Eunice collaris Grube, 1869:494-495 [in part]. 

MATERIAL EXAMINED.—Three syntypes, ZMB F500, F501, 

and F2035, Tor, Red Sea, among corals, coll. Ehrenberg. 

COMMENTS ON MATERIAL EXAMINED.—None of the syntypes 

are complete, two are very soft and generally in bad 

shape, but are sufficiently complete, and so distributional 

features can be identified as can the shapes of the setae. The 

syntype from F500 is in very good condition and is here 

described in detail. 

DESCRIPTION.—Syntype from F500 incomplete with 56 

setigers; length 44 mm; maximal width 6 mm at setiger 10; 

length through setiger 10, 12 mm. Anterior end cylindrical, 

frontally truncate; body becoming dorsally flattened towards 

posterior end of fragment. 



lobes frontally rounded, dorsally flattened; median sulcus deep. 

Eyes present posterior to bases of A-I. Antennae evenly spaced, 


without articulations. Ceratostyles digitiform; maximum 8 

articulations in A-I 11; articulations cylindrical basally, becoming 

drop-shaped, nearly moniliform distally. A-I to setiger 1; 

A-I I to setiger 2 and A-I 11 to setiger 4. Peristomium cylindrical; 

lower lip slightly inflated, distinctly muscular. Separation 

between rings well marked dorsally and ventrally and faintly 

visible laterally; anterior ring - 3 A of total peristomial length. 

Peristomial cirri short and digitiform; articulations absent. 


Branchiae (Figure 330 present, pectinate, distinctly longer 


Branchiae from setiger 17 (18 in other specimens) to end of 

incomplete specimens. All branchiae pectinate with up to 12 

filaments at about setiger 50. Branchial stem weak, flexible. 

Branchial filaments long, slender. 

Neuropodial acicular lobes symmetrically rounded; aciculae 

emerging at midline. All presetal lobes low folds. Anterior 

postsetal lobes distinct ridges behind acicular lobe; median and 

posterior postsetal lobes low, transverse folds. First 3-4 ventral 

cirri thick and tapering. Median ventral cirri slightly inflated 

basally. Inflated bases ovate; narrow tips tapering. Inflated 

bases totally lost by setiger 50; posterior ventral cirri thick and 

tapering. All notopodial cirri distinctly inflated basally, and 

tapering to long tips; basal inflated region especially distinct in 

setigers 5-25, without articulations. 

Limbate setae slender, marginally serrated. Pectinate setae 

(Figure 33e), numbering up to 12 in a parapodium, tapering, 

furled into shallow scoops; 1 marginal tooth distinctly longer 

than other teeth; -15 teeth present. Shafts of compound 

falcigers (Figure 33d) slightly inflated, very finely marginally 

serrated. Appendages long, tapering; head distinct, bidentate.

110 


FIGURE 33.—Eunice collaris (holotype, ZMB F500): a, anterior end, lateral view; b, subacicular hook, 

parapodium 49; c, subacicular hook, parapodium 51; d, compound falciger, parapodium 51; e, pectinate seta, 

parapodium 51; f, parapodium 51, anterior view. (Scale bars in mm.) 

Proximal teeth narrow, directed basally, distal teeth distally 

blunt, curved. Guards symmetrically bluntly pointed, marginally 

serrated, without mucros. Pseudocompound falcigers and 

compound spinigers absent. Aciculae single, dark brown, 

bluntly pointed and straight; cross-section round. Subacicular 

hooks (Figure 33cb,c) brown, bidentate. Hooks first present 

from setiger 27 or 33, present in all setigers thereafter, always 

single (except for replacements). Hooks abruptly tapered, with 

distinct head at right angles to long axis. Proximal teeth very 

large, blunt; distal teeth short and rounded. 




TURES.—Mx III short, forming part of distal arc with left Mx II. 



Characters: 1,2,15, 37, 38, 40,47, 50, 63, 74, 78. 


38,1. 

REMARKS.—Two additional syntypes differ at the species 

level from those listed above and are here referred to E. afra. 

Eunice collaris has been considered a synonym of E. afra since 

Crossland (1904:289) first referred it to that species. As 

indicated above, the type material contains two species, both of


which are similar to E. afra in that they have dark subacicular 

hooks and branchiae starting at a relatively late segment. 

Eunice collaris is listed with similar species in Tables 33 and 

35. It is one of two species in Table 35 with more than six 

branchial filaments: Eunice collaris has 12 filaments and E. 

fauveli has 14 filaments. The latter has mucronate guards in the 

compound falcigers; E. collaris has blunt guards. 

44. Eunice collini Augener, 1906 

FIGURE 34a-k; TABLES 33, 37-39 

Eunice collini Augener, 1906:133-135, pl.4: figs. 66-73. 

Eunice rosaurae Monro, 1939:351-352, fig.28a-f [in part]. 

MATERIAL EXAMINED.—Holotype, MCZ 2011, Blake sta 

288, off Barbados, 730 m. One syntype of E. rosaurae, 

BM(NH) ZK 1941.1.1.217-221, off St. George, Grenada, 

12°05'N, 61°49'W, 720-800 m, 27 Nov 1937, trawled, 

Rosaura Expedition, Atlantic, 1937-1938, sta 34. 

DESCRIPTION.—Holotype of E. collini incomplete with 94 

setigers; length 45 mm; maximal width 4 mm at about setiger 

15; length through sctigcr 10, 7 mm. Syntype of E. rosaurae 

complete with 76 setigers; total length 42 mm; maximal width 

5 mm at about setiger 15; length through setiger 10, 9 mm. 

Body anteriorly inflated; posterior end wide and dorsoventraily 

flattened; anterior and posterior ends abruptly tapered. 

Prostomium (Figure 34a,e) distinctly shorter and narrower 


lobes frontally rounded, dorsally inflated; median sulcus very 

deep. Eyes absent. Antennae in a horseshoe, evenly spaced, 


without articulations. Ceratostyles tapering, without articulations, 

smooth or irregularly wrinkled. A-I to second peristomial 

ring; A-I I to setiger 2; A-I 11 incomplete in both specimens. 

Peristomium flaring anteriorly. Separation between rings 

distinct dorsally and ventrally; anterior ring ~ 3 A of total 

peristomial length. Peristomial cirri to front margin of 

peristomium, slender and digitiform, without articulations. 


Branchiae (Figure 34k) present, palmate, distinctly shorter 


Branchiae from setiger 16 to setiger 85 in holotype; from 

setiger 12 to setiger 70 in syntype of E. rosaurae. Branchiae 

terminating well before posterior end. Most branchiae short, 

thick, single filaments; scattered along branchiated region, 

especially posteriorly, short, nearly tubercular second filaments 

present. 

All neuropodial acicular lobes truncate or gently rounded; 

aciculae emerging at midline. All pre- and postsetal lobes low, 

transverse folds. Anterior ventral cirri thick and tapering. 

Ventral cirri basally inflated from setiger 5. Inflated bases 

ovate, narrow tips tapering. Inflated bases reduced from about 

setiger 50; free tips elongating and becoming digitiform 

(Figure 34d). In far posterior setigers ventral cirri about twice 

as long as notopodial cirri but otherwise similar (Figure 34k). 

Anterior notopodial cirri basally inflated, distally slender; 

posterior notopodial cirri similar in shape but somewhat longer, 


Limbate setae slender, margins smooth. Shafts of pectinate 

setae slender (Figure 34f); blades distinctly flaring, fiat. One 

marginal tooth slightly longer than other teeth; about 10 teeth 

present. Shafts of compound falcigers (Figure 34c,i) tapering 

from wide distal end; margins smooth. Appendages thick, 

tapering; heads distinct, bidentate. Proximal teeth about twice 

as large as distal teeth, directed laterally; distal teeth nearly 

erect or gently curved, tapering. Guards symmetrically 

rounded; cutting edge frayed rather than serrated; mucros 

absent. Pseudocompound falcigers and compound spinigers 

absent. Aciculae single in anterior parapodia, paired in all 

median and posterior parapodia. Superior aciculae (Figure 34h) 

with knife-shaped cross-section, light to dark brown, distally 

pointed. Inferior aciculae (Figure 34g) with round crosssection, 

distally tapering to straight tips, dark brown to 

jet-black. Both aciculae projecting (Figure 34d,k), for nearly 

half their length in median and posterior setigers, nearly half 

width of body represented by projecting aciculae in far 

posterior setigers. Posterior notopodial cirri supported by 

internal black aciculae; notopodial aciculae absent in anterior 

setigers. Subacicular hooks (Figure b, j) black, bidentate. Hooks 

first present from setiger 26-27, present in all setigers 


tapering, with distinct heads; proximal teeth much larger than 

distal teeth, directed laterally; distal teeth directed distally. 

Thin, pergamentaceous tube associated with syntype of E. 

rosaurae. 




TURES.—Mx III short, forming distal arc with left Mx IV. 



Unknown Characters: 1, 2, 6, 23, 38-40,42, 74, 78. 


REMARKS.—Eunice collini resembles E. floridana, but can 

be separated on the structure of the aciculae and the late start of 

the branchiae. Branchiae are present from setigers 7-10 in E. 

floridana and from setiger 12 or not until setiger 16 in E. collini. 

The differences between the syntype of E. rosaurae and the 

holotype of E. collini are minor; some may be due to the state 

of preservation, such as the projecting aciculae in the former, 

and other differences are well within the normal variability 

encountered in the genus. 

Eunice collini is listed with similar species in Tables 33 and 

in Tables 37-39. It is one of the few species in Table 33 in 

which the branchiae terminate well before the posterior end.

112 


.02 

1 

.02 

.5 

• 

2 

b c 

1 

K 

I 

FIGURE 34.—Eunice collini (hololype, MCZ 2011): a, anterior end, lateral view; b, subacicular hook, parapodium 

91; c, compound falciger, parapodium 91; d, parapodium 91, anterior view. Eunice collini (1 syntype of Eunice 

rosaurae BM(NH) ZK 1941.1.1.217-221): e, anterior end, lateral view; f, pectinate seta, parapodium 66; g, 

inferior acicula, parapodium 66; h, superior acicula, parapodium 66; i, compound falciger, parapodium 66; j, 

subacicularhook, parapodium 66; k, parapodium 66. Eunice congesta (holotype, ZMW, Inv. no. 1042): 1, anterior 

end, lateral view; m, pectinate seta, parapodium 40; n, compound falciger, parapodium 40; o, subacicular hook, 

parapodium 40; p, acicula, parapodium 40, q, parapodium 40, anterior view. (Scale bars in mm.)


45. Eunice complanata Grube, 1877 

TABLES 33,37-39 

Eunice complanata Grube, 1877:529-530.—Fauchald, 1986:246-247. 

MATERIAL EXAMINED.—Holotype, ZMB, type 880, Timor, 

Atapupa, coral reef and Salawatti, coral reef. 

COMMENTS ON MATERIAL EXAMINED.—The type is now 

completely dry and clearly has been dry for some time. 

DESCRIPTION.—Holotype with about 100 setigers; reported 

length, 72 mm, probably correct. 

Prostomium completely shrunken. Black eyes present 

between bases of A-I and A-II. All antennae apparently 

outrcaching tip of prostomium, without obvious articulations. 

Separation between peristomial rings distinct dorsally and 

ventrally; anterior ring more than 5 /s of total peristomial length. 

Peristomial cirri at least as long as antennae, reaching well 

beyond frontal margin of peristomium, without obvious 



forming distal arc with short, curved Mx IV. Mx VI missing. 


cirri, not reduced in mid-body region. Branchiae from 

setiger 19, well developed on last seugers present (setiger 100). 

First 2 pairs possibly with single filaments only; all other 

branchiae pectinate; maximum 6 filaments. Branchial stems 

strong, erect. 

Ventral cirri apparently basally inflated in some anteromedial 

setigers; narrow tips tapering. 

Limbate and pectinate setae present. Appendages of compound 

hooks bidentate. Guards without rnucros. Aciculae and 

subacicular hooks dark brown. Aciculae tapering to blunt tips; 

cross-section of aciculae round. Subacicular hooks bidentate; 

first present from setiger 28, rather thick in relation to aciculae. 

UNKNOWN MORPHOLOGICAL FEATURES.—Most prostomial 

and peristomial features, jaw structure; most parapodial 

structures. 


TURES.—Species too poorly known to make predictions. 



Characters: 3, 12, 15-20, 23, 24, 28, 34, 36-38, 40, 42-52, 

54, 57-68, 74, 78. 


38,1. 

REMARKS.—Unless the branchial distribution was highly 

unusual, the branchiae must have continued for at least another 

20 setigers beyond the end of the incomplete specimen. 

The parapodia are well developed and distinct, but are so 

distorted that they cannot be described in any additional detail. 

Due to the distribution of branchiae, color of the aciculae and 

subacicular hooks, and the dentition of the latter, E. complanata 

is here considered related to E. afra and similar species and 

is listed with these in Table 33 and Tables 37-39. It is too 

poorly known to be included in the key. 

Eunice concinna (Verrill, 1900) 

Leodice concinna Verrill, 1900:643.—Hartman, 1942:8. 

REMARKS.—The type is not present in the collections of the 

Peabody Museum and has not been present since before 1940 

(Hartman, 1942; W. Hartman, in litL). The only information of 

any value in the original description is the mention of the 

strongly beaded antennae and the presence of branchiae only on 

the anterior part of the body starting at setiger 3. 

Verrill stated that the species resembled E. stigmatura, 

which in turn has been considered synonymous with E. vittata 

(but see below). The branchial distribution is consistent with a 

membership in group C-l, but the strongly beaded antennae 

mentioned for E. concinna would differentiate the species from 

E. vittata and E. stigmatura. 

The information is clearly insufficient to identify E. 

concinna and, whereas the types are lost, cannot be improved. 

The species is here considered indeterminable. 

46. Eunice congesta Marenzeller, 1879 

FIGURE 341-q; TABLES 41,45 

Eunice congesta Marenzeller, 1879:134-135, pi. 4: fig. 2a-d. 

MATERIAL EXAMINED.—Holotype, ZMW Inv. no. 1042, 

East coast of Eno-shima, Japan, coll. C. Koerbl. 

COMMENTS ON MATERIAL EXAMINED.—The type lot consists 

of an anterior end of 50 setigers, here considered the 

holotype, and one anterior fragment missing the head and 

perhaps the first 10 setigers and two midposterior fragments. 

Only the anterior end is here described in detail; the other 

fragments present agree with this anterior end, but neither of 

the two mid-posterior fragments can unequivocally be associated 

with the anterior end. 

DESCRIPTION.—Holotype with 50 setigers; length 22 mm; 

maximal width 4 mm at setiger 10; length through setiger 10,6 

mm. Body cylindrical, truncate anteriorly. 

Prostomium (Figure 341) distinctly shorter and narrower than 

peristomium, as deep as l /2 of the peristomium. Prostomial 

lobes frontally rounded, dorsally inflated; median sulcus 

shallow. Dark purple eyes present between bases of A-I and 

A-II. Antennae in a horseshoe. Ceratophores ring-shaped in all 


digitiform, without articulations. A-I missing; A-II to setiger 1; 

A-III to setiger 2. Peristomium cylindrical. Separation between 

rings distinct on all sides; anterior ring - 2 /3 of total peristomial 

length. Peristomial cirri missing, but scar visible. 

Maxillary formula 1+1, 6+8,6+0, 6+7, and 1+1 according to 

Marenzeller (1879:135, pi. 4: fig. 2C). Mx III long, located 

behind left Mx II. Mx VI missing. 

Branchiae (Figure 34q) present, pectinate, distinctly longer 


Branchiae from setiger 7 to setiger 47. Only first branchia with 

single filaments; all other branchiae pectinate with up to 18 

filaments. Branchial stems strong, erect. Filaments tapering to


slender tips, at least as long as notopodial cirri in all except first 

few branchiated setigers. 

Anterior neuropodial acicular lobes rounded, from about 

setiger 10 becoming increasingly obliquely conical with 

aciculae emerging dorsal to midline. All presetal lobes low, 

transverse folds. Anterior postsetal lobes about as high as 

acicular lobes following outline of acicular lobes closely; 

median and posterior postsetal lobes low folds. First 8 or 9 

ventral cirri thick and tapering. Median ventral cirri basally 

inflated. Inflated bases spherical; narrow tips short, nearly 

button-shaped. Inflated bases reduced posterior to setiger 35, 

absent in mid-posterior fragments; ventral cirri tapering in 

posterior setigers. All notopodial cirri slender and tapering, 


Limbate setae slender, nearly capillary with very narrow, 

smooth limbations. Pectinate setae (Figure 34m) very small, 

flat, flaring. One marginal tooth larger than other teeth; total of 

10 teeth present. Shafts of compound falcigcrs (Figure 34n) 

distally very slightly inflated and marginally serrated. Appendages 

slender, with a very large head, bidentate. Proximal teeth 

triangular, directed laterally; distal teeth sharply bent, truncate. 

Guards symmetrically rounded, with sharply pointed mucros, 

serrated along the cutting edge. Pseudocompound falcigers and 

compound spinigers absent. Aciculae (Figure 34p) paired, 

slender, yellow, tapering distally to slender, slightly curved or 

bent tips; cross-section round. Separation between core and 


Subacicular hooks (Figure 34o) yellow, tridentate with teeth in 



Hooks slender, with large, triangular main fangs and smaller 

teeth decreasing relatively evenly in size distally. 


with posterior setigers; pygidium and anal cirri. 





Characters: 1,2,16, 17, 23, 27-29, 37, 38,42. 


38,1. 

REMARKS.—Eunice congesta was considered a synonym of 

E. vittata by Imajima and Hartman (1964:258); it is listed with 

similar species in Tables 41 and 45. In addition to E. congesta, 

E. mucronata has more than 15 branchial filaments. The two 

species are very similar and cannot be safely separated on the 

characters used in the key. 

47. Eunice conglomerans Ehlers, 1887 

FIGURE 35a-i; TABLES 33,36 

Eunice conglomerans Ehlers, 1887:93-95, pi. 23: figs. 1-9, pi. 24: figs. 1-4. 

MATERIAL EXAMINED.—One syntype, MCZ 706, Blake sta 

10, 24°44X 83°26'W, 68 m. Two syntypes, ZMB 6730, Key 

West, 2-3 m, coll. A. Agassiz. 

DESCRIPTION.—MCZ syntype complete with 166 setigers; 

total length 163 mm; maximal width 3.5 mm at setiger 10; 

length through setiger 10, 6.5 mm. Ventrum flattened in 

anterior 2 h of body; dorsum strongly convex, especially in 

anterior 

x h of body except cylindrical prostomium and 

peristomium. ZMB syntypes both incomplete with 228 and 223 

setigers; a posterior fragment encased in thick, pergamentaceous 

tubes also present. 


about as wide as peristomium, as deep as V2 of the 

peristomium. Prostomial lobes frontally obliquely truncate, 

dorsally flattened; median sulcus deep. Eyes small, black, 

posterior to bases of A-I. Six antennae present; A-III apparently 

duplicated. Antennae arranged in a straight line, evenly spaced, 

similar in thickness. Ceratophorcs ring-shaped in all antennae, 

without articulations. Ccratostylcs digiliform, with 5 articulations 

in A-I I and A-I 11 and 4 in A-I; all articulations cylindrical. 

All antennae similar in length, barely reaching posterior margin 

of peristomium. Pcristomium cylindrical wiih scoop-shaped 

projecting lower lip. Separation between rings distinct dorsally; 

vcntrally pcristomium also divided into 2 rings, but connection 

between dorsal and ventral transverse grooves uncertain; 

anterior ring 4 /s of total pcristomial length. Pcristomial cirri 

short and tapering, without articulations. 

Maxillary formula of both ZMB specimens 1 + 1, 5+5, 6+0, 

2+6, and 1+1. Left Mx IV very small, curving around Mx III, 

appearing nearly fused to Mx III. 


cirri, not reduced in mid-body region. Branchiae from setiger 

29 (from setigers 23 and 27 in other specimens) to setiger 165. 

Branchiae present to near posterior end, present on more than 

65% of total number of setigers. Most branchiae with 2 

filaments; maximum 4 filaments from about setiger 130. 

Branchial stems short, flexible and conical. Filaments long, 

robust, strongly wrinkled with very large internal blood 

vessels. Anterior neuropodial acicular lobes asymmetrically 

rounded with aciculae emerging from distal end above midline; 

median and posterior acicular lobes (Figure 35b,c) becoming 

symmetrically conical posteriorly. All pre- and postsetal lobes 

low folds. First 4 ventral cirri thick and tapering. Ventral cirri 

becoming inflated with large, transverse welt-like glandular 

bases from about setiger 5; narrow tips tapering. From about 

setiger 100, basal inflations decreasing in importance. Far 

posterior ventral cirri thick, digitiform and held curved up 

along the posterior face of parapodia (Figure 35c); glandular 

contents of ventral cirri retained (apparently) even in these 

posterior segments. Anterior notopodial cirri thick, basally 

inflated and tapering, becoming distinctly digitiform from 

beginning of branchial region; far posterior notopodial cirri 

slender and digitiform; all notopodial cirri without articulations. 

Limbate setae long, slender, rare in posterior setigers. Most

NUMBER 523 

FIGURE 35.—Eunice conglomerans (syntype, MCZ 706): a, anterior end, lateral view; b, parapodium 35, anterior 

view; c, parapodium 134, posterior view; d, appendage, compound falciger, parapodium 10; e, subacicular hook, 

parapodium 35; f, compound falciger, parapodium 35; g, pectinate seta, parapodium 35; h, compound falciger, 

parapodium 10; i, acicula, parapodium 35. Eunice contingens (holotype, USNM 19394): j, parapodium 3, anterior 

view; k, parapodium 19, anterior view; 1, parapodium 40, anterior view; m, pectinate seta, parapodium 3; n, 

compound falciger, parapodium 3; o, pectinate seta, parapodium 19; p, pectinate seta, parapodium 40; q, 

subacicular hook, parapodium 40; r, compound falciger, parapodium 40. (Scale bars in mm.)


dorsal fascicles numerically dominated by very long, slender 

pectinate setae. Pectinate setae (Figure 35g) tapering, furled. 

Both marginal teeth longer than other teeth; -15 teeth present. 

Shafts of compound falcigers (Figure 35f,h) distinctly inflated, 

marginally strongly serrated. Appendages of superior hooks 

(Figure 35h) in anterior setigers, apparently nearly unidentate 

in that both teeth, but especially proximal teeth reduced and the 

distal teeth nearly erect. Inferior hooks (Figure 35d) in same 

fascicle distinctly bidentate with small teeth; appendages 

narrower and somewhat longer than in posterior setigers. In 

posterior setigers all appendages (Figure 35f) similar, each 

appendage with both teeth well developed; proximal teeth very 

large. Guards distally symmetrically rounded, marginally 

strongly serrated in all hooks; mucros absent. Pseudocompound 


(Figure 35i) single, black, distally bent dorsally, with slightly, 

but distinctly knobbed appearance, nearly bidentate in some 

setigers; cross-section round. First few subacicular hooks 

increasingly darker brown, from about setigcr 30 hooks black, 

bidentate. Hooks first present from setigcr 22, present in all 


Hooks (Figure 35e) with distinct curved heads with very large, 

parrot-beaked main fangs and much smaller, curved distal 

teeth. Guards distally rounded. 


anal cirri. 





Characters: 39, 40, 74, 78. 


None. 

REMARKS.—Eunice conglomerans is listed with similar 

species in Tables 33 and 36. Two species in Table 36 have 

knobbed aciculae; all other species have tapering, more or less 

pointed acicula. In addition to E. conglomerans, E. cirrobranchiata 

has such aciculae. In E. conglomerans the bases of the 

ventral cirri are inflated only in median setigers; in E. 

cirrobranchiata the bases are inflated also through the posterior 

setigers. 

48. Eunice contingens (Chamberlin, 1919) 

FIGURE 35j-r; TABLES 27,28 

Leodice contingens Chamberlin, 1919a:260-263, pi. 57: figs. 1-5. 

MATERIAL EXAMINED.—Holotype, USNM 19394, Albatross, 

sta 3642, Galapagos Islands, near Hood Island, Ripple 

Point, l°30'5"S, 89°35'W, 549 m, 7 Nov 1904, broken shells 

and Globigerina. 

COMMENTS ON MATERIAL EXAMINED.—The type is a 

complete mature female with large eggs in the body cavity. It 

has been dried out and no meaningful measurements can be 

made. It appears to agree with Chamberlin's description. 

DESCRIPTION.—Anterior end described, but not illustrated 

by Chamberlin. Ceratophorcs ring-shaped in all antennae. 

Ceratostyles without articulations, or vaguely wrinkled. A-I to 

sctiger 1; A-I I to setigcr 4; A-I 11 to sctiger 6. Peristomium 

cylindrical; separation between rings distinct at least vcntrally. 

Pcristomial cirri long and slender, reaching well beyond frontal 

margin of peristomium, without articulations. Maxillary formula 

according to Chamberlin (1919a:263; see also pi. 57: fig. 

1) 1+1,6+7,7+0,4+9, 1+1. 

Branchiae (Figure 35k-l) present, pectinate, shorter than 


Branchiae from sctigcr 6 to end of specimen. Branchiae present 

to near posterior end, present on more than 65% of total number 

of setigers. First branchia single filaments; all others, except 

possibly far posterior ones, pectinate; 18-19 filaments present 

at or near sctigcr 15. 

Ncuropodial acicular lobes (Figure 35j—1 > distally rounded 

or truncate; aciculae emerging at midlinc. All pre- and postsctal 

lobes low folds. Anterior ventral cirri tapering from wide, 

triangular bases. Median ventral cirri basally inflated; inflated 

bases transverse welts; narrow tips digiliform. Posterior ventral 

cirri digiliform, without basal inflations. Anterior notopodial 

cirri long and slender, basally somewhat inflated; median 

notopodial cirri somewhat shorter; the basal inflated region 

more distinct. Far posterior notopodial cirri slender and shorter 

than in anterior end. Notopodial cirri without articulations 

Anterior pectinate setae (Figure 35m) furled, tapering, with 

1 marginal tooth longer than other teeth; -10 teeth present. 

Median and posterior pectinate setae (Figure 35o,p) flaring, 

furled; -15 teeth present. Shafts of compound falcigers (Figure 

35n/) distinctly to slightly flaring; anterior shafts slightly 

serrated; beaks present. Appendages short with both teeth well 

developed and directed laterally, bidentate. Proximal teeth 

tapering, tilted distally. Guards asymmetrically bluntly pointed 

in anterior setigers, symmetrically bluntly pointed in posterior 

setigers; mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae paired, black, straight and 

tapering; inferior one somewhat heavier than superior one; 

cross-sections round. Subacicular hooks black, bidentate; first 

present from setiger 30. Hooks (Figure 35q) tapering evenly 

from the middle; head indistinct; both teeth directed distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Structure of 

prostomium; relationship between Mx III and left Mx IV; 




IV. 



Characters: 1-17, 19, 20, 23, 25, 26, 28, 36, 40, 51, 

54,65-68,74,78,81,82. 


None.


REMARKS.—Eunice contingens is listed with similar species 

in Tables 27 and 28; it has not been included in the key. 

Numbers of teeth for Mx III and left Mx IV are listed in the 

opposite order by Chamberlin, which would lead to a highly 

unusual distribution of teeth. It is here assumed that the more 

customary arrangement was present, because in other descriptions 

that could be checked by specimens, Chamberlin had 

confused the order of the jaws. 

Eunice contingens is the only species listed in Table 28 with 

branchiae shorter than the notopodial cirri. 

49. Eunice crassitentaculata (Treadwell, 1922) 

Leodice crassilentaculata 

1-5. 

FIGURE 36a-e; TABLES 33. 38 

Treadwell. 1922:146-148, figs. 30-33, pi. 4: figs. 

MATERIAL EXAMINED.—Holotype, AMNH 1526, Pago 

Pago, Samoa. 

COMMENTS ON MATERIAL EXAMINED.—The holotype is an 

anterior fragment and a median fragment of 88 segments, 

which may have belonged to the same specimen. The anterior 

end is described in detail. 

DESCRIPTION.—Anterior fragment with 158 setigers; 135 

mm long; maximum width, near posterior end of fragment, 3.5 

mm; length through setiger 10, 9.5 mm; width at setiger 10, 2 

mm. Anterior part of body cylindrical, becoming dorsoventrally 

flattened and expanded from about setiger 20. 


distinctly narrower than peristomium, less than 1 /2 as 

deep as peristomium. Prostomial lobes frontally rounded, 

dorsally inflated; median sulcus deep. Black eyes present 

between bases of A-I and A-II. Antennae in a horseshoe, evenly 

spaced, similar in thickness. Ceratophores long and cylindrical 


without articulations. A-I to middle of peristomium; A-II to 

setiger 4; A-I 11 to setiger 7. Peristomium about twice as wide as 

prostomium, barrel-shaped. Separation between rings distinct 

dorsally and ventrally; anterior ring slightly more than x li of 

total peristomial length. Peristomial cirri short and tapering, 


Jaws not seen. Maxillary formula according to Treadwell 

(1922:147) 1+1,4+5, 7+0, 5+8, and 1+1. 

Branchiae (Figure 36b) present, palmate, distinctly longer 

than notopodial cirri. Branchiae from setiger 28 as single 

filaments. Maximally 2 filaments present. Branchiae present to 

end of body according to Treadwell, present on more than 65% 

of total number of setigers. 

Neuropodial acicular lobes distally triangular or rounded 

with single prominent aciculae emerging at midline. All 

presetal lobes low folds. Anterior postsetal lobes project as free 

lobes; median and posterior postsetal lobes low, transverse 

folds. First 4 ventral cirri tapering or digitiform. Ventral cirri 

basally inflated from setiger 5; inflated bases ovate, distinct in 

all setigers present in fragment; narrow tips tapering. Notopodial 

cirri thick and basally inflated in anterior setigers, 

becoming medially inflated in branchial region. Notopodial 

cirri without articulations. 

Limbate setae slender with very narrow limbation. Pectinate 

setae (Figure 36c) flat, flaring. Both marginal teeth longer than 

other teeth; about 15 teeth present. Shafts of compound 

falcigers (Figure 36e) distally inflated, smooth. Appendages 

large; heads distinct, bidentate. Both teeth of same size, 

directed laterally. Guards symmetrically rounded; mucros 


absent. Aciculae and subacicular hooks with black cores and 

clear sheaths. Aciculae single, tapering to blunt tips; crosssection 

round. Subacicular hooks (Figure 36d) bidentate. 


thereafter, always single (except for replacements). Hooks with 

both teeth well developed; proximal teeth directed laterally. 





IV. 

CHARACTERS 

USED IN PREPARATION OF KEY NOT 

SCORED.—Inappropriate Characters: 22, 60. Unknown Characters: 

1,2,6,33,34,36,40,42,63. 


None. 

REMARKS.—Eunice crassitentaculata resembles E. afra as 

indicated by Hartman (1956); however, it has much longer 

antennae than that species and the shape of the subacicular 

hooks is also distinctly different. It is listed with similar species 

in Tables 33 and 38. The long, cylindrical ceratophores in all 

antennae are rare among species of the genus. 

Eunice culebra Treadwell, 1901 

Eunice culebra Treadwell, 1901:197, fig. 37.—Hartman, 1956:249. 

REMARKS.—The original description was based on a small, 

incomplete specimen with about 50 anterior setigers. 

The antennae were slender. A-I 11 was three times the length 

of the prostomium; A-II 3 A of the length of the prostomium and 

A-I only a little shorter than A-II. A-I 11 was thus described as 

being considerably longer than the other antennae. A single 

branchia is present from one of the last segments present. 

The type was deposited in the USNM according to Hartman 

(1956); this is not the case and there is no evidence to show that 

it was ever present. The USNM collections contains a specimen 

identified by Treadwell as E. culebra, but it was collected about 

15 years after this species was described. This specimen 

belongs to E. cariboea as defined in this paper and disagrees 

with the description of £. culebra in the shape and size of the 

antennae. 

Eunice culebra is here considered indeterminable.

118 


\ 

.02 

.02 

.1 

2 

.1 

1 

c - e 

f - h 

i 

. a 

FIGURE 36.—Eunice crassilentaculata (holotype, AMNH 1526): a, anterior end, lateral view; b, parapodium 69 

anterior view; c, pectinate seta, parapodium 69; d, subacicular hook, parapodium 69; e, compound falciger, 

parapodium 69. Eunice curlicirrus (hololype): f, compound falciger, parapodium 35; g, subacicular hook, 

parapodium 35; h, pectinate seta, parapodium 35; i, parapodium 35, anterior view; j, anterior end, lateral view. 

(Scale bars in mm.)


50. Eunice curticirrus Knox, 1960 

FIGURE 36f-j; TABLES 33,40 

Eunice (Nicidion) curticirrus Knox, 1960:125-126, figs. 190-195. 

Eunice curticirris [sic].—Fauchald, 1986:247, figs. 24-28. 

MATERIAL EXAMINED.—Holotype, Canterbury Museum, 

Chatham Islands, 43°36.2'S, 176°48.5'W, S of the Sisters, 69 

m, coarse shell, sand, gravel, Chatham Island Expedition sta24. 


length 12 mm; maximal width 1 mm at setiger 10; length 

through setiger 10,2.25 mm. Body cylindrical, tapering slowly 

posteriorly. 

Prostomium (Figure 36j) about as long as peristomium, 



flattened; median groove invisible dorsally; ventral longitudinal 

groove present. Large, reddish eyes between bases of A-I 

and A-I I. Antennae in a straight line; with A-I 11 isolated by a 

gap, similar in thickness. Ceratophores ring-shaped in all 

antennae, without articulations. Ceratostyles digitiform; each 

with 3 or 4 very large, angular articulations. All antennae of 

same length, reaching setiger 1. Peristomium about as long as 

prostomium, flaring, especially anteroventrally. Separation 

between rings distinct dorsally and visible ventrally; anterior 

ring roughly 3 A of total peristomial length. Peristomial cirri to 

middle of peristomium, digitiform, without articulations. 

Maxillary formula unknown. 


First 20 neuropodial acicular lobes symmetrically truncate, 

thereafter reduced so aciculae emerge directly from body-wall; 

aciculae emerging at midline (Figure 36i). All pre- and 

postsetal lobes low folds. Anterior ventral cirri thick, tapering. 

Ventral cirri basally inflated from about setiger 12. Bases 

ovate; narrow tips tapering. Posterior to setiger 25 ventral cirri 

again becoming tapering, nearly triangular in outline. Anterior 

notopodial cirri thick and digitiform, becoming slender and 

digitiform posterior to setiger 20. Notopodial cirri without 


Pectinate setae (Figure 36h) tapering, flat. Marginal teeth no 


falcigers (Figure 36f) slightly inflated, marginally smooth. 

Appendages bidentate. Proximal teeth large, triangular, directed 

laterally; distal teeth smaller than proximal teeth, 

angularly bent and laterally directed. Guards symmetrically 

truncate, without mucros. Pseudocompound falcigers and 

compound spinigers absent. Aciculae and subacicular hooks 

with black cores and clear sheaths; cross-section round. 

Aciculae paired, smoothly tapering and distally pointed. 

Subacicular hooks (Figure 36g) bidentate. Hooks first present 


(except for replacements). Hooks distinctly S-shaped with 

tapering necks and very large, curved distal and proximal teeth. 

UNKNOWN 

MORPHOLOGICAL FEATURES.—Jaw structure; 



TURES.—Mx III part of distal arc with left Mx IV. 



Unknown Characters: None. 


None. 

REMARKS.—Knox (1960) mentioned that tridentate hooks 

with the distal tooth divided laterally were present in this 

species. No such hooks were observed in the specimen, but 

several parapodia are missing and such hooks could have been 

present in at least a few parapodia. Eunice curticirrus can be 

separated from all other abranchiate species of the genus by the 

extremely strongly beaded antennae; it is listed with similar 

species in Tables 33 and 40. 

The species name was misspelled by Fauchald (1986:247). 

51. Eunice denticulata Webster, 1884 


Eunice denticulata Webster, 1884:316-317. pi. 10: figs. 41, 41a-b, 42-45. 

MATERIAL EXAMINED.—Two syntypes, plus 9 slide preparations 

of parapodia, USNM 4790, Bermuda, coll. G. Brown 

Goode, 1876 or 1877. 

DESCRIPTION.—Large syntype incomplete, with 125 setigers; 


through setiger 10, 7 mm. Small syntype incomplete, with 76 

setigers; length 25 mm; maximal width 3 mm at setiger 10; 

length through setiger 10, 6 mm. Anterior end inflated; rest of 

body slightly dorsoventrally flattened, but basically cylindrical. 

Prostomium (Figure 37d) distinctly shorter and narrower 

than peristomium, less than x /i as deep as peristomium. 


sulcus deep. Dark eyes posterior to bases of A-I. Ceratophores 

long in all antennae, without articulations. Ceratostyles 

tapering, without articulations; A-III slightly longer than other 

antennae; none reaching beyond peristomium. Peristomium 

massive, cylindrical, with distinct, muscular lower lip. Separation 

between rings distinct ventrally, visible, but poorly marked 

dorsally; anterior ring 3 A of total peristomial length. Peristomial 

cirri to middle of anterior peristomial ring, without 

articulations, slender and tapering. 

Maxillary formula of large syntype 1+1, 6+6, 6+0, 3+9, and 

1+1. Mx III part of distal arc with left Mx IV. Left Mx IV with 

reduced dentate portion and relatively long wing overlapping 

Mx III. 

Branchiae (Figure 37h) present, palmate, distinctly longer 

than notopodial cirri, not reduced in mid-body region. 

Branchiae from setiger 27 in both specimens. First 10-12 pairs 

single filaments; where best developed, towards end of 

fragments, branchiae with up to 3 filaments. Filaments 

digitiform, about as long as notopodial cirri.

120 


.01 

1 

,1 

.5 

.1 

.5 

b, c, g, i. i. k, I 

d 

h 

, e 

f 

FIGURE 37.—Eunice denliculata ((holotype, USNM 4790): a, anteromedian parapodium; anterior view; b, 

appendage of compound falciger, anteromedian parapodium; c, subacicular hook, anterio-median parapodium; d, 

anterior end, lateral view; e, parapodium 17, anterior view; f, acicula, parapodium 17; g, pectinate seta, 

parapodium 17; h, posterior parapodium, anterior view; i, compound falciger, posterior parapodium; j, 

subacicular hook, posterior parapodium; k, pectinate seta, posterior parapodium; 1, acicula, posterior parapodium. 

(Scale bars in mm.)


Anterior neuropodial acicular lobes (Figure 37e) broadly 

asymmetrically truncate with aciculae emerging near upper 

edge; small elevated tabs present superior to acicula; median 

and posterior acicular lobes without the elevated tabs (Figure 

37a). All presetal and postsetal lobes low folds. Anterior 

ventral cirri thick and tapering, becoming basally inflated from 

about setiger 10. Inflated bases elongated transverse welts; 

narrow tips tapering. Inflated bases reduced from about setiger 

65 and absent from about setiger 85. Posterior ventral cirri short 

and digitiform. Anterior notopodial cirri long and digitiform, 

becoming slender in branchial region, but retaining similar 

length throughout, without articulations. 

Limbate setae narrow and marginally frayed. Pectinate setae 

(Figure 37g Jc) narrow, tapering and furled. Both marginal teeth 

thicker and slightly longer than other teeth; -12 teeth present. 

Shafts of compound falcigers (Figure 37i) distally inflated, 

some marginally serrated, others with smooth margins. 

Appendages (Figure 37b,i) slender, varying in length, bidentate. 

Proximal teeth very much larger than distal teeth, directed 

laterally or slightly basally. Distal teeth short and bent. Guards 

distally symmetrically rounded; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. All 

aciculae single; anterior aciculae dark yellow, darkening to 

brown from about setiger 15, distally slightly expanded, 

slightly hammer-headed (Figure 37f,l), bent towards anterior 

end; cross-section round. Subacicular hooks (Figure 37c,j) 

brown, bidentate. Hooks first present from setiger 18-19, 


replacements). Shafts strongly curved; head very distinct; 

proximal teeth large, curved, directed laterally or basally. Distal 

teeth smaller, strongly curved and directed laterally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Posterior termination 

of branchiae; pygidium and anal cirri. 


TURES.—Branchiae continued to near posterior end. 



Unknown Characters: 1, 2, 36-38,40,42, 74, 78. 


38,1. 

REMARKS.—Eunice denticulata belongs to group B-4 and is 

compared to similar species in Tables 33 and 39. Among the 

species in Table 39, it most closely resembles E.flavapunctata 

in that the inflated bases of median ventral cirri form thick, 

transverse welts in both species. Eunice denticulata has 

expanded, slightly hammer-headed aciculae; the aciculae are 

tapering in E. flavapunctata. 

Eunice depressa Schmarda, 1861 

Eunice depressa Schmarda, 1861:127-128,11 figs. 

Marphysa depressa.—Grube, 1878a:101—Augener, 1924:409. 

REMARKS.—This species was referred to Marphysa by 

Grube (1878a: 101). Augener (1924:409) redefined the species. 

The original description clearly indicates that Schmarda had a 

species of Marphysa. 

52. Eunice dilatata Grube, 1877 


Eunice dilatata Grube, 1877:530-531.—Fauchald, 1986:248-249, figs. 

29-34. 

MATERIAL EXAMINED.—Holotype, ZMB 896, Salavatti, 

Timor, coll. Exp. Gazelle. 

COMMENTS ON MATERIAL EXAMINED.—The prostomium 

had been laterally dissected, so the lower outline of the 

peristomium has been reconstructed in the illustration. 

DESCRIPTION.—Holotype incomplete, of unknown sex, with 

92 anterior setigers; length 70 mm; maximal width 10 mm at 


10, 5 mm. Anterior end cylindrical, becoming strongly 

dorsoventrally flattened by setiger 30; segments becoming very 

short and crowded near posterior end. 



Prostomial lobes frontally obliquely truncate, dorsally flattened; 

median sulcus deep. Eyes between bases of A-I and A-II, 

black. Antennae in a horseshoe; A-I and A-II separated by gap 

from A-III; A-III located well forward of other antennae; A-III 

half as thick as A-I and A-II. Ceratophores ring-shaped in all 

antennae, without articulations. Ceratostyles digitiform, without 

obvious articulations. A-I to posterior peristomial ring; A-II 

to end of setiger 1, and A-III to end of setiger 2. Peristomium 

about twice as long as prostomium, cylindrical. Separation 

between peristomial rings visible, but indistinct dorsally, 

possibly also ventrally, but specimen damaged; anterior ring 3 A 

of total peristomial length. Peristomial cirri short, digitiform, 


Maxillary formula 1+1, 5+5, 8+0, 6+7, and 1+1; Mx II with 

unusually large and heavy teeth compared to other maxillae. 

Branchiae (Figure 38c) present, pectinate, distinctly longer 


Branchiae from setiger 19 to end of fragment. First 5 pairs 

single filaments; maximum 6 filaments; most branchiae with 5 

filaments; this number of filaments continued to end of 

fragment. Branchial stems slender, longer than notopodial cirri. 

Filaments digitiform, longer than notopodial cirri, increasing in 

length posteriorly. 

Anterior neuropodial acicular lobes (Figure 38a) symmetrically 

rounded; median and posterior acicular lobes distally 

truncate; aciculae emerging at midline. Presetal lobes low, 

transverse folds. Anterior postsetal lobes free, rounded, about 

as high as acicular lobes, reduced to low folds from median 

setigers. Median and posterior parapodia on high ridges thus all 

parapodial structures, including aciculae, free of body wall, 

resembling large, flattened paddles with parapodial features 

carried at distal end. Anterior ventral cirri large, tapering from

122 


. d, t, h 

r 

a, c, 

FIGURE 38.—Eunice dilatata (holotype, ZMB 896): a, parapodium 6, anterior view; b, anterior end, lateral view; 

c, parapodium 81, anterior view; d, compound falcigcr, parapodium 6; e, subacicular hook, parapodium 81; f, 

pectinate seta, parapodium 81. Eunice djiboutiensis (syntype, MNHN, Paris): g, subacicular hook, parapodium 

260; h, pectinate seta, parapodium 260; i, parapodium 260, anterior view. (Scale bars in mm.)


a wide triangular base, nearly scoop-shaped. Ventral cirri 

basally inflated in branchial region; bases ovate; narrow tips 

tapering. Anterior notopodial cirri basally distinctly inflated 

with a narrowed attachment to body wall. Notopodial cirri 

decreasing in length in branchial region, retaining similar shape 

in all setigers. Notopodial cirri without articulations. 

Limbate setae slender, nearly capillary. Pectinate setae 

(Figure 38f) in thick fascicles. Shafts wide, cylindrical. Blades 

narrow, tapering, furled. Both marginal teeth longer than other 

teeth; with -15 teeth. Prebranchial setigers with compound 

falcigers in thick fascicles; numbers of falcigers decreasing 

posteriorly; in last segments present only 4 or 5 in each 

parapodium. Shafts (Figure 38d) tapering and marginally 

serrated, with distinct beaks. Appendages large, head distinct, 

bidentate. Proximal teeth smaller than distal teeth, narrowly 

tapering, directed laterally. Distal teeth long, slender, blunttipped, 

directed obliquely distally. Guards symmetrically 

bluntly pointed, marginally coarsely serrated; mucros absent. 


Aciculae single, dark yellow in anterior setigers, darkening to 

dark brown in last segments, tapering to smooth, sharp tips; 

cross-section round. Subacicular hooks (Figure 38e) brown, 

bidentate. Hooks first present from setiger 28, present in all 


Hooks distally abruptly tapered. Proximal teeth larger than 

distal teeth, directed laterally. Distal teeth erect. 


Mx III and left Mx IV; features associated with far 

posterior setigers, including branchial distribution; pygidium 

and anal cirri. 


Mx III forms part of distal arc with left Mx IV; branchiae 

continued to near posterior end. 



Characters: 1,2,36,38,40,42,47,50,51,57-60,63,74,78. 


38,1. 

REMARKS.—The strongly flattened body and the large 

numbers of compound hooks in anterior setigers are features 

often present in the genus Marphysa. Other features, most 

importantly the presence of peristomial cirri, align the species 

with the genus Eunice. It is listed with similar species in Tables 

33 and 38 and can be separated from all other species in these 

tables by the body form. 

53. Eunice djiboutiensis Gravier, 1900 

FIGURE 38g-i; TABLES 27,28 

Eunice aphroditois djiboutiensis Gravier, 1900:224-229, figs. 73-77, pi. 13: 

figs. 63, 67. 

MATERIAL EXAMINED.—One syntype, MNHN, Paris, Djibouti, 

coll. Jousseaume and Goudieu, 1897. 


has been flattened and details of the prostomium and 

peristomium are difficult to distinguish. Gravier originally had 

two specimens. 

DESCRIPTION.—Syntype with 318 setigers, missing a few 

setigers posteriorly; length 320 mm; maximal width 10 mm at 

setiger 10; length through setiger 10,20 mm. Body cylindrical, 

very stiff, dark reddish brown. 


lobes apparently frontally rounded, dorsally slightly inflated; 

median sulcus deep. Eyes at bases of A-I. Antennae in a straight 

line, evenly spaced, similar in thickness. Ceratophores ringshaped 


incomplete, apparently digitiform, rather thick, without articulations. 

No antennae projecting beyond peristomium. Peristomium 

massive, cylindrical; lower lip and nuchal folds 

scalloped. Separation between rings distinct dorsally and 


cirri to middle of anterior peristomial ring, without 

articulations, basally inflated, tapering to thick, digitiform tips, 


Maxillary formula according to Gravier (1900:227-228) 

1+1,5+6, 6+0,4+8, and 1+1. 



Branchiae from setiger 5 to end of specimen. All branchiae 

with two or more filaments; maximum 25 filaments; number of 

filaments reduced slowly posteriorly to about 5. Branchiae 

longer than notopodial cirri in anterior 2 /3 of body; posterior 

notopodial cirri clearly longer than branchiae. Total length of 

branchiae is less than l /2 body width in all setigers. Anterior 

stems short, thick, tapering stiff; posterior stems slender, 

digitiform. Filaments thick, digitiform, shorter than notopodial 

cirri. 

Anterior neuropodial acicular lobes wide and truncate; 

posterior acicular lobes also truncate, but narrower with 

aciculae emerging above midline. Pre- and postsetal lobes low 

folds. First 5 ventral cirri thick, tapering, thereafter modestly 

basally inflated through next 20-30 setigers. Inflated bases 

ovate, narrow tips digitiform. Inflated bases gradually reduced; 

ventral cirri broadly transversely attached tapering to digitiform 

tips in last 150 setigers. Notopodial cirri basally inflated, 

tapering to thick, digitiform tips, without articulations. 

Limbate setae frayed marginally. Pectinate setae (Figure 

38h) numbering up to 25 in a parapodium. Shafts wide, 

cylindrical. Blades flat, flaring. One marginal tooth longer than 

other teeth; with 15-25 teeth. Shafts of compound falcigers at 

least as thick as aciculae, tapering, marginally smooth, 

copper-colored. All appendages lost, illustrated by Gravier 

(1900, fig. 76) as short, triangular, bidentate. Proximal teeth 

short, directed obliquely distally; distal teeth nearly erect; small 

bosses apparently present between proximal and distal teeth. 

Guards asymmetrically bluntly pointed; mucros absenL Pseudocompound 

falcigers and compound spinigers absent. Acicu-


lae single in most parapodia, with dark brown to black cores 

and clear sheaths, distally tapering to slender, gently curved 

tips; cross-section round. Subacicular hooks (Figure 38g) with 

dark brown to black cores and clear sheaths, bidentate. Hooks 

first present from setiger 81, present in less than half the next 

100 setigers, becoming more frequent in posterior half of body 

and present in all setigers near posterior end of specimen, 


tapering smoothly, with very small heads. Proximal teeth 

triangular and directed laterally; distal teeth nearly erect and 

sharply tapered; at junction between teeth small, sharply 

pointed bosses on both sides of hooks, making hooks appear 

quadridentate in frontal view. 


Mx III and left Mx IV; all features associated with far 




TURES.—Mx III forms a distal arc with left Mx IV; branchiae 

continued to near the posterior end. 



Characters: 1,2,11,12, 36-38,40,42. 


12,2; 37,1; 38,1. 

REMARKS.—Eunice djiboutiensis was distinguished from E. 

aphroditois on the structure of the subacicular hooks. It is listed 

with similar species in Tables 27 and 28. In contrast to E. 

aphroditois, it lacks inflated bases to the posterior ventral cirri. 

As usual in the genus, the very large specimens often lose the 

subacicular hooks over large fractions of the body regions in 

which they would be expected to be present. Gravier had two 

specimens; the other specimen must have dissected for the jaw 

structure because the specimen examined for this study had not 

been dissected. 

Eunice dubia Woodworth, 1907 

Eunice dubia Woodworth, 1907:11-12, pi. 1: figs. 4-6, pi. 2: fig. 15. 

Lysidice falax Woodworth, 1903:881. 

REMARKS.—The name Eunice dubia was proposed for the 

epitokous spawning ends of a palola by Woodworth. He had 

previously referred to the same specimens as Lysidice falax. No 

material is available and the illustration of the parapodia 

indicates that no subacicular hooks or pectinate setae were 

present. The species is here considered belonging to the genus 

Palola. 

54. Eunice dubitata Fauchald, 1974 

FIGURE 39a-g; TABLES 4,24,25 

Eunice dubitalus Fauchald, 1974b:18-21, fig. 2a-f [note original, incorrect 

ending of species name]. 

MATERIAL EXAMINED.—Holotype and 3 paratypes, ZM 

Bergen, 55042 and 55043, Hardangerfjorden, S of Huglhamaren, 

59°48'42"N, O5°35'1O"E, 260-180 m, coral bottom, 

Hardangerfjords-undcrsOkelscne St. Z13/59, Square H48-18, 6 

Jun 1959. 

COMMENTS ON MATERIAL EXAMINED.—Body shape and 

parapodial structures arc difficult to identify properly because 

of the poor fixation of the specimens. 


-113 setigers; length 120 mm; maximal width 7 mm at setiger 

10; length through setiger 10, 16 mm. Paratypes of unknown 

sex; with 48 and 92 setigers; length 52 and 134 mm; maximal 

width and length through setiger 10 about as in holotype. All 

specimens basically cylindrical, with slightly flattened dorsum. 

Prostomium (Figure 39c) distinctly shorter and narrower 

than peristomium, as deep as '/2 of the pcristomium. Prostomial 


shallow. Eyes between bases of A-I and A-II. Antennae in 

shallow horseshoe, evenly spaced; A-I thicker than A-II and 

A-I 11. Ccratophorcs ring-shaped in all antennae, without 

articulations. Ceratostylcs of A-I basally slightly inflated; other 

ccratostyles tapering, with up to 10 irregular articulations; 

articulations basally cylindrical, becoming drop-shaped distally. 

A-I to posterior pcristomial ring; A-II to setiger 1 in 

holotype and sctigcr 3 in one paratype; A-I 11 to setiger 3 in 

holotype and setiger 5 in one paratype. Antcnnal length of other 

paratype resembling holotype closely. Peristomium cylindrical; 

lower lip deeply scalloped. Separation between rings 

distinct dorsally and ventrally; anterior ring 5 /6 of total 

peristomial length. Peristomial cirri to posterior edge of 

prostomium in holotype and middle of prostomium in 

paratypes, slender and digitiform, with 4 to 7 articulations. 

Maxillary formula of large paratype 1+1, 7+7, 6+0, 6+8, 

1+1, and 1+1. Mx VI present. Mx III relatively long, but 

forming distal arc with left Mx IV. 

Branchiae (Figure 39g) present, palmate, distinctly shorter 


Branchiae from setiger 3 (4 and 5 in paratypes) to setiger 38 (38 

and 48 in paratypes). Branchiae terminating well before 

posterior end, on less than 55% of total number of setigers. First 

several branchiae very short, button-shaped; occurrence of 

branchiae from about setiger 25 very irregular. First 5-6 and 

last 10-15 branchiae single filaments; maximum 3 filaments at 

about setiger 10-20; even in this region most branchiae with 2 

filaments only. Branchial stems reduced to short, basal area. 

Filaments digitiform, shorter than notopodial cirri. 

Neuropodial acicular lobes truncate or rounded; aciculae 

emerging at midline. Pre- and postsetal lobes low, transverse 

folds. Ventral cirri without inflation in -10 setigers. Bases 

inflated from about setiger 10. Bases ovate; narrow tips 

tapering. Inflated bases reduced from about setiger 35 and lost 

by setiger 40. Remainder of ventral cirri thick, digitiform. 

Anterior notopodial cirri basally inflated, tapering to long, 

slender tips, with 5 to 7 articulations. Notopodial cirri without 

articulations by setiger 20, reaching nearly middle of body in 

most setigers.


FIGURE 39.—Eunice dubitata (hololypc, ZM Bergen, 55042): a, compound falciger, parapodium 110; b, 

subacicular hook, parapodium 110; c, anterior end, lateral view; d, acicula, parapodium 14; e, pectinate seta, 

parapodium 110; f, compound falciger, parapodium 14; g, parapodium 14, anterior view. Eunice edwardsi 

(holotype, BM(NH) ZK. 1885.12.1.3): h, anterior end, lateral view. (Scale bars in mm.)


TABLE 4.—Variable and invariable features in the type lot of Eunice dubitala (N = number of individuals 


VARIABLE FEATURES Max. Min. Mean SD 

Maximal width 




Branchial start 

Last branchia on setiger no. 

Max. no. of branchial filaments 


No. of articulations in notopodial cirri 


4 7 

4 17 

4 12 

4 7 

4 5 

4 48 

4 3 

4 40 

4 6 

4 38 

6 

15 

10 

4 

3 

35 

2 

38 

4 

35 

6.6 

15.8 

10.4 

5.2 

4 

39.6 

2.8 

39.2 

5.4 

36.2 

0.55 

0.84 

0.89 

1.30 

0.71 

4.93 

0.45 

1.10 

0.89 

1.64 


Anterior and posterior peristomial rings distinct 


Notopodial cirri articulated 

Shape of pectinate setae 



Acicular tips 


No. of teeth in subacicular hooks 


N=4 

both dorsally and vcntrally 

mid-prostomium 

anteriorly only 

tapering; flat 

20 

dark brown 

tapering; bent 

dark brown 

2 

indistinct 

Limbate setae marginally serrated. Shafts of pectinate setae 

(Figure 39e) wide, cylindrical. Blades tapering, flat. One 

marginal tooth distinctly longer than other and both marginal 

teeth longer than other teeth, with about 20 very fine, slender 

teeth. Shafts of compound falcigers (Figure 39a,0 slightly 

inflated, internally striated at least in posterior setigers, without 

serrations; distal beaks indistinct. Appendages tapering with 

rather large head, bidentate. Proximal teeth shorter than distal 

teeth, triangular, directed laterally or slightly distally. Distal 

teeth gently curved or nearly erect. Guards tight fitting, 

symmetrically rounded or bluntly pointed; margin serrated near 

bases; mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae and subacicular hooks with 

indistinctly marked dark brown to black cores and clear 

sheaths. Aciculae (Figure 39d) at least paired, tapering with 

slightly bent tips; cross-section round, sometimes with a 

subdistal boss. Subacicular hooks (Figure 39b) bidentate. 

Hooks first present from setiger 35-38, present in all setigers 


gently inflated subdistally, tapering to narrow necks and small 

heads. Proximal teeth larger than distal teeth, tapering, directed 

laterally. Distal teeth triangular, nearly erect. 


anal cirri. 





Characters: 1,2. 

and 


None. 

REMARKS.—The species name as originally proposed is 

grammatically incorrect as pointed out by Inger Winsnes (in 

litt.). The species is listed with similar species in Tables 24 and 

25. Species in this group with poorly developed, palmate 

branchiae include E. palauensis, E. stanleyi, and E. thomasiana, 

in addition to E. dubitata. Among these species E. 

palauensis lacks ceratostyle articulations; the other three 

species have articulated ceratostyles with the distalmost 

articulations drop-shaped or moniliform. The type of E. 

dubitata is about twice as large as those of the two remaining 

species. The last branchiae are present on setiger 79 in E. 

thomasiana, on setiger 33 in E. stanleyi, and on setigers 38-48 

in the much larger E. dubitata. Eunice stanleyi has maximally 

two branchial filaments; E. dubitata has three; other differences 

between these two species are discussed below in the 

"Remarks" section on E. stanleyi. 

Eunice ebranchiata Quatrefages, 1866 

Eunice ebranchiata Quatrefages, 1866:316-317. 

Eunice siciliensis.—Ehlers, 1868:353.—Grube, 1870a:296. 

REMARKS.—The original description is brief, but includes 

the information that branchiae are missing; that the mandibles 

are very large and encloses the other jaws; and that the antennae 

are short. Grube (1870a:296) examined the type and declared it 

completely in agreement with many specimens he had seen of 

Eunice siciliensis. The species is here considered a member of 

the genus Palola.


55. Eunice edwardsi Mclntosh, 1885 

FIGURE 39h; TABLES 19, 21 

Eunice edwardsi Mclntosh, 1885:280-282, figs. 39-40, pi. 38: figs. 14, 15, pi. 

20A: figs. 4-7. 

MATERIAL EXAMINED.—Holotype, BM(NH) ZK. 

1885.12.1.3., off Prince Edward Island, Indian Ocean, 46°40'S, 

37°5O'E, 567 and 274 m, 27 Dec 1873, Challenger sta 145. 

COMMENTS ON MATERIAL EXAMINED.—The holotype has at 

one time been dried out and the setae are all broken; it is in two 

pieces and has been deeply dissected frontally; the jaw 

apparatus is now missing. 


118 sctigcrs; total length about 70 mm; maximal width 3.5 mm 

wide at sctigcr 10; length through setiger 10, 8 mm. Body 

circular in cross-sec lion, tapering posteriorly. 

Prostomium (Figure 39h) distinctly shorter and narrower 

than pcristomium, less than x li as deep as peristomium. 


sulcus shallow. Eyes between bases of A-I and A-II, black. 

Antennae in transverse row; A-I and A-II grouped together, 

separated by gap from A-I 11; all antennae similar in thickness. 


Ceratostyles digitiform, with up to 5 cylindrical articulations in 

A-I 11. A-I to middle of anterior peristomial ring; A-II to setiger 

1; A-III to setiger 3. Peristomium about twice as long as 

prostomium, cylindrical. Separation between rings especially 

distinct dorsally and ventrally, but also visible laterally; 


middle of prostomium, digitiform, without articulations. 

Maxillary formula (1+1), 8+8, 10+0, 6+12, and 1+1 

according to Mclntosh (1885, fig. 39). Mx I missing in 

illustration. 



setiger 3 to setiger 34; to setiger 41 according to Mclntosh 

(1885). Branchiae terminating well before posterior end, 

present on less than 55% of total number of setigers. First 5 and 

last 2 branchiae single filaments, between setiger 10 and 20 

number of filaments increasing to 5 or 6. 

Neuropodial acicular lobes distally rounded, becoming 

conical in last segments present; aciculae emerging at midline. 

Pre- and postsetal lobes visible only as low folds. First 5 ventral 

cirri thick, tapering, becoming distinctly inflated basally from 

about setiger 6. Inflated bases ovate; narrow tips tapering. 

Inflated bases reduced from about setiger 25. Posterior ventral 

cirri somewhat shorter and thicker than notopodial cirri, 

tapering. Notopodial cirri basally somewhat inflated, rather 

thick in first 20 setigers, becoming increasingly digitiform 

posteriorly, without articulations. 

All setae broken. According to Mclntosh's illustrations 

(1885, pi. 20A: figs. 5-8), pectinate setae with few teeth and 1 

marginal tooth distinctly longer than other teeth. Compound 

falcigers bidentate; both teeth well developed. Guards blunt 

without marginal serrations; mucros absent. Aciculae yellow, 

apparently bluntly rounded distally; cross-section round. 

Subacicular hooks yellow, bidentate. Hooks first present from 

setiger 28. Hooks tapering. Both teeth directed distally. 

Proximal teeth very much larger than distal teeth. 


Mx III and left Mx IV; details of setal structures; 



TURES.—Mx III long and located behind left Mx II. Pseudocompound 

falcigers and compound spinigers absent. 



Characters: 65-68, 71, 72, 81, 82. 


72,2. 

REMARKS.—Eunice edwardsi is listed with similar species 

in Tables 19 and 21. It does not appear to be characterized by 

a single unique feature, but rather by a combination of a whole 

set of minor features. 

46. Eunice ehlersi Gravier, 1900 

FIGURE 40a-e; TABLES 33-35 

Eunice ehlersi Gravier, 1900:248-251, figs. 110-113,pl. 13: figs. 75,76. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, Djibouti, 

1897, coll. Coutiere, no. 29. 


53 setigers; length 25 mm; maximal width 4 mm at setiger 10; 

length through setiger 10, 6 mm. Body cylindrical. 




median sulcus deep. Eyes present posterior to A-I, dark. 

Antennae in shallow horseshoe, evenly spaced, similar in 

thickness. Ceratophores ring-shaped, without articulations. 

Ceratostyles digitiform, with up to 8 short, cylindrical 

articulations in A-III. A-I to posterior peristomial ring; A-II and 

A-III to setiger 3. Peristomium flaring slightly ventrally. 

Separation between rings distinct on all sides; anterior ring 2 h 

of total peristomial length on dorsal side and ~ 3 A on ventral 

side. Peristomial cirri to front edge of peristomium, digitiform, 


Maxillary formula 1+1,5+6, 8+0, 3+9, and 1+1. Distal tooth 

on right Mx II very small. Mx III long, located in part behind 

Mx II, however forming arc with very short left Mx IV. Mx VI 

absent. 


than notopodial cirri, flexible. Branchiae from setiger 14 to end 

of fragment. First branchiae single filaments, maximum 6

128 


FIGURE 40.—Eunice ehlersi (holotype, MNHN, Paris): a, anterior end, lateral view; b, pectinate seta, parapodium 

32; c, subacicular hook, parapodium 32; d, compound falciger, parapodium 32; e, parapodium 32, anterior view. 

Eunice eimeorum, new species (holotype, part of RM 310): f, anterior end, lateral view; g, parapodium 49, 

anterior view; h, compound falciger, parapodium 49; i, pectinate seta, parapodium 49; j, acicula, parapodium 49; 

k, subacicular hook, parapodium 49. (Scale bars in mm.) 

filaments. Branchial stems very slender, pliable, longer than 

notopodial cirri. Filaments slender, longer than notopodial cirri. 



folds. First 4 ventral cirri thick, tapering. Median ventral cirri 

with inflated bases forming elongated ventral ridges along 

lower edge of neuropodia; narrow tips short and button-shaped. 

Posteriormost ventral cirri distinctly tapering inflated base


reduced, but still visible as a ridge. Anterior notopodial cirri 

basally inflated, becoming tapering in branchial region; 

articulations absent 

Limbate setae slender, marginally smooth. Pectinate setae 

(Figure 40b) with narrow, cylindrical shafts. Blades slightly 

furled, flaring. One marginal tooth distinctly longer than other 

teeth, with -12 teeth. Shafts of compound falcigers (Figure 

40d) thick, slightly inflated, marginally smooth; beaks indistinct. 

Appendages short, tapering, with very large heads, 

bidentate. Proximal and distal teeth similar in size. Proximal 

teeth tapering, directed obliquely distally. Distal teeth tapering, 

nearly erect. Guards distally symmetrically bluntly pointed, 

nearly rounded, marginally serrated; mucros absent. Pseudocompound 


single, anteriorly pale honey-colored, becoming brown to 

dark brown, sharply and symmetrically pointed, straight; 

cross-section round. Notopodial aciculae honey-colored. Separation 

between core and sheath distinct in both aciculae and 

subacicular hooks. Subacicular hooks (Figure 40c) bidentate. 

Hooks first present from setiger 27. Proximal teeth larger than 

distal teeth, directed laterally, upturned distally. Distal teeth 

narrowly triangular, erect. Guards distally rounded. 

UNKNOWN MORPHOLOGICAL FEATURES.—Parapodial and 

branchial features associated with far posterior setigers; 






1,2, 36-38, 40, 42, 57-60, 81, 82. 


37,1; 38,1. 

REMARKS.—Eunice ehlersi is listed with similar species in 

Tables 33-35. Of the species in these tables, E. ehlersi and E. 

sonorae have the separation between the peristomial rings 

distinct on all sides. Eunice ehlersi has bidentate subacicular 

hooks; E. sonorae has simple, spine-like subacicular hooks. 

Eunice ehlersi is otherwise characterized by a combination of 

features, including strongly articulated antennae and honeycolored 

to brown aciculae and subacicular hooks, rather than 

the black hooks common in the group. 

57. Eunice eimeorum, new species 

FIGURE 40f-k; TABLES 27,31 

Eunice pacifica Kinberg, 1865:562 [in part]; 1910:41-42, pi. 15: fig. 15 [in 

part].—Hartman, 1948:75-76 [in part]. 

MATERIAL EXAMINED.—Holotype, part of RM 310, Eimeo 

(= Moorea), Society Islands (Tahiti Islands), Eugenie Expedition. 


been dried at one time and the last 20 setigers have been 

strongly flattened; thus an accurate count of setigers is 

impossible. 

DESCRIPTION.—Holotype incomplete with -110 setigers; 

length 70 mm; maximal width 3.5 mm; length through setiger 

10, 7.5 mm. 

Prostomium (Figure 40f) distinctly shorter and narrower 

than peristomium, less than l /i as deep as pcristomium. 

Prostomial lobes frontally rounded, dorsally flattened; median 

sulcus deep. Eyes posterior to bases of A-I, dark. Antennae in 

a horseshoe, evenly spaced, similar in thickness. Ceratophores 


slender and tapering, with up to 8 long, cylindrical, wellmarked 

articulations in A-III. Three median antennae to 

posterior peristomial ring; A-III longer than A-I I; A-I to middle 

of anterior peristomial ring. Peristomium massive, cylindrical. 



middle of anterior peristomial ring, medially inflated, without 


Jaws missing. 



Branchiae from setiger 5 to end of fragment. Maximum 5 

filaments from about setiger 20; in last setigers present 

branchiae with 2-3 filaments. Branchial stems short, tapering. 

Filaments long, slender. 

Anterior neuropodial acicular lobes distally truncate; posterior 

acicular lobes rounded; aciculae emerging at midline. All 

pre- and postsetal lobes very low, transverse folds, considerably 

shorter than acicular lobes. Anterior ventral cirri thick, 

tapering, becoming basally inflated in median setigers. Bases 

ovate, narrow; tips long and digitiform. Inflated bases 

apparently reduced posterior to setiger 60-65. All notopodial 

cirri medially inflated, relatively long, less distinctly basally 

inflated posteriorly, digitiform in last setigers present, without 



pectinate setae (Figure 40i) thick, cylindrical. Blades narrow, 

flat, tapering. Marginal teeth no longer than other teeth; about 

10 teeth present. Compound falcigers very large (Figure 40h). 

Shafts tapering, marginally smooth. Appendages very short, 

truncate, with large heads, bidentate. Proximal teeth very much 

smaller than distal teeth, slender, tapering, directed distally. 

Distal teeth distinctly curved or bent, thick, directed obliquely 

distally. Guards short, asymmetrically bluntly pointed, frayed 

in most hooks; mucros absent. Pseudocompound falcigers and 

compound spinigers absent. Aciculae (Figure 40j) paired, clear 

amber, barely tapering, blunt, curved; cross-section of aciculae 

round. Subacicular hooks (Figure 40k) clear amber, bidentate. 

Hooks first present from setiger 38. Hooks slender; neck 

distinct; head large. Proximal teeth much smaller than distal 

teeth, triangular, directed obliquely laterally. Distal teeth 

triangular, erect. 


features associated with posterior setigers; pygidium and anal 

cirri.




IV. 



Characters: 1,2,4, 6, 33, 36-40, 51, 58, 59, 74, 78, 81, 82. 


37,1; 38,1. 

REMARKS.—Eunice eimeorum differs clearly from E. pacifica 

in the distribution of branchiae, which are present from 

setiger 5 in E. eimeorum and from setiger 17-21 in E. pacifica. 

Clear amber subacicular hooks are present from setiger 38 in E. 

eimeorum and dark brown, nearly black subacicular hooks 

present from setigers 23-28 in E. pacifica. The specimens of 

the two species are similar in size. 

Eunice eimeorum is listed with similar species in Tables 27 

and 31. The appendage of compound hooks resembles the 

compound hooks of E. pelamidis. 

58. Eunice elegans (Verrill, 1900) 

FIGURE 41a-k; TABLES 24,25 

Leodice elegans Verrill, 1900:640-641. 

Leodice longicirrata.—Treadwell, 1921:11-14, figs. 3-12, pi. 1: figs. 1-4 

[not Eunice longicirrata Webster, 1884]. 

Eunice longicirrata.—Hartman, 1942:9 [not Eunice longicirrata Webster, 

1884]. 

MATERIAL EXAMINED.—Holotype, YPM 2730, Bermuda, 

low [shallow] water, Apr 1898, coll. A.E. Verrill and party. 



setiger 10; length through setiger 10, 8 mm. Body cylindrical 

anteriorly, becoming ventrally flattened posteriorly. 


than peristomium, as deep as l li of peristomium. Prostomial 

lobes frontally obliquely truncate, flattened dorsally and 

ventrally; median sulcus very shallow. Eyes between bases of 

A-I an A-II. Antennae in a horseshoe, evenly spaced. 

Ceratophores long in all antennae, without articulations. Only 

left A-II present (Figure 41a), now detached, digitiform, with 9 

short, but distinctly cylindrical articulations, to setiger 2. 

Peristomium cylindrical, about twice as long as prostomium. 

Separation between rings distinct on all sides, but especially 

well marked dorsally and ventrally; anterior ring 3 /4 of total 


tapering, with 4 indistinct articulations. 






of total number of setigers. First and last branchiae single 

filaments; maximum 10 filaments at setiger 10-15. Branchiae 

covering dorsum where best developed. Stems slender, erect. 

Filaments trim, slender, shorter than notopodial cirri. 

Anterior neuropodial acicular lobes rounded, becoming 

increasingly conical posteriorly (Figure 4Id); aciculae emerging 

above midline. Pre- and postsetal lobes low, transverse 

folds. First 4 ventral cirri tapering. Ventral cirri basally inflated 

from setiger 5 through about setiger 50. Inflated bases ovate; 

narrow tips tapering. Posterior ventral cirri broadly attached, 

tapering to blunt tips, forming very shallow, open scoops 

around ventral margin of neuropodia. Notopodial cirri supported 

by aciculae; anterior notopodial cirri basally inflated and 

indistinctly separated into 4 articulations. Articulations lost in 

first few setigers of branchial region; notopodial cirri becoming 

slender and tapering from first postbranchial setigcrs, retaining 

that shape to the end. 

Limbate setae slender, marginally finely serrated. Pectinate 

setae (Figure 41c,j) tapering, flat. One marginal tooth longer 

than other teeth, about, 10 teeth present; anterior pectinate setae 

somewhat asymmetrical, becoming symmetrical in postbranchial 

region. Shafts of anterior compound falcigers (Figure 

41k) tapering, becoming gently inflated (Figure 410 in 

postbranchial setigcrs, marginally smooth; beaks indistinct. 

Anterior appendages tapering; heads indistinct, bidentatc. 

Proximal teeth very much shorter than distal teeth, triangular. 

Distal teeth tapering, nearly erect. Guards asymmetrically 

bluntly pointed. Postbranchial appendages short, with large 

heads. Proximal teeth and distal teeth similar in size; proximal 

teeth tapering, directed basally. Distal teeth smoothly curved. 

Guards symmetrically rounded; all guards marginally smooth; 


spinigers absenL Neuropodial aciculae paired, dark yellow to 

amber-colored; anterior aciculae (Figure 41 i) distally expanded 

into rounded knobs; postbranchial aciculae (Figure 41h) 

sharply tapered; superior aciculae gently curved dorsally; 

cross-section round. Separation between core and sheath 


hooks (Figure 41g) amber-colored, bidentate. Hooks first 

present from setiger 30, present in all setigers thereafter, 

usually 2-3 hooks in a parapodium. Hooks with narrow necks 

and distinct heads. Proximal teeth twice as large as the distal 

teeth, triangular, directed laterally. Distal teeth tapering, erect. 

Guards distally truncate. 




Mx III long, located behind left Mx II. 





None. 

REMARKS.—Eunice elegans resembles E. websteri (= E. 

longicirrata Webster) but differs in the changing shape of the 

aciculae and compound hooks in different parts of the body, 

and also in the maximum number of branchial filaments and


1 

2 

.5 

1 

1 

a h 

d 

h, i. o, p 

c. 1 

FIGURE 41.—Eunice elegans (holotype, YPM 2730): a, detached ceratostyle; b, anterior end, lateral view; c, 

parapodium 7, anterior view; d, parapodium 53, anterior view; e, pectinate seta, parapodium 53; f, compound 

falciger, parapodium 53; g, subacicular hook, parapodium 43; h, aciculae, parapodium 53; i, aciculae, parapodium 

7; j, pectinate seta, parapodium 7; k, compound falciger, parapodium 7. Eunice elseyi (syntype, BM(NH) ZH 

1857.11.18.249): 1, parapodium 103, anterior view; m, anterior end, lateral view; n, pectinate seta, parapodium 

103; o, subacicular hook, parapodium 103; p, aciculae, parapodium 103; q, appendage of compound falciger, 



numbers of branchial setigers present and in the much darker 

color of the subacicular hooks and aciculae. The latter makes it, 

by definition, a member of group B-l rather than of group A-l. 

It is listed with similar species in Tables 24 and 25. It is the only 

species listed in Table 25 that has two or three subacicular 

hooks in most segments and has the aciculae distally expanded 

and tab-shaped at least in some setigers. 

59. Eunice elseyi Baird, 1869 

FIGURE 411-q; TABLES 46,47,52.53 

Eunice elsyi Baird, 1869:344. 

Eunice elseyi—Mclntosh, 1885:286-287, figs. 46, 47, pi. 39: figs. 1-3, pi. 

20A: figs. 14-16.—Fauchald, 1986:249-250, figs. 35-40. 

MATERIAL EXAMINED.—Two syntypes, BM(NH) ZH 

1857.11.18.249, and in addition 2 mounted slides with 

parapodia and 1 slide with the jaws. North Australia, coll. J.R. 

Elsey, North Australia Exploring Expedition, 1855-56. 

COMMENTS ON MATERIAL EXAMINED.—One syntype is in 

two pieces; the anterior end has been deeply dissected and the 

jaws removed. The other syntype consists of 48 anterior 

setigers. Three short fragments in the vial do not belong to the 

same species; they were briefly described by Fauchald (1986). 

DESCRIPTION.—Syntype described complete, of unknown 

sex, with 120 setigers; last 16 in regeneration; total length 64 


Prostomium (Figure 41m) distinctly shorter than and 

narrower than peristomium, less than x /i as deep as peristomium. 

Prostomial lobes frontally rounded, dorsally inflated; 

median sulcus deep. Antennae in shallow horseshoe, evenly 

spaced, similar in thickness. Ceratophores ring-shaped in all 


tapering, with up to 22 moniliform articulations in A-III. A-I to 

posterior edge of anterior peristomial ring; A-I I to setiger 3; 

A-III, when stretched, to setiger 5. Peristomium massive, 

cylindrical. Separation between rings well marked ventrally 

and less distinct dorsally; anterior ring 4 /s of total peristomial 


and tapering, with -10 articulations. 


Branchiae (Figure 411) present, pectinate, distinctly longer 


Branchiae from setiger 6 to setiger 104, missing in regenerating 

portion. Branchiae present to near posterior end, present on 

more than 65% of total number of setigers. All branchiae 

strongly pectinate; most branchiae with at least 10 filaments, 

maximum 18 filaments. Branchial stems erect, tapering, shorter 

than notopodial cirri, decreasing in length posteriorly. Filaments 

slender, digitiform, shorter than notopodial cirri. 

Numbers of filaments remaining high even in last branchiated 

segments. 

Neuropodial acicular lobes symmetrically truncate; aciculae 


folds. First 5 ventral cirri digitiform. Median ventral cirri 

basally inflated. Inflated bases ovate; narrow tips digitiform. 

Ventral cirri slender and digitiform from the last branchiated 

segments. Notopodial cirri (other than those in the regenerate) 

long, basally somewhat inflated, with up to 9 articulations. 

Most notopodial cirri reaching nearly midway across body. 

Limbate setae slender. Shafts of pectinate setae (Figure 4In) 

wide, cylindrical. Blades tapering, flat. One marginal tooth 


falcigers inflated, marginally smooth. Appendages (Figure 

41q) tapering, tridentate with small third teeth closely 

appressed to second teeth. Proximal teeth triangular, directed 

laterally. Second teeth larger than proximal teeth, directed 

obliquely distally. Guards bluntly pointed, marginally smooth; 

mucros absent. Pscudocompound falcigers and compound 

spinigcrs absent. Aciculae (Figure 41p) yellow, bent dorsally, 

shallowly bidentate in antcropostcrior axis; cross-section 

round. Separation between core and sheath indistinct in both 

aciculae and subacicular hooks. Subacicular hooks (Figure 

41o) yellow, tridentate. Hooks first present from setiger 30, 


replacements). Primary fangs large; secondary fangs small; 

tertiary fangs small, appended laterally to secondary fangs 

rather than emerging dorsally. 




IURES.—Mx III long and located behind left Mx II. 



Characters: 4,6,13,14,42. 


None. 

REMARKS.—The status of this species was explored in 

Fauchald (1986); it is listed with similar species in Tables 46 

and 47. It has bidentate or bifid aciculae and the subacicular 

teeth are tridentate with the teeth in tandem; this combination is 

unique in Table 47. It is compared to other species with the 

teeth in tandem in Tables 52 and 53. 

Eunice enteles (Chamberlin, 1918) 

Leodice enteles Chamberlin, 1918:175-176. 

Eunice enteles.—Hartman. 1938:97. 

lEunice antennata.—Hartman, 1944:115-117, pi. 7: figs. 154-156 [in part, 

not Eunice antennata Lamarck, 1818]. 

REMARKS.—No material is currently available of this 

species at MCZ where Chamberlin indicated that his types were 

deposited. 

It was originally described with branchiae from setiger 6-7 

to the end of the body with a maximum of five filaments and no 

indication of an intermediate region with a reduced number of 

filaments. All antennae had moniliform articulations. A-III 

reached setiger 4 and A-I I reached setiger 3. The peristomial


cirri did not reach beyond the peristomium and were nearly or 

completely smooth. The notopodial cirri had weakly indicated 


The maxillary formula was 1+1, 13+13, 11+0, 10-11+11, 

and presumably 1+1. The number of teeth indicated is 

confused; if the above interpretation is correct, the species 

would have a maxillary formula similar to species in group C-2, 

with which it was associated by Hartman (1944). 

No information is given about the structure or distribution of 

the setae. 

The species may belong to group C-2, but is insufficiently 

characterized to allow accurate designation. The type has been 

lost for a long time. Hartman (1938) did not include it in her 

annotated list of types in the Museum of Comparative Zoology, 

nor is there any evidence available that she saw the type at a 

later stage. The species is here considered indeterminable. 

Hartman (1938) indicated that the type was collected at 

Mendocino, California; however, in Chamberlin (1918) the 

type locality is not specified. A statement in the introduction to 

Chamberlin's paper indicates that his material came from 

Pacific Grove, California, in shallow water or intertidally, 

rather than from Mendocino. 

Eunice equibranchiata Mclntosh, 1885 

FIGURE 42 

Eunice equibranchiata Mclntosh, 1885:291-292, pi. 39; fig. 11, pi. 20A: figs. 

23,24. 


1885.12.1.202. Brazil, 9°5'-9°10'S, 34°49'-34°53'W, 640, 

219, 59, and 732 m, mud, 10 Sep 1873, Challenger sta 202. 

DESCRIPTION.—Holotype and only known specimen is a 

middle piece of about 300 segments; length 250 mm. 

Prostomium, peristomium and jaws unknown. 

Branchiae long, slender; up to 3 filaments of similar lengths; 

all segments present branchiated. 

Neuropodial acicular lobes (Figure 42d) conical. Pre- and 

postsetal lobes follow outline of acicular lobes closely. Ventral 

cirri digitiform with distinct, but low basal inflations on 

adjoining body wall. Notopodial cirri tapering, slender, without 

articulations, at least twice as long as ventral cirri. 

Most setae broken. Shafts of compound hooks distally 

inflated. Appendages (Figure 42a) short. Both teeth very well 

developed, directed laterally. Guards symmetrically rounded, 

marginally serrated. Aciculae (Figure 42b) single, with dark 

cores and clear sheaths, distally asymmetrically hammerheaded. 

Subacicular hooks (Figure 42c) with dark cores and 

clear, translucent heads. Hooks strongly curved, with large 

heads. Both teeth curved, directed laterally. 

REMARKS.—The structures of E. equibranchiata that can be 

examined resemble the corresponding features in E. filamentosa 

and related species, but the exact relationship cannot be 

determined. The species is here considered indeterminable. 

60. Eunice eugeniae, new species 

FIGURE 43a-h. TABLES 41, 45 

Eunice havaica Kinbcrg, 1865:562 [in part]. 

MATERIAL EXAMINED.—Holotype, part of RM 430, Honolulu, 

Hawaii, Eugenie Expedition 1087-89 (one specimen from 

type lot of E. havaica). 



length through setiger 10,4 mm. 

Prostomium (Figure 43a) about as long as peristomium, 

about as wide as peristomium, deeper than V2 of the 


inflated; median sulcus shallow. Peristomial fold forming 2 

transverse yokes across back of prostomium. Eyes absent. 

Antennae in shallow horseshoe, evenly spaced; A-I thicker than 

A-II and A-I 11. Ceratophores ring-shaped in all antennae, 

without articulations. Ceratostyles slightly tapering, without 

articulations. A-I to middle of anterior peristomial ring; A-II 

antennae to setiger 1; A-I 11 to setiger 4. Peristomium 

cylindrical. Separation between rings distinct on all sides; 

anterior ring little more than x /i of total peristomial length. 

Peristomial cirri to front margin of peristomium, digitiform, 





setiger 4 through setiger 44. Branchiae terminating well before 

posterior end. First 4 and last 2 or 3 branchiae single filaments; 

maximum 5 filaments. Stems strong, erect, shorter than 

notopodial cirri. Filaments digitiform, shorter than notopodial 

cirri. 

Anterior neuropodial acicular lobes distally truncate, 

rounded in last segments present; aciculae emerging at midline. 

Pre- and postsetal lobes low, transverse folds. First 3 ventral 

cirri thick, tapering. Ventral cirri basally inflated from first 

branchial segments through about setiger 25. Bases ovate; 

narrow tips tapering. Ventral cirri digitiform in last setigers 

present. Prebranchial notopodial cirri basally inflated and 

tapering, becoming increasingly digitiform through branchial 

region and in postbranchial setigers. 

Limbate setae slender. Pectinate setae (Figure 430 flaring, 

flat. Both marginal teeth longer than other teeth, with about 15 

teeth. Shafts of compound falcigers (Figure 43b,g) inflated 

distally, marginally finely or coarsely serrated; distal beaks 

distinct Appendages tapering, with small heads, bidentate. 

Proximal teeth smaller than distal teeth, triangular to reduced 

triangular, directed laterally. Distal teeth short, gently curved or 

erect. Guards distally pointed, with distinct mucros, marginally 

serrated. Pseudocompound falcigers and compound spinigers 

absent. Aciculae (Figure 43d,h) single, yellow, tapering, gently 

curved; cross-section round. Separation between core and 

sheath indistinct in both aciculae and subacicular hooks.

134 


b. c 

FIGURE 42.—Eunice equibranchiata (holotype, BM(XH) ZK. 1885.12.1.202): a, appendage of compound 

falciger, b, acicula; c, subacicular hook; d, parapodium; all from medium setigers. (Scale bars in mm.) 

Subacicular hooks (Figure 43c,e) yellow, tridentate with teeth 

in a crest. Hooks first present from setiger 18, present in all 


Hooks with large curved main fangs. Secondary fangs 

triangular. Tertiary fangs very small, erect. 







Unknown Characters: 1, 2, 38, 42, 63. 


REMARKS.—Eunice eugeniae is listed with similar species in 

Tables 41 and 45. It is the only species in Table 45 in which the 

branchiae are no longer than the notopodial cirri; in the other 

species the branchiae distinctly outreach the notopodial cirri. 

61. Eunice excariboea, new species 

FIGURE 43i-o; TABLES 33,36 

Eunice cariboea Grube, 1856:57 [in part]. 

MATERIAL EXAMINED.—Holotype and 1 paratype, ZMC 

(uncataloged), St. Croix, West Indies, coll. H. Kroyer (part of 

original material of E. cariboea Grube). 

COMMENTS ON MATERIAL EXAMINED.—The holotype is the 

larger of the two specimens examined. 



length through setiger 10, 3.7 mm. Anterior part of body 

including prostomium and peristomium cylindrical. Body 

widening rapidly through first few setigers, becoming nearly 

twice as wide as prostomium by setiger 10, slightly dorsoventrally 

flattened with parapodia elevated on low lateral ridges.


FIGURE 43.—Eunice eugeniae, new species (holotype, part of RM 310): a, anterior end, lateral view; b, compound 

falciger, parapodium 27; c, subacicular hook, parapodium 27; d, acicula, parapodium 27; e, subacicular hook, 

parapodium 40; f, pectinate seta, parapodium 40; g, compound falciger, parapodium 40; h, acicula, parapodium 

40. Eunice excariboea, new species (holotype, ZMC): i, parapodium 10, anterior view; j, compound falciger, 

parapodium 10; k, subacicular hook, parapodium 133; 1, compound falciger, parapodium 133; m, pectinate seta, 

parapodium 133; n, parapodium 133, anterior view; o, anterior end, lateral view. (Scale bars in mm.)


Prostomium (Figure 43o) ~ x ji as deep, very nearly as wide as 


flattened; median sulcus shallow. Eyes behind bases of A-I, 

dark. Antennae in a horseshoe; A-I and A-II emerging close 

together, near outer edges of prostomium, separated by gap 

from A-III, similar in thickness. Ceratophorcs ring-shaped in 

all antennae, without articulations. Ceratostyles slender and 

digitiform, with up to 14 closely spaced, indistinct articulations 

in A-III, articulated to bases of ceratostyles. A-I to posterior 

pcristomial ring; A-II and A-III to sctiger 2. Peristomium 

slightly flaring anteriorly. Separation between rings distinct 

dorsally and ventrally; anterior ring 3 A of total pcristomial 

length. Peristomial cirri to middle of anterior peristomial ring, 

slender and digitiform, without articulations. 


Branchiae (Figure 43n) present, palmate, about as long as 

notopodial cirri, not reduced in mid-body region. Branchiae 

from setiger 78-79 to end of fragments. Most branchiae with 2 

long, narrow filaments emerging from joint bases at dorsal edge 

of notopodial cirri; up to 3 filaments present. Stems reduced. 

Filaments longer than notopodial cirri. 

Neuropodial acicular lobes (Figure 43i) rounded; anterior 

setigers with small rounded tabs dorsal to emergence of 

aciculae; tabs absent in posterior setigers; aciculae emerging at 

midline. Lateral body wall forming low ridges with acicular 

lobes at distal ends from about setiger 25. Pre- and postsetal 

lobes low folds; anterior presetal lobes distinctly lower than 

acicular lobes; in posterior setigers all 3 lobes paralleling each 

other closely. First 4 ventral cirri thick and tapering. Ventral 

cirri basally inflated from about setiger 5. Inflated bases 

distinct, inflated ridges along ventral side of neuropodia in all 

later setigers; narrow tips tapering or digitiform. Anterior 

notopodial cirri tapering with 4 articulations; number of 

articulations reduced to one in posterior setigers; notopodial 

cirri becoming digitiform in far posterior setigers. 

Limbate setae slender, very finely serrated. Pectinate setae 

(Figure 43m) very small, with slender shafts. Blades furled, 

flaring. Both marginal teeth longer than other teeth, with -12 

teeth. Shafts of compound falcigers distinctly inflated and 

marginally serrated; beaks distinct. Anterior appendages 

(Figure 43j) tapering, bidentate. Proximal teeth larger than 

distal teeth; both teeth tapering, distally curved towards each 

other, forming a crabclaw-like distal end. Posterior appendages 

(Figure 431) shorter and more abruptly tapering than anterior 

ones, bidentate. Proximal teeth much larger than distal teeth, 

triangular, directed laterally. Distal teeth slender, tapering and 

nearly erect. Guards symmetrically rounded and marginally 

serrated in all setigers; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculae paired 

anteriorly, single posteriorly, amber-colored, tapering to 

straight tips; cross-section round. Subacicular hooks (Figure 

43k) pale amber, bidentate. Hooks first present from setiger 87 

(78 in paratype), missing in many setigers, always single 

(except for replacements). Hooks gently sigmoid, tapering, 

with distinct head. Proximal teeth larger than distal teeth, 

directed laterally. Distal teeth very small, directed obliquely 

distally. Guards distally rounded. 




Mx III short, forming part of distal arc with left Mx IV. 



Characters: 1, 2, 36-40, 42, 74, 78. 


38,1. 

REMARKS.—The original material of E. cariboea consisted 

of three species; most of the type lots, four specimens in ZMC 

and five in ZMB, belong to an abranchiate species with dark 

brown to black subacicular hooks and aciculae and smooth 

antennae. These specimens arc here considered as E. cariboea, 

for which a lectotypc was designated above. Some specimens, 

currently in ZMH, were designated as types of E. gagzoi by 

Augcncr (1922b). Finally, two specimens from ZMC arc here 

described as E. excariboea. 

Eunice excariboea is listed with similar species in Tables 33 

and 36. Perhaps the most outstanding feature of this species is 

the very late start of both branchiae and subacicular hooks. It 

can be separated from all other species listed in Table 36 by 

having the notopodial cirri articulated throughout; the other 

species in this table lack notopodial articulations. 

Eunice fasciata (Risso, 1826) 

Leodice fasciata Risso, 1826:421. 

Eunice fasciata.—Heider, 1925:55-59, figs. 3-6. 

Eunice torquata.—Grube, 1878a:99.—Fauvel, 1923:401. 

REMARKS.—No material is available; the species was 

referred doubtfully to Eunice torquata by Grube (1878:99) and 

Fauvel (1923:401). Heider (1925), in a brilliant study of the 

anterior anatomy of two species from the Mediterranean, gave 

a detailed description of a species he called E. fasciata. His 

arguments for identifying his material as Risso's species 

amounts to an assertion that "there can be no doubt that his 

(Ehlers, 1868:312, of E. harassii) description... refers loEunice 

fasciata" (Heider, 1925:59). There is nothing in Risso's 

original description to support this assertion. The material 

Heider studied belongs to Eunice torquata. 

Risso's species is insufficiently known to assign it even to 

group and is here considered wholly indeterminable. 

62. Eunice fauveli Gravier, 1900 


Eunice fauveli Gravier, 1900: 236-239, figs. 89-93, pi. 12: fig. 60. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, Obock, 

Red Sea, coll. J. Jousseaume, 1894.


J 

.02 

.1 

1 

a 

c, e 

d 

h 

FIGURE 44.—Eunice fauveli (holotype, MNHN): a, anterior end, lateral view; b, parapodium 75, anterior view; 

c, compound falciger, parapodium 75; d, subacicular hook, parapodium 75; e, pectinate seta, parapodium 75. 

(Scale bars in mm.)




about setiger 20; length through setiger 10, 18 mm; width at 

setiger 10, 7.5 mm. Body through setiger 10 cylindrical; 

remainder dorsoventrally flattened with short, crowded, wide 

segments. Anal cirri as long as last 10 setigers, without 




Prostomial lobes frontally rounded, dorsally slightly flattened; 

median sulcus deep. Eyes on lateral sides of bases of A-II, dark. 

Antennae in deep horseshoe, evenly spaced, similar in 


articulations. Ceratostyles tapering to relatively slender tips, 

with up to 7 cylindrical articulations in one A-II. A-I to setiger 

1; A-II to setiger 3; A-III to setiger 4 (incomplete). Peristomium 

cylindrical. Separation between rings distinct dorsally 

and ventrally; anterior ring ~ 4 /5 of total peristomial length. 

Peristomial cirri to anterior x h of anterior peristomial ring, 

slender and tapering, without articulations. 

Maxillary formula 1+1, 5+5, 6+0,4+9, and 1+1. Left Mx IV 

short with reduced teeth, forming a distal arc with Mx III. Mx 

VI missing. Jaws strongly calcified, rather brittle. 




near posterior end, more than 65% of total number of setigers. 

First branchiae single filaments; maximum 14 filaments from 

about setiger 35. First 5-6 pairs shorter than notopodial cirri; 

all other branchiae, including last ones, clearly longer than 

notopodial cirri. Stems slender, tapering, longer than notopodial 

cirri. Filaments slender, nearly filiform, longer than 

notopodial cirri. Branchiae increasingly conspicuous towards 


Neuropodial acicular lobes distally truncately rounded, 

becoming slightly conical near posterior end; aciculae emerging 

at midline. Presetal lobes low, transverse folds. Postsetal 

lobes higher than acicular lobes and rounded in first 20 setigers, 

becoming low, transverse folds by setiger 25. First 9 ventral 

cirri tapering. Ventral cirri becoming modestly basally inflated 

from about setiger 10. Inflated bases ovate; narrow tips 

tapering. Inflated bases decreasing in importance posteriorly. 

From about setiger 100 ventral cirri tapering. Anterior 

notopodial cirri slender, basally distinctly inflated, with 3-4 

articulations. Articulations are lost by first branchial segments; 

basally inflations and slender, tapering tips retained in all 

setigers. 


(Figure 44e) tapering, furled. One marginal tooth longer than 

all other teeth, with -16 teeth. Shafts of compound falcigers 

(Figure 44b) very gently inflated, or tapering without inflated 

region, marginally smooth; internal striations present; beaks 

distinct. Appendages rather large, tapering, with large, distinct 

heads and distinct basal bosses, bidentate. Teeth similar in size, 

tapering. Proximal teeth directed slightly distally. Distal teeth 

nearly erect. Guards symmetrically bluntly pointed, rarely with 

very short mucros, marginally frayed. Pseudocompound falcigers 

and compound spinigers absent. Aciculae mostly single, 

rarely paired in posterior setigers, brown, deepening to dark 

brown in posterior setigers, tapering, distally straight, blunt; 

cross-section round. Subacicular hooks (Figure 44d) light to 

medium brown, bidentate. Hooks first present from setiger 41, 


replacements). Hooks tapering to small heads. Teeth small. 


teeth directed obliquely distally. 


anal cirri. 


None. 





None. 

REMARKS.—Eunice fauveli was originally erroneously described 

as having unidentate subacicular hooks; the hooks are 

distinctly, if shallowly bidentate; the species belongs to group 

B-4 rather than to D-2 as assumed by Fauchald (1970). 

Eunice fauveli is listed with similar species in Tables 33 and 

35. It is the only species in Table 35 other than E. collaris with 

more than six branchial filaments. It has sharply pointed guards 

in the compound falcigers rather than blunt guards as present in 

E. collaris. 

Eunice fijiensis Baird, 1869 

Eunice fijiensis Baird, 1869:347. 

Eunice antenna ta.—Grube, 1878a:99. 

REMARKS.—No material labeled as E. fijiensis is currently 

present in the British Museum (Natural History) (now the 

Natural History Museum, London) (Alex Muir, in litt.). The 

species was briefly characterized by Baird as having branchiae 

from setiger 7 and a total of 98 setigers, bidentate compound 

hooks, and tridentate subacicular hooks. The aciculae were 

described as being stout, swollen in the middle of their length 

and slightly curved at the point. Furthermore, Baird stated that 

"it approaches somewhat to the Eunice gracilis of Grube, from 

Tahiti." This statement is presumably the reason why Grube 

listed E. fijiensis as a possible synonym of E. antennata. The 

species is clearly insufficiently characterized and is here 


63. Eunice ftlamentosa Grube, 1856 

Eunice fdamentosa Grube, 1856:56. 

FIGURE 45a-g; TABLES 33,37

NUMBER 523 

139 

.0 2 

1 

2 

1 

.1 

.5 

1 

h r, ri 

a 

h 

a 

f 

FIGURE 45.—Eunice fdamentosa (lectotype, ZMC): a, anterior end, lateral view; b, subacicular hook, parapodium 

29; c, pectinate seta, parapodium 29; d, compound falciger, parapodium 29; e, parapodium 29, anterior view; f, 

aciculae, parapodium 29. Eunice fdamentosa (paralectotype, ZMB 1855): g, anterior end, lateral view. Eunice 

fvnbriata (paratype, ZMB Q2262): h, anterior end, lateral view; i, parapodium 33, anterior view; j, compound 

falciger, parapodium 33; k, subacicular hook, parapodium 33; 1, acicula, parapodium 33; m, appendage of 

compound falciger, parapodium 33; n, pectinate seta, parapodium 33. (Scale bars in mm.)


MATERIAL EXAMINED.—Lectotype, ZMC (uncatalogcd), 

and paralectotype, ZMB 1855, St. Croix, Virgin Islands, coll. 

KrOycr. 

COMMENTS ON MATERIAL EXAMINED.—The lectotype is 

very soft and nearly translucent. Because of the condition of the 

specimen, some parapodial features were not clearly visible. 

The paralectotype from Berlin is dry and has been dry for some 

time; it is nevertheless in recognizable condition. 

DESCRIPTION.—Lectotype complete, of unknown sex, with 

119 setigers; length 25 mm; maximal width 1.5 mm at about 

setiger 10; length through setiger 10, 3 mm. Anterior part of 

body is strongly inflated with highly convex dorsum and 

flattened ventrum; median and posterior part of body is 

flattened and evenly wide from about setiger 20 to near 

abruptly tapered posterior end. 

Prostomium (Figure 45a,g) distinctly shorter and narrower 

than peristomium, less than l /2 as deep as pcristomium. 

Prostomial lobes frontally obliquely truncate, tapering, dorsally 

flattened; median sulcus deep. Eyes between bases of A-I and 

A-II. Antennae in horseshoe; A-I and A-I I emerging close 

together near sides of prostomial lobes, well separated from 

bases of A-III; all similar in thickness. Ccratophorcs ringshaped 


digitiform, without articulations. A-I to middle of anterior 

peristomial ring; A-II and A-III to setiger 1. Peristomium 

tapering anteriorly. Separation between rings distinct dorsally 

and ventrally; anterior ring 1 I% of total peristomial length. 

Peristomial cirri to middle of anterior peristomial ring, slender 

and digitiform, without articulations. 




23 (27 in paralectotype) to setiger 119. Branchiae present to 


of setigers. Most branchiae single, flattened filaments (Figure 

45e); some posterior branchiae with 2 filaments. Filaments 

increasing in length posteriorly, being nearly as long as body is 

wide in several far posterior setigers, and distinctly longer than 


Anterior neuropodial acicular lobes rounded; posterior ones 

conical; aciculae emerging above midline. Pre- and postsetal 

lobes are low folds. First 4 ventral cirri thick, tapering. Ventral 

cirri basally inflated from about setiger 5. Inflated bases thick 

ventral ridges in setigers 15-50; narrow tips short and 

button-shaped. Inflated bases rapidly reduced in setigers 

50-60. Ventral cirri digitiform in posterior setigers. Notopodial 

cirri slender and tapering, probably originally basally 

inflated in anterior setigers, definitely becoming more slender 

in posterior setigers, articulations absent. 


(Figure 45c) with very long, thick, cylindrical shafts. Blades 

distinctly furled; tapering. One marginal tooth longer than other 


45d) very long, strongly inflated, marginally serrated; distal 

beaks distinct. Appendages very short with nearly parallel 

sides, bidentate. Proximal teeth much larger than distal teeth, 

triangular, directed basally. Distal teeth very short, strongly 

bent, tapering, directed obliquely distally. Guards symmetrically 

rounded, marginally serrated; mucros absent. Pseudocompound 


(Figure 450 paired, amber, slender, distally symmetrically 

hammer-headed; cross-section round. Subacicular hooks (Figure 

45b) amber, bidentate. Hooks first present from setiger 17 

(22 in paralectotype), present in all setigers thereafter, always 

single (except for replacements). Hooks strongly beaked with 

inflated region below distinct heads. Proximal teeth larger than 

distal teeth, strongly curved. Distal teeth bent or curved, 

directed laterally. Guards truncate. 







Unknown Characters: 39-40, 42, 74, 78. 


None. 

REMARKS.—A second specimen present in the vial from 

ZMC is an unidentifiable member of the genus Palola. The 

described specimen from ZMC is here named as lectotype and 

the dried specimen from ZMB as paralectotype. Eunice 

filamentosa has been reported widely from the western Atlantic 

Ocean; some of the many records may belong to related 

species; these species are listed in Tables 33 and 37. It is the 

only species in Table 37 with hammer-headed aciculae. 

64. Eunice fimbriata Grube, 1870 

Eunice fimbriataGrube, 1870b:55-56. 

FIGURE 45h-n; TABLES 27,30 

MATERIAL EXAMINED.—Holotype, ZMH V-793; paratype 

ZMB Q2262, Viti Islands (Fiji) Mus. Godeffroy. 


originally described as consisting of 322 segments and being 

230 mm long. The Hamburg specimen is complete with 362 

setigers and is 230 mm long and is here considered the 

holotype. The Berlin specimen is incomplete and consists of 72 

setigers. 

DESCRIPTION.—Holotype of unknown sex, with 362 setigers, 

total length 230 mm; maximal width 7 mm at setiger 30; 

length through setiger 10, 11 mm; width at setiger 10, 6 mm. 

Paratype incomplete with 72 setigers; length 42 mm; maximal 

width 7 mm around setiger 30; length through setiger 10, 10 

mm. Body cylindrical, dorsally covered with rugosities, mostly 

as short, longitudinal folds. 

Prostomium (Figure 45h) distinctly shorter than peristomium, 

nearly as wide as peristomium, as deep as x li of



excavate with thickened rim; median sulcus shallow. Antennae 

in transverse row, evenly spaced, similar in thickness, 

emerging from shallow depressions on prostomial face. 


Ceratostyles short, digitiform, with 5 distinct articulations. All 

antennae to front edge of setiger 1. Peristomium cylindrical, 


distinct on all sides; anterior ring ~ A /s of total peristomial 


digitiform, with 3 articulations. 

Maxillary formula of paratype 1+1, 444, 7+0, 5+8, and 1+1. 



Branchiae from sctigcr 9 to setiger 122, thus absent on posterior 

half of body, present on less than 55% of total number of 

setigers. All branchiae except last 10-12 pairs with 2 or more 

filaments, maximum 5 filaments at about setiger 15. Branchial 

stems pliable, shorter than notopodial cirri. Filaments digitiform, 

much longer than notopodial cirri. Number of filaments 

reduced to 3 from about setiger 30, retaining same length and 

shape in all remaining branchial setigers. 

Neuropodial acicular lobes asymmetrically rounded with 

high point above midline; aciculac emerging above midline. 

Presetal lobes low, transverse folds. Anterior postsetal lobes 

truncate folds, becoming symmetrically rounded in median and 

posterior setigers, outreaching acicular lobes. First 4 to 5 

ventral cirri thick, tapering. Ventral cirri basally inflated in 

branchial region. Inflated bases transverse welts; narrow tips 

short and button-shaped. Postbranchial ventral cirri short, 

nearly tubercular, nearly invisible in last 100 setigers. Anterior 

notopodial cirri thick, digitiform, with 3 indistinct articulations. 

Articulations lost in branchial region. Notopodia decreasing in 

size towards posterior end of body, retaining similar shape. 

Limbate setae slender, nearly straight; limbation marginally 

fraying in most setae. Pectinate setae (Figure 45n) short, with 

slender, cylindrical shafts. Blades flaring, flat. Both marginal 

teeth longer than other teeth, with -15 teeth. Shafts of 

compound falcigers (Figure 45j) long, slender, tapering, 

marginally smooth; beaks indistinct. Appendages (Figure 

45j,m) short; heads distinct, bidentate. Proximal and distal teeth 

similar in length. Proximal teeth slender, tapering, directed 

obliquely distally, upturned near tips. Distal teeth curved, 

directed obliquely distally, nearly claw-like. Guards symmetrically 

rounded, marginally finely serrated; mucros absent. 


Aciculae (Figure 451) single, black, distally sharply bent 

ventrally; cross-section round. Separation between core and 

sheath distinct in both aciculae and subacicular hooks. 

Subacicular hooks (Figure 45k) black, bidentate. Hooks first 


single (except for replacements). Hooks tapering towards large 

heads. Proximal teeth larger than distal teeth; both teeth 

directed distally. 









None. 

REMARKS.—Eunice fimbriata has remained unreported 

since its brief mention by Grube (1870b); it belongs, by 

definition, to group B-2 and is listed with similar species in 

Tables 27 and 30. In E. fimbriata, E. coccinea, and E. reducta, 

the branchiae terminate well before the posterior end, in 

contrast to all other species listed in Table 30. Eunice reducta 

has -21 branchial filaments, E. coccinea has 10, and E. 

fimbriata has five. In E. coccinea the bases of the ventral cirri 

are inflated to the posterior end; in the other two species, they 

are inflated only in the median body region. 

65. Eunice flaccida Grube, 1869 

FIGURE 46a-e; TABLES 41,44,46,49, 52,53 

Eunice flaccida Grube, 1869:491-492. 

MATERIAL EXAMINED.—Holotype, ZMB 496, Tor, Red Sea, 

coll. Ehrenberg, Sep year. 

COMMENTS ON MATERIAL EXAMINED.—The pectinate setae 

were somewhat difficult to study due to the amount of debris 

attached. 


43 setigers; length 27 mm; maximal width 2.5 mm at setiger 

10; length through setiger 10, 8.5 mm. Anterior part of body 

through setiger 15 cylindrical; remainder dorsoventrally flattened. 




shallow. Eyes not observed. Antennae in a horseshoe with A-I 

anterior to A-II and III, evenly spaced, similar in thickness. 


Ceratostyles tapering, articulated to bases, with up to 18 short, 

moniliform articulations in A-III. A-I to bases of peristomial 

cirri; A-II to setiger 1; A-III to setiger 2. Peristomium 

cylindrical with strongly jutting, muscular lower lip. Separation 

between rings distinct only ventrally; anterior ring 5 /6 of total 


peristomial ring, digitiform, with 6 articulations. 



than notopodial cirri, erect. Branchiae from setiger 7 to end of 

fragment. All branchiae pectinate; maximum 9 filaments at 

setiger 12; numbers of filaments decreasing posteriorly, at end 

of fragment 2 or 3 filaments. Branchial stems stout, cylindrical,

142 


.c. d. g. h, i 

. e 

_b 

FIGURE 46.—Eunice JIaccida (holotype, ZMB 496): a, anterior end, lateral view; b, parapodium 35, anterior view; 

c, subacicular hook, parapodium 35; d, compound fakigcr, parapodium 35; e, aciculae, parapodium 35. Eunice 

flavapunctata (holotype, AMNH 1532): f, anterior end, lateral view; g, subacicular hook, parapodium 29; h, 

pectinate seta, parapodium 29; i, compound falciger, parapodium 29; j, parapodium 29, anterior view. (Scale bars 

in mm.)


shorter than filaments. Filaments digitiform, shorter than 



becoming asymmetrically rounded with aciculae emerging 

above midline. Prcsetal lobes low, somewhat asymmetrical 

folds with low side ventrally. Postsetal lobe low asymmetrical 

folds with low side dorsally. Ventral cirri thick and rounded in 

all setigers, developing long digitiform tips in posterior setigers 

present, possibly distorted in fixation. Anterior notopodial cirri 

short, inflated, becoming longer posteriorly, but retaining 

distinct inflated basal section, without articulations. 

Limbate setae slender, marginally smooth. About 5 pectinate 

setae in a parapodium. Pectinate setae tapering, flat. Marginal 

teeth about as long as other teeth, with 15 teeth. Shafts of 

compound falcigcrs (Figure 46d) gently inflated, marginally 

smooth; beaks indistinct. Appendages large; heads large, 

bidentate. Teeth similar in size. Proximal teeth narrowly 

triangular, directed laterally. Distal teeth tapering, directed 

obliquely distally. Guards asymmetrically bluntly pointed, 

marginally serrated; mucros absent. Pseudocompound falcigers 

and compound spinigers absent. Aciculae (Figure 46e) paired, 

yellow; exposed portion twisted in double curve, sharp tipped; 

distal end when worn slightly knobbed; cross-section round. 


and subacicular hooks. Subacicular hooks (Figure 46c) yellow, 

tridentate with teeth in tandem. Hooks first present from setiger 


replacements). Shafts strongly curved distally, tapering. Two 

lower fangs, including largest and smallest arranged on either 

side of middle fang, which clearly outreach other fangs. 


characters associated with posterior setigers, including branchial 

distribution; pygidium and anal cirri. 





Characters: 1, 2, 13, 14, 33, 36-38,40,47, 50, 57-60,63. 


None. 

REMARKS.—Eunice flaccida was considered a synonym of 

E. antennata by Crossland (1904:312). It may be a member of 

either group C-l or C-2, because the type is posteriorly 

incomplete; thus it has been included in Tables 41 and 44 as 

well as in Tables 46 and 49. It has tridentate hooks with the 

teeth in tandem as does E. elseyi in addition to two species with 

dark subacicular hooks; these are compared in Tables 52 and 

53. It differs from E. elseyi in that it has distally entire aciculae 

and bidentate compound falcigers; in E. elseyi the aciculae are 

bidentate and the compound hooks distally tridentate. 

66. Eunice flavapunctata (Treadwell, 1922) 


Leodice flavapunctata Treadwell, 1922:136-138, figs. 8-11, pi. 2: fig.1-7. 

Eunice qfra.—Hartman, 1956:282 [not Eunice afra Peters, 1854]. 

MATERIAL EXAMINED.—Holotype, AMNH 1532, Breaker 

Point, Pago Pago, Samoa, 1920. 


has been deeply dissected frontally and the illustration of the 

anterior end is a reconstruction (all antennae were present 

however). 




Prostomium (Figure 460 about as long as peristomium, 

about as wide as peristomium, as deep as peristomium. 


sulcus deep. Eyes not observed. Antennae in transverse row; 

A-I separated by gap from A-II and A-III, similar in thickness. 


Ceratostyles digitiform, irregularly wrinkled, without articulations. 

A-II and A-III similar in length, barely outreach 

peristomium. Peristomium cylindrical. Separation between 

rings distinct on all sides; anterior ring ~ 2 h of total peristomial 

length. Peristomial cirri barely reaching beyond posterior 

peristomial ring, digitiform, without articulations. 

Jaws now missing. 



from setiger 13 to end of fragment. Maximum 6 filaments, 

reached at about setiger 50. Most segments with 2 or 3 long, 

digitiform filaments (Figure 46j). Branchial stems cylindrical, 

shorter than notopodial cirri. Branchiae located well above 

neuropodial cirri on body wall. 

Neuropodial acicular lobes truncate; aciculae emerging 

above midline. Pre- and postsetal lobes low folds. First 4 

ventral cirri tapering, nearly digitiform. Ventral cirri basally 

inflated from about setiger 5; bases inflated in all remaining 

parapodia. Inflated bases thick, transverse welts; narrow tips 

tapering. Anterior notopodial cirri relatively short, digitiform, 

becoming increasingly slender and longer in branchial region; 

articulations absent. 

Limbate setae long, narrowly limbate, marginally dentate. 

Shafts of pectinate setae (Figure 46h) wide, cylindrical. Blades 

narrow, tapering, flat. One marginal tooth very much longer 

than other teeth, with 10 narrow, slender teeth. Shafts of 

compound falcigers (Figure 46i) inflated, marginally serrated; 

beaks distinct. Appendages short; heads large, bidentate. 

Proximal teeth larger than distal teeth, narrowly triangular, 

directed obliquely distally. Distal teeth abruptly tapering, 

directed obliquely distally. Guards symmetrically rounded, 


and compound spinigers absent. Aciculae black, tapering to 

narrow tips, straight; cross-sections round. Subacicular hooks 

(Figure 46g) black, bidentate. Hooks first present from setiger 


replacements). Hooks tapering, curved. Proximal teeth very 

much larger than distal teeth, directed laterally. Distal teeth 

blunt, directed nearly laterally. 

UNKNOWN 

MORPHOLOGICAL FEATURES.—Jaw structure;

144 


features associated with posterior setigcrs, including branchial 


EXPECTED STATES OF SELECTED UNKNOWN FEATURES — 

Mx III short and forming part of distal arc with left Mx IV. 

Branchiae terminating well before the posterior end. 



Characters: 1, 2, 13, 14, 36-40,47, 50, 57-60, 63, 74, 78. 


38,1. 

REMARKS.—Eunice flavapunctata was considered a synonym 

of E. afra by Hartman (1956). It is listed with similar 

species in Tables 33 and 39. Of the species in Table 39 only E. 

denticulata and E. flavapunctata have the inflated bases of the 

median ventral cirri forming thick transverse welts. The former 

has hammer-headed aciculae; the latter has pointed aciculac. 

Treadwell stated that branchiae were missing on the last 100 

segments; this statement cannot be confirmed because the 

holotype is incomplete; Treadwell may have had another 

fragment of the holotype or additional material at his disposal. 

67. Eunice flavocuprea Grube, 1869 

FIGURE 47a-c; TABLES 41-42 

Eunice flavocupreaGrube, 1869:493-494. 

MATERIAL EXAMINED.—Holotype, ZMB 498, Tor, Red Sea, 

coll. Ehrenberg. 

COMMENTS ON MATERIAL EXAMINED.—The holotype is in 

rather poor condition; it appears to have been in posterior 

regeneration. 


length 17 mm; maximal width 1 mm; length through setiger 10 

about 2 mm. 




median sulcus shallow. Eyes not observed. Antennae in 

shallow horseshoe, evenly spaced, similar in thickness. 


Ceratostyles digitiform, with 2 or 3 long, cylindrical articulations. 

A-I to setiger 1; A-II to setiger 3; A-III missing. 


all sides; anterior ring ~2fr of total peristomial length. 

Peristomial cirri very short, nearly ovoid, without articulations. 




from setiger 4 to setiger 34. Branchiae terminating well before 

posterior end. First 5 and last 10 branchiae single filaments; 

maximum 4 filaments at about setiger 15. All branchiae 

outreaching notopodial cirri, even in first branchiated segment. 

Branchial stems reduced. Filaments very long, slender. 

Shape of neuropodia indeterminable in detail without 

destroying specimen. Ventral cirri basally inflated from about 

setiger 5 through setiger 40. Inflated bases ovate; narrow tips 

short and button-shaped. Notopodial cirri distinctly medially 

inflated anteriorly, becoming digitiform in posterior setigcrs, 

articulations absent. 

Limbatc setae marginally serrated. Pectinate setae not 

observed in detail. Shafts of compound falcigers distinctly 

inflated, marginally smooth. Appendages (Figure 47a) very 

long and narrow, bidentate. Proximal teeth very short, 

triangular projections, directed basally. Distal teeth longer than 

proximal teeth, slender, curved. Guards symmetrically 

rounded; some guards with distinct, narrow mucros. Pscudocompound 


tapering, yellow. Separation between core and sheath 

indistinct in both aciculac and subacicular hooks. Subacicular 

hooks (Figure 47b) yellow, tridentatc. Hooks first present from 

setiger 17, present in all setigcrs thereafter, sometimes paired. 

Main fangs large, curved; 2 distal fangs in crests; very small 

tertiary fangs closely appended to secondary fangs. 


most parapodial features; pygidium and anal cirri. 





1,4,6, 13, 14, 38,43-50, 52, 57-60, 65-68. 


REMARKS.—In the vial, a small handwritten label reads as 

follows: "Eunice flavocuprea Gr. (nomen nudum) Nach v. 

Marenzeller Brief = E. murrayi M'Int." Eunice flavocuprea 

belongs to the same group as E. murrayi but differs sharply 

from that species in the structure of the antennae, which are 

nearly smooth in the former and strongly articulated in the 

latter. Grube's description, while brief, is certainly adequate to 

avoid labeling the name as a nomen nudum. Eunice flavocuprea 

is listed with similar species in Tables 41 and 42. It can be 

identified only by a combination of character states, rather than 

by a single, unique character state. 

Eunice flavofasciata Grube, 1878 

Eunice flavo-fasciataGrube, 1878b: 155-156, pi. 9: fig. 2. 

REMARKS.—Grube had a single complete specimen, which 

now has been lost. The following is a translation of the 

description supplemented with interpretations of Grube's 

illustrations. 

Specimen chestnut-colored with lighter anterior transverse 

bands and cirri. Antennae without articulations. A-III distinctly 

longer than other antennae, reaching roughly setiger 5. 

Peristomial cirri reaching posterior part of prostomium. The 

maxillary formula is 1+1, 3+3, 6+0, 2+7, and 1+1. Branchiae 

present from setiger 5 to far posterior end, where best 

developed, from about setiger 5 through setiger 24, with up to

NUMBER 523 

145 

FIGURE 47.—Eunice flavocuprea (holotype, ZMB 498): a, appendage of compound falciger, median parapodium; 

b, subacicular hook, median parapodium; c, anterior end, lateral view. Eunice floridana (complete specimen from 

MCZ 674): d, anterior end, lateral view; e, parapodium 21, anterior view; f, parapodium 2, anterior view; g, 

compound falciger, parapodium 2; h, compound falciger, parapodium 21; i, pectinate seta, parapodium 21; j, 

parapodium 60, anterior view; k, pectinate seta, parapodium 60; 1, compound falciger, parapodium 60; m, 

subacicular hooks, parapodium 60. (Scale bars in mm.)


14 filaments. Notopodial cirri articulated; ventral cirri inflated 

from about setiger 3 through setiger 28. Aciculae and 

subacicular hooks black. Subacicular hooks bidentate. 

Grube clearly had a specimen belonging to group B-2; it is 

well described, but without access to the types, it is difficult to 

differentiate it from a series of similar species. The species is 

here considered indeterminable. 

68. Eunice flavopicta Izuka, 1912 

TABLES 27,29,30 

Eunice flavopicta Izuka, 1912:121-123, pi. 14: figs. 1-5. 

REMARKS.—The type material of this species has been lost, 

according to Miura (1986:269), who considered the species 

synonymous with E. aphroditois. Considering the complex 

synonymy of the latter species it appears best to keep the two 

apart, until the variability of E. aphroditois has been 

established. The following represents a summary of the 

information in the original description. 

Body with 166 segments; length 215 mm; maximal width 6 

mm at setiger 10. Body tapering relatively abruptly anteriorly 

and slowly posteriorly. Anal cirri rather long. Color lightly 

iridescent deep purple, with 3 bright yellow spots on dorsum of 

each segment as transverse band on setiger 4. 

Palpal region set off by frontal horizontal grooves. Eyes at 

bases of A-I. Antennae faintly articulated. A-I to posterior end 

of posterior peristomial ring; A-II to setiger 1; A-III to setiger 

3. Peristomial rings distinct at least dorsally. Peristomial cirri 

short, presumably not outreaching peristomium. 

Maxillary formula 1+1,4+5, 6+0, 4+9, 1+1, and 1+1. 

Branchiae from setiger 5 to end of body. All() branchiae 

pectinate; maximum 18 or 19 filaments at setiger 14, this 

number maintained to about setiger 50, decreasing slowly 

posteriorly. Branchiae longer than notopodial cirri. Stems erect, 

tapering, longer than notopodial cirri. Filaments digitiform, 

shorter than notopodial cirri. 

Ventral cirri basally inflated. Notopodial cirri basally 

inflated, vaguely articulated, tapering to blunt tips. 


tapering, flat. Marginal teeth no longer than other teeth, with 9 

teeth. Shafts of compound falcigers tapering, marginally finely 

serrated, with distinct internal striations. Appendages broadbased, 

triangular, short, bidentate. Proximal teeth triangular, 

directed laterally. Distal teeth slender, erect; 6 small teeth 

present between proximal and distal teeth. Guards bluntly 

pointed, marginally serrated. Aciculae and presumably subacicular 

hooks dark. Subacicular hooks tapering, bidentate; 

heads bent. Proximal teeth larger than distal teeth, directed 

laterally. Distal teeth short, curved. 

Eunice flavopicta is compared to similar species in Tables 

27, 29, and 30. The presence of small denticles between the 

proximal and distal teeth of the compound falciger is unusual 

and would need to be confirmed on fresh material. 

69. Eunice floridana (Pourtales, 1867) 

FIGURE 47d-m; TABLES 5, 27.32 

Marphysa floridana Pourtales, 1867:108. 

Eunice floridana— Khlcrs, 1887:88-90, pi. 22: figs. 1-7. 

MATERIAL EXAMINED.—MCZ 674, off the Sambos, 229 m, 

8 specimens. 


based on one of the complete specimens; the information about 

the other specimens is summarized in Table 5. 

DESCRIPTION.—Specimen illustrated complete, of unknown 

sex, with 111 setigers; length 80 mm; maximal width 6 mm at 

setiger 10; length through setiger 10, 10.5 mm. Anterior 

dorsum strongly convex; anterior ventrum slightly flattened; 

further posteriorly body dorsovcntrally flattened. Long anal 

cirri tapering, as long as last 5 setigers together, without 



than peristomium, less than '/2 as deep as peristomium. 

Prostomial lobes frontally obliquely truncate, slightly dorsally 

flattened; median sulcus shallow. Eyes between bases of A-I 

and A-II, large purple. Antennae in shallow horseshoe; A-I 

separated by gap from A-II and A-III, similar in thickness. 

Ccratophores ring-shaped in all antennae, without articulations. 

Ceratostyles slender and nearly digitiform, tapering only 

slightly distally, with up to 7 long, irregular, cylindrical 

articulations. Articulations distally drop-shaped in A-I. A-I to 


Other specimens with A-II to setiger 6-7 and A-III to setigers 

12-13. Peristomium short, flaring anteriorly; lower lip muscular. 


anterior ring A /s of total peristomial length. Peristomial cirri at 

least to middle of prostomium, slender, tapering, with 4-5 


Maxillary formula 1+1,6+6, 7-9+0, 6+8, 1+1, and 1+1. Mx 

III part of distal arc with left Mx IV. 

Branchiae (Figure 47e, j) present, pectinate, distinctly longer 

than notopodial cirri, reduced in mid-body region, erect. 

Branchiae from setigers 9 (8-10) to 111. Branchiae present to 


of setigers. All branchiae pectinate, except in 2 cases first 

branchiae. Maximum 8 filaments. Branchiae reduced to 3-4 

filaments in median region, increasing to 7-8 filaments in 

penultimate setigers. Branchial stems thick, tapering, shorter 

than notopodial cirri. Filaments digitiform, shorter than 


Anterior neuropodial acicular lobes (Figure 470 symmetrically 

rounded, becoming obliquely truncate with aciculae 

emerging near superior, higher end. Presetal lobes low, 

transverse folds. Postsetal lobes symmetrically rounded free 

lobes through about setiger 60-75, thereafter reduced to low 

folds following outline of acicular lobes closely. Anterior 

ventral cirri thick and tapering, becoming basally strongly


TABLE 5.—Variable and invariable features in the type lot of Eunice floridana (N = number of individuals 



No. of seligers 

Total length 

Maximal width 


No. of antcnnal articulations 

Branchiae first present from setiger no. 

Ventral cirri inflated to setiger no. 



N 

4 

4 

8 

8 

8 

8 

8 

8 

8 

Max. 

138 

95 

6 

10.5 

8 

10 

55 

4 

40 

Min. 

100 

53 

4.5 

7 

5 

8 

40 

3 

29 

Mean 

117.5 

70.75 

5.31 

8.81 

7.38 

9.00 

47.50 

3.13 

33.25 

SD 

16.13 

20.30 

0.59 

1.22 

1.19 

0.93 

5.35 

0.35 

3.62 


Separation of pcristomial rings 

Pcristomial cirri reach 




Shaft of compound falcigcrs 

Color of aciculac 

Shaft of aciculae 



Core-sheath separation 

N=8 

visible dorsally and ventrally 


8 

furled, flared 

12 

inflated 

black 

flattened and knife-like 

black 

2 

distinct 

inflated from about setiger 8-10. Inflated bases thick, ventral 

ridges along lower edge of parapodia; narrow tips tapering. 

Inflated bases reduced posterior to setiger 30-40, missing from 

setiger 40-50. Posterior ventral cirri nearly triangular in 

outline, tapering, broadly attached ventrally, nearly scoopshaped. 

Anterior notopodial cirri digitiform, with 3 to 5 

indistinct articulations. Median and posterior notopodial cirri 

basally slightly inflated, tapering to thick, digitiform tips, 


Limbate setae frayed. Shafts of pectinate setae (Figure 47i,k) 

slender, cylindrical. Blades slightly flaring, gently furled. One 

marginal tooth distinctly longer than all other teeth; 15-18 

teeth present. Shafts of all compound falcigers (Figure 47g,h,l) 

slightly inflated, marginally serrated in anterior and median 

setigers and marginally smooth further posteriorly; beaks 

distinct. Anterior appendages (Figure 47g) slender, tapering; 

heads small, bidentate. Proximal teeth smaller than distal teeth, 

triangular, directed laterally. Distal teeth nearly erect, tapering. 

Anterior guards asymmetrically sharply pointed; mucros 

absent. Posterior appendages (Figure 471) short, thick, distinctly 

triangular in outline; heads distinct, bidentate. Proximal 

teeth larger than distal teeth, narrowly triangular, directed 

laterally. Distal teeth short, directed obliquely distally. Posterior 

guards symmetrically sharply pointed; mucros absent. 


Anterior aciculae dark brown, tapering to straight tips; 

cross-section round. Posterior aciculae black, extremely shiny, 

flattened anteroposteriorly to form slightly curved knife-shaped 

blades (Figure 47j). Median and posterior aciculae very thick, 

numbering at least 3 in a parapodium. Subacicular hooks 

(Figure 47m) first dark brown, becoming black and shiny in 

later setigers, bidentate. Hooks first present from setiger 

29-40, present in all setigers thereafter, usually paired. Hooks 

tapering abruptly to narrow necks. Proximal teeth larger than 

distal teeth, narrowly triangular, directed laterally. Distal teeth 

very small, triangular, erect 

UNKNOWN MORPHOLOGICAL FEATURES.—None. 


None. 



Characters: 42, 74, 78. 


None. 

REMARKS.—The original description mentioned only the 

thick, pergamentaceous tubes in which the species lives and its 

association with deep-water hard-bottom environments. The 

first complete description of the species was given by Ehlers 

(1887) and the description given above is based on material 

used by Ehlers in his description; thus, the material, although 

not type material, is certainly authoritative in terms of the 

definition of the species. 

The species has been widely reported but appears to have 

been confused with a number of other members of group B-2, 

with which it is listed in Tables 27 and 32. 

Eunice floridana has articulated ceratostyles in which the 

articulations become increasingly moniliform distally. In 

addition to E. floridana, the following species in Table 32 have 

the notopodial cirri articulated only in the anterior end: E. 

frauenfeldi, E. grubei, E. northioidea, and E. pyenobranchiata.


In E. floridana the peristomial cirri reach the middle of the 

prostomium; in the other species, the peristomial cirri do not 

outreach the peristomium. 

70. Eunice franklini Monro, 1924 

FIGURE 48a-e; TABLES 27,31 

Eunice franklini Monro, 1924:56-57, figs. 14-16.—Fauchald, 1986:250-251. 

figs. 41-45. 


1924.1.28.81, the Franklin Shoal, Arafura Sea, 9°53'S, 

129°19'E. 


92 setigers; length 105 mm; maximal width 7 mm at sctigcr 10; 





sulcus shallow. Antennae in transverse row, evenly spaced, 



irregular articulations in A-III. A-I to setiger 1; A-II to setiger 

4 (only left present); A-III to setiger 7. Peristomium cylindrical. 

Separation between rings distinct dorsally, barely visible 


cirri to posterior x h of prostomium, slender and tapering, 

with 5 indistinct articulations. 

Maxillary formula 1+1, 6+7, 8+0, 6+9, and presumably 1+1 

according to Monro (1924, fig. 16). 


than notopodial cirri, not reduced in mid-body region (not 

apparent in illustration), erect. Branchiae from setiger 6 to end 

of fragment. First branchia with 5 filaments, maximum about 

25 filaments. Branchial stems long, erect, shorter than 

notopodial cirri. Filaments short, digitiform. 

Neuropodial acicular lobes flattened conical; aciculae 

emerging at midline. Anterior pre- and postsetal lobes distinct, 

distally rounded in first 15 setigers, thereafter reduced to low 

transverse folds. Anterior ventral cirri tapering, blunt. Ventral 

cirri basally inflated in median setigers. Inflated bases 

indistinct, ovate; narrow tips tapering. Notopodial cirri supported 

by dark aciculae, very long, digitiform, without 


Shafts of pectinate setae (Figure 48b) wide, cylindrical. 

Blades flat, slightly flaring. Both marginal teeth longer than 

other teeth, with -12 teeth. Shafts of compound falcigers 

(Figure 48d) slender, smoothly tapering, marginally smooth, 

without beaks or internal striations. Appendages slender; heads 

distinct, bidentate. Proximal and distal teeth similar in length; 

proximal teeth narrowly triangular, directed laterally. Distal 

teeth tapering, slimmer than proximal teeth, directed obliquely 

distally. Guards symmetrically bluntly rounded, marginally 

smooth; mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae, including 2 of 4 notopodial 

aciculae, black, tapering, straight, pointed; cross-section round. 

Subacicular hooks black (Figure 48c), bidentate. Hooks first 

present from setiger 34; present in all setigers thereafter, always 

single (except for replacements). Hooks slender, gently curved. 

Both teeth similar in size. Proximal teeth triangular, directed 

obliquely distally. Distal teeth slightly narrower than proximal 

teeth, nearly erect. 


Mx III and left Mx IV; all features associated with far 






SCORED.—Inappropriate characters: 56. Unknown Characters: 

1, 2, 13, 14, 36-38,40,42,47, 50, 51, 57-60. 74, 78. 


38,1. 

REMARKS.—Eunice franklini is listed with similar species in 

Tables 27 and 31; justification for including the species in 

group B-2 was given by Fauchald (1986). 

The combination of very short, numerous branchial filaments 

and extremely long notopodial cirri is unique to the 

genus. In addition E. franklini has compound falcigers with 

mucronate guards, a feature shared only with E. amphiheliae of 

the species listed in Table 31. The two species are readily 

separated on the relatively length of the branchiae; despite their 

length, these are outreached by the notopodial cirri in E. 

franklini and are distinctly longer than the notopodial cirri in E. 

amphiheliae. 

71. Eunice frauenfeldi Grube, 1866 

FIGURE 48f-k; TABLES 27,32 

Eunice frauenfeldiGrube, 1866c:175; 1867:11-12, pi. 1: fig.3. 

MATERIAL EXAMINED.—Holotype, ZMB 4008, St. Paul's 

Island, South Atlantic. 



length through setiger 10, 7 mm. Body dorsally inflated and 

ventrally flattened, with parallel sides and short parapodia. 

Prostomium (Figure 480 distinctly shorter and narrower than 

peristomium, as deep as V2 of the peristomium. Prostomial 

lobes frontally obliquely transverse, dorsally excavate with 

thickened rim; median sulcus shallow. Eyes not seen. Antennae 

very large, in shallow horseshoe, evenly spaced, similar in 


articulations. Ceratostyles tapering, with up to 14 articulations 

in A-III; inner articulations cylindrical, becoming drop-shaped 

distally. A-I to posterior peristomial ring; A-II to setiger 1; 

A-III to setiger 2. Peristomium cylindrical, with distinct, 

muscular lower lip. Separation distinct dorsally and ventrally;


FIGURE 48.—Eunice franklini (holotype, BM(NH) ZK 1924.1.28.81): a, anterior end, lateral view; b, pectinate 

seta, parapodium 69; c, subacicularhook, parapodium 69; d, compound falciger, parapodium 69; e, parapodium 

69, anterior view. Eunice frauenfeldi (holotype, ZMB 4008): f, anterior end, lateral view; g, parapodium 63, 

anterior view, h, subacicular hook, parapodium 63; i, aciculae, parapodium 63; j, pectinate seta, parapodium 63; 

k, compound falciger, parapodium 63. (Scale bars in mm.) 

anterior ring 3 A of total peristomial length. Perislomial cirri to 

middle of anterior peristomial ring, tapering, with 4 or 5 





to end of fragment. First branchia single filaments; maximum 

3 filaments in sctigers 15-25; all other branchiae with 2 

filaments (Figure 48g). Stems reduced to short buttons. 

Filaments thick, distally blunt, posteriorly as long as notopodial 

cirri. 

Anterior neuropodial acicular lobes distally truncate; postc-


rior acicular lobes rounded; aciculae emerging at midline. 

Pre- and postsetal lobes low, transverse folds. First 4 ventral 

cirri thick, tapering. Ventral cirri basally inflated from about 

setiger 5. Inflated bases short welts where best developed, 

decreasing in size from about setigcr 45; narrow tips tapering. 

Posterior ventral cirri thick, tapering, directed posteriorly. 

Anterior notopodial cirri tapering and articulated with up to 5 

articulations, farther posteriorly cirri decreasing in length, 


Limbate setae slim, marginally smooth. Shafts of pectinate 

setae (Figure 48j) cylindrical. Blades slightly flaring, flat. One 

marginal tooth longer than other teeth; -12 teeth present. 

Compound falcigers (Figure 48k) very large, long-shafted. 

Shafts tapering, marginally smooth; distal beaks absent. 

Appendages with distinctly angled back edge, tapering, 

bidentatc. Proximal teeth longer than distal teeth, tapering, 

directed laterally. Distal teeth gently curved, nearly erect. 

Guards longer than appendages, angular, but distinctly symmetrically 

bluntly pointed; mucros absent. Pscudocompound 

falcigers and compound spinigcrs absent. Aciculae (Figure 48i) 

paired, with light brown or chestnut-colored cores and clear 

sheaths, tapering, slender, gently curved tips; tips curved 

dorsally; cross-sections round. Subacicular hooks (Figure 48h) 

with chestnut-colored cores and clear sheaths, bidentatc. Hooks 

first present from setiger 27, present in all sctigcrs thereafter, 

paired in some sctigers. Hooks slender, not distinctly larger 

than compound falcigers, tapering, with small heads. Proximal 

teeth larger than distal teeth, triangular, directed laterally. 

Distal teeth short, nearly erect. Paired bosses present on distal 

bases of proximal teeth. Bases of proximal teeth much narrower 

than shaft at that point. 

UNKNOWN MORPHOLOGICAL FEATURES.—Jaw structure; all 

features associated with posterior setigers, including branchial 






Characters: 1, 2, 13, 14, 36-38,40, 47, 50, 63. 


38,1. 

REMARKS.—Eunice frauenfeldi is assumed to belong to 

group B-2 and is listed with similar species in Tables 27 and 32; 

the reduction in size of the branchiae towards the posterior end 

of the incomplete fragment makes this assumption somewhat 

suspect. The problem cannot be resolved without access to 

more material from the type locality. The synonymy list given 

in Hartman (1964:118) must be revised: Eunice magellanica is 

here treated as a distinct species. The only other species with 

palmate branchiae listed in Table 32 is E. rubella. The two 

species differ in the proportions of the prostomium and in 

various shape-related characters best elucidated by comparing 

the illustrations of the two species. 

72. Eunice fucata Ehlers, 1887 

FIGURE 49a-d; TABLES 27,28, 50 

Eunice fucata Ehlers. 1887:91-93, pi. 25: figs. 8-20. 

Staurocephalus gregaricus Mayer, 1900:1. 

Mayeria gregaricus.—Vcrrill, 1900:650. 

MATERIAL EXAMINED.—MCZ, 3 posterior fragments from 

type lots, Tortugas, Florida. 

COMMENTS ON MATERIAL EXAMINED.—The rc-description 

is based on the original description, complemented where 

possible by observations on the original material. 

DESCRIPTION.—One specimen complete, with 250 segments; 

total length 195 mm; width ~5 mm; length through 

sctiger 10 -10 mm (based on illustration of anterior end). 

Posterior body roughly circular in cross-section with no 

indication of dorsovcntral flattening. 

Eyes present. Ccratostylcs without articulations. A-I to 

posterior pcristomial ring; A-I I and A-I 11 to setiger 3. Anterior 

per is torn ial ring 3 /" of total pcristomial length. Peristomial cirri 

to middle of prostomium, tapering, without articulations. 

Maxillary formula 1+1, 4+5, 6+0, 3 + 8, 1 + 1, and 1+1. 

Branchiae (Figure 49d) present, pectinate, distinctly longer 


Branchiae from setigcr 5, absent on last 70-80 sctigcrs. 

Branchiae terminating well before posterior end, present on 

more than 65% of total number of setigcrs. Last 30 pairs single 

filaments; maximum 12 primary filaments; primary filaments 

sometimes pectinate, especially towards posterior end, yielding 

up to 24 or more terminal filaments. Branchiae longer than 

notopodial cirri except in last 15 branchial setigers. Filaments 

digitiform, shorter than notopodial cirri. 

Neuropodial acicular lobes obliquely truncate with aciculae 

emerging at upper, high end of lobes. Pre- and postsetal lobes 

low, transverse folds. Median ventral cirri basally inflated. 

Inflated bases missing posteriorly, in fragments available, all 

ventral cirri thick, digitiform. Anterior and median notopodial 

cirri basally inflated, tapering to digitiform tips, without 

articulations. Posterior notopodial cirri short, digitiform. 

Limbate setae very narrowly limbate, marginally smooth. 

Pectinate setae (Figure 49b) very long in posterior setigers. 

Shafts cylindrical. Blades strongly flaring, furled. Marginal 

teeth no longer than other teeth, with 20 teeth. Compound 

falcigers broken in all sctigers examined. Shafts described as 

slightly inflated, marginally smooth. Appendages very short, 

triangular. Cutting edge with series of small, irregular teeth, 

rather than single proximal teeth. Distal teeth nearly erect. 

Guards asymmetrically bluntly to sharply pointed; mucros 


absent. Aciculae (Figure 49c) paired, with dark brown cores 

and clear sheaths, tapering to slender lips, gently curved or 

straight; cross-sections round. Subacicular hooks (Figure 49a) 

with light to medium brown cores and clear sheaths, unidentate. 

Hooks distally gently curved, without any traces of other teeth.


.02 

1 

1 

.1 

f. g, h 

b 

d 

a, c 

FIGURE 49.—Eunice fucala (MCZ, fragments of type lot): a, subacicular hook; b, pectinate seta; c, aciculae; d, 

Tposterior parapodium. Eunice fuscafasc'uxta (syntype, AMNH 1920-1529): e, anterior end, lateral view; f, 

pectinate seu, parapodium 60; g, compound falciger, parapodium 60; h, subacicularhook, parapodium 60. (Scale 

bars in mm.)



Mx III and left Mx IV; all features associated with 

anterior setigers, including branchial distribution; pygidium 






Characters: 4, 6-12, 14-20, 24, 25, 39, 42, 51, 52, 54-56, 

65,66, 80-82. 


None. 

REMARKS.—Eunice fucata was synonymized with E. schemacephala 

by Augener (1925:28-29) after an examination of 

the types of the latter. The latter is here considered to be 

indeterminable. The two are here considered separable. Eunice 

fucata is listed in Tables 27 and 28 and listed with other species 

with simple spine-like subacicular hooks in Table 50. Of the 

species listed in Table 28, E. sebastiani as well as this species 

has these kinds of subacicular hooks. Eunice fucata has up to 

12 branchial filaments; E. sebastiani has 40 filaments, and the 

relative lengths of the antennae are very different in the two 

species. 

73. Eunice fuscafasciata (Treadwell, 1922) 

FIGURE 49e-h; TABLES 33, 40 

Nicidionfusca-fasciata Treadwell. 1922:156-157, figs. 47-50. pi. 7: fig. 5. 

MATERIAL EXAMINED.—Syntype, AMNH 1920-1529, Pago 

Pago, Samoa. Syntype, AMNH 1920-1537, Pago-Pago, Samoa. 

COMMENTS ON MATERIAL EXAMINED.—The syntype from 

1529 is labeled Nicidionflava-fasciata; the syntype from 1537 

is labeled Nicidion fusca-fasciata. The latter is the specimen 

illustrated by Treadwell (1922, pi. 7: fig. 5). The present 

description and illustrations are based on the specimen from 

1529. 

DESCRIPTION.—Syntype described incomplete, of unknown 

sex, with 84 setigers; length 33 mm; maximal width 1.3 mm at 

setiger 10; length through setiger 10, 3 mm. 

Prostomium (Figure 49e) distinctly shorter and narrower 

than peristomium, as deep as 1 /2 of the peristomiurn. Prostomial 


Antennae large, in transverse row, evenly spaced, similar in 


articulations. Ceratostyles medially inflated, with up to 8 

cylindrical articulations in A-III; articulations distinct on distal 

half of ceratostyles only. A-I to middle of anterior peristomial 

ring; A-I I to anterior edge of posterior peristomial ring; A-III to 

posterior edge of peristomium. Peristomium cylindrical with 

distinctly muscular lower lip. Separation between rings barely 

visible dorsally, invisible elsewhere; anterior ring 4 /5 of total 

peristomial length. Peristomial cirri short, slender and digitiform, 


Maxillary formula 1+1, 5+6, 6+0, 4+9, and presumably 1+1 

according to information in Treadwell (1922). 


Neuropodial acicular lobes distally truncate; aciculae emerging 

at midline. Pre- and postsetal lobes low, transverse folds. 

Anterior ventral cirri thick and tapering. Ventral cirri basally 

inflated in median setigers. Inflated bases ovate; narrow tips 

tapering. Posterior ventral cirri slender. Anterior notopodial 

cirri thick, tapering, becoming distinctly medially inflated in 

posterior setigers, without articulations. 

Pectinate setae (Figure 490 slightly flaring, flat. One 

marginal tooth distinctly longer than the teeth, with -10 teeth. 

Shafts of compound falcigcrs (Figure 49g) distally inflated, 

marginally smooth. Appendages short with distinct head, 

bidentate. Proximal teeth larger than distal teeth, pointed, 

tapering, directed basally. Distal teeth slender, curved. Guards 

distally symmetrically rounded; mucros absent. Pseudocompound 

falcigcrs and compound spinigers absent. Aciculae with 

black cores and clear sheaths, tapering distally to blunt, straight 

tips; cross-section round. Subacicular hooks (Figure 49h) with 

black cores and clear sheaths, bidentate. Hooks first present 


(except for replacements). Hooks with distinctly bent, narrowed 

necks, distinct heads. Both teeth directed obliquely 

distally; proximal teeth larger than distal teeth. 


Mx III and left Mx IV; setal characters associated with 

far posterior setigers; pygidium and anal cirri. 


TURES.—Mx III short and forming part of distal arc with left 

Mx IV. 



Unknown Characters: 1, 2, 13, 14,47, 50, 63. 


None. 

REMARKS.—The structure of the subacicular hooks of £. 

fuscafasciata resembles those in E. afra and the species is 

compared to other species in group B-4 in Table 33 and with 

other abranchiate species in Table 40. Only two abranchiate 

species have articulated ceratostyles; of these, E. fuscafasciata 

has cylindrical articulations and E. curticirris has distinctly 

moniliform articulations. The differences between this species 

and E. brevis was discussed above. 

Eunice fusicirris Grube, 1878 

Eunice fusicirris Grube, 1878a:102-103. 

REMARKS.—Grube described E. fusicirris as having short, 

thick antennae and peristomial cirri without articulations. The 

notopodial cirri were thick and stumpy in anterior setigers, 

becoming more slender in posterior setigers. Branchiae were 

present from setiger 5 to setiger 37 (of 50 setigers in the


complete specimen). Aciculae were black and the maxillary 

formula was given as 1+1, 6+5, 8+0,4+7, and 1+1. 

No locality information was available. The species appears 

to belong to group B-2, or possibly group B-l. Without access 

to the type it cannot be further characterized and is here 


74. Eunice gagzoi Augener, 1922 

Eunice gagzoi Augener, 1922b:45. 

FIGURE 50a-h; TABLES 33,37 

MATERIAL EXAMINED.—Lectotype ZMH V-9755, paralectotype, 

ZMH V-6812, St Thomas, coll. KrOyer and Orsted (part 

of original material of Eunice cariboea Grube, 1856); 

paralectotype, ZMB 6286, St. Thomas, coll. Kukenthal and 

Hartmeyer. 

COMMENTS ON MATERIAL EXAMINED.—The lectotype has 

been frontally dissected and the jaws are now missing as are 

several parapodia along the body. 


160 setigers; total length 32.5 mm; maximal width 1.5 mm at 

setiger 10; length through setiger 10, 2.75 mm. ZMH 

paralectotype complete, of unknown sex, with 105 setigers. 

ZMB paralectotype incomplete, of unknown sex, with 167 

setigers. 


about as wide as peristomium, as deep as l /2 of the 


flattened, tapering from junction to peristomium; median 

sulcus distinct ventrally and marked as shallow notch frontally, 

invisible dorsally. Eyes between bases of A-I and A-II. 

Antennae in shallow horseshoe, evenly spaced, similar in 


articulations. Ceratostyles digitiform, without articulations. A-I 

to middle of anterior peristomial ring; A-II and A-I 11 to setiger 

1. Peristomium tapering anteriorly, with distinct muscular 

lower lip. Separation between rings distinct on all sides; 

anterior ring ~ 2 /3 of total peristomial length. Peristomial cirri to 

middle of anterior peristomial ring, without articulations. 

Maxillary formula of ZMB paralectotype 1+1, 6+6, 6+0, 

4+8, and 1+1. Mx III part of distal arc with left Mx IV. Mx VI 

missing. 

Branchiae (Figure 50g) present as single thick branchial 

filaments from setiger 120 in lectotype, from setiger 85 in ZMH 

paralectotype, and from setiger 115 in ZMB paralectotype. 

Branchiae increasing in length posteriorly up to last 5 setigers, 

about as long as body is wide where best developed, reduced 

over last 5 setigers to short, indistinct buttons on last setigers 

present. Branchiae present on less than 55% of the total number 

of setigers. 

Anterior neuropodial acicular lobes (Figure 500 asymmetrically 

rounded, becoming symmetrically rounded in posterior 

setigers; aciculae emerging at midline. Anterior presetal lobes 

low, transverse folds, considerably shorter than acicular lobes, 

increasing in relative length posteriorly and from about setiger 

50 follow outline of acicular lobes closely. Anterior postsetal 

lobes higher than acicular lobes, symmetrically rounded, by 

setiger 25 postsetal lobes reduced to low folds following 

outline of acicular lobes closely. Anterior ventral cirri thick and 

tapering. Ventral cirri basally strongly inflated in setigers 10 to 

about setiger 30. Inflated bases thick, transverse welts; narrow 

tips tapering. Inflation rapidly reduced rapidly from setiger 30, 

from about setiger 45 ventral cirri short, slender and digitiform. 

Anterior notopodial cirri basally inflated, becoming shorter and 

more slender in posterior setigers, but always longer than 

ventral cirri, without articulations. 


setae (Figure 50e) very large, tapering, flat. Both marginal teeth 

longer than other teeth, -12 teeth present. Shafts of compound 

falcigers (Figure 50c,d) inflated, marginally coarsely and 

shallowly serrated. Appendages slender. Anterior appendages 

(Figure 50c) with nearly parallel sides and large heads, 

bidentate. Teeth similar in size. Proximal teeth tapering, 

directed obliquely distally. Distal teeth curved, directed 

obliquely distally. Guards asymmetrically bluntly pointed, 

marginally serrated; mucros absent. Posterior appendages 

(Figure 50d) tapering with small heads, bidentate. Proximal 

teeth smaller than distal teeth, tapering, directed obliquely 

distally. Distal teeth curved, directed obliquely distally. Guards 

increasingly symmetrically rounded in posterior setigers, 


and compound spinigers absent. Aciculae (Figure 50b) single, 

amber-colored, thick, abruptly tapering, distally bent. Subacicular 

hooks (Figure 50h) amber-colored, bidentate. Hooks 

first present from setiger 27-38, present in all setigers 


distinctly inflated subdistal region and large, parrot-beak 

shaped heads. Proximal teeth very large, strongly curved. 

Distal teeth small, distinctly curved. 




TURES.—Mx III short and forming part of distal arc with left 

MxIV. 


SCORED.—Inappropriate Characters: 22, 33, 34, 39-42, 56, 

58, 59. Unknown Characters: 74, 78. 


None. 

REMARKS.—This species was considered a junior synonym 

of E. cariboea by Hartman (1959) and it certainly resembles 

that species. The two species have, however, distinctly different 

subacicular hooks, pectinate setae, and aciculae, in addition to 

having differently shaped prostomia and peristomia. The 

presence of distinct branchiae on E. gagzoi also clearly 

separates the two species. Eunice gagzoi has been compared to 

similar species in Tables 33 and 37. The very late start of the

154 


.02 b, c, d. e. h, j, k, I, m 

f. 9 

a, i 

m 

FIGURE 50.—Eunice gagzoi (lectotype ZMH V-9755): a, anterior end, lateral view; b, acicula, parapodium 12; c, 

compound falciger, parapodium 12; d, compound falciger, parapodium 150; e, pectinate seta, parapodium 150; f, 

parapodium 12, anterior view; g, parapodium 150, anterior view; h, subacicular hook, parapodium 150. Eunice 

gooctei, new name (holotype of Nicidion kinbergi, USNM 4800): i, anterior end, dorsal view; j, pectinate seta, 

median parapodium; k, compound falciger, median parapodium; 1, subacicular hook, median parapodium; rn, 

compound falciger, median parapodium. (Scale bars in mm.)


branchiae is rare in the genus, and unique to the species listed 

in Table 37. 

Eunice gaimardi Quatrefages, 1866 

Eunice gaimardi Quatrefages, 1866:321.—Fauchald, 1986:251. 

REMARKS.—The information available about this species 

was reviewed by Fauchald (1986). The species is indeterminable 

without access to the type material. 

Eunice gallica (Lamarck, 1818) 

Leodice gallica Lamarck, 1818:322.—Savigny, 1820:50-51. 

Eunice tail tea — Auduuin and Milne Edwards, 1833:218; 1834:144. 

REMARKS.—Lamarck gave a brief Latin diagnosis and an 

even briefer comment in French. The combined information is 

summarized in the next paragraph. 

Specimen with 71 segments. Antennae without articulations. 

Branchiae first present from segment 6 [setiger 4, K. Fauchald 

comment]. Anterior branchiae single filaments; other branchiae 

bifid. Branchiae missing on last 18 setigers. 

Savigny added that the antennae were shorter than in E. 

antennata and lacked articulations and that branchiae in 

segments 6-8 were simple filaments, that the branchiae of 

segment 9 had two filaments, and that branchiae were missing 

in the last 18 segments of a total of 72 segments. 

Audouin and Milne Edwards (1834) stated that they had not 

seen any material, that the species resembles E. harassii, except 

for the absence of articulations from the ceratostyles, and that 

the branchiae started on segment 6. They otherwise quoted 

Savigny's description in toto. 

No specimens are known and no type locality other than 

what is implied by the species name is known. The species is 

here considered indeterminable, in agreement with Hartman 

(1959). 

Eunice gigantea auctores 

REMARKS.—This specific name has been used several times, 

in a variety of contexts. The specimens to which the different 

versions of the name were originally applied are no longer in 

existence, and the name, in all its many guises, is invalid. Part 

of the discussion has referred to the designation of type species 

for the genus; this aspect was reviewed above. However, a brief 

review of the various uses of this name may clarify some of the 

confusion at the species level. 

1. Nereis gigantea Linnaeus, 1758:654. The total description 

is the following line: 

Nereis teniaculorum fasciculis triplici online. 

As a synonym for this species, Linnaeus lists Millepora 

marina amboinensi and as habitat "Mare Indico." The 

information in the Linnean description is clearly inadequate to 

identify the taxon even to family. 

2. "Eunice gigantea Cuvier" sensu auctores. According to 

Savigny (1820:49-50), Cuvier used this name for specimens in 

the collections in Paris and published in Cuvier (1817:525). 

Cuvier (1817:525) named the genus Eunice, but no species 

named gigantea was mentioned. He listed aphroditois, pinnata, 

norwegica, tubicola, and cuprea as members of his new genus 

and stated that he "had seen a giant specimen [of Eunice]" from 

the Indian Ocean. That informal mention may have been the 

source of the reference by later authors. Lamarck (1818:322) 

made no reference to a species named gigantea authored by 

Cuvier. In a later edition, Cuvier (1830:200) included gigantea 

in a list of synonyms similar to the one issued by Savigny 

(1820). Cuvier (1830) reviewed material from both the Indian 

and Atlantic oceans. 

3. Leodice gigantea Lamarck, 1818. Lamarck (1818:322) 

gave a brief Latin diagnosis of the species and a rather 

extensive description in French. His description is an excerpt of 

the one published by Savigny two years later and clearly refers 

to the same specimens. 

4. Leodice gigantea.—Savigny, 1820:49-50. Savigny 

listed Nereis aphroditois Pallas (= Terebella aphroditois, sensu 

Gmelin) as a possible synonym for his new species (which 

actually had been published already by Lamarck). The quote 

reads "Varie*te d'age ou espece tres-voisine." In addition, 

Savigny quoted the museum collections and gave reference to 

Cuvier as listed above. The material studied by Savigny was 

from the Indian Ocean and the fairly detailed description is 

consistent with Eunice aphroditois (Pallas). Because the latter 

name is valid (see above), in this sense the name is a junior 

synonym of E. aphroditois. 

5. Quatrefages (1866:311) reserved the name gigantea for 

material from the Indian Ocean. He gave a new name, Eunice 

roussaei, to Atlantic specimens, including some he had 

collected; as a synonym for this species he quoted "Eunice 

gigantea Cuvier Regne anim. L III, p. 200." This reference is to 

the second edition of Cuvier's book, issued in 1833, where the 

species occurs on the page indicated and where Cuvier included 

specimens from both the Atlantic and Indian oceans. Quatrefages 

is correct in separating the Atlantic species from the 

Indian Ocean species; thus he was also correct in giving the 

Atlantic material a new name. As the original author for the 

name gigantea, Quatrefages quoted Nereis gigantea Linnaeus 

with no date. As indicated above, the Linnaean species, as 

originally described in 1758, is wholly unidentifiable, even to 

family. In summary, where identifiable, the name gigantea in 

combination with the generic names Eunice or Leodice used 

about material from the Indian Ocean refers to the species 

validly named E. aphroditois (Pallas). The Linnaean name 

Nereis gigantea cannot be identified even to family. Cuvier did 

not use the name E. gigantea in the 1817 edition of his "Regne 

animal"; he used it first in 1833; thus Lamarck (1818) first used 

the name Leodice gigantea and in his sense the name can be 

identified as a junior synonym of Pallas' Nereis aphroditois.


75. Eunice goodei, new name 

FIGURE 50i-m; TABLES 33,40 

Nicidion kinbergi Webster, 1884:320-321, pi. 12: figs. 81-88 [noc Eunice 

kinbergi Ehlers, 1868:306]. 

Eunice (Nicidion) kinbergi—Hartman, 1944:124; 1959:313. 

MATERIAL EXAMINED.—Holotype of Nicidion kinbergi, 

USNM 4800, 1 specimen plus 3 slides of prostomium and 

parapodia, Bermuda, coll. G. Brown Goode. 


now in three pieces; the anterior end and two first setigers are 

mounted on a slide; two other slides contain various parapodia. 


56 setigers; estimated length 16 mm; maximal width 0.75 mm; 

length through setiger 10, 4 mm. Body dorsally convex, 

ventrally slightly flattened, basically cylindrical with little 

indication of taper posteriorly. 

Prostomium (Figure 50i) distinctly shorter and narrower than 


inflated; median sulcus shallow. Eyes posterior to bases of A-I 

and A-II. Antennae in horseshoe; A-I and A-II close together, 

separated by gap from A-I 11, similar in thickness. Ccratophorcs 


digitiform, without articulations. A-I and A-II to posterior 

peristomial ring; A-II I to setiger 1. Peristomium tapering 

anteriorly. Separation between rings distinct on all sides; 

anterior ring 2 /3 of total peristomial length. Peristomial cirrus 

on right side lost; left cirrus short, ovate, possibly in 

regeneration. 




conical in posterior setigers; aciculae emerging at midline. 

Presetal lobes low, transverse folds. Anterior postsetal lobes 

rounded, longer than acicular lobes, becoming low, transverse 

folds at about setiger 20. Anterior ventral cirri tapering, 

becoming basally inflated by setiger 10. Inflated bases large, 

nearly hemispherical; narrow tips short and button-shaped. 

Inflated bases rapidly reduced posteriorly, absent by setiger 40. 

Posterior ventral cirri slender and digitiform. Anterior notopodial 

cirri basally slightly inflated, tapering to slender tips; 

inflated bases lost posteriorly; last notopodial cirri present 

slender, digitiform. Notopodial cirri without articulations. 

Limbate setae marginally smooth with long, thin, filamentous 

tips. Pectinate setae (Figure 50j) tapering, flat. One 

marginal tooth longer than other teeth, with 10 teeth. Shafts of 

compound falcigers (Figure 50k,m) inflated, marginally 

smooth. Appendages tapering, bidentate. Proximal teeth tapering, 

directed slightly distally in most hooks. Distal teeth 

tapering to sharp or blunt points, directed obliquely distally. 

Guards symmetrically rounded, marginally smooth; mucros 


absent. Aciculae with dark yellow to dark brown cores and 

clear sheaths, sharply pointed, straight; cross-sections round. 

Subacicular hooks (Figure 501) with light to dark brown cores 

and clear sheaths, bidentate. Hooks first present from setiger 

23, present in all sctigers thereafter, always single (except for 

replacements). Hooks curved, with distinctly narrowed necks 

just below heads. Proximal teeth larger than distal teeth, 

triangular, directed laterally. Distal teeth triangular, nearly 

erect. 


setal features associated with posterior setigers; pygidium and 

anal cirri. 





Unknown Characters: 1, 2, 4, 6, 12, 24,47, 50,63. 


None. 

REMARKS.—This species was originally described as Nicidion 

kinbergi. Because of the fusion of the genus Nicidion 

with Eunice, the species name ends up being preoccupied in the 

combination Eunice kinbergi Ehlers (1868:306), proposed for 

a very different species (see below). The new specific name 

honors G. Brown Goodc, who at the lime of collecting this 

material was a professor at Wesleyan University, he later 

became Assistant Secretary of the Smithsonian Institution. 

Eunice goodei is listed with similar species in Tables 33 and 

40. It can be separated from the other abranchiate species as 

suggested in the discussion of E. cariboea. 

207. Eunice goodsiri (Mclntosh, 1885) 

TABLES 33,34 

Marphysa goodsiri Mclntosh, 1885:299-301, figs. 56-58, pi. 38: figs. 6-8. pi. 

19A: figs. 18-20. 

DESCRIPTION.—No material is available. The following 

description is based on Mclntosh's description and interpretation 

of his illustrations. 

Maximum width 2.5 mm. 

Median sulcus deep. Eyes present Antennae similar in 

thickness. Ceratostyles tapering, articulated with distal articulations 

drop-shaped. Length of A-II and A-III similar. Peristomial 

cirri to anterior end of peristomium. 

Maxillary formula (after Mclntosh, 1885:300 and fig. 56) 

1+1,4+4,3+0,3+6,1+1. 



setiger 25 or 26 to end of fragment. First 11 pairs single 

filaments, maximum 2 filaments continued to end of fragment. 

Median acicular lobes conical; aciculae emerging close to 

midline. Pre- and postsetal lobes unknown. Ventral cirri 

inflated in median setigers (at least in setigers 20 and 40). 

Notopodial cirri tapering in all setigers present, without 

articulations.


Limbate and pectinate setae unknown. Shafts of compound 

falcigcrs distally inflated, without marginal serrations, with 

internal striations. Appendages tapering, bidentate. Proximal 

teeth larger than distal teeth, tapering, directed laterally. Distal 

teeth tapering, curved, distally directed obliquely laterally. 

Guards of compound falcigers distally symmetrically rounded, 

marginally smooth; mucros absent. Pseudocompound falcigers 

and compound spinigers absent. Up to 3 aciculae in anterior 

setigers, single in median setigers (setiger 40), dark, pointed, 

bent in anterior setigers. Subacicular hooks with dark core and 

clear sheath, bidentate. Distribution unknown, single. Proximal 

teeth larger than distal teeth, directed laterally, tilted upward 

distally. Distal teeth short, thick, tapering, directed distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Most features 

remain unknown. 


Species too poorly known to allow any predictions. 



Characters: 1, 2, 4-8, 10-12, 14-16; 18, 19, 24-26, 28, 29, 

36-38,40,45-52, 57-60,63, 65-68,80. 


None. 

REMARKS.—Described from St. Thomas in the West Indies, 

this species was compared to E. filamentosa by Mclntosh. It 

has not been reported since it was first recorded. It belongs to 

group B-4 and has been compared to similar species in Tables 

33 and 34. The only other species in Table 34 with distally 

drop-shaped articulations in the antennae is E. collaris. The two 

species differ in many characters; perhaps most noticeable are 

the differences in the branchial structures. Eunice goodsiri has 

palmate branchiae with up to two filaments; E. collaris has 

pectinate branchiae with up to 12 filaments. 

Eunice goodsiri is too poorly known to be included in the 

key. 

76. Eunice gracilicirrata (Treadwell, 1922) 

FIGURE 51a-g; TABLES 22,23 

Ltodice gracilicirrata Treadwell, 1922:149-150, figs. 36-38, pi. 5: figs. 1-8. 

Eunice arndllata.—Hartman, 1956:282 [not Eunice armillata Treadwell, 

1922]. 

MATERIAL EXAMINED.—Holotype, AMNH IV-1920-1535, 

Suva Harbor, Fiji. 

COMMENTS ON MATERIAL EXAMINED.—The antennae are 

all bent over and cannot be illustrated as stretched without 

damage to the specimen; the sizes of the antennae given below 

are low estimates. 



10; length through setiger 10,9 mm. 


than peristomium, as deep as x /i of the peristomium. Prostomial 

lobes firontally rounded, dorsally inflated; median sulcus deep. 

Antennae in shallow horseshoe; A-I and A-II close together, 

separated by gap from A-I II; A-I 11 slimmer than A-I and A-II. 


Ceratostyles tapering, slender, with up to 5 indistinct, long, 

cylindrical articulations; articulations limited to distal half of 

styles. A-I to middle of anterior peristomial ring; A-II to setiger 

2; A-I 11 to front edge of setiger 4. Peristomium about twice as 

long as prostomium, cylindrical. Separation between rings 

distinct dorsally and ventrally and less well marked, but visible 

also laterally; anterior ring ~ 4 /s of total peristomial length. 

Peristomial cirri to tip of prostomium, slender, with 3 or 4 

indistinct articulations. 


Branchiae (Figure 510 present, pectinate, distinctly longer 




number of setigers. First 2 pairs single filaments; maximum 7 

filaments in setiger 20-60, thereafter numbers decreasing 

gradually. From setiger 145 to setiger 181 only single filaments 

present. Branchial stems slender, tapering, shorter than notopodial 

cirri. Filaments very long, slender, digitiform. Last few 

branchiae short, nearly button-shaped protrusions. 


at midline. Presetal lobes low, transverse folds. Anterior 

postsetal lobes about as long as acicular lobes, rounded, 

becoming reduced to low, transverse folds by setiger 20. 

Ventral cirri digitiform through setiger 10, thereafter basally 

inflated through next 40 setigers, retaining long, digitiform 

narrow tips. Inflated bases ovate, decreasing rapidly from 

setiger 60, essentially lost by setiger 70. Posterior to setiger 

100, ventral cirri tapering and digitiform. Anterior notopodial 

cirri slender and tapering, similar in length to anterior ventral 

cirri. Notopodial cirri increasing rapidly in length, by setiger 10 

longer than half of body width, with 2 to 5 indistinct 

articulations. Median and posterior notopodial cirri without 

articulations. Notopodial cirri long in all branchial segments, 

thereafter decreasing rapidly to about setiger 240. Posterior to 

setiger 240 ventral cirri and notopodial cirri again of about 

same length (Figure 51g); far posterior notopodial cirri less 

than half as wide as ventral cirri. 


(Figure 5 le) large, tapering, flat. Either 1 or both marginal teeth 


falcigers (Figure 51c) inflated, with fine marginal serrations. 

Appendages slender, bidentate. Proximal teeth smaller than 

distal teeth, triangular, directed laterally. Distal teeth tapering, 

curved. Guards symmetrically rounded, marginally finely 

serrated; mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae paired, yellow, straight; 

cross-sections round. Superior aciculae (Figure 51b) tapering 

abruptly, distally covered with a fine hood or mucro. Inferior 

aciculae bluntly rounded. Separation between core and sheath

158 


FIGURE 51.—Eunice gracilicirrata (holotype, AMNH IV-1920-1535): a, anterior end, lateral view; b, acicula, 

parapodium 74; c, compound falciger, parapodium 74; d, subacicular hook, parapodium 275; e, pectinate seta, 

parapodium 275; f, parapodium 74, anterior view; g, parapodium 275, anterior view. Eunice gracilis (syntype, 

ZMB 1856): h, anterior end, lateral view; i, parapodium 30, anterior view; j, subacicular hook, parapodium 30; 

k, compound falciger, parapodium 6; 1, parapodium 6, anterior view; m, acicula, parapodium 30. (Scale bars in 

mm.)



hooks (Figure 5Id) yellow, bidentate. Hooks first present from 


(except for replacements). Hooks distally strongly curved; 

heads small. Proximal teeth larger than distal teeth, triangular, 

directed laterally. Distal teeth less than half as massive as 

proximal teeth, nearly erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—Jaw 


structure; 


Mx III long and located behind left Mx II. 





None. 

REMARKS.—Presumably through a lapsus, E. gracilicirrata 

was referred to E. armillata by Hartman (1956). Eunice 

gracilicirrata belongs to group A-2 (Fauchald, 1970). 

It is a very long, slender species with long, slender 

ceratostyles and notopodial cirri. The aciculae are hooded in 

some anterior setigers. It is listed with similar species in Tables 

22 and 23. Of the species listed in Table 23, only E. 

gracilicirrata and E. stigmatura have articulated peristomial 

cirri. The latter has tridentate subacicular hooks, in addition to 

bidentate ones; E. gracilicirrata has only distinctly bidentate 


77. Eunice gracilis Grube, 1866 

FIGURE 51h-m; TABUES 46-48 

Eunice antennata gracilis Grube, 1866a:65. 

Eunice gracilis.—Grube, 1866c: 174. 

Eunice gracilis Gr. var. antennata.—Grube, 1867:9-11, pi. 1: fig. 2. 

MATERIAL EXAMINED.—Two syntypes, ZMB 1856 and 

ZMH V-787, Tahiti, Novara Exp. 

COMMENTS ON MATERIAL EXAMINED.—Both specimens are 

now rather flaccid, but are otherwise in good condition; but the 

shape of the parapodial lobes is difficult to describe in detail. 

The description is based on the Berlin syntype with notes where 

the Hamburg syntype differs. 

DESCRIPTION.—Berlin syntype complete, of unknown sex, 

with 81 setigers; total length 66 mm; maximal width 2 mm at 

setiger 10; length through setiger 10, 6 mm. Hamburg syntype 

incomplete, of unknown sex, with 31 setigers; length 25 mm; 


mm. 


than peristomium, as deep as l /2 of the peristomium, sloping 

steeply ventrally. Prostomial lobes frontally obliquely truncate, 

dorsally flattened; median sulcus shallow. Eyes between bases 

of A-I and A-II in Hamburg syntype, reddish. Antennae in 

horseshoe, evenly spaced; A-I thicker than other antennae. 


Ceratostyles digitiform with up to 23 moniliform articulations; 

styles articulated to bases. A-I to middle of anterior peristomial 

ring; right A-II to setiger 2; left A-II and A-III incomplete. A-I II 

probably longest antenna, in incomplete condition as long as 

right A-II. Peristomium cylindrical. Separation between rings 

faintly visible dorsally, indiscernible ventrally and laterally. 

Anterior ring 4 /s of total peristomial length. Peristomial cirri to 

front edge of peristomium, very slender and digitiform, with 6 


Maxillary formula 1+1, 6+7, 8+0, 5+10, and 1+1. Teeth 

slender and delicate. Mx III long and located behind left Mx II. 

Mx VI missing. 


than notopodial cirri, reduced in mid-body region, erect. 

Branchiae from setiger 5 (4 in Hamburg specimen) to setiger 

81. Branchiae present to near posterior end, present on more 

than 65% of total number of setigers. First branchia with 3 

filaments, maximally 6 filaments present. Number of filaments 

decreasing rapidly from setiger 20, by setiger 35 single 

filaments. At setiger 65 number of filaments increasing to 2; 

last 10 branchiae with 3 filaments. Anterior and posterior 

branchiae longer than notopodial cirri; median and posterior 

branchiae shorter than notopodial cirri. Stems tapering, about 

as long as notopodial cirri. Anterior and posterior filaments 

digitiform; median filaments filiform; anterior and posterior 

filaments shorter than notopodial cirri. 

Anterior neuropodial acicular lobes apparently rounded; 

posterior acicular lobes triangular or perhaps conical; aciculae 

emerging at midline. Pre- and postsetal lobes apparently low, 

transverse folds. Anterior ventral cirri thick and tapering. Bases 

inflated base from about setiger 10. Bases ovate; narrow tips 

tapering. From setiger 30 ventral cirri reduced to short, sharply 

tapering lobes (Figure 51i). Anterior notopodial cirri long, 

tapering or digitiform, with 4 or 5 articulations. Number of 

articulations decreasing posteriorly, and completely lost at 

about setiger 50. Posterior notopodial cirri retaining shape but 

shorter than in anterior part of body. 

Limbate setae slender. Shafts of compound falcigers (Figure 

51k) distally gently inflated, marginally smooth. Appendages 

short, tapering; head distinct, bidentate. Proximal teeth shorter 

than distal teeth, tapering, directed distally. Distal teeth 

tapering, directed obliquely distally. Guards asymmetrically 

bluntly pointed, marginally finely serrated; mucros absent. 


Aciculae (Figure 51m) paired, yellow, tapering to bluntly 

pointed, straight tips; cross-sections round. Separation between 


hooks. Subacicular hooks (Figure 51j) yellow; tridentate with 

teeth in a crest. Hooks first present from setiger 28, present in 


Hooks very large; fangs in a row, decreasing evenly and rapidly 

in size from large main fangs to small tertiary fangs; all fangs 

curved.

160 



anal cirri. 


None. 



Characters: 23,42, 65-68. 


None. 

REMARKS.—Described on material collected during the 

Novara Expedition, this species was named in a brief note. In 

this note Grube stated that the species described from the 

Novara material under the name E. gracilis should be 

considered a variety of E. antennata Sav., differing basically in 

the greater articulations of the notopodia. This note was 

published before the expedition report came out, so the official 

original name for the species becomes E. antennata gracilis. 

The types differ clearly from E. antennata and is here 

considered a distinct species. 

Eunice gracilis is listed with similar species in Tables 46 and 

48. It is the only species in Table 48 with very poorly marked 

separation between the anterior and posterior ring of the 

peristomium; in this species it is barely visible dorsally and 

indiscernible laterally and ventrally; in the other species in this 

table, the separation is at the very least distinct dorsally and 

ventrally and in some species is distinct on all sides. 

Eunice gracilis Moore, 1903 

Eunice gracilis Moore, 1903:440-441, pi. 25: figs. 46-48. 

REMARKS.—The name E. gracilis is preoccupied in the 

combination E. gracilis Grube, 1866. Moore's species, which 

clearly differs from Grube's, is here renamed E.japonica and is 

described below. 

Eunice gracilis (Crossland, 1904) 

Nicidion gracilis Crossland, 1904:327-329, fig. 66, pi. 22: figs. 10, 11. 

Eunice (Nicidion) gracilis.—Hartman, 1959:313. 

REMARKS.—With the fusion of Nicidion and Eunice, the 

specific name becomes preoccupied in the combination E. 

gracilis Grube, 1866. Crossland's species is here renamed E. 

wasinensis and is described below. 

78. Eunice gravieri Fauvel, 1911 

TABLES 24,25 

Eunice gravieri Fauvel, 1911:14-15, figs. la-b. 2a-d; 1914:145-146, pi. 10: 

figs. 1-4 and 11-18. 

DESCRIPTION.—The types are not available. Fauvel's two 

descriptions are detailed and suffice to characterize the species. 

Types two small specimens, one incomplete, other 28 mm 

long and 2 mm wide (presumably with parapodia); numbers of 

setigers not mentioned but must have exceeded 50. 

Prostomium frontally rounded. Two pairs of eyes; 1 between 

bases of A-I and A-I I and 1 pair posterior and slightly medial to 

other pair, hidden under peristomial fold. Ceratophores 


slender and digitiform, with up to 6 long, cylindrical 

articulations in A-I 11 (as illustrated, not moniliform as stated by 

Fauvel). A-I I and III to setiger 8. Peristomium apparently a 

single ring. Peristomial cirri very slender and digitiform, 

reaching well beyond frontal edge of peristomium, articulated 

(Fauvel's illustrations conflict with his text). 

Maxillary formula 1+1,6+7,7+0,6+10,1+1, and 1 + 1. Mx V 

and VI reduced. Mx III part of distal arc with left Mx IV. 

Branchiae present, single filaments, distinctly shorter than 

notopodial cirri. Branchiae from sctigcr 5 to setiger 20 or 26. 

Branchiae terminating well before posterior end. Filaments 

digitiform. 

Ventral cirri basally inflated in branchial region, tapering in 

anterior and far posterior setigers. Notopodial cirri digiliform, 

vaguely articulated, at least through branchial region. 

Pectinate setae narrow. One marginal tooth very long, stout, 

with -10 teeth. Shafts of compound falcigcrs lapcring or 

slightly inflated, marginally smooth. Appendages relatively 

short, with large heads, bidentate. Proximal teeth triangular, 

slightly tapering, directed at various angles. Distal teeth either 

gently curved and slender or thick with distinct bends. Guards 

symmetrically bluntly pointed; mucros absent. Pseudocompound 

falcigcrs and compound spinigers absent. Aciculae dark, 

tapering, pointed, straight in anterior setigers, bent in posterior 

setigers; cross-sections round. Subacicular hooks dark, bidentate. 

Hooks first present from setiger 25. Hooks strongly 

curved, with distinct inflated region just below heads. Proximal 

teeth larger than distal tooth, directed laterally. Distal tooth 

directed obliquely distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Numerous features 

from all parts of the body. 


Species too poorly known to allow predictions. 


SCORED.—Inappropriate Characters: 26, 33, 34, 39,40, 56, 

58, 59. Unknown Characters: 1, 4-6, 8-12, 15-17, 24, 38, 

43-51, 54, 55, 65, 67, 74, 78, 81, 82. 


None. 

REMARKS.—Some puzzling features described by Fauvel 

need confirmation. The presence of four eyes is rare in the 

family. An entire peristomium, as far as I know, has never been 

reported for any other species. The distribution of branchiae 

and the structure of the branchiae are similar to what is present 

in certain juveniles, but these features are by no means uniquely 

juvenile features. The species is listed with similar species in 

Tables 24 and 25; it is not characterized by a single unique 

feature; other than those noted above. It is too poorly known to 

be included in the key.


79. Eunice grubei Gravier, 1900 

FIGURE 52a-«; TABLES 27,32 

Eunice grubei Gravier, 1900:258-261. figs. 125-129, pi. 14: figs. 87, 88. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, Djibouti, 

H. Coutiere no. 29, 1897. 


been mildly dehydrated at one time. 


55 setigers; length 42 mm long; maximal width 6 mm at setiger 

10; length through setiger 10, 12 mm. Body cylindrical 

throughout 


than peristomium, as deep as */2 of peristomium. Prostomial 


shallow. Eyes not observed. Antennae in shallow horseshoe, 



with up to 8 cylindrical articulations, articulations basally 

cylindrical becoming drop-shaped distally. A-I to posterior 

peristomial ring; A-I I and A-I 11 to setiger 1. Peristomium 

cylindrical with distinct muscular lower lip. Separation 

between rings distinct dorsally and ventrally, and invisible on 

short lateral section; anterior ring 5 /6 of total peristomial length. 

Peristomial cirri to middle of anterior peristomial ring, slender 

and digitiform with 5 long, drop-shaped articulations. 

Maxillary formula 1+1, 5+5,7+0,6+9, and 1+1. Mx III part 

of distal arc with very short left Mx IV. Mx VI absent. 



from setiger 3 to end of fragment. First branchiae single 

filaments, maximum 8 filaments at about setiger 20; number of 

filaments decreasing posteriorly; 4 or 5 filaments in each 

branchia in last segments. Branchial stems thick, tapering, as 

long as notopodial cirri. Filaments digitiform, shorter than 



at midline. Pre- and postsetal lobes transverse folds; 

presetal lobes shorter than acicular lobes; postsetal lobes nearly 

as long as acicular lobes. First 5 ventral cirri large, nearly 

triangular, tapering to blunt tips. Ventral cirri basally inflated 

from setiger 6 through setiger 30. Inflated bases modest, ovate, 

in relation to large, tapering tips. Posterior to setiger 30 ventral 

cirri very broadly attached ventral to neuropodial acicular 

lobes, tapering to blunt tips; attachment point slightly posterior 

to emergence of subacicular hooks, forming incomplete scoops 

around lower edge of neuropodia. Notopodial cirri basally 

inflated, tapering to digitiform tips. Prebranchial notopodial 

cirri with 4 or 5 indistinct articulations; all other notopodial 


Limbate setae marginally smooth or slightly frayed. Pectinate 

setae (Figure 52b) tapering, gently furled. Both marginal 

teeth thicker than other teeth; one marginal tooth distinctly 


falcigers (Figure 52c,d) tapering, marginally smooth. Appendages 

short, tapering, with large heads, bidentate. Proximal teeth 

smaller than distal teeth, conical to nearly triangular, directed 

laterally or slightly distally. Distal teeth tapering, curved, 

directed laterally. Guards asymmetrically bluntly pointed, 

marginally smooth; mucros absent. Pseudocompound falcigers 

and compound spinigers absent Aciculae numbering 2 or 3, 

amber to dark brown, thick, distally blunt, gently curved or 

straight; cross-sections round. Subacicular hooks (Figure 52a) 

amber to dark brown, bidentate. First few hooks light amber 

and easily overlooked. Hooks first present from setiger 24, 

present in all setigers thereafter, usually single, paired in some 

setigers. Hooks with narrow neck and distinct head. Proximal 

teeth large, directed laterally. Distal teeth smaller than proximal 

teeth, directed obliquely distally. 







Characters: 1, 2, 13, 14, 36-38,40,47, 50, 63, 74, 78. 


38,1. 

REMARKS.—Eunice grubei is listed with similar species in 

Tables 27 and 32. It is the only species listed in Table 32 that 

combines branchiae starting from setiger 3 with articulated 

notopodial cirri limited to the anterior end; in the other species 

in the table with branchiae from setiger 3, the notopodial cirri 

are articulated throughout. 

Eunice grunwaldi (Risso, 1826) 

Leodice GrunwaldiRisso, 1826:423— Fauvel, 1923:451. 

Eunice Grunwaldi.—-Grube, 1850:292. 

REMARKS.—Grube (1850) referred this species to his section 

of Eunice without tentacular cirri (his "Leodicae Marphysae 

Sav."); Fauvel (1923) referred the species to Marphysa. The 

species will be further considered in a planned review of that 

genus. 

80. Eunice guanica (Treadwell, 1921) 

FIGURE 52f-i; TABLES 33,36 

Leodice guanica Treadwell, 1921: 39-40, figs. 107-116, pi. 2: figs. 9-12. 

Eunice afra.—Hartman, 1956:282 [not Eunice afra Peters, 1854]. 

MATERIAL EXAMINED.—Holotype, AMNH 1915-1342, 

Sand Key, Key West, Florida. 



setiger 10; length through setiger 10, 7 mm. Body anteriorly 

cylindrical with very short parapodia.

162 


FIGURE 52.—Eunice grubei (holotype, MNHN, Paris): a, subacicular hook, parapodium 42; b, pectinate seta, 

parapodium 42; c, compound falciger, parapodium 49; d, compound falciger, parapodium 42; e.anterior end, 

lateral view. Eunice guanica (holotype, AMNH 1915-1342): f, anterior end, lateral view; g, posterior parapodium, 

anterior view; h, compound falciger, posterior parapodium; i, pectinate seta, posterior parapodium; j, subacicular 

hook, posterior parapodium. (Scale bars in mm.)



than peristomium, as deep as V2 of peristomium. Prostomial 

lobes short, frontally rounded, dorsally inflated; median sulcus 

short. Antennae in a nearly complete circle and directed 

forwards, evenly spaced, similar in thickness. Ceratophores 


digitiform, with 2 or 3 cylindrical articulations in A-II and 

A-III. A-I to middle of anterior peristomial ring; A-II to middle 

of anterior peristomial ring; A-III to posterior edge of anterior 

peristomial ring. Peristomium more than twice as long as 

prostomium, cylindrical with a ventral depression anteriorly. 

Peristomium divided ventrally into 3 equally long parts; 

posteriormost of these parts distinct also dorsally representing 

posterior peristomial ring, separation indistinct laterally. 

Peristomial cirri barely reach front edge of posterior peristomial 

ring, tapering, without articulations. 

Jaws missing. 

Branchiae present, distinctly longer than notopodial cirri, not 

reduced in mid-body region. Branchiae from setiger 19 to 

sctigcr 224. Branchiae present to near posterior end, present on 

more than 65% of total number of setigers. First 10 branchiae 

single filaments, thereafter some segments with 2 filaments, 

others with single filaments, up to 'A of segments in branchial 

region, without branchiae. Branchiae very long (Figure 52g), 

slender, digitiform, outreaching notopodial cirri in all but a few 

posterior branchial segments. 

Anterior neuropodial acicular lobes distally symmetrically 

rounded; posterior acicular lobes triangular with prominently 

projecting aciculae; aciculae emerging at midline. All presetal 

lobes low transverse folds. Anterior postsetal lobes higher than 

acicular lobes, distally rounded; postsetal lobes reduced to low 

folds by beginning of branchial region. First 6-7 ventral cirri 

digitiform. Ventral cirri becoming basally inflated in median 

setigers. Bases ovate; narrow tips short and button-shaped. 

Inflated bases continued through rest of body. Notopodial cirri 

tapering, similar in length except shorter in first 2 segments. 

Notopodial cirri without articulations. 

Limbate setae slender, marginally frayed. Pectinate setae 

(Figure 52i) relatively large, but very delicate, flaring, flat. 

Both marginal teeth distinctly longer than other teeth, with 15 

teeth. Shafts of compound falcigers (Figure 52h) distally 

inflated, marginally smooth. Appendages tapering; head small, 

bidentate. Proximal teeth considerably larger than distal teeth, 

tapering, directed slightly distally. Distal teeth gently curved, 

tapering. Guards symmetrically rounded; mucros absent. 


Aciculae single, black, distally tapering to blunt, straight tips; 

cross-sections round. Subacicular hooks (Figure 52j) black, 



Hooks abruptly tapered with small heads; both teeth directed 

distally, very small, blunt. 







Characters: 13, 14,40, 74, 78. 


None. 

REMARKS.—Eunice guanica is listed with similar species in 

Tables 33 and 36. It is one of three species in Table 36 in which 

the bases of the ventral cirri remain inflated through posterior 

setigers; the other two species are E. cirrobranchiata and E. 

excariboea. In the latter the branchiae do not start until setigers 

78-79; in the two former, branchiae are present from about 

setigers 19 and 28 respectively. In E. cirrobranchiata at least 

some of the subacicular hooks are tridentate with the teeth in 

tandem; all subacicular hooks are bidentate in E. guanica. 

Eunice guildingi Baird, 1869:351. 

81. Eunice guildingi Baird, 1869 

FIGURE 53a -e; TABLES 27,31 

MATERIAL EXAMINED.—Holotype, BM(NH) 1839.12.27.5, 

St. Vincent, West Indies. 







sulcus deep. Eyes not observed. Antennae in horseshoe, evenly 


antennae, without articulations. Ceratostyles digitiform; A-I 

without articulations; A-II and A-III with up to 7 long, 

cylindrical articulations. A-I to posterior peristomial ring. A-II 

to posterior edge of setiger 1; A-III to setiger 2. Peristomium 

slightly flaring anteriorly, with distinct muscular lower lip. 

Separation between rings distinct on all sides; anterior ring ~ 3 A 


prostomium, slender and digitiform, without articulations. 





well before posterior end, more than 65% of total number of 

setigers. First branchia with 3 filaments; maximum 11 

filaments at setiger 15. From about setiger 40 number of 

filaments reduced rapidly and by setiger 60 2 filaments 

remaining. Last 20 branchiae single filaments. Posterior to 

setiger 50 filaments short and easily overlooked. Branchial 

stems slender, cylindrical, longer than notopodial cirri. Filaments 

filiform, longer than notopodial cirri. 

Anterior neuropodial acicular lobes broadly rounded; median 

and posterior acicular lobes asymmetrically rounded;

164 


FIGURE 53.—Eunice guildingi (holotype, BM(NH) 1839.12.27.5): a, anterior end, lateral view; b, subacicular 

hook, parapodium 37; c, compound falciger, parapodium 37; d, pectinate seU, parapodium 37; e, parapodium 37, 

anterior view. Eunice guttata (holotype, BM(NH) ZB 1972.71): f, anterior end, lateral view; g, parapodium 34, 

anterior view; h, compound falciger, parapodium 34; i, pectinate seta, parapodium 34; j, subacicular hook, 

parapodium 34. (Scale bars in mm.) 

aciculae emerging above midline. Presetal lobes low, transverse 

folds. Anterior postsetal lobes following outline of 

acicular lobes closely, becoming low transverse folds in 

median and posterior setigers. First 4 ventral cirri tapering. 

Ventral cirri basally inflated from about setiger 5 through 

remainder of body. Bases ovate, narrow tips tapering. Anterior 

notopodial cirri digitiform, becoming slightly inflated medially 

in anterior branchial segments. Notopodial cirri decreasing in 

length from about setiger 5 to setiger 20, thereafter short, 

slender, digitiform. Notopodial cirri without articulations. 

Pectinate setae (Figure 53d) flat, flaring. One marginal tooth 

distinctly longer than other teeth, with -10 coarse teeth. Shafts 

of compound falcigers (Figure 53c) distally slightly inflated, 

marginally frayed or serrated. Appendages short, tapering with 

very large head, bidentate. Proximal teeth shorter than distal 

teeth, tapering, directed obliquely distally. Distal teeth very 

nearly erect, tapering. Guards symmetrically bluntly pointed, 


and compound spinigers absent Notopodia supported by 

paired aciculae. Neuropodial aciculae single, black, tapering, 

distally straight; cross-sections round. Subacicular hooks 

(Figure 53b) black, bidentate. Hooks first present from setiger



replacements). Hooks tapering to small heads; both teeth 

directed distally and similar in size. 









None. 

REMARKS.—The holotype was examined by Arthur Willey, 

who referred it to E. floridana. Baird stated that the posterior 

half of the type was abranchiate; as indicated above, this is not 

the case: about 80% of the body is branchiated. 

Eunice guildingi has been reported only once. It is listed in 

Tables 27 and 31. Two species listed in Table 31 have the bases 

of the ventral cirri inflated to near the posterior end. Eunice 

bornecnsis has articulated peristomial cirri reaching the middle 

of the peristomium; E. guildingi has peristomial cirri without 

articulations reaching the middle of the prostomium. 

Leodice gunnertStorm. 1881:92-95. 

"Eunice" gunneri (Storm, 1881) 

REMARKS.—Three species of Nereis were described in the 

late 1700s from the ahermatypic coral reefs of Norwegian 

fjords; two by O.F. Miiller, Nereis pennata and Nereis pinnata, 

were found in the Oslofjord near DrCbak. The third species, 

named by Linnaeus on material sent him by Gunnerus from 

Trondheims-fjorden, was called Nereis norvegica. 

The characterization of these species were adequate for the 

time, and especially as illustrated by Gunnerus (1768), N. 

norvegica appears to be well characterized as having branchiae 

starting at about setiger 6 and continued to the far posterior end. 

Nereis pinnata and N. norwegica (note spelling) were 

transferred to Eunice by Cuvier (1817:525), who appears to 

have overlooked N. pennata. Lamarck (1818:323) synonymized 

N. pennata with N. norwegica under the name 

Leodice norwegica and maintained L. pinnata as a distinct 

species. Orsted (1845:402 and 406) complicated the issue by 

reporting a single species, which he called E. norvegica, from 

the Oslofjord and described it as having branchiae limited to 

the anterior end and light-colored aciculae and subacicular 

hooks. 

Storm (1881) recognized the presence of two species 

associated with the ahermatypic reef at Statsbygd, Trondheimsfjorden, 

Norway. One species, which he, in accord with 

Orsted, called Leodice norvegica, had branchiae from setiger 7 

and limited to the anterior end, with up to 17 branchial 

filaments where best developed. The other species, which he 

newly described as Leodice gunneri, had branchiae from setiger 

6 and continued to the posterior end, with up to nine branchial 

filaments where best developed. The aciculae and subacicular 

hooks were described as black. This description matches the 

original description of N. norvegica by Linnaeus (1767) and the 

illustration given of that species by Gunnerus (1768) and 

cannot match either of Muller's two species. 

Storms name, Leodice gunneri (= Eunice gunneri), is here 

considered a junior synonym of Eunice norvegica. No material 

of E. gunneri identified by Storm exists in the collections in 

Trondheim or Oslo, Norway (Tor StrOmgren, Inger Winsnes, in 

litt.). 

The name Eunice pennata (Miiller, 1776) is here, as in most 

of the literature, applied to a species with branchiae limited to 

the anterior end and light-colored aciculae and subacicular 

hooks (a member of group Al, see below). Muller's second 

species, Nereis pinnata, is unidentifiable (see below). 

Eunice guttata Baird, 1869:350. 

82. Eunice guttata Baird, 1869 


MATERIAL EXAMINED.—Holotype, BM(NH) ZB 1972.71, 

taken between Bombay and Singapore, Indian Ocean. 




Prostomium (Figure 530 distinctly shorter and narrower 

than peristomium, as deep as l /2 of peristomium. Prostomial 

lobes frontally obliquely truncate, dorsally flattened; median 

sulcus deep. Eyes not observed. Antennae in shallow horseshoe, 

evenly spaced, similar in thickness. Ceratophores 


long, tapering, without articulations. A-I to posterior edge of 

peristomium; A-II to setiger 2; A-III to setiger 3. Peristomium 

slightly tapering anteriorly. Separation between rings distinct 

only dorsally; anterior ring 4 /s of total peristomial length. 

Peristomial cirri to middle of anterior peristomial ring, 







of setigers. Where best developed branchiae distinctly pectinate 

with up to 10 filaments. Branchial stems nearly digitiform, 

longer than notopodial cirri. Filaments slender, digitiform, as 

long as or shorter than notopodial cirri. 

Neuropodial acicular lobes conical or depressed conical; 

aciculae emerging at midline. Presetal lobes low, transverse 

folds. Anterior postsetal lobes rounded, flattened, as high as 

acicular lobe, becoming reduced to low, transverse folds in 

median and posterior setigers. Anterior ventral cirri tapering. 

Ventral cirri basally inflated from about setiger 20; ventral cirri


remaining inflated through rest of body. Basal inflation very 

large, nearly spherical, very small tapering tips present in all 

setigers. Notopodial cirri basally inflated, distally tapering, 


Limbate setae tapering, gently geniculate, frayed. Pectinate 

setae (Figure 53i) numbering 5 or more in a fascicle. Shafts 

wide, flat Blades tapering, flat. One marginal tooth clearly 


falcigers (Figure 53h) distally inflated, marginally smooth. 

Appendages very slender, tapering, with large heads, bidentate. 

Proximal teeth about as large as distal teeth, sharply directed 

laterally. Distal teeth flattened, nearly geniculate and directed 

laterally. Guards asymmetrically bluntly pointed, marginally 


spinigers absent Aciculae numbering 1 to 3, dark, 

tapering, straight; cross-sections round. Notopodial aciculae 

paired, dark. Subacicular hooks (Figure 53j) dark, bidentate. 


thereafter, paired in some setigers. Hooks smoothly tapering to 

small head. Proximal teeth larger than distal teeth; both teeth 

directed obliquely distally. Guards short and blunt. 







Characters: 13, 14, 39,40,42, 74, 78. 


None. 

REMARKS.—The specimen may have been highly colored; a 

ghost of dark pigmentation remains. The specimen may have 

been dark brown or red, with white circular spots scattered 

densely over the anterior end. A white band may have been 

present. 

As far as known, this species has never been reported since 

it was briefly described by Baird (1869); it belongs to group 

B-2 and is listed with similar species in Tables 27 and 28. It is 

the only species, other than E. aphroditois, listed in Table 28 

that has the ventral cirri basally inflated through the posterior 

end. It differs from that species mainly in that it has far fewer 

branchial filaments and in the proportions of the prostomium. 

Nothing is known about its habitat other than that it was 

collected somewhere in the Indian Ocean/Indonesian region, 

presumably in shallow water. 

83. Eunice harassii Audouin and Milne Edwards, 1833 


Eunice harassii Audouin and Milne Edwards, 1833:215-218, pi. 11: figs.5-7, 

10, 11; 1834:141-144, pi. 3: figs. 5-7. 10, 11.—Quatrefages, 1866:307- 

309, pi. 10: fig. 3. 

Eunke pimctata.—Heider, 1925:55-89, figs. 1, 2, 7-16 [not Leodice 

punctata Risso, 1826]. 

MATERIAL EXAMINED.—One specimen, MNHN, Paris 

A.l(R.)-1868-no. 46a. lies Chausey, Quatrefages. 

COMMENTS ON MATERIAL EXAMINED.—The type has been 

lost; justification for using the current specimen to represent the 

species is given below. The jaws are half everted and the 

maxillary formula given is incomplete because detailing the 

formula would destroy the jaws completely. 

DESCRIPTION.—Specimen incomplete, of unknown sex, 

with 63 setigers; length 22 mm; maximal width 2.5 mm at 

setiger 10; length through setiger 10, 5 mm. Body cylindrical. 

Prostomium less than x li as thick as peristomium, narrow. 




sulcus deep. Eyes at bases of A-I, faded. Antennae in shallow 

horseshoe, evenly spaced, similar in thickness. Ccratophorcs 


tapering, with up to 5 indistinct, cylindrical articulations in 

A-I 11. A-I to posterior pcristomial ring; A-I I to setiger 2; A-I 11 

to setiger 4. Peristomium cylindrical. Separation between rings 

distinct on all sides; anterior ring 4 /s of total pcristomial length. 

Pcristomial cirri to middle of anterior pcristomial ring, slender 

and tapering, without articulations. 

Incomplete maxillary formula 1 + 1, +, 6+0, 3+8, 1+1. Mx 

VI missing. Mx III long, but part of distal arc with left Mx IV. 



Branchiae from setiger 4 to end of fragment. First 3 branchiae 

single filaments; all others pectinate with maximally 10 

filaments at about setiger 15; last branchia present, on setiger 

63, with 2 filaments only. Stems tapering, shorter than 

notopodial cirri. Filaments long, digitiform, as long as 

notopodial cirri where best developed. 

Anterior neuropodial acicular lobes truncate; posterior 

acicular lobes triangular; aciculae emerging at midline. Pre-and 


tapering. Ventral cirri basally inflated from setiger 6 through 

about setiger 35; inflated base ovate; narrow lips tapering. 

Posterior ventral cirri broadly transversely attached; enfolding 

lower edge of parapodia, nearly scoop-shaped. Anterior 

notopodial cirri basally slightly inflated, retaining same size in 

all setigers, but less distinctly inflated in posterior setigers, 



tapering, flat. One marginal tooth longer than other teeth, with 

-12 teeth. Shafts of compound falcigers (Figure 54c) distally 

tapering, marginally smooth. Appendages relatively short, 

tapering with large, distinct heads, bidentate. Proximal teeth as 

large as distal teeth, triangular, directed laterally. Distal teeth 

distinctly bent, sharply tapering. Guards asymmetrically 

bluntly pointed, marginally smooth; mucros absent. Pseudocompound 


paired; anterior aciculae honey colored, becoming darker 

brown in posterior setigers, distally tapering to sharp points,


FIGURE 54.—Eunice harassii (MNHN, Paris, A.l(R.)-1868-no. 46a): a, anterior end, lateral view; b, subacicular 

hook, parapodium 46; c, compound falciger, parapodium 46; d, parapodium 46, anterior view. Eunice havaica 

(syntype, RM 430): e, appendage of compound falciger, median parapodium; f, compound falciger, median 

parapodium; g, subacicular hook, median parapodium; h, anterior end, lateral view. (Scale bars in mm.) 

straight; cross-sections round. Separation between core and 


Subacicular hooks (Figure 54b) light brown, bidentate. Hooks 


always single (except for replacements). Hooks with distinct 

neck. Proximal teeth large, curved, directed laterally. Distal 

teeth bluntly triangular, nearly erect. 


anal cirri. 


None. 



Characters: 1, 2, 36-38,40. 

and 


38,1. 

REMARKS.—Eunice harassii was described from lies 

Chausey and St. Malo. Audouin and Milne Edwards 

(1833:215) mentioned that the species was relatively common 

in these areas. Quatrefages (1866:309) mentioned that he 

described the species based on a specimen reported from lies 

Chausey by Audouin and Milne Edwards. The specimen 

described above appears to be the specimen used by Quatrefages 

(1866) in his review and thus potentially part of the 

original material. At the very least, the specimen comes from 

one of the two type areas and may be part of the type material. 

It is here considered an authoritative specimen, but not as a 

type. 

Heider (1925) gave an excellent description of the anatomy 

of the anterior end of a species he called E. punctata (Risso,


1826); he cited E. harassii as a synonym. Risso's species is not 

identifiable (see below); Heider's description fits E. harassii, 

thus his detailed anatomical work is assumed to be of this 

species. 

Eunice harassii is a member of B-2, but differs from most 

species in that group by having brown aciculae and subacicular 

hooks rather than black ones as usual in that group. It is listed 

with similar species in Tables 27 and 31. One unique character 

state of this species appear to be the broadly attached, flattened, 

nearly leaf-shaped ventral cirri in posterior setigers. The 

detailed branchial distribution may also be characteristic, in 

that the number of filaments is reduced from middle segments, 

but branchiae are retained through the rest of the body. 

84. Eunice havaica Kinberg, 1865 

FIGURE 54e-g; TABLES 46-48 

Eunice havaica Kinberg, 1865: 562; 1910:41, pi. 15: fig. 14. 

Eunice antennata.—Hartman, 1948:78 [in part, not Eunice antennata Lamarck, 

1818]. 

MATERIAL EXAMINED.—14 syntypes, RM 430, Honolulu, 

Hawaii, Eugenie Expedition 1087-89. 

COMMENTS ON MATERIAL EXAMINED.—All but one of the 

15 specimens of the original material of E. havaica belong to 

this species; the last one differs from the others at the specific 

level and is described above as E. eugeniae. All the syntypes 

are poorly preserved and most are anterior fragments only. Due 

to the poor preservation parapodial shape can be described only 

tentatively. 

DESCRIPTION.—Complete syntype of unknown sex, with 78 

setigers; total length 25 mm; maximal width 1 mm at setiger 

10; length through setiger 10,4 mm. 


about as wide as peristomium, less than l /2 as deep as 

peristomium. Prostomial lobes frontally obliquely truncate, 

dorsally flattened; median sulcus shallow. Eyes barely visible 

well behind bases of A-I. Antennae in horseshoe, evenly 


antennae, without articulations. Ceratostyles tapering, with up 

to 22 moniliform articulations in A-III, articulated to bases. A-I 

to middle of first peristomial ring; A-I I to setiger 2; A-I 11 to 

setiger 3. Peristomium cylindrical. Separation between rings 

distinct on all sides; anterior ring 3 A of total peristomial length. 

Peristomial cirri to frontal margin of peristomium, slender, with 

9-10 moniliform articulations. 

Summary maxillary formula for 2 syntypes 1+1, 7+7, 7+0, 

6-8+8, and 1+1. Max III long, located behind left Mx II. Mx VI 

missing. 


cirri, reduced in mid-body region, erecL Branchiae from setiger 

6 to setiger 78. Branchiae present to near posterior end, on more 

than 65% of total number of setigers. In complete syntype 

branchial occurrence interrupted between setigers 28 and 48 

leaving median segments abranchiate. Maximum number of 

filaments 5 at about setiger 20; maximum number of filaments 

in posterior branchiated region 3. Stems slender, tapering, 

shorter than notopodial cirri. Filaments slender, digitiform, 


Neuropodial acicular lobes distally apparently truncate; 

aciculae emerging at midline. Prc- and postsetal lobes low, 

transverse folds. Anterior and posterior ventral cirri digitiform, 

basally inflated between setigers 5 and 40. Inflated bases ovate; 

narrow tips tapering. Notopodial cirri tapering, with up to 6 

articulations at least in first 25 setigers; articulations indistinct 

posteriorly; far posterior notopodial cirri without articulations. 

Limbate setae slender. Pectinate setae flat, tapering. Both 

marginal teeth slightly longer than other teeth, with -15 teeth. 

Shafts of compound falcigers (Figure 540 distally inflated, 

usually marginally smooth. Appendages short with large head, 

bidentatc or tridcntatc (Figure 54c). Proximal and distal teeth of 

same size. Proximal teeth narrowly triangular, directed laterally. 

Distal teeth distinctly bent, split in some hooks, forming 

distinctly tridcntatc appendages. Guards symmetrically bluntly 

pointed; mucros absent. Pscudocompound falcigers and compound 

spinigers absent. Aciculae yellow, tapering to bluntly 

conical tips, straight; cross-sections round. Separation between 


hooks. Subacicular hooks (Figure 54g) yellow, tridentate with 

teeth in a crest. Hooks first present from setiger 19-23, present 

in all setigers thereafter, always single (except for replacements). 

Main fang large, curved. Smaller fangs decreasing 

evenly in size. 


anal cirri. 


None. 





None. 

REMARKS.—Eunice havaica was considered a synonym of 

E. antennata by Hartman (1948). It is listed in Tables 46 and 

48. Of the species listed in Table 48 only E. havaica and E. 

stigmatura have complete cinctures marking off the separation 

between the two peristomial rings. The compound falcigers 

have both bi- and tridentate appendages in E. havaica and only 

bidentate appendages in E. stigmatura. 

Hartman (1948:78) indicated that she had seen only one of 

the syntypes of this species, but later indicated a range of values 

for first occurrence of subacicular hooks, showing that she must 

have examined at least several specimens. The specimens are 

very soft, making study of variability difficult. 

and


85. Eunice hawaiensis Treadwell, 1906 

FIGURE 55»-f; TABLES 19,21 

Eunice hawaiensis Treadwell, 1906: 1166-1167, figs. 42-44. 

Eunice hawaiiensis.—Fauchald, 1970:206 [note misspelling of species name]. 


sta 4134, off Hawaii Island, 225-324 fathoms, fine coral and 

volcanic sand. 

COMMENTS ON MATERIAL EXAMINED.—The lower half of 

the peristomium and first setiger have been sliced off; the cut is 

indicated by a dashed line in the illustration. The specimen is 

otherwise in good condition; it is in part covered with a loose 

mucoid-appearing material and may have been tubicolous in 

life. 


118 sciigcrs; length 115 mm; maximal width 7 mm at setiger 

10; length through setiger 10, 13 mm. 


than peristomium, as deep as x li of peristomium. Prostomial 

lobes frontally rounded, dorsally slightly inflated; median 

sulcus deep. Palpal areas set off by frontal, horizontal grooves. 

Eyes between bases of A-I and A-II, brown. Antennae in 

shallow horseshoe, evenly spaced; A-I thinner than A-II and 

A-I 11. Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostyles slender and tapering, without articulations. 

A-I to middle of anterior peristomial ring; A-II to 

setiger 5; A-I 11 to setiger 8. Peristomium apparently cylindrical, 

considerably wider than prostomium. Separation between rings 

barely visible dorsally, indeterminable ventrally; anterior ring 

-*h of total peristomial length. Peristomial cirri to front edge of 

peristomium, tapering to slender tips, without articulations. 

Maxillary formula 1+1, 8+9, 8+0, 12+12, 1+1 according to 

Treadwell (1906:1167). 



Branchiae from setiger 3 to 54. Branchiae terminating well 

before posterior end. Last 4 pairs single filaments; all other 

branchiae strongly pectinate with up to 30 filaments. Stems 

strong, erect, tapering, longer than notopodial cirri. Filaments 

slender, filiform, shorter than notopodial cirri. 

Anterior neuropodial acicular lobes distally rounded, somewhat 

more pointed in last setigers present; aciculae emerging at 

midline. Pre- and postsetal lobes low, transverse folds. First 3 

ventral cirri thick, tapering, becoming basally inflated in 

branchial region. Inflated bases ovate; narrow tips long, 

tapering. Postbranchial ventral cirri less distinctly inflated 

basally, becoming thick and tapering in last setigers present. 

Notopodial cirri distinctly inflated basally, tapering to slender, 

digitiform tips, similar in shape, but shorter in postbranchial 

region. Notopodial cirri without articulations. 

Limbate setae long, slender. Pectinate setae (Figure 55e) 

very short, slender, tapering, flat. One marginal tooth distinctly 

longer than other teeth, with -12 very long, slender teeth. 

Shafts of compound falcigers (Figure 55f) distally inflated, 

marginally smooth. Appendages long, narrow, with nearly 

parallel sides and small head, bidentate. Proximal teeth smaller 

than distal teeth, reduced triangular, directed laterally. Distal 

teeth erect, tapering. Guards symmetrically sharply pointed 

with short, slender mucros, marginally serrated. Pseudocompound 


(Figure 55c) paired, yellow. Inferior aciculae blunt or truncate, 

slightly curved. Superior aciculae tapering to blunt tips, 



Subacicular hooks (Figure 55d) yellow, bidentate. Hooks first 


single (except for replacements). Hooks tapering to very small 

heads. Proximal teeth larger than distal teeth, tapering, directed 

laterally. Distal teeth very short, thick, sharply tapered. Guards 

short and truncate. 


Mx III and left Mx IV; features associated with far 

posterior setigers; pygidium and anal cirri. 





Characters: 1,2,24, 25, 38,42,47, 50, 63. 


REMARKS.—Eunice hawaiensis was considered, incorrectly, 

to be a member of group C-l by Fauchald (1970), who also 

misspelled the species name; it is in fact a member of group 

A-l and closely resembles E. megabranchia and E. validobranchiata. 

These and related species are listed in Tables 19 and 21. 

Eunice hawaiensis has branchiae ending at setiger 54, with up 

to 30 filaments, mucronate compound falcigers and the 

notopodial cirri lack articulations throughout the body. Eunice 

megabranchia has branchiae ending at setiger 54, up to 47 

branchial filaments, mucronate compound falcigers and the 

notopodial cirri are articulated in anterior setigers. The 

separation between the peristomial rings is barely visible 

dorsally in E. hawaiensis and forming a distinct cincture in E. 

megabranchia. Eunice validobranchiata has branchiae through 

setigers 37-40 with up to 33 filaments; amucronate compound 

falcigers and the notopodial cirri lack articulations throughout 

the body; the separation between the peristomial rings is 

indistinct laterally in this species. 

86. Eunice heterochaeta Quatrefages, 1866 

FIGURE 55g-l; TABLES 19,21 

Eunice heterochaeta Quatrefages. 1866:314, pi.10: fig. 3.—Grube, 1870a:295- 

-296. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, A.I (R.)- 

1868-no. 48a, Guettary, Quatrefages. 

DESCRIPTION.—Holotype complete, of unknown sex, with

170 


02 

i 

I 

2 

.1 

e. t. h i 

b 

i 

c a. i k 

FIGURE 55.—Eunice hawaiensis (holotype, USNM 5210): a, anterior end, lateral view; b, parapodium 34, anterior 

view; c, aciculae, parapodium 34; d, subacicular hook, parapodium 34; e, pectinate seta, parapodium 34; f, 

compound falciger, parapodium 34. Eunice helerochaela (holotype, MNHN, Paris A.l(R.)-1868-no. 48a): g, 

anterior end, lateral view; h, compound falciger, parapodium 28; i, pectinate seta, parapodium 28; j, subacicular 

hook, parapodium 50; k, aciculae, parapodium 28; 1, parapodium 28, anterior view. (Scale bars in mm.)



setiger 10; length through setiger 10, 4 mm. Body cylindrical 

throughout. 




sulcus shallow. Eyes absent. Antennae in horseshoe, evenly 

spaced; A-I slimmer than A-II and A-III. Ceratophores 


slender and tapering, with up to 5 long, cylindrical articulations 

in A-III. A-I to posterior peristomial ring; A-II to setiger 1; 

A-III to setiger 5. Peristomium very much wider than 

prostomium, with distinct, muscular lower lip. Separation 

between rings distinct dorsally and ventrally; anterior ring 2 h of 

total peristomial length. Peristomial cirri to middle of anterior 

peristomial ring, slender and tapering, without articulations. 

Jaws missing. 




before posterior end, present on more than 65% of total number 

of setigers. First 3 and last 15 branchiae single filaments, 

maximum 7 filaments at about setiger 25. Stems short, 

tapering, shorter than filaments. Filaments slender, as long as 

notopodial cirri in all except first 2 and last 10 branchial 

setigers. 

Neuropodial acicular lobes truncate to rounded; aciculae 

emerging at midline. Pre- and postsctal lobes low, transverse 

folds. First 4 ventral cirri tapering, digitiform, becoming 

basally inflated from setiger 5. Inflated bases thick, short 

transverse welts, lost by setiger 30; narrow tips tapering. 

Posterior ventral cirri tapering, becoming increasingly digitiform 

in last setigers present. Anterior notopodial cirri basally 

inflated, with 3-4 long, cylindrical articulations, losing 

articulations in branchial region, retaining inflated bases in all 

setigers. Limbate setae slender, marginally smooth. Pectinate 

setae (Figure 55i) tapering, flat. One marginal tooth slightly 

longer than other teeth, less than 10 teeth present. Shafts of 

compound falcigers (Figure 55h) tapering, marginally smooth; 

beaks indistinct. Appendages tapering; heads large, bidentate. 

Proximal teeth about twice as large as distal teeth, triangular. 

Distal teeth distinctly curved, tapering. Guards symmetrically 


falcigers and compound spinigers absent. Usually 

3 or more aciculae (Figure 55k) in anterior or medial parapodia, 

yellow, slender, gently curved or bent; cross-sections round. 


and subacicular hooks. Subacicular hooks (Figure 55j) yellow, 



Hooks slender, tapering, curved. Proximal teeth about twice as 

large as distal teeth; both teeth directed laterally. 




None. 





None. 

REMARKS.—Quatrefages original description and illustrations 

are inadequate to characterize the species. Grube 

(1870a:295-296) gave some detail, but saw no reason to retain 

the species and referred it to E. pennata, despite the fact that 

branchiae are continued to setiger 85 and not terminated at 

setiger 39 as in the latter. Fauvel (1923:400, 451) referred E. 

heterochaeta to E. pennata apparently under some doubt. 

Eunice heterochaeta clearly belongs to group A-l and is 

listed with similar species in Tables 19 and 21. Only one 

species in this list, E. rubrocincta, resembles E. heterochaeta in 

having better than 70% of the total numbers of setigers 

branchiated; most species in this group have less than 50% of 

the setigers branchiated. The antennae increase in length from 

A-I through A-III in E. heterochaeta; all antennae are similar in 

length and very short in E. rubrocincta. 

87. Eunice hirschi, new name 


Leodice articulata Hoagland, 1920:615-616, pi. 50: figs. 9-12. 

MATERIAL EXAMINED.—Holotype and 1 paratype of Leodice 

articulata Hoagland, USNM 18955, Albatross sta 5159, 

Tinakta Island, Philippines, 5°1l'50"N, 119°54'E, 22 Feb 1908, 

22 m, sand, dredged. One additional specimen of Leodice 

articulata Hoagland, USNM 19024, Albatross sta 5205, off 

Caguayan Point, Leyte Island, Philippines, 1F19'3O"N, 

124°58'05"E, 13 Apr 1908, 15 m, Agassiz trawl. 

COMMENTS ON MATERIAL EXAMINED.—The paratype from 

station 5205 has at one time been dry and no meaningful 

measures can be taken; it does not appear to differ from the two 

other specimens. 



setiger 10; length through setiger 10, 7 mm. Paratype from sta 

5159 incomplete, of unknown sex, with 71 setigers; length 37 




lobes frontally truncate, dorsally flattened; median sulcus deep. 

Eyes not observed. Antennae in a horseshoe, evenly spaced; 

A-III distinctly thinner than other antennae. Ceratophores 


very slender, tapering, articulated to bases, with up to 35 

moniliform articulations in A-III. A-I to setiger 1; A-II to 

setiger 4; A-III to setiger 6. Peristomium cylindrical, with 

distinct muscular lower lip. Separation between rings distinct

172 


FIGURE 56.—Eunice hirschi, new name (holotype of Leodice articulata Hoagland, USNM 18955): a, anterior 

end, lateral view; b, subacicular hook, parapodium 66; c, compound falciger, parapodium 66; d, parapodium 66, 

anterior view; e, subacicular hook, parapodium 102; f, compound falciger, parapodium 102; g, pectinate seta, 

parapodium 102; h, aciculae, parapodium 66; i, parapodium 102, anterior view. Eunice imogena (holotype of 

Nicidion imogena, BM(NH) ZK 1924.1.28.91): j. anterior end, lateral view; k, subacicular hook, parapodium 113; 

1, compound falciger, parapodium 113; m, pectinate seta, parapodium 113; n, parapodium 113, anterior view. 

(Scale bars in mm.)


dorsally and especially ventrally; anterior ring 3 A of total 

pcristomial length. Peristomial cirri to middle of prostomium, 

tapering, slender, with 10 articulations. 

Maxillary formula 1+1, 8+9, 9+0, 6+10, 1+1 according to 

Hoagland (1920, pi. 50: fig. 10). Teeth very small and 

indistinct. Mx III long, located behind left Mx II. 

Branchiae (Figure 56d) present, pectinate, distinctly shorter 


Branchiae from setiger 4 to setiger 102 (Figure 56i). Branchiae 

present to near posterior end, on more than 65% of total number 

of setigers. First and last 7 branchiae single filaments. 

Maximum 7 filaments from setiger 10 through setiger 30. From 

about setiger 35 through setiger 90, 3 filaments. Stems shorter 

than filaments, tapering. Filaments short, digiliform. 


emerging at midlinc. Pre-and postsetal lobes low, transverse 

folds. First 4 ventral cirri thick, tapering, basally inflated from 

about setiger 5. Inflated bases modest, ovate, reduced by setiger 

50; narrow tips tapering. Posterior ventral cirri increasingly 

long and slender, tapering, less than half as long as notopodial 

cirri. Notopodial cirri long, basally somewhat inflated, with 6 

articulations anteriorly, decreasing to 2 in far posterior setigers. 

Articulations moniliform in anterior setigers, becoming cylindrical 

in posteriormost setigers. 

Limbate setae longer than all other setae, slender, marginally 

serrated. Pectinate setae (Figure 56g) very small; shafts slender, 

cylindrical. Blades flaring, flat. One marginal tooth distinctly 

longer than other teeth, with 10 long, slender teeth. Shafts of 

anterior and median compound falcigers (Figure 56c) inflated, 

marginally serrated; distal beak absent. Anterior appendages 

very narrow, tapering; head small, bidentate. Proximal teeth 

reduced triangular, directed laterally. Distal teeth longer than 

proximal teeth, directed obliquely laterally. Shafts of posterior 

compound falcigers (Figure 560 inflated, but less so than in 

median setigers, marginally serrated; distal beak present 

Posterior appendages short, tapering; head distinct, bidentate. 

Proximal teeth large, triangular, directed laterally. Distal teeth 

smaller than proximal teeth, tapering, directed obliquely 

distally. Guards of all compound hooks asymmetrically bluntly 

pointed, marginally serrated in anterior and median hooks, 

smooth in posterior hooks; mucros absent. Pseudocompound 

falcigers and compound spinigers absent Aciculae (Figure 

56h) paired, yellow; cross-sections round. Inferior aciculae 

curved, bluntly pointed, about twice as thick as superior 

aciculae. Superior aciculae straight, sharply pointed. Separation 

between core and sheath indistinct in both aciculae and 

subacicular hooks. Subacicular hooks (Figure 56b,e) yellow, 

tridentate with teeth in a crest Hooks first present from setiger 

22 or 25, present in all setigers thereafter, always single (except 

for replacements). Proximal teeth large, directed laterally, 

straight; two distal teeth erect, emerging from common shaft. 

UNKNOWN 

anal cirri. 

MORPHOLOGICAL FEATURES.—Pygidium and 


None. 





None. 

REMARKS.—The specific name articulata is preoccupied in 

the combination Eunice articulata Ehlers, 1887, and must be 

replaced. 

Eunice hirschi is listed with similar species in Tables 46 and 

48. Among the species in Table 48, E. hirschi, E. panamena, 

and E. stigmatura lack a middle region with reduced branchiae; 

such a region is present in some specimens of E. ornata, but is 

missing in others. Among these species, E. panamena and E. 

stigmatura have the three median antennae similar in length; 

the other two species have A-III distinctly longer than A-II. In 

E. ornata the branchiae are distinctly longer than the 

notopodial cirri; in E. hirschi the branchiae are shorter than the 


The species is named for Dr. Leonard P. Hirsch for all his 

support and encouragement during this study. 

Eunice hispanica (Lamarck, 1818) 

Leodice hispanica Lamarck. 1818:323.—Savigny, 1820:51. 

Eunice hispanica.—Audouin and Milne Edwards. 1833:219; 1834:145.— 

Gnibe, 1850:292.—Quatrefages. 1866:324.—Fauvel, 1923:451. 

REMARKS.—This species was described as having smooth 

antennae and chestnut-colored aciculae; anterior notopodial 

cirri long; branchiae with up to three filaments present from 

setiger 3; only 15 or 16 pairs of branchiae present; branchiae 


Audouin and Milne Edwards (1833:219) suggested that the 

species was primarily characterized by the poor development of 

the branchiae. 

Grube (1850:292) listed the species as valid without 

comment. Quatrefages (1866:324) found it to be incompletely 

described. Fauvel (1923:451) listed it as incertae sedis without 

comment. 

No material is available; the precise origin of Lamarck's 

material (other than presumably Spain) is unknown and the 

available information is wholly inadequate to characterize the 

species. The species is here considered indeterminable. 

88. Eunice imogena (Monro, 1924) 

FIGURE 56j-n; TABLES 33,40 

Nicidion imogena Monro, 1924:61-62, figs. 22-24. 


1924.1.28.91, Hotspur Bank, off the coast of Brazil, 16°S, 

36°W. 


originally described as consisting of 108 setigers and being 25


mm long; the current specimen, labeled as type, is larger, as 

indicated below. It is unclear how this discrepancy has arisen: 

it is here assumed that a simple lapsus has taken place and that 

specimen here described in fact is the holotype. 



length through setiger 10, 3.5 mm. 

Prostomium (Figure 56j) distinctly shorter and narrower than 

peristomium, as deep as x li of peristomium. Prostomial lobes 

frontally rounded, somewhat flattened dorsally; median sulcus 

very deep. Eyes behind bases of A-I, kidney-shaped, black. 



Ceratostyles medially inflated with irregular notches, but 

without articulations. A-I to middle of anterior peristomial ring; 

A-II and A-III to posterior end of peristomium. Peristomium 

longer than prostomium, cylindrical. Separation between rings 

distinct on all sides; anterior ring ~ 2 h of total peristomial 

length. Peristomial cirri barely to front edge of posterior 


Maxillary formula 1+1,5+5, 7+0,6+9, and presumably 1+1; 

Mx V apparently absent in preparation (see comment below). 

Distal tooth of both Mx II with small subdistal notches, 

indicating possible presence of reduced sixth tooth. 


Anterior neuropodial acicular lobes short, distally rounded, 

becoming sessile in median and posterior setigers (Figure 56n); 

aciculae emerging above midline. Pre- and postsetal lobes low, 

transverse folds. Anterior ventral cirri digitiform, becoming 

basally inflated by setiger 10. Ventral cirri retaining inflated 

bases through rest of body, but inflated bases increasingly 

retracted into body wall. Narrow tips short and button-shaped 

in median setigers, becoming distinct and tapering in far 

posterior setigers. Anterior notopodial cirri tapering and about 

twice as long as ventral cirri, increasing in length through 

median setigers. Notopodial cirri reduced in length from about 

setiger 50, becoming short and tapering in far posterior setigers, 

located well above neuropodia on body wall. Notopodial cirri 


Pectinate setae (Figure 56m) tapering, flat. Both marginal 

teeth longer than other teeth; -10 teeth present. Shafts of 

compound falcigers (Figure 561) slender, distally strongly 

inflated, marginally strongly serrated; distinct beak present. 

Appendages very short, with nearly parallel sides; heads small, 

bidentate. Both teeth similar in size. Proximal teeth slender, 

tapering, directed laterally. Distal teeth tapering, directed 

laterally. Guards symmetrically rounded, marginally serrated; 


spinigers absent. Aciculae (Figure 56n) single, with black core 

and clear sheath, thick, distally tapering, straight; crosssections 

round. Subacicular hooks (Figure 56k) with black core 

and clear, nearly translucent heads, bidentate. Hooks first 


single (except for replacements). Hooks with distinctly bent 

necks; heads distinct. Proximal teeth larger than distal teeth, 

curved, directed laterally. Distal teeth triangular, tapering, 










None. 

REMARKS.—Eunice imogena is compared to similar species 

in Tables 33 and 40. It is unique among the abranchiate species 

in that subacicular hooks are first present from sctigcr 50; in the 

other species, subacicular hooks arc present from sctigcrs 

16-31. 

The maxillary formula was given by Monro (1924:62) in the 

following words: "formula as follows: 4-4:7+57-9." The 

illustration given by Monro (1924, fig. 24) shows the formula 

to be 1+1, 5+5, 7+0, 6+9. The jaws arc now missing; Monro's 

illustration suggests a less aberrant formula than the text and is 

here assumed to approximate the condition in the species. 

89. Eunice impexa Grube, 1878 

FIGURE 57i-m; TABLES 52,53 

Eunice impexa Grube, 1878b: 159-160. pi. 9: Tig. 6. 

MATERIAL EXAMINED.—Holotype, ZMB 4001, Cape of 

Lapinig, Philippines, coll. Semper. 

COMMENTS ON MATERIAL EXAMINED.—Four parapodia 

removed from the holotype were also available; two of these 

were previously examined by Grube, the other two by Augener. 



10; length through setiger 10, 11 mm. 


distinctly narrower than peristomium, less than x li as 

deep as peristomium. Prostomial lobes frontally rounded, 

dorsally flattened; median sulcus shallow. Eyes present 

posterior to bases of A-I, faded purple. Antennae in horseshoe, 



with 1 or 2 irregular, cylindrical articulations. A-I to anterior 

peristomial ring; A-II to posterior peristomial ring; A-III to 

setiger 1. Peristomium massive, expanded anteriorly due to 

eversion of jaws; muscular lower lip present. Separation 

between rings distinct ventrally and visible dorsally; anterior 

ring 3 /4 of total peristomial length. Peristomial cirri to middle of 

anterior peristomial ring, digitiform, without articulations. 


short, forming a distal arc with very short left Mx IV and MX 

V. Mx VI absent.


FIGURE 57.—Eunice impexa (holotype, ZMB 4001): a, anterior end, lateral view; b, pectinate seta, parapodium 

45; c, compound spiniger, parapodium 15; d, adcula, parapodium 15; e, parapodium 15, anterior view; f, inferior 

subacicular hook, posterior parapodium; g, median subacicular hook, posterior parapodium; h, superior 

subacicular hook, posterior parapodium; i, parapodium 45, anterior view; j, notopodium and branchia, 

parapodium 60-70; k, notopodium and branchia, parapodium 65, drawn in situ; 1, pectinate seta, posterior 

parapodium; m, compound falciger, posterior parapodium. Eunice indica (holotype, RM 435): n, anterior end, 

lateral view; o, subacicular hook, parapodium 27; p, compound falciger, parapodium 27; q, compound falciger, 


176 


Branchiae (Figure 57jjc) present, palmate, distinctly longer 


Branchiae from setiger 22 to end of fragment. First 10 

branchiae short, button-shaped. From setiger 31 branchiae 

long, with slender filaments. Branchial distribution irregular 

with most branchiae having 4-5 filaments clustering at tip of 

short stems. Several branchiae with 6 to 7 long filaments and up 

to 3 short, truncate filaments emerging near bases. Some 

branchiae with filaments in 2 groups emerging from common 

stems. Relation between length of filaments and branchiae and 

notopodial cirri variable. 

Neuropodial acicular lobes (Figure 57e,i) distally obliquely 

rounded with aciculae emerging above midline. Anterior 

presetal lobes high dorsally, forming rounded tabs visible 

dorsal to aciculae. At level of aciculae presetal lobes deeply cut 

away leaving bases of compound hooks exposed. Median and 

posterior presetal lobes low, transverse folds. Postsetal lobes 

low folds following outline of acicular lobes. First 4 ventral 

cirri thick, tapering. Ventral cirri basally inflated from setiger 5. 

Inflated bases thick; prominent ventrolateral ridges in all later 

setigers. Narrow tips short and button-shaped in median 

setigers, tapering in posterior setigers. Notopodial cirri short, 

basally distinctly inflated, tapering towards slender tips, 


Limbate setae stout, tapering, marginally smooth. Pectinate 

setae (Figure 57b,l) tapering, furled or funnel-shaped. One 

marginal tooth distinctly longer than other teeth; -12 teeth 

present. First 30 setigers with compound spinigers (Figure 

57c). Shafts distally inflated, marginally serrated. Appendages 

tapering, knife-shaped. Compound falcigers (Figure 57m) 

replacing compound spinigers from about setiger 30, considerably 

smaller than spinigers. Shafts inflated, marginally serrated. 

Appendages short, narrow, with nearly parallel sides, 

with small, distinct heads, bidentate. Proximal teeth larger than 

distal teeth, narrow, tapering, curved, directed laterally. Distal 

teeth gently curved, directed obliquely distally. Guards longer 

than appendage, symmetrically rounded, marginally smooth; 

mucros absent Pseudocompound falcigers absent. Aciculae 

(Figure 57d) single or paired, with chestnut to copper-colored 

cores and clear sheaths, tapering, gently curved dorsally; 

cross-sections round. Separation between core and sheath 

distinct in both aciculae and subacicular hooks. Subacicular 

hooks (Figure 57f-h) honey colored anteriorly, becoming 

darker posteriorly, but always lighter in color than aciculae, 

bidentate or tridentate, the latter with teeth in a crest Hooks 

first present from setiger 60, present in all setigers thereafter; 

three hooks in most setigers. Most hooks tapering distally 

towards tip with distinct inflated subdistal region. Proximal 

teeth large, directed laterally. Distal teeth short, nearly erect. 

Some hooks distally tridentate with small distal, medium-sized 

intermediate tooth and large main fangs. 


anal cirri. 


None. 

and 



Characters: 1-2, 36-38,40, 42,47, 50, 63. 


38,1. 

REMARKS.—Eunice impexa is listed with other species with 

compound spinigers in Tables 52 and 53. It has articulated 

ceratostyles, in contrast to the two other species, E. tubicola 

and E. tubifex. 

Eunice incerta (Hansen, 1882) 

Nicidion incerta Hansen. 1882:8, pi. 2: figs. 19-21. 

Eunice (Nicidion) incerta.—Hartman. 1959:313. 

REMARKS.—An examination of the type material shows that 

this species is a junior synonym of E. cariboea and is treated as 

such above. 

90. Eunice indie a Kinberg, 1865 

FlOURE 57n-q; TABLES 41, 42 

Eunice indica Kinberg. 1865:562; 1910:41, pi. 15: fig.12. 

MATERIAL EXAMINED.—Holotype, RM 435, Bangka Strait, 

Indian Ocean. Kinberg (1865:562) gives locality information 

as: "Pars australis Freti Bangka, in fundo maris." 

COMMENTS ON MATERIAL EXAMINED.—Holotype is now 

incomplete, in three pieces, but is reasonably well preserved. A 

posterior end in the same vial may belong to the same 

specimen, but association is not unequivocal due to the 

different state of preservation of the posterior end. 


38 setigers; length 11 mm; maximal width 2 mm; length 

through setiger 10,4 mm. 

Prostomium (Figure 57n) distinctly shorter than peristomium, 



flattened; median sulcus shallow. Eyes posterior to bases of 

A-I, dark. Antennae in horseshoe; A-I separated from A-II and 

A-III by gap, similar in thickness. Ceratophores ring-shaped in 


tapering, with up to 14 regular, cylindrical, long articulations in 

A-III. A-I to posterior edge of peristomium; A-II to setiger 4; 

A-III to setiger 14. Peristomium tapering slightly anteriorly 

(possibly distorted by removal of jaws). Separation between 

rings distinct on all sides; anterior ring 2 /3 of total peristomial 

length. Peristomial cirri to middle of prostomium, tapering, 


Jaws missing. 



from setiger 3 to 21. Branchiae terminating well before 


setigers. First branchiae single filaments; all other branchiae 

pectinate; maximum 8 filaments. Stems tapering, erect.


Filaments slender. 

Neuropodial acicular lobes distally rounded; aciculae emerging 


First 2 ventral cirri thick, tapering. Ventral cirri basally inflated 

in next 20 setigers. Inflated bases ovate; narrow tips tapering. 

From about setiger 23 inflated bases decreasing. Posteriormost 

ventral cirri tapering evenly. Notopodial cirri medially inflated, 

becoming slender and digitiform in posterior setigers, without 


Limbatc setae slender, marginally serrated. Shafts of anterior 

compound falcigers (Figure 57m,n) tapering evenly, becoming 

distally inflated in median setigers, marginally smooth. 

Appendages very long and slender, especially in anterior 

setigers, bidentate. Anterior appendages with proximal teeth 

very much smaller than distal teeth, forming a small, triangular 

lateral projection. Distal teeth nearly erect, tapering. In median 

and posterior appendages proximal teeth larger, triangular, 

usually directed basally. Distal teeth distinctly curved, directed 

laterally. All guards tapering to fine slender tips, mucronate. 

Guards very long, especially in anterior setigers. Pseudocompound 


single, yellow, tapering, distally blunt, straight or gently 

curved; cross-sections round. Separation between core and 


Subacicular hooks (Figure S7o) yellow, tridentate with teeth in 


setigers thereafter, at first single, but all posterior setigers with 

at least 3, usually 4 hooks. Main fangs large, curved; median 

fang similar in shape but smaller, distal fang very small, but 

usually distinctly pointed. 

UNKNOWN MORPHOLOGICAL FEATURES.—Jaw structure; all 

features associated with posterior setigers; pygidium and anal 

cirri. 





Characters: 1, 2,4, 6,42,47, 50, 63,65-68. 


None. 

REMARKS.—Eunice indica is listed with similar species in 

Tables 41 and 42. In addition to E. indica, E. tentaculata and E. 

vittata have multiple subacicular hooks in most setigers. Eunice 

indica and E. vittata both have the first branchia on setiger 3; E. 

tentaculata has the first branchia on setiger 6. Eunice indica has 

mucronate guards on the compound falcigers; E. vittata has 

distally blunt guards. 

Kinberg indicated that A-III should reach setiger 19; this 

cannot be confirmed and would be unusually long, even for this 

species. 

Eunice indica has been widely reported from Indian Ocean 

and western Pacific Ocean; a number of similar species have 

been reported from this area, and the distribution of E. indica 

proper must be considered unsettled. 

91. Eunice interrupta Treadwell, 1906 

FIGURE 58a-e; TABUES 41,44,46,47 

Eunice interrupta Treadwell. 1906:1167-1168, figs. 45, 46. 

MATERIAL EXAMINED.—Holotype, USNM 5211, and 5 

paralectotypes, USNM 5417, Albatross sta 3850, 79-84 m, 

coarse sand and broken shells; 1 paralectotype, USNM 5418 

and 1 paralectotype, CAS TY74 (IZ No. 018795), Albatross sta 

3940, Hawaii, Leeward lands, Laysan Island Light S84°, E7.0', 

16 May 1902, 108-128 m, white sand and broken shells. 

COMMENTS ON MATERIAL EXAMINED.—Both paratypes 

from station 3940 have been dried out and are not considered 

further. The description is based on the holotype, but comments 

about variable features are included. 



through setiger 10,9 mm. Paratypes from sta 3850 2 complete 

specimens with 79 and 69 setigers, respectively; other 

paratypes incomplete. 



lobes frontally rounded, dorsally flattened; median sulcus deep. 

Eyes at base of A-I, dark. Antennae in horseshoe, evenly 

spaced, similar in thickness. Ceratophores long in all antennae, 


moniliform articulations in A-I I and A-III, articulated to bases 

of styles. A-I to middle of anterior peristomial ring; A-I I and 

A-III to setiger 2. A-III incomplete in holotype, complete in 

several paratypes. Peristomium cylindrical, long; anterior 

margin flared in holotype, but not in paratypes; muscular lower 

lip distinct. Separation between rings distinct only ventrally; 


middle of anterior peristomial ring, slender and tapering, with 

5 to 7 articulations. 

Maxillary formula of paratype from USNM 5417 1+1, 4+5, 

6+0, 4+7, and 1+1. Jaws completely decalcified, soft and 

fragile. 

Branchiae present, palmate, distinctly shorter than notopodial 

cirri, reduced in mid-body region in some specimens. 

Branchiae from setiger 5-7 either to setiger 24-26 or 

continued to near the posterior end. Present on more than 65% 

of total number of setigers in some specimens. Maximum 3 

filaments at or near setiger 15. Most specimens with branchiae 

to setigers 24-26; holotype and one of 2 complete paratypes 

with single branchial filaments also present in posterior end 

starting at setiger 50 (Figure 58b) or 55, continuing to posterior 

end. Other complete paratype with branchiae only near anterior 

end. Filaments digitiform. 

Anterior neuropodial acicular lobes distally truncate; median 

acicular lobes rounded; posterior acicular lobes triangular; 

aciculae emerging at midline. Pre- and postsetal lobes low, 

transverse folds. First 6 ventral cirri thick, tapering. Ventral 

cirri basally inflated from about setiger 7. Inflated bases ovate; 

narrow tips tapering. Ventral cirri increasingly slender and

178 


a. g 

d. e. t. h, i 

FIGURE 58.—Eunice interrupta (holotype, USNM 5211): a, anterior end, lateral view; b, parapodium 53, anterior 

view; c, acicula, parapodium 53; d, compound falciger, parapodium 53; e, subacicular hook, parapodium 53. 

Eunice jagori (holotype, ZMB F2003): f, compound falciger, parapodium 53; g, anterior end, lateral view; h, 

subacicular hook, parapodium 44; i, pectinate seta, parapodium 44; j, parapodium 44, anterior view. (Scale bars 

in mm.) 

digitiform from about setiger 35. Notopodial cirri slender, with 

up to 5 distinct articulations; articulations present in all 

setigers. 

Shafts of compound falcigers (Figure 47d) inflated, marginally 

smooth. Appendages thick, relatively short, with large 

heads, bidentate. Proximal teeth smaller than distal teeth, 

tapering, directed laterally. Distal teeth thick, bent, with small 

notch sometimes appearing as rudimentary third teeth. Guards 

nearly symmetrically sharply pointed; mucros absent. Pseudocompound 


(Figure 48c) paired, yellow, tapering, gently curved; 



hooks (Figure 57e) yellow, tridentate with teeth in a crest. 



3 very large fangs decreasing evenly from main to distal fangs. 

UNKNOWN MORPHOLOGICAL FEATURES.—Relationship be-


tween Mx III and left Mx IV; pygidium and anal cirri. 





Characters: 1, 2,4, 6, 39, 40, 42,47, 50, 63. 


None. 

REMARKS.—Eunice interrupta was named for the interrupted 

branchial distribution in the holotype; this kind of 

branchial distribution is also present in one paratype, but other 

paratypes of sufficient length do not show this feature. 

Consequently the species has been listed among species in 

group C-l (Tables 40 and 44) and C-2 (Tables 46 and 47). 

Non-type specimens identified by Treadwell include very 

small, juvenile specimens and one larger incomplete specimen 

in which the branchial distribution could not be determined. 

Among species in group C-l with moniliform articulations in 

the ceratostylcs, E. interrupta is the only one to have the 

notopodial cirri articulated throughout the body. Only E. 

interrupia and E. oliga, of all the species in group C-2 with all 

notopodial cirri articulated, have as few as three branchial 

filaments where the branchiae are best developed. In E. 

interrupta the three median antennae are similar in length; in E. 

oliga the median antenna is distinctly longer than the other four 

antennae. 

Treadwell designated only a single specimen as type; the rest 

of the material here examined is part of his original material 

and have here been designated as paralectotypes. 

92. Eunice investigatoris Fauvel, 1932 

TABLES 27,28 

Eunice investigatoris Fauvel, 1932:137-138, fig. 19a-f. 

DESCRIPTION.—The type of this species, described from 45 

m depth in the Persian Gulf, is not available. The description is 

sufficiently detailed to allow some comments. 

Numbers of setigers not indicated. Specimen complete; total 

length 110 mm; width 7 mm. 

Antennae similar in thickness. Ceratophores ring-shaped in 


tapering, without articulations. A-I presumably to setiger 1-2; 

A-II and A-I 11 to setiger 6 or 7. 


cirri, reduced in mid-body region, erect. Branchiae from 

setiger 6 to near posterior end, present on more than 65% of 

total number of setigers. Maximum 18-20 filaments at about 

setiger 14. Branchiae "reduced in size" in middle of body, 

"increase very much" in posterior end. Branchiae in median 

and especially posterior regions dichotomously or trichotomously 

regularly branched. 

Anterior ventral cirri tapering, becoming basally inflated in 

early branchial setigers. Narrow tips tapering. Ventral cirri 

without inflated bases in posterior half of body, increasingly 

long and slender in far posterior setiger, up to twice as long as 

neuropodial lobes. Notopodial cirri long in anterior setigers, 

becoming shorter than branchiae in branchial region, without 


Pectinate setae with either 1 or both marginal teeth longer 

than other teeth, with 8-10 teeth. Shafts of compound falcigers 

inflated, marginally smooth. Appendages thick, tapering, 

bidentate. Proximal teeth larger than distal teeth, triangular, 

directed laterally. Distal teeth very short, slender, bent. Guards 

symmetrically rounded; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculae black, 

tapering, straight; cross-sections round. Subacicular hooks 

black, bidentate. Hooks first present from setiger 44-45, 


replacements). Hooks with inflated median shafts; heads small. 

Proximal teeth directed laterally. Distal teeth smaller than 

proximal teeth, directed obliquely distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Many features of 

all parts of the body. 


None. 



Characters: 1, 4-16, 24-29, 36, 39, 40, 43-51, 54, 

65-68, 74, 78. 


None. 

REMARKS.—Eunice investigatoris is listed with similar 

species in Tables 27 and 28; it is very unusual among species 

with dark aciculae and subacicular hooks in that is has a 

reduced number of branchial filaments in a mid-body region. 

This characters state is other limited exclusive to species with 

yellow, tridentate subacicular hooks. It was not included in the 

key, because it is relatively poorly known. 

Eunice jagori Grube, 1878a: 103. 

93. Eunice jagori Grube, 1878 

FIGURE 58g-j; TABLES 33,35 

MATERIAL EXAMINED.—Holotype, ZMB F2003, Manila, 

Philippines, coll. Semper. 


58 setigers; length 30 mm; maximal width 6 mm near setiger 

15; length through setiger 10, 6 mm. Body dorsoventrally 

flattened; segments crowded with parapodia on lateral ridges, 

especially in posteriormost setigers present. 




flattened; median sulcus deep. Eyes lateral to bases of A-II, 

faded. Antennae in horseshoe; A-I distinctly isolated from A-II 

and A-III, similar in thickness. Ceratophores ring-shaped in all

180 



to 20 short, cylindrical articulations in A-III. A-I to posterior 

peristomial ring; A-I I to setiger 6; A-III to setiger 12 when 

folded back over dorsum. Peristomium slightly wider than 

prostomium, tapering anteriorly. Separation between rings 


peristomial length. Peristomial cirri to posterior edge of 

prostomium, digitiform, with 3 to 4 cylindrical articulations. 

Maxillary formula 1+1,4+4,8+0,5+8,and 1+1. Mx III part 

of distal arc with left Mx IV. Mx VI absent. Jaws friable; right 

Mx IV damaged so number of teeth uncertain. 

Branchiae (Figure 58j) present, pectinate, distinctly longer 



filaments in setigers 25 to 30. Stems long, erect in median 

setigers, shortening rapidly posteriorly. Filaments relatively 

thick, folded up against branchial stems, relative length of 

filament increasing posteriorly, but shorter than notopodial cirri 

in all setigers. 

Neuropodial acicular lobes rounded; aciculae emerging 

above midline. Presetal lobes low, transverse folds. Postsetal 

lobes free, distinct lobes, higher than acicular lobes, becoming 

reduced in size, following outline of acicular lobes in posterior 

end of fragment. Ventral cirri thick, tapering; inflated bases 

totally imperceptible. Notopodial cirri basally inflated, prominent, 

with 2 to 3 long, cylindrical rings. 


(Figure 58i) gently furled, slightly flaring. One marginal tooth 

longer than other teeth, with 13 to 14 teeth. Shafts of compound 

falcigers (Figure 58f) slender, tapering, marginally smooth. 

Appendages narrow, tapering; heads distinct, bidentate. Proximal 

teeth slightly longer than distal teeth, tapering, directed 

obliquely distally. Distal teeth tapering, curved. Guards 

symmetrically rounded, marginally smooth; mucros absent. 


Aciculae single, with brown cores and clear sheaths, tapering, 

straight; cross-sections round. Subacicular hooks (Figure 58h) 

with brown cores; heads completely clear, nearly translucent, 



Hooks tapering, with distinct head. Proximal teeth larger than 

distal teeth, triangular, directed laterally. Distal teeth triangular, 

erect. 



setigers, including branchial distribution; pygidium and anal 

cirri. 


None. 



Characters: 1,2,6, 36-40,47, 50, 57-60,63. 

ASSUMED STATES FOR PURPOSE OF PREPARING KEY.—371- 

38,1. 

REMARKS.—Eunice jagori has remained incompletely 

known since it was originally described; it is listed with similar 

species in Tables 33 and 35. The ventral cirri are not basally 

inflated in median setigers, which is an unusual feature in the 

genus, but perhaps most characteristic is the crowding of the 

posterior setigers, with the parapodia carried on fleshy ridges. 

Eunice januarii Grube, 1881:111. 

Eunice januarii Grube, 1881 

REMARKS.—The material of this species currently present in 

Zoologisches Museum, Berlin, consists of a large specimen and 

two parapodia. It lacks peristomial cirri and has we 11-developed 

fascicles of compound spinigers and lacks compound falcigers. 

The species belong to the genus Marphysa and will be treated 

in a review of that genus. 

94. Eunice japonica, new name 

FIGURE 59a-c; TABLES 19. 20 

Eunice gracilis Moore, 1903:440-441. pi. 25: figi. 46-48. 

Eunice longicirrala.—Imajima and Hartman, 1964:256 Inol Eunice longicirraw 

Webster. 1884]. 

MATERIAL EXAMINED.—Holotype of E. gracilis Moore, 

1903, USNM 15724, Albatross sta 3730, off Omae Zaki, 

Honshu Island, Japan, approximate position 34°36'N, 

138°14'E, 16 May 1900, 62-67 m, mud, gravel, rock. 

COMMENTS ON MATERIAL EXAMINED.—Station 3730 was a 

surface tow, according to the published station list; several 

dredge and beam trawl hauls were taken off Omae Zaki on the 

same day in approximately the same depths. The specimen may 

have come from any one of these hauls. The position indicated 

is that of the cape, Omae Zaki proper. 


110 setigers; total length 50 mm; maximal width 2.5 mm; 

length through setiger 10, 7 mm. Body cylindrical with 

relatively long segments. Two pairs of anal cirri present; 

ventral pair long; dorsal pair very short. All anal cirri with long, 





sulcus shallow. Eyes between bases of A-I and A-I I. Antennae 

in horseshoe, evenly spaced, similar in thickness. Ceratophores 


slender, tapering; articulations long, cylindrical in A-I I and 

A-III, nearly moniliform in A-I. Maximum 10 articulations in 

A-III; articulations indistinct towards bases of 3 longer 

antennae. A-I to middle of anterior peristomial ring; A-II to 

setiger 3; A-III to setiger 5. Peristomium cylindrical. Separation 

between rings distinct on all sides, but especially distinct 

dorsally; anterior ring ~ 3 A of total peristomial length. Peristomial 

cirri to posterior half of prostomium, slender and tapering,

NUMBER 523 

181 

FIGURE 59.—Eunice japonica, new name (holotype of E. gracilis, USNM 15724): a, subacicular hook, 

parapodium 75; b, acicula, parapodium 75; c, compound falciger, parapodium 75; d, secondary subacicular hook, 

parapodium 75; e, anterior end, lateral view. Eunice johnsoni (holotype, USNM 24717): f, anterior end, lateral 

view; g, subacicular hook, parapodium 45; h, pectinate seta, parapodium 45; i, compound falciger, parapodium 

45; j. parapodium 45, anterior view. (Scale bars in mm.)


with 3 or 4 indistinct articulations. 

Maxillary formula 1+1, 8+9, 8+0, 7+11, and 1+1; maxillae 

slender and teeth even in size in each maxilla. 



from setiger 4 to 46, terminating well before posterior end, 


last 2 or 3 pairs single filaments. All other branchiae pectinate 

with up to 11 filaments. Stems slender, erect. Filaments 

slender. 


acicular lobes increasingly triangular; posterior acicular lobes 

bluntly pointed; aciculae emerging at midline. Pre- and 


tapering. Inflated bases never very distinct, ovate; narrow tips 

large and tapering. Postbranchial ventral cirri gradually 

becoming tapering and finally digitiform. Notopodial cirri 

prominent in anterior setigers, becoming shorter in far posterior 

setigers. Anterior notopodial cirri basally somewhat inflated, 

with 2 or 3 articulations. Postbranchial notopodial cirri less 

distinctly basally inflated. Articulations missing from early 

branchial setigers. 

Most setae broken. Shafts of compound falcigers (Figure 

59c) inflated, marginally smooth. Appendages narrow, slightly 

tapering; heads indistinct, bidentate. Proximal teeth shorter 

than distal teeth, forming a wide-based, low triangular 

projection. Distal teeth tapering, narrow, nearly erect Guards 


falcigers and compound spinigers absent. Up to 3 

aciculae (Figure 59b) present, yellow, pointed, very nearly 

straight or very gently curved ventrally; cross-sections round. 


and subacicular hooks. Subacicular hooks (Figure 59a,d) 

yellow, bidentate. Hooks first present from setiger 34, present 


tapering; heads large. Proximal teeth very much larger than 

distal teeth, triangular, directed slightly basally. Distal teeth 

triangular, erect. Secondary subacicular hooks (Figure 59d), 

when present, resembling primary hooks, except proximal teeth 

strongly parrot-beaked. 


Mx III and left Mx IV; structure of pectinate setae. 


None. 



Characters: 4, 6,42,65-68. 


None. 

REMARKS.—The original name, E. gracilis, is preoccupied 

in the combination E. gracilis Grube, 1866. The new name 

refers to the country of origin of the type material. 

Eunice japonica was considered a synonym of E. longicirrata 

Webster (Imajima and Hartman, 1964:256; see E. 

websteri). It belongs to group A-l and is listed with similar taxa 

in Tables 19 and 20. It can be separated from E. websteri and 

similar species by having branchiae from setiger 4 rather than 

from setiger 3. It is very similar to E. kobiensis, but in the 

former the proximal teeth of the subacicular hooks are directed 

laterally; both teeth are directed distally in £. kobiensis. Eunice 

kobiensis can be separated from E. websteri as indicated below. 

Eunice jeffreysii Mclntosh, 1903 

Eunice Jeffreys UMclnlosh, 1903:137-140, fig. 1. pi. 11: figs. 15-20. 

REMARKS.—Examination of the type material demonstrates 

that this species belongs to the genus Euniphysa; a review of 

that genus is in preparation. 

95. Eunice johnsoni Hartman, 1954 

FIGURE 59f-j; TABU* 22,23 

Eunice johnsoni Uanman. 1954:633-634. fig. 175a-c,e,f. 

MATERIAL EXAMINED.—Holotypc, USNM 24717, Bikini 

Atoll, Marshall Islands, Operation CROSSROADS, 1946, coll. 

M. W. Johnson. 



through setiger 10, 7 mm. Anterior body cylindrical, farther 

posteriorly dorsally flattened, flat and broad, Marphysa-Mkc, 

with crowded segments. 




sulcus deep. Eyes between bases of A-I and A-II. Antennae in 



Ceratostyles distally club-shaped, with up to 12 poorly marked, 

cylindrical articulations in A-I 11. A-I to posterior peristomial 

ring; A-II to setiger 3; A-II I to setiger 4. Peristomium 

cylindrical. Separation between rings distinct on all sides, but 

best marked dorsally and ventrally; anterior ring 3 A* of total 


peristomial ring, slender and digitiform, without articulations. 

Maxillary formula 1+1,4+5, 7+0, 4+8, and 1+1. 




before posterior end, present more than 65% of total number of 

setigers. First 3 and last 20 pairs single filaments. Other 

branchiae with up to 10 primary filaments. Stems cylindrical, 

about as long as notopodial cirri. Filaments thick, often 

somewhat folded, some branching distally with short terminal 

filaments, longer filaments as long as notopodial cirri. 

Anterior neuropodial acicular lobes truncate; median acicular 

lobes rounded; posterior acicular lobes increasingly triangular, 

aciculae emerging at midline. Pre- and postsetal lobes low,


transverse folds. All ventral cirri thick, tapering, not distinctly 

inflated anywhere, an inflation on body wall proximal to 

ventral cirri present in most setigers, may represent remnant 

inflated bases. Anterior notopodial cirri digitiform, about as 

long as peristomial cirri. Notopodial cirri tapering with slightly 

inflated bases in branchial region, becoming digitiform in last 

20 setigers. Notopodial cirri without articulations. 

Limbate setae long, slender. Pectinate setae (Figure 59h) 

number at least 10 in a parapodium, tapering, furled. Both 

marginal teeth longer than other teeth, with -20 teeth. Shafts of 

compound falcigers (Figure 59i) inflated, marginally serrated. 

Appendages short, tapering; heads distinct, bidentate. Both 

teeth equally well developed. Proximal teeth narrowly tapering, 

directed laterally. Distal teeth tapering, nearly erect. Guards 

short, symmetrically bluntly pointed, nearly rounded, marginally 

serrated; mucros absent. Pseudocompound falcigers and 

compound spinigcrs absent. Aciculae single, yellow, smoothly 

tapering, straight; cross-sections round. Separation between 


hooks. Subacicular hooks (Figure 59g) yellow, bidentate. 


thereafter, always single (except for replacements). Proximal 

teeth much larger than distal teeth, triangular, directed 

obliquely laterally. Distal teeth triangular, erect. Guards short 

and truncate. 




None. 



Unknown Characters: 4, 6, 42. 


None. 

REMARKS.—Eunice johnsoni is listed with similar species in 

Tables 22 and 23. Of the species listed in Table 22, E. johnsoni 

and E. schizobranchia have a very late start of the branchiae; 

the other species have branchiae from setigers 3-6. The 

subacicular hooks and aciculae are both yellow in E. johnsoni. 

In E. schizobranchia, the aciculae are black, whereas the 

subacicular hooks are yellow. 

96. Eunice kerguelensis Averincev, 1974 

FIGURE 60a-c; TABLES 41,43,46,49 

Eunice kerguelensis Averincev, 1974:172, pi. 31: figs. 11-14. 

MATERIAL EXAMINED.—Holotype, ZIL, 1/15808, Antarctic 

Ocean near Kerguelen Island, 49°38'7"S, 70 o 43'7",OB sta 121, 

141 m, 20 May 1956, coll. Averincev, Ushakov, Belayev. 


has been deeply dissected and especially the peristomium has 

been partially reconstructed in the illustration. 


26 setigers; length 4 mm; maximal width 0.3 mm; length 

through setiger 10, 1.2 mm. Body cylindrical throughout 

Prostomium (Figure 60c) about as long as peristomium, 

about as wide as peristomium, as deep as l /2 of peristomium. 

Prostomial lobes frontally rounded, dorsally very slightly 

inflated; median sulcus shallow. Eyes between bases of A-I and 

A-II, large reddish. Antennae in horseshoe, evenly spaced, 



angular articulations in A-III. A-I to posterior peristomial ring; 

A-II (only 1 present) to setiger 4; A-III to setiger 4. 

Peristomium cylindrical, with distinct muscular lower lip. 

Separation between rings distinct on all sides; anterior ring ~ 2 /3 


anterior peristomial ring, digitiform, without articulations. 

Maxillary formula 1+1,8+11,8+0,7+10, and 1+1 according 

to Averincev (1974:172). 

Branchiae present, palmate, longer than notopodial cirri, not 

reduced in mid-body region. Branchiae from setiger 3 

(according to Averincev, both parapodia on setiger 3 now 

missing) to end of fragment. Where best developed, branchiae 

with 2 slender tapering filaments. Filaments of same length as 


Neuropodial acicular lobes distally rounded; aciculae emerging 


First 3 ventral cirri thick, tapering. Ventral cirri becoming 

basally inflated from setiger 4-5. Inflated bases ovate; narrow 

tips tapering. Inflated bases retained to end of fragment 

Notopodial cirri basally slightly inflated, tapering to narrow 

tips, without articulations. 

Limbate setae marginally smooth. Pectinate setae now 

missing, illustrated by Averincev (1974, pi. 31: fig. 13) as 

tapering, with 1 marginal tooth very much longer than other 

teeth, with -10 teeth totally. Shafts of compound falcigers 

(Figure 60a) inflated, marginally serrated. Appendages slender, 

tapering; heads large, bidentate. Proximal teeth smaller than 

distal teeth, tapering, directed basally. Distal teeth curved, 

directed nearly laterally. Guards symmetrically sharply 

pointed, sometimes appearing mucronate. Pseudocompound 

falcigers and compound spinigers absent. Aciculae mostly 

paired, yellow, tapering, straight; cross-sections round. Separation 


subacicular hooks. Subacicular hooks (Figure 60b) yellow, 

tridentate with teeth in a crest Hooks first present from setiger 

19, present in all setigers thereafter always single (except for 

replacements). Hooks with large curved main fangs. Fangs 

decreasing evenly in size distally. 



posterior parapodia; pygidium and anal cirri. 


TURES.—The high number of teeth in Mx III suggests that this 

jaw piece was long and located behind left Mx II rather than 

part of a distal arc.

184 


FIGURE 60.—Eunice kerguelensis (holotype, ZML, 1/15808): a, compound falciger, parapodium 19; b, 

subacicularhook, parapodium 19; c, anterior end, lateral view. Eunice kinbergi (syntypes, RM Type 1255, d-j of 

incomplete syntype; k and 1 of complete syntype): d, anterior end, lateral view; e, compound falciger, parapodium 

27; f, pectinate seta, parapodium 27; g, parapodium 27, anterior view; h, parapodium 275, anterior view; i, 

compound falciger, parapodium 275; j, pectinate seta, parapodium 275; k, subacicular hook, parapodium 350; 1, 

parapodium 350, anterior view. (Scale bars in mm.)




Characters: 1, 2, 4, 6, 36-40, 42, 47, 50, 57-60, 63, 65, 67. 


38,2. 

REMARKS.—The type is not in very good condition; most of 

the setae are now broken. The specimen is very small and 

appears juvenile. It has the structure of a specimen of group 

C-2; the pectinate setae as illustrated by Averincev resemble 

those present in E. vittata and related species in group C-l. The 

specimen is sufficiently incomplete that the branchial distribution 

cannot be accurately determined. The species is compared 

both to species in group C-l and C-2 in Tables 41 and 43 as 

well as in Tables 46 and 49. It is the only species in Table 43 

with only single branchial filaments. Of the species listed in 

Table 49 only E. eugeniae and E. kerguelensis have mucronate 

guards on the compound falcigers. The former lacks eyes and 

antenna] articulations; E. kerguelensis has articulated ceratostyles 

and eyes are prominent. 

97. Eunice kinbergi Ehlers, 1868 

FIGURE 60d-l; TABLES 27.29 

Eriphyle capensis Kinbcrg, 1865:561; 1910:41. pi. 15: fig. 16. 

Eunice kinbergi Ehlers. 1868:306. 

MATERIAL EXAMINED.—Two syntypes, RM Type 1255, 

Cape of Good Hope, coll. Wahlberg. 

COMMENTS ON MATERIAL EXAMINED.—The complete syntype 

is currently in four pieces. Most illustrations are of the 

incomplete syntype as indicated in the figure legends. 

DESCRIPTION.—One syntype complete, of unknown sex, 

with 380 setigers; length 398 mm; maximal width 15 mm; 

length through setiger 10,16 mm. Other syntype incomplete, of 

unknown sex, with 282 setigers; length 655 mm; maximal 

width 15 mm; length through setiger 10, 22 mm. 


than peristomium, as deep as x li of peristomium. Each 

prostomial lobe distinctly divided longitudinally into 2 parts; 

lateral parts large, frontally rounded, dorsally inflated. Medial 

parts narrow, frontally angularly rounded. Median sulcus deep, 

separating prostomium to more than half its length and 

continued as distinct suture to base of A-III. Palpal region set 

off by horizontal, frontal grooves. Eyes between bases of A-I 

and A-I I; black. Antennae in transverse row, evenly spaced, 


without articulations. Ceratostyles tapering, relatively slender, 

with up to 13 irregular, cylindrical articulations in A-III. A-I to 

middle of anterior peristomial ring; A-I I to setiger 1; A-III to 

setiger 1 in one syntype and to setiger 4 in other syntype. 

Peristomium about twice as wide as prostomium, overall 

cylindrical, flaring anteriorly, with paired large ventrolateral 

cushions forming lower lip. Separation between rings distinct 


length. Peristomial cirri to front end of peristomium, slender 

and gently tapering, with 5 articulations. 

Maxillary formula 1+1,5+5, +0,4+9,1+1, and 1+1. Mx III 

now missing, probably short, forming part of distal arc with left 

Mx IV-VI judging from compressed shape of left Mx IV. 

Branchiae (Figure 60g,h,l) present, pectinate, about as long 

as notopodial cirri, not reduced in mid-body region, erect. 

Branchiae from setiger 8-9 to 365. Branchiae present to near 

posterior end, more than 65% of total number of setigers. First 

2 pairs single filaments, maximum 22 filaments from setiger 

25, slowly reduced in length and numbers of filaments 

posteriorly. Stems strong, erect anteriorly and medially, usually 

held curved posteriorly, shorter than notopodial cirri. Filaments 

slender, nearly filiform, shorter than notopodial cirri. 

Neuropodial acicular lobes asymmetrically triangular with 

aciculae emerging dorsal to midline. Pre- and postsetal lobes 

forming low folds around dorsal edge of acicular lobes, 

especially cut away frontally. First 4 ventral cirri thick, 

tapering. Ventral cirri basally inflated from about setiger 5. 

Inflated bases continued to end of body, forming thick, 

elongated welts along ventral edge of neuropodia. Upper edge 

of inflated bases wrapped around lower edges of neuropodia, 

forming shallow scoops; free edge produced as short, thick 

tapering narrow tips. Notopodial cirri slightly inflated basally, 

tapering to long, digitiform tips; anterior cirri with 5 cylindrical 

articulations. Posterior notopodial cirri shorter than anterior 

ones, but otherwise similar, articulations indistinct, but present. 

Limbate setae very slender, marginally smooth. Anterior 

pectinate setae (Figure 60f) with cylindrical shafts. Blades 

tapering, furled into open scoops; marginal teeth barely longer 

than other teeth, with 12 teeth. Posterior pectinate setae (Figure 

60j) with flat shafts. Blades completely flattened, otherwise 

similar to anterior pectinate setae. Shafts of compound falcigers 

(Figure 60e,i) tapering, marginally smooth, with distinct 

internal striations; distal peak present. Appendages tapering; 

heads small, bidentate. Proximal and distal teeth similar in size. 

Proximal teeth tapering, directed obliquely distally. Distal teeth 

slender, tapering, distinctly curved. Guards slightly asymmetrically 

rounded, marginally smooth; mucros absent Slender, 

honey-colored notopodial aciculae present in all setigers. 

Pseudocompound falcigers and compound spinigers absenL 

Neuropodial aciculae paired, with brown cores and clear 

sheaths, tapering, blunt-tipped, straight; cross-sections round. 

Subacicular hooks (Figure 60k) with brown cores and clear 

sheaths, bidentate. Shorter syntype without subacicular hooks; 

other syntype with first subacicular hook in setiger 123, next in 

setiger 135 and scattered thereafter, becoming more frequent in 

far posterior setigers, where most frequent paired in some, but 

not all parapodia. Hooks very slender (no thicker than 

compound hooks), abruptly tapering distally, with distinct 

heads. Proximal teeth larger than distal teeth, triangular, 

directed laterally. Distal teeth triangular, nearly erect. Guards 

symmetrically rounded. 


anal cirri.

186 



None. 





None. 

REMARKS.—Kinberg (1865) erected the genus Eriphyle for 

species of Eunice with quadrilobate prostomia, including E. 

gigantea Savigny (actually Lamarck, K. Fauchald comment), E. 

aphroditois Pallas, and E. violacea Grube. The genus has not 

been recognized by later authors, including Ehlers (1868), who 

renamed Kinberg's species, because Kinberg's species name 

was preoccupied in the combination Eunice capensis Schmarda 

(1861). 

Eunice kinbergi has been considered a synonym of E. 

aphroditois: two forms both belong to group B-2 and are 

similar, as indicated in Table 27. However, E. kinbergi has 

articulated notopodial cirri and cylindrical articulations in the 

ceratostyles; E. aphroditois lacks articulations of both structures. 

Eunice kinbergi is one of two species in Table 29 with 

basally inflated ventral cirri continued to the posterior end; the 

other species is E. plicata. The latter has only about five 

branchial filaments; the former has 22 filaments. 

Eunice kinbergi (Webster, 1884) 

Nicidion kinbergi Webster, 1884:320, pi. 12: figs. 81-88 [not Eunice kinbergi 

Ehlers, 1868:306]. 

Eunice (Nicidion) kinbergi.—Hartman, 1944:124; 1959:313. 

REMARKS.—Originally described in Nicidion, this species 

must be renamed, because the name, once the genus Nicidion 

was fused with Eunice, became pre-occupied in the combination 

E. kinbergi Ehlers, 1868. The species is here renamed E. 

goodei. 

98. Eunice kobiensis Mclntosh, 1885 


Eunice kobiensis Mclntosh, 1885:278-280, figs. 37, 38, pi. 38: figs. 12, 13, pi. 

20A: figs. 1, 3.—Fauchald, 1969: 4-6. fig. 2a-g. 


1885.12.1.197, Dredged off Kobe, Japan, 8-50 fathoms. 

Challenger. 

COMMENTS ON MATERIAL EXAMINED.—The specimen has 

been frontally dissected and the jaw apparatus removed; it is cut 

into two pieces. Two sets of maxillae are present in a small vial 

accompanying the specimen; the maxillary formula is the same 

for both sets, even if one set appears too large to fit the 

holotype. 



through setiger 10, 6.5 mm. 


than peristomium, less than x ti as deep as peristomium. 


median sulcus deep. Antennae in shallow horseshoe, evenly 


antennae, without articulations. Ceratostyles slender, tapering, 

with up to 5 long, cylindrical articulations. A-I to posterior 

peristomial ring; A-I I to sctigcr 3; A-I 11 to sctiger 8. 


all sides; anterior ring ~ 2 h of total peristomial length. 

Peristomial cirri to middle of anterior peristomial ring, slender, 



nearly as long as left Mx II; located behind Max II. 



Branchiae from sctigcr 3 to sctigcr 41. Branchiae terminating 


number of setigers. Three first and last 2 pairs single filaments, 

all others pectinate with up to 8 filaments. Stems slender, 

tapering, about as long as notopodial cirri. Filaments slender, 

about as long as notopodial cirri. 

Prc- and postbranchial ncuropodial acicular lobes (Figure 

61b, 0 distally rounded; acicular lobes in branchial region 

distally bluntly conical; aciculac emerging at midline. Pre- and 

postsetal lobes low, transverse folds. Prc- and postbranchial 

ventral cirri thick, tapering. Ventral cirri basally inflated from 

about setiger 4. Inflated bases ovate; tips thick and tapering. 

Anterior notopodial cirri long, medially inflated, becoming 

increasingly slender and tapering in branchial and postbranchial 

setigers. Prebranchial and branchial notopodial cirri with 

up to 3 long, cylindrical articulations; postbranchial notopodial 


Limbate setae slender. Pectinate setae (Figure 6Id) narrow, 

tapering, flat. One marginal tooth distinctly longer than other 


61c,g) gently inflated, marginally finely dentate at least in some 

hooks. Appendages narrow, tapering, bidentate. Proximal teeth 

smaller than distal teeth, triangular, directed laterally or slightly 

distally. Distal teeth gently curved, blunt or pointed. Guards 

asymmetrically pointed; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculae (Figure 61 i) 

paired, yellow, pointed, straight or gently curved; crosssections 

round. Separation between core and sheath indistinct 

in both aciculae and subacicular hooks. Subacicular hooks 

(Figure 61 h) yellow, bidentate. Hooks first present from setiger 


replacements). Hooks tapering; head distinct. Proximal and 

distal teeth similar in size. Proximal teeth triangular, directed 

laterally. Distal teeth curved, distinctly narrower than proximal 

teeth. 


with far posterior parapodia; pygidium and anal cirri. 


None. 

CHARACTERS 

USED IN PREPARATION OF KEY NOT

NUMBER 523 

187 

• 02 

.5 

] 

.1 

2 

.5 

1. m 

c. d. g. h 

e, t 

a 

FIGURE 61.—Eunice kobiensis (holotype, BM(NH) ZK. 1885.12.1.197): a, anterior end, lateral view; b, 

parapodium 2, anterior view; c, compound falciger, parapodium 19; d, pectinate seta, parapodium 19; e, 

parapodium 19, anterior view; f, parapodium 127, anterior view; g, compound falciger, parapodium 127; h, 

subacicular hook, parapodium 127; i, aciculae, parapodium 127. Eunice langi (holotype, AMHN 3739): j, 

parapodium 51, anterior view; k, parapodium 20, anterior view; 1, compound falciger, parapodium 20; m, 

pectinate seta, parapodium 51; n, acicula, parapodium 51; o, subacicular hook, parapodium 51. (Scale bars in 

mm.)



Characters: 1,2,4,6, 13, 14, 42,47, 50,63. 


None. 

REMARKS.—Eunice kobiensis was considered a synonym of 

E. websteri (as E. longicirrata Webster) by Imajima and 

Hartman (1964:256); as already indicated by Fauchald (1969), 

the proposed synonymy cannot be correct. Eunice kobiensis is 

compared to similar species in Tables 19 and 20. Fauchald 

(1969) compared it to a number of similar species; at this time 

nothing can be added to that comparison and to discussions of 

other species listed in Table 20. 

99. Eunice langi (Treadwell, 1943) 

FIGURE 61j-o; TABLES 24,26 

Leodice langi Treadwell, 1943:3, figs. 14-18. 

Eunice savignyi.—Hartman, 1956:283 [not Eunice savignyi Grube, 1878]. 

MATERIAL EXAMINED.—Holotype, AMHN 3739, Cape 

Town, South Africa. 


has been badly mangled and partly dissected, and all antennae 

are lost, so no illustrations were attempted of the anterior end. 



through setiger 10, 5.5 mm. 

Prostomial lobes frontally obliquely truncate, dorsally 

flattened; median sulcus shallow. Eyes not observed. Ceratophores 

ring-shaped in all antennae. Ceratostyles missing. 

Peristomial rings distinct dorsally and ventrally; anterior ring 

3 A of total peristomial length. Peristomial cirri to setiger 3, with 

6 distinct articulations. 

Maxillary formula 1+1,6+5,9+0,6+8, and presumably 1+1 

according to Treadwell (1943, fig.18). 

Branchiae (Figure 61k) present, pectinate, distinctly longer 


Branchiae from setiger 3 to setiger 39. Maximum 8 filaments at 

or near setiger 15; most branchiae with 4 or 5 filaments. Stems 

stout, strongly tapered, shorter than notopodial cirri. Filaments 

digitiform, about as long as notopodial cirri. 

Anterior neuropodial acicular lobes distally truncate; posterior 

acicular lobes (Figure 61j) obliquely conical; aciculae 


folds. First 4 ventral cirri very long, digitiform, as long as 

notopodial cirri. Ventral cirri basally inflated from setiger 5 to 

setiger 36, with distinct, digitiform, obliquely attached tips. 

Inflated bases increasing evenly from setiger 5 to setiger 15, 

ovate, decreasing from setiger 30, missing posterior to setiger 

36. Posterior ventral cirri short, nearly tubercular. Anterior 

notopodial cirri slender, digitiform; posterior notopodial cirri 

long, tapering. Notopodial cirri without articulations. 

Shafts of pectinate setae (Figure 61m) slender, cylindrical. 

Blades flat, strongly flaring. One marginal tooth longer than 

other teeth, with -15 teeth. Shafts of compound falcigers 

(Figure 611) gently inflated, marginally smooth. Appendages 

long, tapering; heads very small, bidentate. Proximal teeth 

much smaller than distal teeth, very short, triangular, directed 

laterally. Distal teeth tapering, nearly erect. Guards asymmetrically 

sharply pointed; mucros absent. Pscudocompound falcigers 

and compound spinigers absent. Aciculae (Figure 61n) 

single, amber-colored, tapering, distinctly bent in median and 

posterior setigers; cross-sections round. Subacicular hooks 

(Figure 61o) amber colored, bidentate. Hooks first present from 


(except for replacements). Hooks tapering; heads small, 

distinctly curved. Proximal and distal teeth similar in size. 

Proximal teeth directed laterally, blunt. Distal teeth truncate, 


UNKNOWN MORPHOLOGICAL FEATURES.—Prostomial features; 

relationship between Mx III and left Mx IV; pygidium 



Mx III short; forming a distal arc with left Mx IV. 



Characters: 4, 6, 10-17, 19-24, 39-40,74, 78. 


REMARKS.—The compound falcigers of E. langi resemble 

those of E. tubifex, but compound spinigers are absent in E. 

langi. The aciculae and subacicular hooks are amber colored, 

rather than clear yellow; by definition this places the species in 

B-l as defined by Fauchald (1970). It is listed with similar 

species in Table 24 and 26. 

Eunice langi was considered synonymous with E. savignyi 

by Hartman (1956); it differs by having considerably darker 

subacicular hooks and acicula. In addition the latter have 

branchiae from setiger 4 through setiger 45 with a maximum of 

12 filaments; E. langi has branchiae from setiger 3 through 

setiger 39 with a maximum of eight filaments. Due to the 

indeterminate status of E. savignyi, other differences or 

similarities cannot be detailed at this time. 

None of the character states listed in Table 26 are unique to 

this species. Among species listed in Table 24, it is 

characterized first and foremost by the very poor development 

of the head of the compound falcigers and the sharply pointed, 

but not mucronate, hood of these hooks. 

100. Eunice laticeps Ehlers, 1868 


Eunice tentaculata Quatrefages, 1866:317-318. 

Eunice laliceps Ehlers, 1868:312.—Fauchald, 1986:251-252, figs. 46-50. 

MATERIAL EXAMINED.—Holotype of E. tentaculata Quatrefages, 

MNHN, Paris, A.l(R.)-1868-no. 52b. Port Western, 

Australia, coll. Quoy and Gaimard, 1839. 

COMMENTS ON MATERIAL EXAMINED.—The holotype of E.


02 

FIGURE 62.—Eunice laticeps (holotype of E. tentaculata Quatrcfages, MNHN, Paris, A.l(R.)-1868-no. 52b): a, 

anterior end, lateral view; b, pectinate seta, parapodium 50; c, parapodium 50, anterior view; d, compound 

falciger, parapodium 50; e, subacicular hook, parapodium 50. (Scale bars in mm.) 

tentaculata is an incomplete specimen that currently is in three 81 segments; length 126 mm; maximal width 10 mm; length 

fragments. In addition a small posterior fragment with a through setiger 10, 21 mm. Anterior fragment with first 38 

pygidium is present; its relation to the type cannot be setigers. 

determined. 


DESCRIPTION.—Holotype incomplete, of unknown sex, with than peristomium, less than x /i as deep as peristomium.

190 SMITHSONIAN CONTRIBUTION'S TO ZOOLOGY 

Prostomial lobes frontally rounded, dorsally strongly inflated; 

median sulcus deep. Antennal bases overlapped by peristomial 

fold. Antennae in transverse row, evenly spaced, similar in 


articulations. Ceratostyles tapering, with up to 10 distinct, 

cylindrical articulations in A-II and A-III. A-I to middle of 

anterior peristomial ring; A-II and A-III to middle of posterior 

peristomial ring. Left A-I doubled and left A-II missing. 

Peristomium massive, cylindrical. Separation between rings 

distinct dorsally and visible ventrally; anterior ring 3 A of total 

peristomial length. Peristomial cirri to anterior ! /3 of anterior 

peristomial ring, digitiform, with 6 rings. 

Partial maxillary formula 1+1, 4+, 6+0, 6+9, and 1+1. 

Mandibles massive, strongly calcified, presently very friable. 



Branchiae from setiger 4 to end of fragments. All branchiae 

pectinate. First branchiae with 4 filaments, maximum 18 

filaments reached at about setiger 20, most median segments 

with 10 or fewer filaments. Stems tapering, shorter than 

notopodial cirri. Filaments digitiform, as long as notopodial 

cirri. 

Ncuropodial acicular lobes obliquely truncate; aciculae 

emerging from high dorsal section. Pre- and postsetal lobes 

low, transverse folds. First 3 ventral cirri digitiform. Median 

ventral cirri basally inflated. Inflated bases modest, ovate; 

narrow tips tapering. Posterior ventral cirri with less distinctly 

inflated bases, increasingly tapering. Notopodial cirri digitiform; 

anterior cirri with 2 or 3 articulations; posterior cirri 


Limbate setae narrow, marginally smooth. Pectinate setae 

(Figure 62b) present in large numbers, most tapering, some 

slightly flaring, flat. One marginal tooth longer than other teeth, 

with 15-20 teeth. Shafts of compound falcigers (Figure 62d) 

tapering, marginally smooth, internally striated. Appendages 

narrow, bidentate. Proximal teeth smaller than distal teeth, 

bluntly triangular, directed laterally. Distal teeth blunt, erect. 

Guards short, symmetrically rounded (not illustrated). Pseudocompound 


at least paired, black, pointed, straight; cross-sections 

round. Subacicular hooks (Figure 62e) black, bidentate. Hooks 


paired in some setigers. Hooks tapering. Proximal teeth much 

larger than distal teeth, triangular, directed laterally. Distal 

teeth triangular, nearly erect. 





Mx III short; forming part of distal arc with left Mx IV. 



Characters: 1, 2, 4, 6, 13, 14, 36-38, 40, 47, 50, 63, 70 74 

78. 


38,1; 70,2. 

REMARKS.—The use of the name E. laticeps for this species 

was explored by Fauchald (1986). The species belongs to group 

B-2 and is listed with similar species in Tables 27 and 30. In 

numbers of branchial filaments, E. laticeps most closely 

resembles two poorly known species, E. flavopicta and E. 

macrobranchia; two better-known species, E. reducta and £. 

violaceomaculata, have slightly higher number of filaments. Of 

these, E. reducta has branchiae terminating well before the 

posterior end. The inflated bases of the ventral cirri are ovate in 

E. laticeps and thick, transverse welts in E. violaceomaculata. 

A-III is distinctly longer than A-II in E. macrobranchia and of 

about the same length as A-II in E. laticeps, and a similar 

difference separates the latter from E. flavopicta. 

101. Eunice laurillardi Quatrefages, 1866 

FIOURH 63; TABU'S 27, 32 

Eunice laurillardi Quaircfagci, 1866:314-315, pi. 10: fig. 3—Grube, 

1870a:294. 

MATERIAL EXAMINED.—Syntype, MNHN, Paris, A.1(R.)- 

1868-no. 49b, Nice. Syntype, MNHN-Paris, A.l(R.)-1868-no. 

49c, Palermo. Syntype, MNHN-Paris, A.l(R.)-1868-no. 49a, 

Marseille, coll. M. Barbour (posterior fragment only). 

COMMENTS ON MATERIAL EXAMINED.—One additional 

specimen identified as E. laurillardi, MNHN Al (R) 1868 no 

49d from Palermo, Italy, is a posterior fragment of an arabellid 

and cannot be further identified. The description is based 

primarily on the specimen from Nice, except as noted. 

DESCRIPTION.—Syntype from Nice complete, of unknown 

sex, with 189 setigers; total length 150 mm; maximal width 5 

mm; length through setiger 10, 9 mm. Syntype from Palermo 

complete, of unknown sex, with 170 setigers; total length 143 

mm; maximal width 4.5 mm; length through setiger 10,9 mm. 




median sulcus deep. Eyes between bases of A-I and A-II. 

Antennae in transverse row, evenly spaced, similar in thickness. 

Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostyles tapering, articulated to bases, with 

up to 13 articulations in A-III; innermost articulations 

cylindrical, gradually becoming drop-shaped distally. A-I to 


slightly flaring anteriorly. Separation between rings distinct 



with 5 cylindrical articulations. 

Maxillary formula 1+1, 6+6, 5+0, 5+8, and 1+1. Left Mx IV 

short; part of distal arc with Mx III. 

Branchiae (Figure 63b,c) present, pectinate, distinctly 

shorter than notopodial cirri, not reduced in mid-body region,


.02 

1 

.1 

e. t, q 

h. c 

FIGURE 63.—Eunice laurillardi (syntype, MNHN, Paris, A.l(R.)-1868-no. 49b): a, anterior end, lateral view; b, 

parapodium 37, anterior view; c, first parapodium of posterior fragment from Marseille, anterior view; d, 

compound falciger, first parapodium of Marseille fragment; e, pectinate seta, parapodium 37; f, compound 

falcigcr, parapodium 37; g, subacicular hook, parapodium 37. (Scale bars in mm.)


erect. Branchiae from setiger 3 to setiger 184 (five setigers 

before pygidium). Branchiae present to near posterior end, 

present on more than 65% of total number of setigers. First 

branchia single filaments; maximum 10 filaments at about 

setiger 15. Stems short, flexible. Filaments slender, longer than 

stems. 

Neuropodial acicular lobes rounded; aciculae emerging at 


ventral cirri tapering, becoming basally inflated by setiger 10. 

Inflated bases modest, ovate, completely reduced by setiger 35; 

narrow tips tapering. Posterior ventral cirri broadly attached 

forming shallow open scoops covering lower edge of neuropodia. 

Notopodial cirri long, tapering to slender digiuform tips, 

with 4 to 5 articulations anteriorly and 3 to 4 articulations 

posteriorly. 


(Figure 63e) tapering, flat. One marginal tooth longer than 

other teeth, with -15 very slender teeth. Shafts of compound 

falcigers (Figure 63d, 0 tapering, marginally coarsely serrated, 

internally striated. Appendages narrow, tapering towards small 

head, bidentate. Proximal teeth very much smaller than distal 

teeth, nearly button-shaped, directed laterally. Distal teeth 

tapering, directed obliquely distally. Guards asymmetrically 



mostly paired, dark brown or black, tapering to blunt tips, 

slightly curved; cross-sections round. Subacicular hooks 

(Figure 63g) dark brown to black, bidentate. Hooks first present 

from setiger 32-33, present in all setigers thereafter, always 

single (except for replacements). Hooks tapering to small head. 

Proximal teeth very much larger than distal teeth. Both teeth 

directed distally. Subacicular hooks darker than aciculae. 


anal cirri. 


None. 



Characters: 4,6, 74, 78. 


None. 

REMARKS.—Grube (1870a) indicated that the type material 

contained specimens of two previously known taxa. As 

indicated above, one lot consists of an unidentifiable arabellid; 

the remaining material currently present under this name in 

MNHN, Paris, appears to belong to a single species in group 

B-2. The fragment from no. 49a agrees with the other two 

specimens in structure of parapodia and setae. 

Eunice laurillardi is listed with similar species in Tables 27 

and 32. Of the three species with 10 or more branchial filaments 

listed in Table 32, E. bowerbanki has A-III distinctly longer 

than A-II; A-II and III are similar in length in E. laurillardi; the 

length of A-III is unknown in E. multipectinata. The 

peristomial cirri reach the middle of the prostomium in E. 

laurillardi and the middle of the peristomium in E. multipectinata. 

102. Eunice leptocirris Grube, 1870 

Eunice leptocirris Grube, 187Ob:55. 

FIGURE 64a-f; TABLES 22.23 

MATERIAL EXAMINED.—Holotype, ZMH V-795, Fiji Islands. 



through setiger 10, 9 mm. Body cylindrical, with very 

prominent notopodial cirri. 


than peristomium, as deep as '/2 of peristomium. Prostomial 

lobes frontally obliquely truncate, dorsally slightly flattened; 

median sulcus deep. Eyes not observed. Antennae in horseshoe; 

A-I and A-II separated from A-III by gap, similar in 

thickness. Ccratophorcs long in all antennae, without articulations. 

Ccratostylcs slender and digitiform, without articulations. 

A-I to posterior peristomial ring; A-II to setiger 6; A-III 

to setiger 9. Peristomium cylindrical. Separation between 

peristomial rings distinct on all sides; anterior ring - 2 /s of total 

peristomial length. Peristomial cirri reaching well beyond tip of 






filaments at about setiger 15. Stems tapering, slender, shorter 

than filaments. Filaments slender, nearly filiform, shorter than 


Neuropodial acicular lobes rounded or slightly triangular, 



cirri basally inflated from about setiger 5. Inflated bases 

modest, ovate, very long, distally obliquely truncate tips 

present through median seugers. Inflation gradually lost from 

about setiger 60; posteriormost ventral cirri slender and 

digitiform, distally obliquely truncate. Notopodial cirri very 

long, distally tapering to very slender tips; bases modestly 

inflated, especially in anteriormost setigers; articulations 

absent. 


(Figure 64e) tapering, flaL One marginal tooth longer than 

thicker than other teeth, with -10 teeth. Shafts of compound 

falcigers (Figure 64c) slightly inflated, marginally smooth; 

beaks distinct. Appendages slender, with distinct heads, 

bidentate. Proximal teeth smaller than distal teeth, narrowly 

triangular, directed laterally. Distal teeth slender, strongly 

curved, directed laterally. Guards asymmetrically bluntly 

pointed, marginally smooth; mucros absent. Pseudocompound 

falcigers and compound spinigers absenL Aciculae (Figure

NUMBER 523 

193 

FIGURE 64.—Eunice Uptocirris (holotype, ZMH V-795): a, anterior end, lateral view; b, parapodium 34, anterior 

view; c, compound falciger, parapodium 34; d, aciculae, parapodium 34; e, pectinate seta, parapodium 134; f, 

subacicular hook, parapodium 134. Eunice leucosticta (slide from holotype, ZMB F-2044): g, posterior 

parapodium, anterior view; h, compound falciger, posterior parapodium; i, subacicular hook, posterior 

parapodium. (Scale bars in mm.)


64d) paired, yellow, most tapering distally to nearly straight, 

blunt tips, some very abruptly tapering with short, nearly 

nipple-shaped tips; cross-sections round. Separation between 


hooks. Subacicular hooks (Figure 64f) yellow, bidentate. 



teeth much larger than distal teeth, very thick, short, triangular, 

directed laterally. Distal teeth short, blunt, erect. 




None. 



Characters: 1, 2,4, 6, 13, 14, 36-40. 


38,1. 

REMARKS.—Eunice leptocirris has not been reported since it 

was originally described by Grubc (1870b:55); the original 

description was so brief that the species has been considered 

indeterminable by most authors. The species belong to group 

A-2 and is listed with similar species in Tables 22 and 23. The 

only other species listed in Table 23 without articulated 

ceratostyles is E. schizobranchia. Among other differences, the 

latter has dark aciculae and yellow subacicular hooks; E. 

leptocirris has both the aciculae and subacicular hooks 

light-colored. 

Eunice leucodon Ehlers, 1901:261. 

Eunice leucodon Ehlers, 1901 

REMARKS.—Examination of the type material demonstrated 

that this species has the jaw structure and setal complement of 

the genus Palola. It is included in a treatment of that genus 

currently in preparation. 

Eunice leuconuchalis Benham, 1900 

Eunice leuconuchalis Benham, 1900:21-22.—Fauchald, 1986:252-253. 

Eunice australis.—Ehlers, 1907:12.—Fauchald, 1986:244-245, figs. 9-14. 

REMARKS.—Justification for considering this species a 

synonym of E. australis was reviewed by Fauchald (1986). 

103. Eunice leucosticta Grube, 1878 

Eunice leucosiicia Grube, 1878a:103. 

FIGURE 64g-i; TABLES 33,37 

MATERIAL EXAMINED.—Slide of parapodium from holotype, 

Zool. Mus. Berlin, F-2044, East Africa, coll. v.d. Decken. 

DESCRIPTION.—Slide of a posterior parapodium, presumably 

near segment 120, without branchiae, but apparently not a 

segment immediately in front of the pygidium. 

Species described by Grube (1878a: 103) with 131 segments; 

total length 130 mm. Color dark red, densely studded with 

white spots. Three median antennae of same length, reaching 

segment 4 (including peristomium). 

Maxillary formula (rewritten from Grube) 1+1, 5+5, 10+0, 

4+6, and 1+1. 

Branchiae from setiger 14 and 15 (presumably on either side 

of body) and continued to setiger 113, terminating well before 

end of specimen. Where best developed with 4 filaments, but 3 

filaments by far most common number. Filaments short, no 

longer than short notopodial cirri. 

Neuropodia (Figure 64g) distally evenly rounded; aciculae 

emerging above midline. Pre- and postsctal lobes low folds. 

Posterior ventral cirri large, distally tapering, basally moderately 

inflated. Notopodial cirri short, digiliform in parapodium 

examined. 

Parapodium with a thick bundle of limbatc setae, a number 

of compound falcigcrs, and a few pectinate setae. Shafts of 

compound falcigcrs (Figure 64h) inflated, marginally smooth. 

Appendages short triangular, bidcnlatc. Tcelh similar in size. 

Proximal teeth tapering, curved, directed laterally. Distal teeth 

tapering, nearly erect. Guards symmetrically rounded; mucros 

absent. Pscudocompound falcigcrs and compound spinigers 

absent. Aciculae single, with dark brown cores and clear 

sheaths, tapering to straight lips; cross-sections round. Subacicular 

hooks (Figure 64i) with dark brown cores and clear 

sheaths. Hooks distally abruptly tapered. Proximal teeth larger 

than distal tcelh, tapering, directed laterally. Distal teeth 

abruptly tapering, directed obliquely distally. 


with anterior and median parts of the body. 


None. 



Characters: 3-22, 24-29, 31-34, 38-40, 42, 45, 46, 

48,49, 51-56, 60-62, 65-68, 80-82. 


None. 

REMARKS.—Eunice leucosiicia was considered a synonym 

of E. afra by Hartman (1959). Eunice leucosticta appears to 

differ from E. afra in that the antennae are much longer than in 

that species and in that the two teeth on the subacicular hooks 

are directed laterally rather than distally as in E. afra. It is listed 

with similar species in Tables 33 and 37. It is too poorly known 

to be included in the key or to be discussed in detail in relation 

to similar species listed in Table 37. 

104. Eunice levibranchia (Hoagland, 1920) 


Leodice levibranchia Hoagland, 1920:614-615, pi. 50: figs. 1-8. 

MATERIAL EXAMINED.—Holotype, USNM 19018, Alba-

NUMBER 523 

195 

FlOURE 65.—Eunice levibranchia (holotype, USNM 19018): a, anterior end, lateral view; b, appendage of 

compound falciger, parapodium 13; c, pectinate seta, parapodium 13; d, subacicular hook, approximately 

parapodium 80. Eunice lita (holotype, USNM 19395): e, parapodium 106, anterior view; f, anterior end, lateral 

view; g, subacicular hook, parapodium 106; h, pectinate seu, parapodium 106; i, compound falciger, parapodium 

106. (Scale bars in mm.)

196 


tross sta 5348, Point Tabonan, Palawan Passage, Philippine 

Islands, 10°57 / 45'TSf, 118°38'15"E,27 Dec 1908, 686 m, coral 

and sand bottom, beam-trawl. 


poorly preserved and especially the different parapodial lobes 

are difficult to distinguish. The type has been dissected for the 

jaws, and details of the ventral side of the prostomia and 

peristomia are difficult to discern. 



through setiger 10, 9 mm. Body anteriorly dorsally strongly 

convex, tapering abruptly to a ragged posterior end. 


than peristomium, less than l /i as deep as peristomium. 

Prostomial lobes short, frontally rounded, dorsally inflated. 

Eyes between bases of A-I and A-II, black. Antennae in 



Ceratostyles slender and digitiform, without articulations. A-I 

to middle of posterior peristomial ring; A-I I and A-I 11 to setiger 


dorsally and probably ventrally; anterior ring ~ 2 /3 of total 


peristomial ring, without articulations. 

Maxillary formula (combining observations of Mx I and II, 

which are present, with Hoagland's observations) 1+1, 8+7, 

+0,10+10, and 1+1. 



setiger 69 to setigers 85-95. Branchiae present to near 

posterior end to terminating well before posterior end, present 

on less than 55% of total number of setigers. Branchiae small 

tufts with 2 to 3 filaments. 

Parapodia all very small compared to size of specimen, 

especially in anterior end. Neuropodial acicular lobes distally 

truncate; aciculae emerging at midline. Pre- and postsetal lobes 

low, transverse folds. First 1-2 ventral cirri tapering. Median 

ventral cirri with inflated bases. Inflated bases transverse welts; 

narrow tips short and button-shaped. Inflated bases decreasing 

posteriorly. Posterior ventral cirri tapering. Notopodial cirri 

short, tapering, slender, without articulations. 

Limbate setae marginally smooth. Pectinate setae (Figure 

65c) short, relatively stout, slightly flared, furled. One marginal 

tooth distinctly longer than other teeth, with -10 coarse teeth. 

Shafts of compound falcigers inflated, marginally serrated. 

Appendages (Figure 65b) thick, tapering; heads large, bidentate. 

Teeth similar in size. Proximal teeth narrowly triangular, 

directed laterally. Distal teeth thick, with distinct basal bend, 

blunt, directed obliquely distally. Guards asymmetrically 

bluntly pointed, marginally serrated; mucros absent. Pseudocompound 


numbering 2-3, dark, tapering, straight or very gently 

curved; cross-sections round. Subacicular hooks (Figure 65d) 

dark, bidentate. Hooks first present from setiger 28, present in 


Hooks slender, distally abruptly tapering to slender necks; 

heads distinct Proximal teeth very much larger than distal 

teeth, directed laterally. Distal teeth blunt, very short, nearly 

erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—Various details 

of the jaw apparatus; pygidium and anal cirri. 


None. 



Unknown Characters: 4, 6,42, 74, 78. 


None. 

REMARKS.—Eunice levibranchia is characterized by antennae 

without articulations and dark subacicular hooks and by the 

very late start of the very poorly developed branchiae. 

Hoagland's illustration of the jaws (1920, pi. 50: fig. 2) 

indicates a maxillary formula of 1+1, 8+7, 10+10, and 8+0. A 

symmetric development of Mx III appears unlikely; the 

match-up of the different jaw pieces appears to have been 

confused. 

Branchiae do not appear to have been present as far forward 

as setiger 50 as indicated by Hoagland, but the specimen is very 

poorly preserved and the branchial distribution is difficult to 

determine. Two features are certain: the branchiae do not start 

until very late, and the branchial filaments are few and very 

short. 

Eunice levibranchia is listed with similar species in Tables 

33 and 37. Other species listed in 33 with a very late start of the 

branchiae include E. excariboea and E. gagzoi. The former has 

articulated ceratostyles; the two latter lack articulations. Eunice 

gagzoi has single branchial filaments; E. levibranchia has as 

many as three filaments. There are other shape differences best 

evaluated by comparing the illustrations of the two species. 

105. Eunice limosa Enters, 1868 

TABLES 41,43 

Eunice limosa Ehlers, 1868:348-352, pi. 15: figs. 15-22. 

Eunice vittata.—Fauvel, 1911:11 [ not Eunice vittata Chiaje, 1828]. 

DESCRIPTION.—This species was originally described from 

Kvarner Gulf in the Adriatic Sea. The types are no longer 

available, but the description given by Ehlers is sufficiently 

detailed to characterize the form. 

Specimens with 81-106 segments; total length 32-65 mm; 

width ~2 mm. Body cylindrical anteriorly, tapering to slender 

posterior ends. Anal cirri as long as last 5 setigers, without 


Prostomium frontally rounded; median sulcus shallow. 


Ceratostyles digitiform, with up to 25 cylindrical articulations 


A-II I to setiger 5. Separation between peristomial rings distinct 

on all sides; anterior ring twice as long as posterior ring.


Peristomial cirri to front edge of peristomium, digitiform, 


Maxillary formula 1+1, 9+10, 7+0, 6+10, and presumably 

1+1. Mx III long, located behind left Mx II. 





setigers. Maximum 12 slender, nearly filiform filaments. 

Filaments about as long as notopodial cirri. 

Ventral cirri basally inflated in median setigers. Posterior 

ventral cirri not inflated. Notopodial cirri digitiform, without 

articulations in all setigers. 

Shafts of compound falcigers distally inflated, marginally 

serrated. Appendages short, tapering, bidentate. Proximal teeth 

larger than distal teeth, triangular, directed laterally. Distal 

teeth short, obliquely directed distally. Guards symmetrically 

sharply pointed, with distinct mucros. Pseudocompound 

falcigers and compound spinigers absent. Aciculae yellow, 

slender, tapering, straight; cross-sections round. Separation 


subacicular hooks. Subacicular hooks yellow, tridentate with 

teeth in a crest Distribution unknown, present well before 

setiger 30. 

UNKNOWN MORPHOLOGICAL FEATURES.—Several features 



None. 



Characters: 4-6, 8, 10-17, 24, 39, 40, 42-52, 54-56, 60, 

65-68,80-81 


None. 

REMARKS.—Eunice limosa is listed with similar species in 

Tables 41 and 43. It is the only species in Table 43 to combine 

mucronate guards on the compound falcigers with as many as 

12 branchial filaments. It has usually been considered a 

synonym of E. vittata; the two are very similar and may not be 

separable. A comparison shows a series of minor differences 

and without knowledge of the variability in these characters a 

possible synonymy should be held in abeyance. 

An indication of the distribution of subacicular hooks is 

given by Enters in his illustration of parapodium (1868, pi. 15: 

fig. 17) shown as being a heavily branchiated parapodium with 

nine branchial filaments and subacicular hooks. In the text 

Ehlers indicated that such heavily branchiated parapodia are 

present only before setiger 30. 

106. Eunice lita (Chamberlin, 1919) 

FIGURE 65c-i; TABLES 33,38 

Leodice lita Chwnberlin, 1919a:240-244, pi. 54: figs. 6-10, pi. 55: figi. 1-7. 


(no station indicated), Marshall Islands, on anchor coming 

up from 12 fathoms, Exped. 1899-1900. 

DESCRIPTION.—Holotype complete (in 2 pieces), of unknown 

sex, with 240 setigers; length 78 mm; maximal width 4 

mm; length through setiger 10,4 mm. 


than peristomium, less than l /i as deep as peristomium. 

Prostomial lobes frontally obliquely truncate, dorsally excavate 

with a thickened rim; median sulcus deep. Eyes between bases 

of A-I and A-II, black. Antennae in horseshoe; A-I separated by 

gap from A-II and A-III; A-I thicker than A-II and A-III. 


Ceratostyles tapering, without articulations. A-I to middle of 


anteriorly somewhat flaring. Separation between rings very 

distinct both dorsally and ventrally and visible laterally. 

Anterior ring 5 /6 of total peristomial length. Peristomial cirri to 

middle of prostomium, medially inflated, without articulations. 

Maxillary formula 1+1,4+4,6+0,4+9, and 1+1. Mx III and 

left Mx IV forming distal arc. 



15 to setiger 120. Branchiae terminating well before posterior 

end, present on less than 55% of total number of setigers. Most 

branchiae single, digitiform filaments (Figure 65e); some 

branchiae, especially at about setigers 35-70 with 2 filaments. 

Filaments digitiform, longer than notopodial cirri. 

Anterior neuropodial acicular lobes distally truncate, becoming 

conical posteriorly; aciculae emerging at midline. Pre- and 


tapering, becoming basally inflated from about setiger 5. 

Inflated bases thick, transverse welts; narrow tips short and 

button-shaped. From about setiger 50 inflated bases reduced. 

Ventral cirri emerging increasingly from posterior face of 

parapodium, by last setigers present reduced to very small, 

nearly rudimentary knobs. Anterior notopodial cirri short, 

digitiform, becoming short, slender filaments at bases of much 

larger branchial filaments in branchial region. Notopodial cirri 



(Figure 65i) small. Shafts slender. Blades flaring, flat. One 

marginal tooth longer than other teeth, with 12 teeth. 

Compound falcigers (Figure 65h) long-shafted, shafts gently 

inflated, marginally smooth. Appendages short, sharply tapering; 

heads large, bidentate. Proximal teeth larger than distal 

teeth, tapering, directed laterally, distally upturned. Distal teeth 

thick, blunt, directed obliquely distally. Guards symmetrically 

rounded, marginally smooth; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculae single 

except in a few anterior setigers, dark, pointed, straight; 

cross-sections round. Posterior aciculae projecting well beyond 

acicular lobes. Subacicular hooks (Figure 65g) dark, bidentate. 



curved, distally abruptly tapering; heads large. Proximal teeth

198 


larger than distal teeth, directed laterally, distally upturned. 

Distal teeth nearly erect, triangular. 


anal cirri. 

and 


None. 



Unknown Characters: 4,6,74,78. 


None. 

REMARKS.—Eunice Uta is listed with similar species in 

Tables 33 and 38. An unusual feature of the species is the high 

number of setigers with single branchial filaments, both before 

and after the peak number is reached, and the fact that the 

branchiae terminate well before the posterior end of the body. 

Eunice longicirrata (Kinberg, 1865) 

Nicidion longicirrata Kinberg, 1865:564; 1910:43. pL 16: fig. 20. 

Eunice (Nicidion) cariboea.—Hartman, 1948:80 [not Eunice cariboea Gmbe, 

1856]. 

MATERIAL EXAMINED.—Holotype, RM 415, Honolulu, 

Hawaii, Eugenie Expedition. 

COMMENTS ON MATERIAL EXAMINED.—Kinberg 

(1865:564) gave locality information as "Mare pacificum juxta 

urbem Honolulu insulae Oahu, inter corallia, summa aqua." 

The type lot includes a very poorly preserved anterior end and 

a median fragment. The anterior end has been ripped apart and 

the whole jaw apparatus is missing. 

DESCRIPTION.—A-111 distinctly longer than A-I and A-II. 

Separation between peristomial rings distinct dorsally, but 

probably not laterally; unknown for ventral side. Peristomial 

cirri to frontal margin of peristomium. 

Branchial distribution cannot be determined. Median fragment 

with long, digitiform single branchial filaments. 

Shape of parapodial structures indeterminable. Notopodial 

cirri short, slender in median fragment 

Limbate setae marginally serrated. Shafts of compound 

hooks strongly inflated. Appendages short; both teeth directed 

laterally. Hoods short, symmetrically rounded, marginally 

serrated. Aciculae brown, distally bent Subacicular hooks and 

pectinate setae absent in both fragments. 

REMARKS.—Hartman (1948:80) referred Nicidion longicirrata 

to Grube's species, Eunice cariboea, originally described 

from Caribbean. This synonymy is incorrect. In E. cariboea all 

antennae are similar in size and digitiform, in the type of E. 

longicirrata, A-III is distinctly longer than other antennae and 

all antennae tapering. The absence not only of subacicular 

hooks, but also of pectinate setae points to an affiliation with 

Palola rather than with Eunice, but the specimen is in such poor 

shape that it cannot be accurately assigned to genus. Eunice 

longicirrata Kinberg is here considered indeterminable. 

Eunice longicirrata Webster, 1884 

Eunice longicirrata Webrter, 1884:318-319, pL 12: figi. 75-80. 

Eunice websteri Fauchald, 1969:12-14. fig. 6a-e. 

REMARKS.—Fauchald (1969:12) renamed this species, 

pointing out that the name applied by Webster had been 

preoccupied in combination E. longicirrata (Kinberg, 1865). 

The species is described below. 

107. Eunice longicirris Grube, 1869 

Eunice longicirris Grube, 1869:492. 

FIGURE 66a-f. 

MATERIAL EXAMINED.—Holotype, ZMB 499, Suez, coll. 

Ehrenberg. 


of one anterior end, two posterior ends, and several long 

sections of median setigers. The features of all pieces are 

closely similar and are here assumed to belong to the same 

species', however, it is unclear how the pieces match, so 

measurements were taken on the anterior fragment only. 

DESCRIPTION.—Anterior fragment of unknown sex, with 30 

setigers; length 43 mm; maximal width 5 mm; length through 

setiger 10,16 mm. 


than peristomium, as deep as x fi of peristomium. Prostomial 


shallow. Eyes posterior to bases of A-I; faded. Antennae in 

horseshoe; A-I and A-II separated from A-III by gap; A-III 

slimmer than A-I and A-II. Ceratophores ring-shaped in all 


to 14 long, articulations in A-III; basal articulations cylindrical, 

distal ones drop-shaped. A-I to posterior peristomial ring; all 

other antennae incomplete; A-II to setiger 1; A-III to setiger 3. 


dorsally and ventrally; anterior ring ~ 3 A of total peristomial 

length. Peristomial cirri outreaching prostomium, very slender, 

tapering, with 8 articulations. 

Maxillary formula 1+1,7+7,7+0, 8+10,1+1. Mx III part of 

distal arc with left Mx IV. Left Mx IV with very small teeth. 

Mx VI absent. 

Branchiae (Figure 66a) present, pectinate, distinctly shorter 

than notopodial cirri, erect Branchiae from setiger 3 to end; all 

branchiae pectinate except last 10-15 pairs. Maximum 7 

filaments at about setiger 15. Stems slender, cylindrical, shorter 

than filaments. Filaments slender, digitiform, shorter than 



triangular in posterior setigers; aciculae emerging above 


ventral cirri thick, tapering, becoming basally distinctly 

inflated from about setiger 5. Inflated bases ovate; narrow tips 

tapering. Posterior ventral cirri tapering without inflated bases.

NUMBER 523 

199 

FIGURE 66.—Eunice longicirris (holotype, ZMB 499): a, median parapodium, anterior view; b, compound 

falciger, median parapodium; c, compound falciger, median parapodium; d, anterior end, lateral view; e, pectinate 

seta, median parapodium; f, subacicularhook, median parapodium. Eunice longiselis (holotype, USNM 4791): g, 

anterior end, lateral view; h, pectinate seta, anterior parapodium; i, compound falciger, anterior parapodium; j, 

compound falciger, anterior parapodium; k, anterior parapodium, anterior view; 1, median parapodium, anterior 

view; m, aciculae, median parapodium; n, compound falciger, median parapodium; o, subacicular hook, median 



Notopodial cirri very slender, very long, nearly whip-like, 

longer than branchiae, with 5 or more long, slender articulations. 

Limbate setae slender, marginally smooth. Shafts of pectinate 

setae (Figure 66e) wide flat. Blades tapering, flat. One 

marginal tooth longer than other teeth, with -10 teeth. Shafts of 

compound falcigers (Figure 66b,c) slightly inflated, marginally 

serrated when not worn. Appendages thick, tapering, bidentate. 

Proximal teeth shorter than distal teeth, tapering, directed 

laterally. Distal teeth distinctly thicker than proximal teeth, 

bent, tapering. Many appendages with both teeth distinctly 

worn, rounded or truncated. Guards asymmetrically bluntly 

pointed, marginally serrated; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculac paired, 

chestnut-colored without distinction between cores and 

sheaths, tapering, straight in anterior and median setigers, 

becoming bent ventrally in posterior setigers; cross-sections 

round. Subacicular hooks (Figure 660 w 'th chestnut-colored 

cores and clear sheaths, bidentate. Hooks first present from 


(except for replacements). Hooks tapering, with distinct heads. 

Proximal teeth larger than distal teeth, slender, directed 

laterally. Distal teeth blunt, nearly erect. 




None. 



Characters: 1, 2,4,6, 23, 33, 36-40,47, 50,63. 


37,1; 38,1. 

REMARKS.—Eunice longicirris belongs to group B-2; it is 

listed with similar species in Tables 27 and 32. It is primarily 

characterized by the very long, slender, articulated notopodial 

cirri and peristomial cirri. It is the only species listed in Table 

32 in which the peristomial cirri clearly outreach the 

prostomium. 

Eunice longicornis Grube, 1866 

Eunice longicornis Grube, 1866b:68. 

Eunice longicirris.—Grube, 1878b: 100 [bottom of page, not Eunice longicirris 

Grube, 1869:492]. 

REMARKS.—This species was described with branchiae from 

setiger 6 to the end of the incomplete specimen (145+ setigers), 

where best developed 23 filaments were present, and even in 

the last setiger 19 filaments were present. A-III reached 

segment 7; all antennae and peristomial cirri without articulations. 

The branchiae outreach the notopodial cirri by setiger 12. 

Through a lapsus, this species was referred to as E. 

longicirris in a tabular review of the genus issued by Grube 

(1878b: 100); however, on the same page is also mentioned the 

very different species described as E. longicirris by Grube in 

1869 and redescribed, based on the type above. 

No material is available of E. longicornis and the original 

description is insufficient to characterize the species; it is here 


Eunice longiqua Kinberg, 1865 

Eunice longiqua Kinberg, 1865:563. 

Eunice antennaia.—Hartman, 1948:78-79 [in part, not Lamarck, 1818]. 

REMARKS.—The single specimen reported by Harunan 

(1948:78) is no longer present in the collections of Riksmuseet, 

Stockholm (R. OlertJd, in litL). The available information, 

summarized from Kinberg (1865) and Hartman (1948), is given 

below. 

Antennae, peristomial cirri, and notopodial cirri with 

moniliform articulations. A-III to setiger 5; with 20 articulations. 

Branchiae from setiger 6 to end of body; median region 

with lowered number of branchial filaments. Aciculac yellow; 

tapering with blunt, straight or gently curved tips. Subacicular 

hooks yellow; tridentate. Compound hooks distally bidentate. 

Eunice longiqua was reported from Foua in the Pacific 

Ocean. It belongs to group C-2. Without access to any material, 

the species cannot be separated from a series of similar species 

and it is here considered indeterminable. 

108. Eunice longisetis Webster, 1884 

FIGURE 66g-o 

Eunice longisetis Webster. 1884:317-318, pi. 10: Tigs. 46, 46a,b, 47-49. 

MATERIAL EXAMINED.—Holotype and 4 slides of parapodia, 

USNM 4791, Bermuda, coll. G. Brown Goode, 1876. 



through setiger 10,6.5 mm. Body thickset, cylindrical, tapering 

posteriorly, vaguely dorsoventral flattened. 



lobes frontally rounded, dorsally depressed and flattened; 

median sulcus shallow. Eyes between bases of A-I and A-II, 

purple. Antennae in horseshoe, evenly spaced, with A-I thicker 

than other antennae. Ceratophores ring-shaped in all antennae, 

without articulations. Ceratostyles thick and digitiform, with up 

to 10 distinct, cylindrical articulations in A-II. A-I to middle of 

anterior peristomial ring; A-II to setiger 4; A-III incomplete, at 

least as long as A-II judging from its girth. Peristomium 

cylindrical. Separation between rings distinct dorsally and 

ventrally; anterior ring 5 /« of total peristomial length. Peristomial 

cirri to middle of anterior peristomial ring, digitiform, 





Branchiae from setiger 6 to setiger 104. Branchiae present to



of setigers. First and last 15 branchiae single filaments, 

maximum 8 filaments. Stems slender, tapering, longer than 

notopodial cirri. Filaments digitiform, shorter than notopodial 

cirri. 

Neuropodial acicular lobes obliquely conical with aciculae 

emerging from tips of lobes, dorsal to midline. Presetal lobes 

low, transverse folds. Postsetal lobes free, about as long as 

acicular lobes, visible behind acicular lobes. Anterior ventral 

cirri thick, tapering, becoming ventrally inflated from about 

setiger 10. Inflated bases forming short transverse inflated 

ridges along lower edge of parapodia; narrow tips long and 

digitiform. Posterior to setiger 45 ventral cirri increasingly 

digitiform, elongated. Notopodial cirri digitiform, tapering, 

similar in length. Anterior notopodial cirri articulated (Figure 

66k), loosing articulations in early branchial setigers. 

Limbate setae long, slender, very narrowly limbate, marginally 

smooth. Pectinate setae (Figure 66h) narrow, tapering, flat. 

One marginal tooth longer than other teeth, with 7 teeth in 

anterior setigers and 12 teeth in median and posterior sctigers. 

Shafts of anterior compound falcigcrs (Figure 66i,j) inflated, 

becoming tapering in median and posterior setigers (Figure 

66n), all marginally smooth. Anterior appendages relatively 

short, tapering, with small heads, bidentate. Proximal teeth 

shorter than distal teeth, tapering, directed obliquely distally. 

Distal teeth long, slender, strongly curved. Median appendages 

very slender, with large heads, bidentate. Proximal teeth much 

larger than distal teeth, strongly tapering, directed laterally. 

Distal teeth strongly tapering, directed obliquely distally. 

Anterior guards symmetrically bluntly pointed, becoming 

distinctly rounded in posterior setigers, marginally smooth; 


spinigers absent. Aciculae (Figure 66m) paired, with dark 

brown to black cores and clear sheaths, slender, tapering, 

distinctly curved dorsally; cross-sections round. Subacicular 

hooks (Figure 66o) with dark brown to black cores and clear 

sheaths, bidentate. Hooks first present from setiger 18, present 


Hooks slender, tapering, with distinct heads. Proximal 

teeth larger than distal teeth, directed laterally. Distal teeth 

slender, curved. 




None. 



Characters: 4, 6, 23,42. 


None. 

REMARKS.—Eunice longisetis belongs to group B-2 and is 

listed with similar species in Tables 27 and 30. The only 

unusual feature, compared to the other species in Table 30, 

appears to be the high number of posterior branchiated setigers 

with single filaments; most species listed in this table have a 

maximum of five; E. longisetis and E. fimbriata have 15 and 

10-12, respectively. The latter have branchiae terminating well 

before the posterior end; in E. longisetis the branchiae are 

continued to the far posterior setigers. 

Eunice lucei Grube, 1856:57-58. 

109. Eunice lucei Grube, 1856 



and a posterior fragments, ZMC (uncataloged), Puntarenas, 

Costa Rica, coll. H. KrOyer, 1846. 

DESCRIPTION.—Lectotype of unknown sex, with 170 setigers, 

missing perhaps 15 setigers; length 93 mm; maximal 

width 3 mm at about setiger 40; length through setiger 10, 5.5 

mm. 




sulcus deep. Palpal region marked by vague frontal horizontal 

groove. Eyes posterior to bases of A-I, dark. Antennae in 

horseshoe; A-I separated from other antennae by gap, similar in 

thickness. Ceratophores ring-shaped, in all antennae, without 

articulations. Ceratostyles tapering, with up to 14 very distinct, 

nearly moniliform articulations; inner l /s of A-II and A-IH 


A-II to setiger 1; A-III to setiger 3. Antennae of paralectotypes 

slightly shorter, but otherwise similar. Peristomium basically 

cylindrical with large, inflated lower lip. Separation between 

rings distinct on all sides; anterior ring 4 /s of total peristomial 

length. Peristomial cirri to middle of peristomium, slightly 

inflated basally, tapering to fine, slender tips, with up to 7 

distinct, cylindrical articulations. 

Maxillary formula of 1 paralectotype 1+1, 7+7, 9+0, 4+10, 

and 1+1. Mx III long, located behind left Mx II. Mx VI missing. 

Branchiae (Figure 67a) present, pectinate, distinctly longer 




of setigers. First and at least last 10-15 pairs short, single 

filaments (Figure 67h), maximum 8 filaments. Stems relatively 

slender, tapering, as long as notopodial cirri. Filaments slender, 

as long as notopodial cirri. 


rounded in median setigers and conical in last setigers present; 


transverse folds. Anterior ventral cirri thick, tapering, becoming 

basally inflated from about setiger 5. Inflated bases of 

median region thick, inflated ridges; narrow tips very short and 

button-shaped. From about setiger 80 ventral cirri increasingly 

dominated by a long, tapering distal tip; inflated bases 

decreasing. Last 20 ventral cirri present long, digitiform, flexed

202 


FIGURE 67.—Eunice lucei (lectotype, ZMC): a, parapodium 45, anterior view; b, subacicular hook, parapodium 

45; c, compound falciger, parapodium 45; d, anterior end, lateral view; e, pectinate seta, parapodium 45; f, 

aciculae, parapodium 45; g, aciculae, parapodium 100; h, parapodium 160, anterior view; i, subacicular hook, 

parapodium 160; j, aciculae, parapodium 160; k, compound falciger, parapodium 160. (Scale bars in mm.) 

up behind neuropodial acicular lobe. Notopodial cirri basally 

inflated, with 5 to 6 cylindrical articulations. Notopodial cirri of 

posterior setigers very large, longer than in anterior setigers and 

nearly as wide at base as neuropodial acicular lobe. 


(Figure 67d) tapering, flat. One marginal tooth slightly longer 

than other teeth, with 15 teeth. Shafts of compound falcigers 

(Figure 67c,k) distally slightly inflated, marginally serrated. 

Anterior appendages (Figure 67c) slender, tapering, bidentate. 

Proximal teeth thicker, but shorter than distal teeth, triangular,


TABLE 6.—Variable and invariable features in the type lot of Eunice lucei (N = number of individuals examined; 

SD = standard deviation; measurements in mm). 



Total length 

Maximal width 



Subacicular hooks first present from 

N 

4 

4 

4 

4 

4 

4 

Max. 

195 

93 

3.5 

7.5 

14 

34 

Min. 

108 

22 

2 

3 

8 

24 

Mean 

151 

58.25 

2.75 

4.88 

12.5 

28.75 

SD 

33.71 

32.42 

0.56 

1.78 

2.60 

4.32 


Peristomial rings 


Branchiae first present from sctigcr no. 

End of branchiae 



Guards of compound hooks 


Acicular tip 



N=4 

complete 

5 

5 

prepygidially 

10 

15 

mucronate 

yellow 

hammer-headed 

yellow 

3 

directed laterally. Distal teeth nearly erect. Guards symmetrically 

rounded, with short, distinct mucros, marginally serrated. 

Posterior appendages (Figure 67k) thicker, bidentate. Proximal 

teeth smaller than distal teeth, short, triangular. Distal teeth 

large, thick, distinctly bent. Guards asymmetrically bluntly 

pointed, marginally serrated; mucros absent. Compound hooks 

of paralectotypes with both kinds of appendages as well. 


Aciculae paired, yellow, most distinctly hammer-headed, 

nearly bifid (Figure 67e,g,j), some superior aciculae tapering, 



Subacicular hooks (Figure 67b,i) yellow, tridentate with teeth 

in a crest. Hooks first present from setiger 24-34, present in all 


Main fangs large curved; fangs decreasing evenly in size to 

slender, distinct tertiary fangs. 


anal cirri. 


None. 



Characters: 1,2, 6,40,42. 


None. 

REMARKS.—Eunice lucei has remained poorly understood 

since its first description; it is listed with similar species in 

Tables 46 and 47. Unusual are the long, thick notopodial cirri 

in posterior setigers and the mixture of mucronate and 

amucronate guards on the compound falcigers. Four species in 

Table 47 have modified aciculae. These species include E. 

elseyi, E. makemoana, and E. rubra, in addition to E. lucei. 

Eunice elseyi has tridentate subacicular hooks with the distal 

teeth in tandem; the other three species have crested tridentate 

hooks. In E. lucei the aciculae may be hammer-headed or 

weakly bidentate; in E. makemoana they are pointed or 

hammer-headed, and in E. rubra they are pointed or weakly 

bidentate. Eunice makemoana has single branchial filaments; 

E. lucei has up to eight branchial filaments, and E. rubra has as 

many as 21 filaments. 

110. Eunice macrobranchia Schmarda, 1861 

TABLES 27,30,50 

Eunice macrobranchia Schmarda, 1861:130,7 figs., pi. 32: fig. 258. 

Eunice (Eriphyle) macrobranchia.—Augener, 1918:310-312. 

Eriphyle capensis.—AugeneT, 1918:310-312. 

REMARKS.—No types are currently available of this species; 

however, Augener (1918) did review the types. The following 

comments combines information in Schmarda's original 

description with additional information specified by Augener. 

Type very large of about 200 mm in total length, with about 

150 setigers. Ceratostyles irregularly articulated. A-I to 


Peristomial cirri shorter than peristomium. Maxillary formula 

indeterminable. Branchiae present, pectinate, about as long as 

notopodial cirri, erect. Branchiae from setiger 7-8; first several 

pairs single filaments. Where best developed branchiae with 15 

to 17 filaments. Schmarda illustrated parapodia as having very 

long, thick, tapering notopodial cirri of very nearly same length 

as branchiae. Pectinate setae were illustrated as narrow, 

tapering, with relatively few teeth and no marginal teeth 

obviously longer than other teeth. Appendages of compound 

falcigers bidentate; guards illustrated as symmetrically


rounded; mucros absent. Pseudocompound falcigers and 

compound spinigers absent. Aciculae and subacicular hooks 

black; subacicular hooks present in second half of body and 

either uni- or bidentate. 

Schmarda's illustration of the jaws is so confused that no 

maxillary formula can be recorded. He must also have missed 

the first several pairs of simple branchiae and stated that 

branchiae were from setiger 12 where the strongly pectinate 

complex branchiae start according to his illustration. 


SCORED.—Inappropriate Characters: Unknown Characters: 

3-20, 23-26, 28-29, 33, 36-38, 40, 42-65, 67, 74-76, 

78, 80-82. 


37,1; 38,1. 

REMARKS.—Augener (1918) synonymized Eriphyle capensis 

Kinberg (1865 = Eunice kinbergi Ehlers, 1868) with this 

species, but rejected the synonymy of both with E. aphroditois 

as proposed by Mclntosh (1905). All three species belong to the 

same group, B-2, and are rather similar. Mclntosh's synonyms 

have been generally accepted (e.g., Day, 1967). The species are 

here listed separately in Tables 27 and 30, because resolution of 

this problem will have to await studies of morphological 

variability in material of E. aphroditois from Indian Ocean. 

Eunice macrobranchia has not been included in the key. 

Eunice macrobranchia is supposed to have either simple, 

spine-like or bidentate subacicular hooks. For that reason it is 

also listed with other species with simple subacicular hooks in 

Table 50. Because it is so poorly known, it cannot be clearly 

separated from other species in this table. 

Eunice macrochaeta Schmarda, 1861 

Eunice macrochaeta Schmarda, 1861:128, 6 figs. 

REMARKS.—Schmarda (1861:128) stated that the type and 

apparently only known specimen lacked the anterior end. The 

species is thus wholly unidentifiable. Grube (1878b: 101) 

referred the species under doubt to the genus Marphysa based 

on the illustration of the branchiae. Hartman (1944:107) 

referred the species, again doubtfully, to E. filamentosa, 

following suggestions by Augener (1925). The structure of the 

setae as illustrated by Schmarda (1861:128) agree with E. 

filamentosa, except that the characteristic hammer-headed 

aciculae of that species were not illustrated. The species is here 


Eunice madeirensis Baird, 1869 

Eunice madeirensis Baird, 1869:344-345. 



the species in the genus Pcdola. 

111. Eunice magellanica Mclntosh, 1885 


Eunice magellanica Mclntosh, 1885:265-268, fig. 28, pi. 37: figs. 12-15, pi. 

19A: figs. 6-9. 

MATERIAL EXAMINED.—TWO syntypes, BM(NH) 

ZK.1885.12.1.191, and slide preparation, ZK.1885.12.1.191a. 

Challenger sta 311, Magellan Strait, north of Isla Desolation, 

52°50'S, 73°53'W,448 m, mud, 11 Jan 1876. 

COMMENTS ON MATERIAL EXAMINED.—Unless otherwise 

indicated, the description is based on the larger of the two 

syntypes. 

DESCRIPTION.—Larger of 2 syntypes complete female with 

eggs in body cavity with 130 setigers; total length 140 mm; 

maximal width 7 mm; length through setiger 10, 17 mm. Small 

syntypc complete (in 2 pieces) with 108 sctigcrs; total length 78 

mm; maximal width 5 mm; length through sctigcr 10, 11 mm. 


than pcristomium, less than '/.• as deep as pcristomium. 

Prostomial lobes short, frontal ly rounded; median sulcus deep. 

Eyes posterior to bases of A-I, black. Antennae evenly spaced, 

similar in thickness. Ccratophores ring-shaped in all antennae, 

without articulations. Ceratostylcs digitiform, with up to 5 

irregular articulations in A-I 11. A-I barely to middle of anterior 

peristomial ring; A-I I to posterior peristomial ring; A-I 11 to 

setiger 2. Peristomial fold deep, covering bases of antennae; 

peristomium flaring slightly ventrally; lower lip muscular. 


anterior ring 4 /5 of total peristomial length. Peristomial cirri of 

large syntype very short, ovate, to middle of peristomium in 

small syntype, without articulations. 

Two sets of maxillae present in vial, both of roughly same 

size, either one could belong to large syntype; small syntype 

not dissected. One maxillary formula 1+1, 6+6, 8+0, 8+10, 

1+1, and 1+1. Other maxillary formula 1+1, 7+7, 8+0, 7+9, 

1+1, and 1+1. Both maxillae with Mx III forming part of distal 

arc with left Mx IV. Mx VI with distinct teeth in both sets of 

jaws. 



Branchiae from setiger 7-8 to setiger 120. Branchiae present to 


of setigers. First branchiae with 2 filaments, increasing to 5 

filaments at about setiger 20. From about setiger 70-75 

number of filaments increasing rapidly to 20 at setiger 85; this 

number continued to setiger 115 in large syntype. Some 

filaments irregularly branching. Reduction in size and numbers 

of filaments in far posterior setigers very rapid. Where best 

developed branchial stems thick, somewhat coiled, and so 

branchiae may appear palmately, rather than pectinately 

branched. 

Neuropodial acicular lobes distally truncate or rounded; 

aciculae emerging at midline. All pre- and postsetal lobes low,

NUMBER 523 

205 

FIGURE 68.—Eunice magellanica (syntype, BM(NH) ZK.1885.12.1.191 and slide preparation, 

ZK.1885.12.1.191a): a, anterior end, lateral view; b, subacicular hook, parapodium 102; c, subacicular hook, 

parapodium 100; d, appendage of compound falciger, parapodium 100; e, parapodium 100, anterior view; f, 

pectinate seta, parapodium 100. Eunice magnifica (holotype, ZMB F1993):g, anterior parapodium, anterior view; 

h, pectinate seu, median parapodium; i, compound falciger, anterior parapodium; j, compound falciger, median 

parapodium; k, median parapodium, anterior view. (Scale bars in mm.)


transverse folds. First 5 ventral cirri thick, digitiform. Ventral 

cirri basally inflated from about setiger 6 through setiger 30, 

inflated bases ovate; narrow tips tapering. Posterior ventral cirri 

digitiform. All notopodial cirri with 5 to 6 irregular articulations. 

Anterior notopodial cirri thick, digitiform. Posterior 

notopodial cirri relatively larger than anterior ones, becoming 

very prominent in far posterior setigers. 

Limbate setae slender. Pectinate setae (Figure 68f) tapering, 

flat. Both marginal teeth slightly larger than other teeth, -10 

teeth present. Shafts of compound falcigers inflated, marginally 

smooth. Appendages (Figure 68d) long, bidentate. Proximal 

teeth much larger than distal teeth, triangular, directed laterally. 

Distal teeth tapering, nearly erect or gently curved. Guards 


falcigers and compound spinigers absent Aciculae 

numbering up to 3, with black cores and clear sheaths, tapering 

to blunt tips; ventralmost aciculae gently curved in most 

parapodia; cross-sections round. Subacicular hooks (Figure 

68b,c) with dark brown to black cores and clear sheaths, 

bidentate. Hooks first present from setiger 26 or 30, present in 


Proximal teeth about twice as large as distal teeth, directed 

laterally. 


anal cirri. 







None. 

REMARKS.—Hartman (1964:118, pi. 36: figs. 8-10) referred 

E. magellanica to E. frauenfeldi, which it resembles in many 

respects. Note that Hartman's illustrations were redrawn from 

Mclntosh's original description of E. magellanica and most of 

the text was taken from Mclntosh as well; thus Hartman's 

description does not, except as the two species are identical, 

refer to Grube's E. frauenfeldi. The two species appear to differ 

most clearly in the distribution of branchiae. In E. frauenfeldi 

the maximum number of filaments is -3 at setigers 15-25; In 

E. magellanica the maximum number of filaments is -20 near 

setiger 85. These and other species in group B-2 are compared 

in Tables 27 and 29. 

112. Eunice magnifica Grube, 1866 

Eunice magnifica Grube, 1866a:64. 

FIGURE 68g-k; TABLES 33,37-39 

MATERIAL EXAMINED.—Holotype, ZMB F1993 (marked 

E. bipapillata), Samoa, 2 parapodia. 

COMMENTS ON MATERIAL EXAMINED.—The identity of the 

two parapodia examined was discussed above in the section of 

E. bipapillata. 

REMARKS.—The following is a summary of Grube's original 

description combined with notes on the two parapodia. 

Type incomplete with 102 setigers; about 100 mm in length; 

violet-copper-colored with white spots. 


cirri, erecL Branchiae from setiger 22, with up to 8 

filaments. 

One parapodium from an anterior segment (Figure 68g), 

probably before setiger 15 of a large specimen, with neuropodial 

acicular lobe distally transverse; aciculae emerging at 

midline. Presetal lobe a low fold; postsctal lobe a high ridge 

behind acicular lobe. Notopodial cirrus basally slightly 

inflated, tapering to a digitiform tip; ventral cirrus thick, with a 

thick, blunt tip, without articulations. 

The other parapodium is distorted and cannot be described in 

detail; it docs not appear to have been remarkably different 

from usual pattern in genus. Notopodium is similar to the one 

present in the other parapodium and a very large branchia is 

present (Figure 68k); it has a strong branchial stem and 8 

distinct branchial filaments. Ventral cirrus without basal 

inflation; tapering. The label indicates that this parapodium 

should be from segment 96; this agrees with the shape of 

acicular lobe and ventral cirrus. The branchial development is 

unusual for such a late segment, but may be related to the 

apparent very large size of the specimen. 

Pectinate setae (Figure 68h) tapering in both parapodia. One 

marginal tooth distinctly longer than other teeth; 12 teeth 

present. Shafts of compound falcigers (Figure 68i,j) distally 

inflated, with smooth cutting edge; distal beak indistinct. 

Appendages slender, tapering with large head anteriorly; with 

small heads posteriorly, bidentate. Proximal teeth triangular in 

anterior setigers, slender and tapering posteriorly, directed 

laterally or slightly distally. Distal teeth nearly erect, nearly 

triangular in anterior setigers, slender and tapering in posterior 

setigers. Guards asymmetrically bluntly pointed; mucros 


absent. Aciculae single, dark, bluntly conical, straight; crosssections 

round. Separation between core and sheath distinct in 

both aciculae and subacicular hooks. Subacicular hook present 

in branchiated parapodium, single, dark, distally broken so the 

dentition cannot be determined. 



Characters: 1-29, 33, 36-40, 42, 46, 47, 49-51, 53-56, 62, 

80-82. 


None. 

REMARKS.—The first parapodium described should be from 

setiger 26, according to the label. This would be the beginning 

of the branchial region; it is, however, distinctly from a 

prebranchial setiger. The number of the parapodia may have 

been confused, but the structure of aciculae and compound


falcigers agree well with the other parapodium present. It is 

here assumed that parapodium number indicated is erroneous. 

Eunice magnified belongs to group B-4; it differs from E. afra 

and related species in that the branchiae are very well 

developed as far back as setiger 96; it is listed with similar 

species in Tables 33 and 37-39. It is too poorly known to be 

included in the key and cannot really be discussed in detail in 

relation to other species in tables 37-39. 

113. FM nice make mo ana (Chamberlin, 1919) 

TABLES 46.47 

Leodice makemoana Chamberlin, 1919a:233-236, plate S3: figs. 1-11. 

MATERIAL EXAMINED.—Holotype, USNM 19396, Makemo, 

Paumotu Islands (= Tuamotu Archipelago), 21 Oct 1899. 

COMMENTS ON MATERIAL EXAMINED.—The holotype and 

only known specimen of species has been frontally deeply 

dissected and has been dry at one time; all setae are broken, so 

no meaningful illustrations can be made. The following 

remarks are based on Chambcrlin's original description in 

addition to the few observations that could be made on the 

specimen. 


length 20 mm; maximal width 1 mm; length through setiger 10, 

3.5 mm. 


lobes frontally rounded, dorsally inflated. Eyes behind bases of 

A-I. Ceratophores ring-shaped in all antennae, without articulations. 

Ceratostyles with up to 15 moniliform articulations in 

A-I 11. A-I to middle of peristomium; A-II to setiger 3; A-III to 

setiger 4. 

Maxillary formula (re-written from Chamberlin, 1919a:236): 

1+1, 6+7, 8+0, 8+ll(12), and 1+1. 

Branchiae from setiger 7 to 6 setigers from posterior end. 


65% of total number of setigers. All branchiae single filaments. 

Neuropodial acicular lobes distally rounded or truncate. 

Pre- and postsetal lobes low, transverse folds. First few ventral 

cirri thick, tapering, becoming basally inflated in anterior 

branchial region. Posteriorly inflated bases decreasing in 

importance; far posterior ventral cirri slender, resembling 

notopodial cirri. All notopodial cirri medially inflated, articulated 

with relatively short, vague articulations, shorter and 

more slender in posterior than in anterior setigers. 

Pectinate setae tapering, with -10 coarse teeth. One marginal 

tooth much longer than other teeth. Shafts of compound 

falcigers inflated. Appendages short, bidentate. Both teeth 

distally well curved. Guards asymmetrically bluntly pointed; 


spinigers absent. Aciculae paired, yellow; ventral aciculae 

slender, distally pointed, curved ventrally; dorsal aciculae 

thick, distally blunt, possibly slightly hammer-headed; crosssections 



yellow, tridentate with teeth in a crest. Hooks first present from 

setiger 28. Fangs decreasing in size from large main fang to 

very small distal fang. 






SCORED.—Inappropriate Characters: 33, 34, 39, 40,42, 58, 

59. Unknown Characters: 4, 6, 9, 11, 12, 15-17, 20, 24-29, 

44,51,54-56,65,67,68,81,82. 


None. 

REMARKS.—Eunice makemoana is listed with similar 

species in Tables 46 and 47. It appears unique in the group in 

having single branchial filaments in all setigers; all other 

species in group have branched branchiae at least in some 

setigers. As pointed out by Chamberlin (1919a:236), the 

reduced condition of branchiae might be considered a larval 

feature, but the sexually mature state of the individual argues 

for retaining separate status of specimen. 

114. Eunice manihine Longbottom, 1972 

FIGURE 69a-g; TABLES 19,21 

Eunice manihine Longbottom, 1972:339-344, figs. 1-17. 


western Indian Ocean, Cosmoledo, S of Menai Island, 10°S, 

47°E, foul-hooked, 421 m, coral debris, 6 Feb 1969, coll. M.R. 

Longbottom, Royal Society Indian Ocean Deep Slope Fishing 

Expedition. 



9 mm. 




median sulcus deep. Eyes absent Antennae arranged in 

a horseshoe, evenly spaced. Ceratophores ring-shaped in all 

antennae, without articulations. Only a single A-II attached to 

prostomium, slender and tapering, without articulations, reaching 

setiger 10. Another ceratostyle, presumably A-III, loose in 

vial, same shape as A-II, considerably longer, probably 

reaching setiger 20. Peristomium much longer ventrally than 

dorsally, projecting forward nearly to tip of prostomium, with 

a distinct muscular lower lip. Separation between rings distinct 

dorsally and ventrally; anterior ring 2 h of total length dorsally, 

~ 3 /4 ventrally. Peristomial cirri to middle of prostomium, 


Maxillary formula 1+1, 12+11, 12+0, 11+13, and 1+1 

according to Longbottom (1972). 

Branchiae present, palmate, distinctly longer than notopodial

208 


m 

FIGURE 69.—Eunice manihine (holotype, BM(NH) ZB 1971.1): a, anterior end, lateral view; b, subacicular hook, 

parapodium 34; c, acicula, parapodium 34; d, pectinate seta, parapodium 90; e, parapodium 34, anterior view; f, 

compound falciger, parapodium 34; g, pectinate seu, parapodium 90. Eunice marenzelleri (syntype, MNHN): h, 

anterior end, lateral view; i, compound falciger, parapodium 125; j, subacicular hook, parapodium 125; k, acicula, 

parapodium 125; 1, pectinate seta, parapodium 125; m, parapodium 125, anterior view. (Scale ban in mm.)



to setiger 43. Branchiae terminating well before posterior end, 

present on less than 55% of total number of setigers. Most 

branchiae single, slender filaments (Figure 69e), up to 3 

filaments present 

Neuropodial acicular lobes distally truncate, usually somewhat 

lobed between aciculae; aciculae emerging at midline. All 

pre- and postsetal lobes low, transverse folds. First 4 ventral 

cirri thick, tapering. From setiger 5 bases increasingly inflated, 

by setiger 25 inflated bases projecting as ridges on ventral side 

of neuropodia; narrow tips short and button-shaped Inflated 

bases reduced from about setiger 50, by setiger 60 inflated 

bases completely lost. Ventral cirri thick, tapering through 

remainder of body. Anterior notopodial cirri strongly inflated 

basally; notopodial cirri of branchial region tapering; postbranchial 

notopodial cirri digitiform; all without articulations. 

Limbaic setae present marginally smooth. Pectinate setae 

(Figure 69d,g) flaring, flat. One marginal tooth slightly longer 

than other teeth, -15 teeth present. Shafts of compound 

falcigers (Figure 690 distally inflated, marginally smooth. 

Appendages short, bidentate. Both teeth well developed. 

Guards symmetrically bluntly pointed; mucros absent Pseudocompound 

falcigers and compound spinigers absent Notoaciculae 

present in base of notopodial cirri in middle part of 

body. All neuropodia with multiple aciculae arranged in 

dorsoventral row, at least 3 aciculae, but up to 6 aciculae 

present, yellow, tapering to blunt tip (Figure 69c), especially 

inferior aciculae slightly bent; cross-sections round. Separation 


subacicular hooks. Subacicular hooks (Figure 69b) yellow, 


setigers thereafter; most parapodia with 2 hooks, up to 4 hooks 

present. Both teeth well developed, rather similar in size. 


anal cirri. 




SCORED.—Inappropriate Characters: 22, 34, 39,40, 56, 58, 

59. Unknown Characters: 4,6, 17, 23,42. 


None. 

REMARKS.—Unusual features of this species include the 

multiple aciculae and subacicular hooks, combined with the 

late start of branchiae and well-developed basal inflation of 

ventral cirri. Eunice manihine is listed with similar species in 

Tables 19 and 21. 

Eunice manorae Aziz, 1938 

Eunice manorae Aziz:1938:34-35. pi. 3: fig.3. pi. 6: fig. 34. pi. 7: figs. 61-64. 

REMARKS.—The type is not available. The species apparently 

belongs to group B-4. It has non-articulated, relatively 

long antennae; peristomial cirri do not outreach peristomium. It 

has branchiae from setiger 80 through posterior region; with up 

to 5 filaments. Aziz' illustration indicates that the branchial 

filaments are longer than the notopodial cirri. 

Aciculae dark-colored; pectinate setae and compound hooks 

present, the latter with both teeth well developed. The distal 

tooth is illustrated as being considerably shorter than the 

proximal teeth; guards are symmetrically rounded. Subacicular 

hooks not illustrated nor mentioned in the text. 

The development of branchiae and the presence of pectinate 

setae precludes placement of the species in Palola; however, no 

other related taxa are known to lack subacicular hooks. Without 

access to the type, exact position of this species cannot be 

determined and it is here considered indeterminable. 

115. Eunice marenzelleri Gravier, 1900 

FIGURE 69h-m; TABLES 33,37,50 

Eunice marenzelleri Gravier, 1900:229-232. figs. 78-82, pi. 13: figs. 68,69. 

MATERIAL EXAMINED.—TWO syntypes, MNHN, Paris, 

Djibouti, Red Sea, coll. Coutiere (label marked no. 29). 

COMMENTS ON MATERIAL EXAMINED.—Syntype material 

contains two incomplete specimens; the long syntype has been 

slightly desiccated at one time; illustrations are based on the 

long syntype. 

DESCRIPTION.—Short syntype with 91 setigers. Long syntype 

with 257 setigers; length 150 mm; maximal width 5 mm; 

length through setiger 10,11 mm. 




sulcus shallow. Eyes posterior to bases of A-I, black. Antennae 

in a horseshoe, with A-I 11 emerging near peristomial fold; bases 

of A-I and A-II well forward of A-III; A-I distinctly thicker 

than other antennae. Ceratophores ring-shaped in all antennae, 

without articulations. Ceratostyles digitiform, without articulations. 

A-I to middle of peristomium; A-II and A-III to posterior 

edge of peristomium. Peristomium cylindrical, massively 

muscular. Separation between rings distinct dorsally and 

ventrally; anterior ring 4 /s of total peristomial length. Peristomial 

cirri short, nearly ovate, without articulations. 

Maxillary formula for 2 syntypes 1+1, 444, 5+0, 4+5, and 

1+1. Mx III and left Mx IV part of distal arc. Mx VI absent 

Branchiae (Figure 69m) present single filaments, distinctly 

longer than notopodial cirri. Branchiae from setiger 28 in both 

specimens and present on last setigers present. All filaments 

long. A single medioposterior segment on the smaller syntype 

with 3 filaments in a palmate arrangement 

Anterior neuropodial acicular lobes truncate; median and 

posterior acicular lobes rounded; aciculae emerging at midline. 

All presetal lobes low, transverse folds. Anterior postsetal lobes 

as high as or slightly higher than acicular lobes, becoming low, 

transverse folds from about setiger 25. Anterior ventral cirri


thick; tapering, becoming basally inflated from about setiger 

12; inflated bases retained to about setiger 100. Inflated bases 

long transverse welts; narrow tips tapering. Posterior ventral 

cirri tapering. Anterior notopodial cirri distinctly medially 

inflated, becoming short and strongly tapering from about 

setiger 75, without articulations. 


(Figure 691) large, flaring, flat. Both marginal teeth longer than 

other teeth; -20 teeth present Shafts of compound falcigers 

(Figure 69i) distinctly inflated, marginally serrated. Appendages 

slender, narrow, with nearly parallel sides, bidentate. 

Proximal and distal teeth similar in size. Proximal teeth 

tapering, directed laterally. Distal teeth curved. Guards symmetrically 

rounded; mucros absent. Pseudocompound falcigers 

and compound spinigers absent. Aciculae (Figure 69k) dark 

brown, tapering to blunt, distally rounded tips, straight; 

cross-sections round. Subacicular hooks (Figure 69j) dark 

brown, unidentate. Hooks first present from setiger 38, present 


Tips more or less gently curved, slightly asymmetrical 

in some setigers, but no trace of secondary teeth seen. 

Notopodial aciculae light brown. 


with the far posterior end. 




SCORED.—Inappropriate Characters: 22, 33, 34, 39, 40,42, 

56, 58, 59. Unknown Characters: 1,2,4, 6, 36-38,64, 78. 


38,1. 

REMARKS.—Eunice marenzelleri is listed with similar 


with simple, spine-like subacicular hooks. It is listed with other 

species with simple spine-like subacicular hooks in Table 50. It 

is the only species in Table 50 with exclusively single branchial 

filaments. 

Eunice margaritacea Williams, 1853 

Eunice margaritacea Williams, 1853:408, fig. 4. 

REMARKS.—The article in which this species was named 

concerns mechanisms of aquatic respiration in invertebrates. 

The species name is mentioned in the caption to an illustration 

showing the pectinate branchial structure of a eunicid. The 

discussion in the text gives no additional information about the 

species, other than indicating that it presumably was found in 

Great Britain, nor is the disposition of any material mentioned. 

The species is indeterminable; the name is here considered a 

nomen nudum, making the combination available. 

Eunice margaritacea (Verrill, 1900) 

Leodice margaritacea Verrill, 1900:644-645. 

Leodice longicirrata.—Treadwell, 1921:11-14, figs. 3-12, pi. 1: figs. 1-4 

[not Nicidion longicirrata Kinberg, 1865). 

Eunice longicirrata.—Hartman, 1942:9 (in part, not Nicidion longicirrata 

Kinberg. 1865]. 

REMARKS.—The types of this species are no longer available 

in the collections of the Peabody Museum (W. Hartman, in 

litt.). It is not clear whether Hartman (1942:9) saw any 

specimens of this species when she examined the type 

collections of that museum. She referred to TrcadweH's 

treatment of the species. Treadwell (1921:11-14) clearly did 

not see Verrill's types; on page 14 he stated: "From comparison 

of his descriptions with specimens of longicirrata collected by 

myself in Bermuda, in the locality where [Verrill] records 

margaritacea as abundant, I am convinced that [elegans and 

margaritacea] are identical with longicirrata." I am inclined to 

believe that the spccimcn(s) on which Vcrril! based his species 

were missing already by 1920. 

The only descriptive detail available, namely that branchiae 

arc present from setigers 3-4 and end at sctigcrs 45-50 with up 

to 5 branchial filaments, is consistent with TrcadweH's 

conclusion and with Webster's species, also originally described 

from Bermuda. Webster's name is preoccupied and his 

species was renamed Eunice websteri by Fauchald (1969). The 

synonymy proposed by Treadwell is here accepted. VerriH's 

name is valid because the previously published combination 

Eunice margaritacea Williams (1853) is a nomen nudum (see 

above). 

116. Eunice margariticacea Fischli, 1900 

TABLES 46,47 

Eunice margariticacea Fischli. 1900:104-106. pi. 4: figs. 8, 9. pi. 6: figs. 35, 

36, pi. 7: figs. 55-58. 

COMMENTS ON MATERIAL EXAMINED.—The types, from 

Temate in the Moluccas, are not available. Fischli's description 

and illustrations are quite detailed; the following summary of 

his findings updates his terminology. 

DESCRIPTION.—Medium-size specimen with 95 segments; 

length 14 mm. Ceratostyles with up to 30 moniliform 

articulations in A-III. A-III to setiger 7. Peristomium cylindrical. 

Separation between peristomial rings distinct on all sides. 

Peristomial cirri to posterior edge of prostomium, with up to 7 


Maxillary formula 1+1, 5+5, 7+0, 5+7, and presumably 1+1. 


Branchiae present, pectinate, reduced in mid-body region, 

erect. Branchiae from setiger 6 to setiger 95. Branchiae present 


of setigers. Maximum 5 filaments at about setiger 10. 

Mid-body setigers in 1 specimen with single filaments; 

posterior setiger with 2-3 filaments. 

Ventral cirri basally strongly inflated in median setigers, less 

distinctly inflated in far posterior setigers. Inflated bases nearly 

spherical. Notopodial cirri with up to 5-7 articulations, most 

distinct anteriorly, but present in all setigers.


Limbate setae marginally smooth. Pectinate setae flaring, 

flat Both marginal teeth distinctly longer than other teeth; -15 

teeth present. Appendages of compound falcigers tapering, 

tridentate. Distal tooth well developed. Guards symmetrically 


compound spinigers absent. Aciculae, presumably yellow or 

light-colored, distally gently double-curved, tapering to blunt 

tips; cross-sections round. Subacicular hooks tridentate with 

teeth in a crest. Starting segment for subacicular hooks not 

mentioned. Fangs decreasing evenly in size from large main 

fang to small tertiary fangs. 







Characters: 3-20, 23, 25, 26, 28, 32, 39, 40, 43-51, 

55, 56, 58-63, 80-82. 


None. 

REMARKS.—Eunice margariticacea is incompletely known, 

and without access to material from the type locality, it cannot 

be adequately characterized; it is, however, sufficiently well 

characterized that it can be compared with other species in the 

group in Tables 46 and 47. 

117. Eunice marovoi Gibbs, 1971 

FIGURE 70a-f; TABLES 7,41, 45 

Eunice marovoi Gibbs, 1971:158-160, figs. lOa-h. 


Solomon Islands, sta ML 96, Marovo Lagoon, Vangunu Island, 

dredged, 9 m, sand, 28 Oct 1965, coll. Royal Society 

Expedition to Solomon Islands. Nine paratypes, BM(NH) ZB 

1970.46, Solomon Islands, sta ML 110, Koto Lagoon, 

Vangunu Island, 26 m, mud, 28 Oct 1965, coll. Royal Society 

Expedition to Solomon Islands. One paratype, BM(NH) ZB 

1970.49, Solomon Islands, sta ML 218, Tokavai Lagoon, New 

Georgia, dredged, 18 m, mud, 8 Nov 1965, coll. Royal Society 

Expedition to Solomon Islands. 


based on the holotype and a single paratype from ZB 1970.46. 

The variability is indicated in Table 7. 


length 14.5 mm; maximal width 1 mm wide; length through 

setiger 10, 2.5 mm. 



lobes frontally rounded, dorsally slightly inflated; median 

sulcus represented by very shallow frontal notch; ventral side 

deeply furrowed. Eyes between bases of A-I and A-II, black; 

posterior ! A of eyes overlapped by peristomial fold. Antennae 

in a horseshoe, evenly spaced, similar in thickness. Ceratophores 

ring-shaped in all antennae, without articulations. 

Ceratostyles slender and tapering, without articulations. A-I to 

setiger 1; A-II to setiger 2; A-I 11 to setiger 3. Peristomium 

cylindrical. Separation between peristomial rings distinct on all 

sides of body; anterior ring about twice as long as posterior 

ring. Peristomial cirri about as long as A-I, to middle of 

peristomium, digitiform, without articulations. 

Maxillary formula 1+1, 4-6+10-13, 10-13+0, 10-13+10- 

13, according to Gibbs. Left Mx II with large fang; such fang 

absent on right side. 

Branchiae (Figure 70f) present, pectinate, distinctly longer 




number of setigers. First branchia with 2 or more usually 3 

filaments, maximum 6 filaments. Only last branchia single 

filaments. 

Neuropodial acicular lobes distally obliquely rounded with 

high side superiorly; aciculae emerging above midline. All 


cirri bluntly tapering. Bases of ventral cirri inflated from 

beginning of branchial region. Inflated bases about V2 of cirral 

length, ovate; narrow tips digitiform. Basal inflation less 

distinct in postbranchial region, but even in last setigers 

present, glandular bases of ventral cirri slightly thicker than the 

remainder of the cirri, distinctly set off from tips. Prebranchial 

notopodial cirri increasing in length, tapering. Notopodial cirri 

of branchial region long, tapering, with embedded aciculae 

basally. Postbranchial notopodial cirri decreasing in length, 

becoming slender and digitiform in posterior setigers. Notopodial 

cirri located on dorsal body wall well above bases of 

acicular lobes in posterior setigers. Notopodial cirri without 



not seen. Compound falcigers (Figure 70d,e) slender. Shafts 

distinctly inflated, marginally serrated. Appendages narrow, 

tapering to tips, bidentate. Proximal teeth triangular, directed 

laterally. Distal teeth very nearly erect. Guards tapering to 

distinct, slender mucros. Guards of posterior hooks (Figure 

70d) distally rounded with very small, narrow mucros. 


Aciculae yellow; anterior aciculae tapering to sharp tips; 

postbranchial aciculae distally blunt and slightly bent dorsally; 



hooks (Figure 70c) yellow, tridentate with teeth in a crest. 


thereafter; 3 hooks present in most setigers. Hooks strongly 

curved with large heads. Main fangs large; secondary fangs 

much smaller; tertiary fangs very small and appended to back 

of secondary fangs. 


anal cirri. 


TURES.—None.

212 


FIGURE 70.—Eunice marovoi (holotypc, BM(NH) ZB 1970.42): a, anterior end, lateral view; b, posterior 

parapodium, anterior view; c, subacicular hook, posterior parapodium; d, appendage of compound falciger, 

posterior parapodium; e, compound falciger, parapodium 9; f, parapodium 9, anterior view. Eunice martensi 

(holotype, ZMB 923): g, aciculae, parapodium 45; h, pectinate seta, parapodium 45; i, compound falciger, 

parapodium 45; j, subacicular hook, parapodium 45. (Scale bars in mm.)


TABLE 7.—Variable and invariable features in the type lot of Eunice marovoi (N 


number of individuals 



Total length 

Maximal width 


Last branchiae on setiger no. 





N 

5 

5 

5 

5 

5 

5 

3 

4 

5 

Max. 

61 

13 

1 

3 

24 

5 

25 

3 

19 

Min. 

46 

7 

0.5 

1.5 

19 

3 

20 

2 

10 

Mean 

56.2 

10.7 

0.9 

2.29 

22 

4.2 

23.33 

225 

15.6 

SD 

5.38 

2.18 

0.2 

0.54 

1.90 

0.75 

2.36 

0.43 

3.38 


Antennal articulations 






Color of aciculae 

Acicular tip 



Core-sheath separation 

N=5 

absent 

absent 

3 

tapering 

10 

inflated 

yellow 

tapering; straight 

yellow 

3 

indistinct 



Characters: 4, 6,42, 65-68. 


None. 

REMARKS.—Eunice marovoi was described with bluntly 

rounded guards to compound hooks and with branchiae 

extended to setigers 23-25; as indicated above, guards are 

mucronate and branchiae are not found on any segment later 

than 22 in the specimens examined. The dentition of the jaws is 

remarkable and may represent a juvenile dentition. The shape 

of the subacicular hooks also resembles the condition found in 

juvenile specimens. None of the specimens are sexually 

mature. 

Eunice marovoi is listed with similar species in Tables 41 

and 45. It is the only species known to combined nonarticulated 

ceratostyles with multiple subacicular hooks. 

118. Eunice martensi Grube, 1878 

Eunice martensi Grube, 1878a:102. 

FIOURE 70g-j; TABLES 46,47 

MATERIAL EXAMINED.—Holotype, ZMB 923, Zamboanga, 

Philippines, coll. v. Martens. 

COMMENTS ON MATERIAL EXAMINED.—The specimen is 

very poorly preserved and no meaningful illustrations of soft 

parts are possible. 


length 220 mm; maximal width 6 mm; length through setiger 

10, 14 mm. Body apparently cylindrical, slowly tapering 

posteriorly. 

Prostomium distinctly shorter than peristomium. Median 

sulcus very shallow. Eyes between bases of A-I and A-II. 


Ceratostyles digitiform, with up to 10 cylindrical articulations 

in A-II. Only right A-I and A-II complete. A-I to anterior 

peristomial ring; A-II to setiger 1. Peristomium cylindrical. 



posterior end of prostomium, slender and digitiform, with 5 



long, located behind left Mx II. Mx VI absent. 



from setiger 6 to setiger 163. Branchiae present to near 

posterior end, present on more than 65% of total number of 

setigers. All branchiae pectinate with up to 16 filaments at 

setigers 15-20. Branchial stems very thick, tapering, erect. 

Filaments short, digitiform. Number of filaments decreasing 

posteriorly; 6 filaments in setiger 150. No increase in number 

of filaments, length of filaments, or of branchial stem towards 


All neuropodial acicular lobes apparently truncate or 

rounded; aciculae emerging at midline. All pre- and postsetal 

lobes low, transverse folds. Anterior ventral cirri tapering, 

becoming moderately basally inflated in some median setigers. 

Inflated bases ovate; narrow tips tapering. Posterior ventral cirri 

without basal inflation, tapering. All notopodial cirri basally

214 


inflated, tapering, with at least one articulation. 

Limbate setae short, slender, marginally smooth. Pectinate 

setae (Figure 70h) small, slightly flaring, flat. One marginal 

tooth longer than other teeth; -10 teeth present. Shafts of 

compound falcigers (Figure 70i) tapering, marginally smooth. 

Appendages short, very chunky, tridentate. Proximal teeth 

smaller than distal teeth, triangular. Distal teeth distally split, 

forming tridentate appendages. Guards distally rounded; 


spinigers absent. Aciculae (Figure 70g) yellow, tapering, 

sharply bent; cross-sections round. Separation between core 


Subacicular hooks (Figure 70j) yellow, tridentate with teeth in 



Main fangs large, laterally directed. Two distal fangs forming 

small crests, erect. 


anal cirri. 





Characters: 4, 6-8, 11, 12, 15-17, 23, 24, 51. 


None. 

REMARKS.—Eunice martensi is listed with similar species in 

Tables 46 and 47. Four species listed in Table 47 lack reduced 

branchiae in a median region; these species include E. elseyi, E. 

lucei, and E. rubra in addition to E. martensi. All the other 

three species have at least some hammer-headed or weakly 

bidentate aciculae; in E. martensi all aciculae are pointed. 

Eunice maxima Quatrefages, 1866 

Eunice maxima Quatrefages, 1866:330.—Ehlers, 1868:310-311, pi. 15: figs. 

30-34. 

Nereis gigantea Chiaje, 1825:389, 424, pi. 27: figs. 1-8. 

REMARKS.—Quatrefages introduced the name Eunice maxima 

as a replacement name for delle Chiaje's Nereis gigantea 

described from Gulf of Naples. 

Quatrefages had no specimens available and was clearly in 

considerable doubt that the species could be separated from 

other, similar forms, such as Cuvier's Eunice gigantea and 

Grube's E. magnifica. He commented that he renamed the form 

mainly due to the excellent reputation of delle Chiaje as a 

describer. 

Ehlers described an anterior fragment collected in Naples 

region, present in the collection in GOttingen under this name. 

This specimen is closely similar to Eunice aphroditois, but 

crucial pieces of information are missing and the identity 

cannot be confirmed, nor is the specimen currently available. 

Grube (1878:98) referred the form renamed by Quatrefages 

to Eunice roussaei, another very large species described by 

Quatrefages (1866). 

and 

No type material is available; the species has never been 

adequately characterized and is here considered indeterminable. 

119. Eunice medicina Moore, 1903 

FIGURE 71a-«; TABLES 41, 42 

Eunice medicina Moore, 1903:441-444, pi. 25: figs. 49-51. 

MATERIAL EXAMINED.—Syntype, USNM 15852, Albatross 

sta 3700, Japan, Suruga Wan, 2 miles NE of Seno Umi Bank, 

(approximately 34°45TSf, 138°3O"E), 7 May 1900, 115 m, 

volcanic mud and sand, Blake trawl, grapnels and tangles. 

DESCRIPTION.—Syntype complete with 79 setigers; total 

length 29 mm; maximal width 1.3 mm; length through scliger 

10, 5 mm. Body cylindrical, tapering abruptly anteriorly and 

slowly posteriorly. 

Prostomium (Figure 7Id) distinctly shorter and narrower 

than pcristomium, less than '/2 as deep as pcristomium. 


median sulcus shallow. Eyes posterior to bases of A-I, 

dark. Antennae in a horseshoe, evenly spaced. Ccratophores 

ring-shaped in all antennae, without articulations. Only left A-I 

remaining of ccralostylcs, slender and digitiform, with 3 long, 

cylindrical articulations, reaching sctigcr 1. Pcristomium 


anterior ring 2 /3 of total pcristomial length. Pcristomial cirri to 

middle of prostomium, slender and digitiform, with 3 articulations. 


Branchiae present, pectinate, not reduced in mid-body 

region, erect Branchiae from setiger 3 to setiger 26. Branchiae 


of total number of setigers. First 5 pairs single filaments (Figure 

71a); maximum 7 long, flattened filaments present. Branchial 

stems slender, erect. 


rounded in posterior setigers; aciculae emerging above midline. 

All presetal lobes low, transverse folds. Anterior postsetal lobe 

following outline of acicular lobe closely, becoming low 

transverse fold in postbranchial region. First 2 ventral cirri 

digitiform, becoming basally inflated in branchial region. 

Inflated bases barrel-shaped; narrow tips digitiform. Far 

posterior setigers with less prominent inflated bases; last few 

ventral cirri digitiform. All notopodial cirri slender and 

digitiform; anterior notopodial cirri with 3 articulations; 

posterior notopodial cirri slightly shorter than anterior ones, 

slightly more distinctly inflated basally, without articulations. 

Limbate setae slender. Pectinate setae short, flaring, flat. 

Marginal teeth similar to other teeth; -12 teeth present. Shafts 

of compound falcigers (Figure 71b) inflated, marginally 

serrated. Appendages long, narrow, bidentate. Proximal teeth 

shorter than distal teeth, tapering, directed basally. Distal teeth 

sharply bent. Guards symmetrically sharply pointed, mucro-

NUMBER 523 

215 

FIGURE 71.—Eunice medicina (syntype, USNM 15852): a, parapodium 3, anterior view; b, compound falciger, 

parapodium 3; c, subacicular hook, parapodium 39; d, anterior end, lateral view; e, aciculae, parapodium 39. 

Eunice megabranchia (holotype, AHF Poly 1056): f, anterior end, lateral view; g, compound falciger, parapodium 

18; h, pectinate seta, parapodium 18; i, parapodium 61, anterior view; j, compound falciger, parapodium 61; k, 

subacicular hook, parapodium 61; 1, pectinate seu, parapodium 61; m, parapodium 18, anterior view. (Scale bars 

in mm.)


nate. Pseudocompound falcigers and compound spinigers 

absent. Aciculae (Figure 71e) paired, yellow, tapering, slightly 

bent or straight; cross-sections round. Separation between core 


Subacicular hooks (Figure 71c) yellow, tridentate with teeth in 

a crest. Subacicular hooks first present from setiger 17, present 

in all thereafter, occurring singly (except for replacements). 

Hooks distally bent. Two distal fangs emerging from joint 

bases. 


anal cirri. 





Characters: 4, 6, 17, 23, 32,42. 


None. 

REMARKS.—Eunice medicina is listed with similar species 

in Tables 41 and 42. In addition to E. medicina, two species 

listed in Table 42 lack posterior simple branchiae; these are E. 

indica and E. multicylindri. Subacicular hooks are always 

single in E. medicina and E. multicylindri; in E. indica three or 

more subacicular hooks are present in each segment. The 

maximum number of branchial filaments is seven in E. 

medicina and four in E. multicylindri. Other differences can be 

seen by comparing illustrations and descriptions of the two 

species. 

120. Eunice megabranchia Fauchald, 1970 

FIGURE 71f-m; TABLES 19.21,24,26 

Eunice megabranchia Fauchald, 1970:33-36, pL 4: figs. a-e. 

MATERIAL EXAMINED.—Holotype, AHF Poly 1056, Gulf of 

California, Mexico, 27°03'N, 112°18"W, 894 m, coll. S. 

Calvert, sta L-184. 

DESCRIPTION.—Holotype incomplete mature female with 

large eggs in body cavity with 74 setigers; length 68 mm; 

maximal width 7 mm; length through setiger 10, 12 mm. 

Anterior end of body cylindrical, becoming dorsally and 

ventrally flattened towards posterior end of fragment; crosssection 

nearly quadrangular posteriorly. 

Prostomium (Figure 7If) distinctly shorter and narrower 

than peristomium, as deep as */2 of peristomium. Prostomial 

lobes frontally obliquely rounded; median sulcus shallow, 

separation continued as distinct ridge to base of A-III. Surface 

of prostomium rugose, palps distinctly marked frontolaterally 

by shallow grooves. Eyes posterior to bases of A-I, hidden 

under peristomial fold, purple. Antennae in a horseshoe, evenly 



tapering, without articulations. A-I to setiger 1; A-I I to setiger 

6; A-III to setiger 9. Peristomium cylindrical. Lower lip 

scalloped. Separation between rings distinct on all sides; 


slightly beyond tip of prostomium slender and tapering, 



Branchiae (Figure 71m) present, pectinate, distinctly longer 




number of setigers. Last 7 pairs single filaments; all other 

branchiae strongly pectinate with 47 or more filaments where 

best developed, at setigers 15-20. Branchial stems erect, 

strong, tapering to very narrow tips, and outreaching notopodial 

cirri in all but last 7 branchial segments. Filaments filiform, 

forming tangled masses on sides of specimen; some filaments 

longer than notopodial cirri, but most distinctly shorter than 


Anterior ncuropodial acicular lobes obliquely truncate with 

aciculae emerging dorsal to midline. Postcriormost ncuropodial 

acicular lobes present (Figure 71i) symmetrically rounded with 

aciculae emerging medially. Pre- and postselal lobes low, 

transverse folds. Pre- and postbranchial ventral cirri tapering. 

Ventral cirri modestly basally inflated in branchial region. 

Inflated bases ovate; narrow tips tapering. Anterior notopodial 

cirri, slightly inflated basally, becoming tapering with long, 

slender, filiform tips in branchial region; postbranchial notopodial 

cirri slender, tapering, very much shorter than in branchial 

region. All notopodial cirri with distinct cirrophores; anterior 

notopodial cirri with 3 to 4 irregular articulations; articulations 

lost in first few branchial setigers. 

Limbate setae narrow, marginally smooth. Shafts of pectinate 

setae (Figure 71 h,l) wide, flattened. Blades slightly flaring, 

flat or gently furled. One marginal tooth distinctly longer than 

other teeth; 16 teeth present. Shafts of anterior compound 

falcigers (Figure 71g) distally inflated, becoming tapering in 

posterior setigers (Figure 71j), marginally smooth; internal 

striation distinct, with distinct, narrow beaks. Appendages 

tapering from base to very small distal heads, bidentate. 

Proximal teeth smaller than distal teeth, forming low, triangular 

lateral projections. Distal teeth erect, slender in anterior setigers 

and thick in posterior setigers. Guards tapering to slender, 

distinct mucros, marginally serrated. Pseudocompound falcigers 

and compound spinigers absent. Aciculae single in anterior 

setigers, up to 3 in posterior setigers, honey-colored, tapering to 

blunt tips, straight; cross-sections round. Subacicular hooks 

(Figure 71k) honey-colored, bidentate. Hooks first present 


(except for replacements). Hooks tapering to small heads. 

Proximal teeth much larger than distal teeth; both teeth directed 

distally. 


features associated with posterior parapodia; pygidium and 

anal cirri.






Characters: 1,2,4, 6,47, 50, 63, 74, 78. 


None. 

REMARKS.—Eunice megabranchia is a member of group Al 

and was compared to other deep-water species by Fauchald 

(1970); it is listed with similar species in Tables 19 and 21; it 

is also listed in Tables 24 and 26, because the aciculae and 

subacicular hooks may be a rather dark yellow. A detailed 

discussion of the relation between this species, E. hawaiensis, 

and E. validobranchiata is given as part of the description of 

the former. 

Eunice megabranchia differs from all species in Table 26 by 

having more than five times as many branchial filaments (47 

vs. 9) as the species with the next high number listed in that 

table. 

Eunice megalodus Grubc, 1878 

Eunice megalodus Grubc, 1878b:lS6-lS8.pl 9: fig. S. 

REMARKS.—The unusual jaw apparatus and the structure of 

the anterior end as described by Grube, the setal complement, 

and the structure of the notopodial cirri are all features found in 

the genus Euniphysa. 

121. Eunice mexicana Fauchald, 1970 

FIGURE 72; TABLES 8,19,20 

Eunice biannulata mexicana Fauchald, 1970:27-28, pi. 1: figs, f, g. 

MATERIAL EXAMINED.—Lectotype (AHF Poly 1539) and 10 

paralectotypes (AHF Poly 1540), Mexico, Isla Isabel, 

21°54 / 1(TN, 1O5°53'O5"W, 18-33 m, corallines, dredge, 2 Apr 

1937, Velero sta 747-37. 

DESCRIPTION.—Lectotype complete with 106 setigers; total 

length 87 mm; maximal width 3.5 mm wide at about setiger 15; 

length through setiger 10, 10.5 mm; width at setiger 10, 3.2 

mm. Body cylindrical anteriorly, slightly dorsoventrally flattened 

posteriorly. Anal cirri short, tapering, possibly articulated. 



lobes frontally obliquely truncate, dorsally slightly inflated, 

sloping obliquely laterally; median sulcus very shallow. Eyes 

posterior to bases of A-I, black. Palpal region marked by 

transverse folds on ventral side. Antennae in very open 



slender and slightly tapering; maximum 17 articulations in 

A-III; first several articulations cylindrical, becoming increasingly 

drop-shaped distally; in some paralectotypes distal 

articulations moniliform. A-I to posterior peristomial ring; A-I I 

to setiger 5; A-III to setiger 8. Peristomium with distinctly 

muscular lower lip. Separation between rings distinct dorsally, 

visible ventrally; anterior ring 3 A of total peristomial length. 

Peristomial cirri to middle of prostomium, slender and 

tapering, with 4 to 5 cylindrical articulations. 

Maxillary formula of 5 paralectotypes 1+1, 6-7+7-8, 

7-9+0, 6-8+8-11, and 1+1. Mx III relatively short, slightly 

curved, located behind left Mx II. Mx VI missing. 

Branchiae (Figure 72d,e) present, pectinate, distinctly longer 

than notopodial cirri, not reduced in mid-body region, erect 



number of setigers. Maximum 18 filaments at about setiger 15; 

in lectotype all branchiae with at least 2 filaments; in 

paralectotypes up to 3 posterior segments with single branchial 

filaments. First 2 and last 3 branchiae shorter than notopodial 

cirri; all other branchiae longer. Branchial stems slender, 

tapering. Filaments slender, digitiform, shorter than notopodial 

cirri in all setigers. 

Anterior and median neuropodial acicular lobes asymmetrically 

rounded with aciculae emerging dorsal to midline; 

posterior acicular lobes increasingly lower, merging into body 

wall near posterior end. Pre- and postsetal lobes low, transverse 

folds in all setigers. Anterior ventral cirri thick and tapering, 

becoming distinctly inflated by setiger 10. Inflated bases ovate, 

modest in size; narrow tips digitiform. Inflated bases reduced 

from about setiger 25-30. Ventral cirri thick and digitiform 

from about setiger 45, retaining that shape in remainder of 

body. Anterior and median notopodial cirri thick and tapering, 

with up to 5 cylindrical articulations. Posterior notopodial cirri 

nearly as long as those in median setigers, digitiform, without 


Limbate setae thick, marginally finely serrated, longer than 

compound hooks in all setigers. All pectinate setae (Figure 

72c, f,i) furled, tapering. Both marginal teeth longer than other 

teeth; number of teeth increasing from 8 to -12 posteriorly. 

Shafts of all compound falcigers (Figure 72b,g,j) tapering, 

internally distinctly striated, with distinct beak. All appendages 

relatively wide basally, tapering towards small heads, bidentate. 

Proximal teeth very short; indistinct low knobs in anterior 

and median setigers, distinctly triangular in posterior setigers. 

Distal teeth nearly erect in all setigers, increasing in size from 

anterior to posterior setigers. Anterior and median guards 

asymmetrically bluntly pointed, becoming increasingly symmetrical 

and rounded posteriorly, marginally finely serrated; 

mucros absent. Pscudocompound falcigers and compound 

spinigers absent. Aciculae at least paired, with clear yellow 

cores. Anterior aciculae bluntly pointed, distally straight, 

medially at least superiormost aciculae distally expanded into 

tabs (Figure 72h); posterior aciculae (Figure 72k) tapering, 

distinctly bent dorsally; cross-sections round. Separation 


subacicular hooks. Subacicular hooks (Figure 72k) yellow,

218 


FIGURE 72.—Eunice mexicana (lectotype, AHF Poly 1539): a, anterior end, lateral view; b, compound falciger, 

parapodium 3; c, pectinate seta, parapodium 3; d, parapodium 3, anterior view; e, parapodium 15, anterior view; 

f, pectinate seta, parapodium 15; g, compound falciger, parapodium 15; h, aciculae, parapodium 15; i, pectinate 

seta, parapodium 48; j, compound falciger, parapodium 48; k, aciculae and subacicular hooks, parapodium 48. 

(Scale bars in mm.)


TABLE 8.—Variable and invariable features in the type lot of Eunice mexicana (N = number of individuals 



Maximal width 



Branchiae present to sctiger no. 


Ventral cirri inflated through sctiger no. 

Subacicular hooks first present from 

N 

Max. 

11 4 

11 10.5 

3 21 

11 40 

11 18 

10 30 

11 32 

Min. 

2 

6 

16 

26 

15 

20 

21 

Mean 

3.2 

8.8 

18.67 

34 

16.91 

26.4 

26.45 

SD 

0.64 

1.63 

2.52 

3.61 

1.38 

2.95 

3.11 


Separation of rings 



Branchiae first present from sctiger no. 





Guards of compound falcigers 


Acicular shape 




N=ll 



4 

3 

5 

tapering; furled 

10 

tapering 

bluntly pointed to rounded 

yellow 

flattened tab 

yellow 

2 

visible 


setigers thereafter, usually 3 in a vertical row in posterior 

setigers. Hooks gently curved; necks and heads distinct. 


teeth nearly erect, slender. 


anal cirri. 







None. 

REMARKS.—Through a lapsus, type locality and type 

specimens were not designated when the species was first 

described; I take the opportunity to rectify the error at this time 

by designating lecto- and paralectotypes for the species. As 

indicated above, the species is separated from other species in 

the group by the same kinds of characters used to separate other 

species, thus subspecific status originally designated is unjustified. 

Eunice mexicana is listed with similar species in Tables 19 

and 20. It is very similar to E. pennata in distributional features, 

but can be separated from it by having 18 rather than 12 

branchial filaments and usually three subacicular hooks in a 

vertical row rather than having maximally paired subacicular 

hooks as in E. pennata. 

Eunice elegans (see Table 24) also resembles E. mexicana 

and 

closely, but can be separated as suggested in the discussion of 

the former. 

122. Eunice microprion Marenzeller, 1879 

FIGURE 73a-k; TABLES 27,29 

Eunice microprion Marenzeller, 1879:135-136, pi 5: fig. 1, 1A-1D.—Imajima 

and Hartman, 1964:252. 

MATERIAL EXAMINED.—ZMW 1062, Jaqashima, Japan, 

don. DOderlein, 1 specimen; ZMW 1064, Eno Shima, Japan, 

don. Doderlein, 1 specimen. 

COMMENTS ON MATERIAL EXAMINED.—The specimens 

listed above are not the types, but specimens identified by v. 

Marenzeller from collections taken at or near the type area. 

DESCRIPTION.—Specimen from ZMW 1062 complete with 


through setiger 10, 10 mm. Other specimen incomplete with 

124 setigers; length 85 mm; maximal length 5 mm; length 

through setiger 10,12 mm. Both specimens cylindrical, gently 

tapering. Anal cirri as long as last 10 setigers together. 



Prostomial lobes frontally obliquely truncate, dorsally excavate 

with a thickened rim; median sulcus deep. Eyes near bases of 

A-II, dark. Antennae in a horseshoe; with A-III isolated by a 


antennae, without articulations. Ceratostyles tapering very 

slowly towards tips, with up to 10 poorly marked, cylindrical

220 


FIGURE 73.—Eunice microprion (ZMW 1062): a, anterior end, lateral view; b, pectinate seta, parapodium 9; c, 

compound falciger, parapodium 9; d, parapodium 9, anterior view; e, aciculae, parapodium 46; f, compound 

falciger, parapodium 46; g, pectinate seta, parapodium 46; h, subacicular hooks, parapodium 47; i, parapodium 

46, anterior view; j, parapodium 105, anterior view; k, pectinate seta, parapodium 105. Eunice mindanavensis 

(holotype, BM(NH) ZK.1885.12.1.51a): 1, anterior end, lateral view; m, parapodium 30, anterior view; n, 

pectinate seta, parapodium 30; o, subacicular hook, parapodium 30; p, compound falciger, parapodium 30. (Scale 

ban in mm.l


articulations in A-111. A-I to posterior peristomial ring; A-II and 

A-III to setiger 3; A-I 11 slightly longer than A-II. Peristomium 

cylindrical. Separation between rings distinct nearly all way 

around body, indistinct only for short stretch laterally; anterior 

ring 5 /6 of total peristomial length. Peristomial cirri to middle of 

prostomium, slender and tapering, with 5 indistinct articulations. 

Maxillary formula of specimen from ZMW 1062 1+1, 6+6, 

10+0, 4+8, 1+1, and 1+1. Mx III with very short, small teeth, 

forming an arc with left Mx IV. Mx VI small plate with small 

point on either side. 

Branchiae (Figure 73d,i,j) present, pectinate, distinctly 

shorter than notopodial cirri, reduced in mid-body region, 

flexible. Branchiae from setiger 7-8 to setiger 127. Branchiae 


number of setigers. All anterior and posterior branchiae with 2 

or more filaments, in median region, from about setiger 35 to 

65, branchiae reduced to single, relatively long filaments 

(Figure 73i). Maximum -8 filaments at about setiger 15. 

Filaments short, digitiform. 

Anterior neuropodial acicular lobes rounded; posterior 

acicular lobes triangular or conical; aciculae emerging at 

midline. All prc- and postsetal lobes low, transverse folds. 

Anterior ventral cirri thick, tapering, becoming basally inflated 

from about setiger 10. Inflated bases distinct transverse welts; 

narrow tips tapering. Inflated bases reduced from about setiger 

55-60, completely missing from about setiger 60-65. Posterior 

ventral cirri tapering, broadly attached ventrally, nearly 

scoop-shaped. All notopodial cirri digitiform, with 2 or more 

articulations. Anterior notopodial cirri about as long as 

peristomial cirri, becoming shorter from about setiger 3, but 

very much longer than branchiae in all setigers, retaining about 

same length from setiger 10 through rest of body, becoming 

slowly slimmer. Notopodial cirri articulated throughout body. 

Limbate setae marginally serrated. All pectinate setae 

(Figure 73d,g,k) tapering, flat. Both marginal teeth slightly 

longer than other teeth; number of teeth increasing from -15 to 

-20 posteriorly. Shafts of pectinate setae increasing in length 

posteriorly. Pectinate setae numbering -10 in all parapodia. 

Shafts of compound falcigers (Figure 73c, 0 tapering, with 

distinct distal beak, marginally smooth, with distinct internal 

striations. Appendages elongated with parallel sides anteriorly 

(Figure 73c), becoming shorter and distinctly tapering posteriorly 

(Figure 730, bidentate. Proximal and distal teeth similar in 

size. Proximal teeth triangular, directed laterally. Distal teeth 

gently curved. Anterior guards symmetrically rounded, becoming 

symmetrically bluntly pointed in posterior setigers; mucros 


absent. Aciculae (Figure 73e) at least paired, up to 4 present, 

with dark brown, nearly black cores and clear sheaths, tapering 

with slender tips, bent or curved dorsally; cross-sections round. 

Subacicular hooks (Figure 73h) with dark brown, nearly black 


setiger 38 or 41, present in all setigers thereafter, 2 in most 

setigers; 3 or more in some setigers. Proximal teeth larger than 

distal teeth, triangular, directed laterally. Distal teeth nearly 

erect. 








None. 

REMARKS.—The species belongs to group B-2 and is 

compared to similar species in Tables 27 and 29. A 

characteristic feature is the reduced number of branchial 

filaments in median region of body; this pattern is present in E. 

antennata and a few similar species in group C-2, but is 

otherwise not known to be present in any other species in the 

genus. 

123. Eunice mindanavensis Mclntosh, 1885 

FIGURE 731-p; TABLES 27,30 

Eunice mindanavensis Mclntosh, 1885:289-291, figs. 50,51, pi. 39: figs. 9,10, 

pi. 20A: figs. 21, 22. 


1885.12.1.51a, off Mindanao, Philippines, 7°3'N, \2\°AVE, 

148 and 186 m, stones and gravel, 26 Oct 1874, Challenger sta 

201. 


now in three pieces; an anterior end consisting of 21 setigers 

(cut into two pieces) and a posterior end of 42 setigers. The two 

pieces clearly belong to the same specimen and appear to 

represent a complete specimen. Mclntosh reported only the first 

of the two depths listed above; Alex Muir (in litL) informed me 

that station 201 included hauls at two different depths. 


length -40 mm; maximal width 4 mm; length through setiger 

10, 7 mm. Body thickset anteriorly, circular in cross-section, 

tapering strongly, with relatively few, very distinct segments. 


peristomium, less than l /i as deep as peristomium. Prostomial 

lobes frontally rounded; dorsally flattened lobes; median sulcus 

deep. Eyes not seen. Antennae covered by large overhanging 

peristomial fold, similar in thickness. Ceratophores ring-shaped 

in all antennae, without articulations. Ceratostyles tapering and 

slender, with up to 15 relatively long, rather indistinct, 

cylindrical articulations. A-I to setiger 1, A-II to setiger 3; A-III 

to setiger 7. Peristomium roughly cylindrical. Separation 

between rings distinct dorsally and possibly ventrally; separation 

on ventral side confounded by presence of superficial 

folds; anterior ring ~ 6 fr of total peristomial length. Peristomial 

cirri to tip of prostomium, tapering, with 7 long, cylindrical 

articulations.


Maxillary formula 1+1, 7+7, 10+0, 10+10, 1+1, and 1+1. 

Teeth of Mx III and IV very small, but distinct. Mx III short; 

part of distal arc with left Mx IV. 

Branchiae (Figure 73m) present, pectinate, distinctly shorter 




of setigers. Last 5 pairs single filaments. First few pairs of 

filaments very unequal in length. Maximum 3 filaments from 

about setiger 15 through setiger 30. Filaments very thick, 

digitiform. 


distally rounded posteriorly; aciculac emerging above 

midline. All pre-setal lobes low, transverse folds. Anterior 

postsetal lobes following outline of acicular lobes closely, 

becoming low, transverse folds from about setiger 15. 

Prebranchial ventral cirri thick, tapering. Ventral cirri large, 

flattened, forming scoop around lower edge of acicular lobe 

from early branchial setigers, eventually forming scoops 

around emergent part of subacicular hooks; tips tapering. All 

notopodia medially inflated; anterior notopodial cirri with 5 or 

6 cylindrical articulations; articulations indistinct from mid 

body. 

Limbate setae narrow. Pectinate setae (Figure 73n) flaring, 

flat. Both marginal teeth slightly longer than other teeth; -15 

teeth present. Shafts of compound falcigers (Figure 73p) 

slightly inflated, internally strongly striated, marginally 

smooth. Appendages short, slender, bidentate with thick teeth. 

Proximal teeth triangular, directed laterally. Distal teeth erect. 

Guards asymmetrically bluntly pointed; mucros absent Pseudocompound 


single anteriorly; 2-3 present in median and posterior 

setigers, dark brown, tapering to blunt tips, straight; crosssections 

round. Subacicular hooks (Figure 73o) with dark 

brown cores and clear sheaths, bidentate. Hooks distinct from 

setiger 21; setigers 17-20 with broken remnants of much more 

delicate hooks in subacicular positions, present in all setigers 


distinctly tapering; heads distinct. Proximal teeth directed 

laterally. Distal teeth triangular, rather blunt 


anal cirri. 





Characters: 4,6, 13-16, 74. 


None. 

REMARKS.—Eunice mindanavensis is listed with similar 

species in Tables 27 and 30. It has scoop-shaped ventral cirri 

with inflated bases continued to the far posterior end; the only 

other species in Table 30 with inflated bases through the 

posterior end is E. coccinea, in which the bases are ovate. The 

and 

branchiae are continued to the far posterior setigers in £. 

mindanavensis and terminate well before the posterior end in E. 

coccinea. 

Eunice minuta Grube. 1850:292. 

Eunice vittata.—Grube, 1866b:68. 

Eunice minuta Grube, 1850 

REMARKS.—Originally a nomen nudum, Grube did not issue 

any description of this form before withdrawing it in 1867 as a 

juvenile of E. vittata. Eunice vittata he considered, incorrectly, 

as being indistinguishable from alcohol-preserved specimens 

of E. pennata. Eunice minuta is here considered a junior 

synonym of E. vittata. 

124. Eunice modesta Grub«, 1866 

Eunice modesta Grube, 1866a:64. 

FIOURF. 74; I AMI J s 33. 36 

MATERIAL EXAMINED.—TWO syntypes, ZMH V-801, 

Samoa. ZMB 3998, Samoa, coll. Godcffroy. 

COMMENTS ON MATERIAL EXAMINED.—The Hamburg syntype 

has been completely dried out at one lime and no 

meaningful illustrations of soft parts could be made, so the 

illustration of the anterior end is based on the very soft Berlin 

specimen, but otherwise the description is based on the 

Hamburg syntype with comments as necessary where the 

Berlin syntype differs. 

DESCRIPTION.—Hamburg syntype complete female with 

very large eggs in body cavity, with -80 setigers; total length 


Berlin syntype incomplete with 37 setigers; length 21 mm; 

maximal width 2 mm; length through setiger 10, 5.5 mm. 



Prostomium frontally rounded, dorsally excavate with a 

thickened rim; median sulcus very deep. Eyes between bases of 

A-I and A-II, reddish. Ceratophores long in all antennae, 

confluent with prostomium, rather than ring-shaped, without 

articulations. Ceratostyles digitiform, with up to 6 long, 

cylindrical articulations in A-I 11. All antennae short; none 

reaching beyond the peristomium. Peristomium with large, 

inflated lower lip, set off from rest of peristomium as thickened 

cushion delimited posteriorly by ventral groove separating 

anterior and posterior peristomial rings. Peristomial rings 

distinct dorsally and ventrally; anterior ring 3 A of total 

peristomial length. Peristomial cirri barely to posterior edge of 

anterior peristomial ring, ovate, without articulations. 


Branchiae present, single filaments, distinctly longer than 

notopodial cirri. Branchiae from setiger 17-18 to setiger 70, 

from setiger 11 to end of fragment in Berlin syntype. Branchiae 

terminating before posterior end, present on 65% of total


J32 

J 

b, c, d 

a 

FIGURE 74.—Eunice modesta (syntype, ZMB 3998 (a only), syntype, ZMH V-801 (b-d)): a, anterior end, lateral 

view; b, compound falciger, parapodium 29; c, subacicular hook, parapodium 29; d, pectinate seta, parapodium 


number of setigers. All branchiae relatively long, slender 

filaments. 

Shape of neuropodial acicular lobes cannot be determined 

accurately. Ventral cirri basally inflated from about setiger 5 

through setiger 45, tapering in posterior setigers. Inflated bases 

ovate. All notopodial cirri basally inflated, without articulations, 

very much shorter than branchial filaments in all but first 

few branchial setigers. 


(Figure 74d) flaring, flat. One marginal tooth longer than other 


74b) slightly inflated, marginally smooth. Appendages tapering, 


directed laterally. Distal teeth tapering, distinctly curved. 

Guards symmetrically rounded; mucros absent. Pseudocompound 


light brown, tapering to blunt tips, straight; cross-sections 

round. Subacicular hooks (Figure 74c) light brown, thick, 

tapering, bidentate. Hooks first present from setiger 22, present 


Both teeth directed distally. Proximal teeth about twice 

as large as distal teeth. 


parapodial structures; pygidium and anal cirri. 




SCORED.—Inappropriate Characters: 33, 34, 39, 40, 56, 58, 

59. Unknown Characters: 4,6,43-50, 52, 55, 74,78, 80. 


None. 

REMARKS.—Eunice modesta has not been reported since it


was first described. It is listed with similar species in Tables 33 

and 36. Eunice pruvoti and E. modesta both have single 

filaments in all branchiae. Branchiae are continued to the 

posterior end in the former and are missing on the last 10 

setigers in the latter. Eunice modesta is further characterized by 

the relatively light color of subacicular hooks and aciculae. 

Eunice monilifer (Chamberlin, 1919) 

Leodice monilifer Chamberlin, 1919b:ll-12. 

Eunice enteles.—Hartman, 1938:97 [not Leodice enieles Chamberlin, 1918]. 

Eunice antennata.—Ruiman, 1944:115-117, pi. 7: figs. 154-156 [in part, not 

Leodice antennota Lamarck, 1818]. 

REMARKS.—This species was originally described nestling 

in kelp hold-fasts near Laguna Beach, California. The type was 

deposited at the Museum of Comparative Zoology according to 

Chamberlin, but is now missing (A. Johnston, pers. comm.). 

The description given here summarizes information given by 

Chamberlin, with updated terminology. 

Antennae short, with up to 12 moniliform articulations. 

Penstomial cirri much shorter than pcristomium and wrinkled 

or distinctly articulated. Maxillary formula 1+1,4+6,8+0,9+9, 

and presumably 1+1. Branchiae from setiger 7 to setiger 30 (of 

109 setigers in complete type specimen), with maximum 3 

filaments. 

No mention was made of any parapodial or setal features, 

except that notopodial cirri were characterized as being 

wrinkled, but not distinctly divided into articulations. 

Hartman (1938:97) stated that Chamberlin's information 

about branchial distribution was unreliable. She illustrated 

tridentate hooks in a specimen of Eunice from Laguna Beach, 

but did not specify branchial distribution in the specimen she 

illustrated. In other members of groups C-l and C-2, 

differences in branchial distribution such as the one separating 

E. enteles from E. monilifer in similar-sized specimens is 

considered highly significant The two forms cannot be 

synonymized with each other, however, neither one is 

sufficiently well characterized to allow positive identification. 

Eunice monilifer is here considered indeterminable. 

Eunice mossambica Peters, 1854 

Eunice mossambica Peters, 1854:612. 

Marpkysa mossambica,—-Gravier, 1900:267-270, figs. 137-139, pi. 14: figs. 

89,90.—Crossland, 1903:139-140, pi. 15: figs. 7-10.—Day, 1967:395, fig. 

17.5i-m. 

Nauphanta mossambica.—Fauchald, 1987:376-378, fig. la-e. 

REMARKS.—Originally described in Eunice and usually 

considered a member of Marphysa, this species was moved to 

the genus Nauphanta by Fauchald (1987). 

125. Eunice mucronata Moore, 1903 

FIGURE 75a-e; TABLES 9,41,45 

Eunice mucronata Moore, 1903:437-440, pL 25: figs. 42-45. 

MATERIAL EXAMINED.—Lectotype, USNM 15851, and 11 

paralectotypes USNM 5341, Albatross sta 3698, Japan, Sagami 

Bay, 4.5 miles SE of Manazuru Saki (~35°O8'N, 139° 10Ti), 278 

m, green mud, volcanic ash and sand, beamtrawl. 

COMMENTS ON MATERIAL EXAMINED.—The lectotype was 

isolated as USNM 15851; this specimen and one complete 

specimen with 111 setigers from USNM 5341 had been 

dissected by Moore. The.description is based on the specimen 

from 15851. 

DESCRIPTION.—Lectotype complete, with 116 setigers; total 

length 58 mm; maximal width 4 mm at about setiger 25; length 

through setiger 10, 8 mm. Body cylindrical, tapering relatively 

slowly towards both ends. 


than pcristomium, less than '/2 as deep as pcristomium. 


sulcus shallow. Palpal region distinguished by transverse 

frontal furrows. Eyes posterior to bases of A-1, very large, 

brown, not clearly visible in all specimens. Antennae in a 

horseshoe, with A-I isolated by a gap, similar in thickness. 


Ceratostyles slender and tapering to very narrow tips without 

articulations. A-I apparently articulated in some specimens due 

to irregular muscle contractions rather than structural features. 

A-I to setiger 2; A-I I to setiger 10; A-III to setiger 13. 


all sides; anterior ring ~ 2 h of total penstomial length. 

Peristomial cirri attached at extreme frontal edge of posterior 

penstomial ring, reaching beyond tip of prostomium, same 

length as A-I, slender and tapering, without articulations. 

Summary maxillary formula for 2 syntypes 1+1, 7-8+7-8, 

7-8+0, 6+8, and 1+1. Syntype from USNM 5341 with 

consistently lower numbers of teeth where numbers differ. 



Branchiae from setiger 3 to setiger 34; in other syntypes last 

branchiae on setigers 29-33. Branchiae terminating well 

before posterior end, present on less than 55% of total number 

of setigers. First 2 pairs single filaments; maximum 15 slender 

filaments. Branchial stems slender, erect. 

Anterior neuropodial acicular lobes distally truncate or 

rounded, becoming distinctly triangular posteriorly; aciculae 

emerging above midline. Pre- and postsetal lobes low, 


cirri basally inflated in branchial region. Inflated bases ovate; 

narrow tips tapering. Postbranchial ventral cirri increasingly 

slender, tapering, resembling notopodial cirri closely, but 

consistently shorter. All notopodial cirri slender, tapering, with 

gently inflated bases, without articulations. Notopodial cirri 

long in prebranchial region, becoming shorter through branchial 

region, so in last of branchial setigers notopodial cirri no 

longer than branchial filaments. Postbranchial notopodial cirri 

about half as long as those in prebranchial region. 

Limbate setae slender, marginally smooth. Pectinate setae


1 

10 

b, d, e, g. h, i 

t 

FIGURE 75.—Eunice mucronala (lectotype, USNM 15851): a, anterior end, lateral view; b, subacicular hook, 

parapodium 29; c, parapodium 29, anterior view; d, pectinate seta, parapodium 29; e, compound falciger, 

parapodium 29. Eunice multicylindri (holotype, USNM 63066): f, anterior end, lateral view; g, subacicular hook, 

parapodium 40; h, pectinate seta, parapodium 40; i, compound falciger. parapodium 40. (Scale bars in mm.)


TABLE 9.—Variable and invariable features in the type lot of Eunice mucronata (N = number of individuals 




Total length 

Maximum width 

Max. width at setiger no. 


Width at 10 

Length of A-I 

Length of A-II 

Length of A-III 

Teeth in left MxII 

Teeth in right Mx II 

Teeth in Mx III 

Teeth in left Mx IV 

Teeth in right Mx IV 

Last bronchia on setiger no. 


No. of anterior setigers with single filaments 

No. of posterior setigers with single filaments 

No. of anterior setigers with tapering ventral cirri 


No. of aciculae 


No. of subacicular hooks present 

N 

6 

6 

14 

14 

14 

14 

14 

13 

6 

7 

7 

7 

7 

7 

13 

14 

14 

13 

14 

13 

14 

14 

14 

Max. 

116 

59 

4 

25 

8 

3.5 

2 

12 

15 

9 

10 

12 

9 

11 

34 

15 

3 

1 

5 

40 

3 

22 

2 

Min. 

95 

36 

2 

15 

6 

1.75 

0 

4 

7 

7 

8 

8 

6 

8 

29 

11 

2 

0 

3 

31 

2 

18 

1 

Mean 

SD 

106 8.05 

48.17 9.26 

2.82 0.5 

21.07 4.87 

6.71 0.64 

2.46 0.44 

1.21 0.58 

8.15 2.7 

10.67 :1.14 

8.43 0.79 

8.29 ().76 

9.86 

8.29 

10.14 

32 

12.5 

2.93 

.35 

.11 

.07 

.63 

.34 

().27 

0.15 ( ).3K 

4.86 ( ).53 

36.23 

2.07 

2.59 

()27 

20.07 

1.14 

1.07 

()36 





No. of teeth in Mx V 


Branchiae 

Articulations of notopodial cirri 






Subacicular hooks 

Dorsal edge of pygidium (N=6) 

Anal cirri (N=6) 

N=14 

absent 

middle or front of prostomium 

absent 

1 

3 

outreach notopodial cirri 

absent 

slightly flaring; flat 

10 

slightly inflated 

tapering 

distinct 

in every segment after start 

crenulated 

long; without articulations 

(Figure 75d) gently flared, flat. One marginal tooth longer than 


(Figure 75e) gently inflated, marginally finely serrated. 

Appendages extremely long, slender, tapering to blunt, gently 

curved tips, bidentate. Proximal teeth low, triangular thickenings 

near distal ends of appendages. Guards extremely long, 

tapering to fine mucronate tips. Pseudocompound falcigers and 

compound spinigers absent. Aciculae paired in most parapodia, 

yellow, slender, tapering, straight or gently curved; crosssections 



(Figure 75b) yellow, tridentate. Hooks first present from setiger 

22 (18-21 in other syntypes), present in all setigers thereafter, 

always single (except for replacements). Hooks distally curved. 

Fangs decreasing evenly in size from large main fangs to small 

tertiary fangs. 




TURES.—Mx III long and located behind left Mx II. 


SCORED—Inappropriate Characters: 22, 56, 58, 59. Unknown 



None. 

REMARKS.—Eunice mucronata is listed with similar species 

in Tables 41 and 45. It is the only species in Table 45 with 

peristomial cirri clearly projecting beyond the prostomium. It is 

also characterized by the very long antennae and the extremely 

long, pointed guards in the compound falcigers.


126. Eunice multicylindri Shisko, 1981 

FIGURE 75f-i; TABLES 10,41,42 

Eunice multicylindri Shisko, 1981:971-973. figs. 2a-d, table 1. 

MATERIAL EXAMINED.—Holotype, USNM 63066, and 2 

paratypes, USNM 63067, Southern California, 32°43.87'N, 

119°9.86'W, 81 m, coarse sand, 17 Feb 1956, BLM sta 

24356BFI; 2 paratypes, USNM 63068, Southern California, 

33°51.46H 119°57,13'W, 87 m, coarse sand, 13 Oct 1975, 

BLM sta 22952BFI; 2 paratypes, USNM 63069, Southern 

California, 33°39.99'N, 120°0.1'W, 120 m, sand and gravel, 19 

Oct 1975, BLM sta 23087BFI; 1 paratype, USNM 63070, 

Southern California, 33°59.78TSf, 120°22.33'W, 70 m, 6 Nov 

1975, BLMsta23170BFI. 

COMMENTS ON MATERIAL EXAMINED.—Measurements of 

paratypes, including one juvenile, are summarized in Table 10. 



3.5 mm. Body cylindrical, slender, with short, inconspicuous 

parapodia. Anal cirri as long as last 15 setigers together. 


than pcristomium, as deep as l /2 of peristomium. Prostomial 

lobes frontally rounded, dorsally excavate with a thickened 

rim; median sulcus deep. Eyes lateral to bases of A-II, large, 

dark. Antennae arranged in a horseshoe, with A-III isolated by 

a gap, similar in thickness. Ceratophores ring-shaped in all 

antennae, without articulations. Ceratostyles digitiform, distally 

abruptly tapering, with a few, large cylindrical articulations; 

maximum 4 articulations in A-III. A-I to posterior 

peristomial ring. A-II and A-III to setiger 1. Peristomium 

cylindrical. Separation between rings distinct on all sides, but 

best marked dorsally and ventrally; anterior ring 2 /3 of total 

peristomial length. Peristomial cirri to anteriormost edge of 

peristomium or middle of prostomium, digitiform, with 2 to 4 


Maxillary formula of paratype from USNM 63067 1+1,8+8, 

8+0, 7+10, and 1+1. Mx III long, located behind left Mx II. 

Jaws very lightly sclerotinized without any trace of calcification. 





setigers. First 2 and last 10-15 branchiae single filaments; 

maximum 4 filaments. Filaments digitiform, about as long as 


All neuropodial acicular lobes truncate or rounded; aciculae 

emerging at midline. All pre- and postsetal lobes low, 

transverse folds. First 5-6 ventral cirri thick, tapering, basally 

inflated from setigers 6-7 through setiger 45-50. Inflated 

bases modest, ovate; narrow tips long and tapering. Inflated 

portion lost by setiger 50; posterior ventral cirri tapering. All 

notopodial cirri slightly inflated basally, tapering to digitiform 

tips; notopodial cirri thinner in posterior than in anterior 

setigers, retaining same length. Notopodial cirri without 



75h) tapering, furled. Both marginal teeth longer than other 

teeth; 1 marginal tooth longer than other; -12 teeth present. 

Shafts of compound falcigers (Figure 75i) slightly inflated, 

marginally serrated. Appendages long, slightly tapering, 

bidentate. Proximal teeth very much larger than distal teeth, 

triangular, directed slightly basally. Distal teeth distinctly 

hooked. Guards asymmetrically bluntly pointed; mucros 

absenL Pseudocompound falcigers and compound spinigers 

TABLE 10.—Variable and invariable features in the type lot of Eunice multicylindri (N = number of individuals 



N 

Max. 

Min. 

Mean 

SD 

Maximal width 









8 

8 

8 

8 

8 

8 

8 

8 

8 

1 

3.5 

8 

4 

4 

40 

4 

45 

27 

0.1 

1.2 

4 

0 

3 

20 

1 

30 

13 

0.51 

2.63 

5.63 

1.25 

3.38 

29.63 

2.63 

39.88 

23.50 

0.26 

0.78 

1.30 

1.75 

0.52 

6.55 

0.92 

6.38 

4.66 


Separation between rings 







N=8 

visible on all sides 

furled; tapering 

yellow 

tapering;! straight 

yellow 

3 

indistinct

228 


absent. Aciculae yellow, slender, tapering, straight; crosssections 



(Figure 75g) yellow, tridentate with teeth in crest. Hooks first 


single (except for replacements). Hooks with large curved main 

fang; 2 distal fangs emerging from common base; tertiary fangs 

very much smaller than secondary fangs. 








None. 

REMARKS.—Eunice multicylindri was described from rather 

well-preserved material and enough specimens to allow 

statistical treatment of material along the same lines presented 

by Fauchald (in press). Variability in material is indicated in 

Table 10. 

Eunice multicylindri is listed with similar species in Tables 

41 and 42. It is characterized more by a combination of 

character states than by a single unique one. It is discussed in 

the section on E. medicina. 

127. Eunice multipectinata Moore, 1911 

FIGURE 76a-i; TABLES 27, 32 

Eunice nudtipectinata Moore, 1911:248-251, pi. 15: figs. 20-23. 


sta 4377, Southern California, 10.2 miles southwest of 

Point Loma (~32°35'N, 117°25'W), 17 Mar 1904, 232-547 m, 

green mud and sand, dredge; USNM 16880, Albatross sta 

4431,2 specimens, Southern California, 3.5 to 5.2 miles NE of 

Brockway Point, Santa Rosa Island (approximately 34°06 / N, 

120°04'W), 15 Apr 1904, 69-82 m, green and yellow mud, 

rock and sand, various dredges. 


at one time desiccated by exposure to highly concentrated 

alcohol and is now partially distorted; the two other specimens 

examined are part of the original material, but were not 

designated as types by Moore. 



10,16 mm. Body cylindrical, tapering slowly posteriorly. Anal 

cirri as long as last 12 setigers combined. 


than peristomium, less than 1 /i as deep as peristomium. 

Prostomial lobes frontally rounded, dorsally excavate with a 

thickened rim; median sulcus shallow, but distinct. Eyes 

between bases of A-I and A-II, dark. Antennae evenly spaced, 


without articulations. Ceratostyles tapering, with cylindrical 

articulations except drop-shaped distalmost articulations in 

A-I. A-I to posterior peristomial ring; A-II to setiger 1; A-III 

incomplete in all specimens, apparently at least as long as A-II. 

Peristomium cylindrical; front edge of lower lip coarsely 

scalloped, muscular. Separation between rings distinct dorsally 

and ventrally; anterior ring about 5 A of total peristomial length. 

Peristomial cirri to front of anterior peristomial ring; in 1 

specimen to posterior end of prostomium, slender and 

digitiform, with 5 to 6 articulations. 

Maxillary formula of specimen from USNM 16880 1+1, 

8+8, 9+0, 7+11, 1+1, and 1+1. Mx VI small with barely 

projecting point. Mx III short; part of distal arc with left Mx IV. 

Branchiae (Figure 76a,e) present, pectinate, distinctly longer 


Branchiae from setiger 7-8 to sctiger 1%. Branchiae present to 


of setigers. Single filaments in first 8-10 branchiated setigcrs; 

up to 12 slender filaments in trim scries at sctigcrs 15-20. 

Branchial stems slender. Filaments shorter than notopodial cirri 

in all setigers. Branchiae shorter than notopodial cirri except in 

setigers 15-30. 

All neuropodial acicular lobes asymmetrically obliquely 

rounded; aciculae emerging from upper edge of lobes. All 


cirri thick, digitiform, becoming basally inflated from about 

setiger 5. Inflated bases distinct, elevated ridges along lower 

edge of neuropodia; narrow tips tapering. Inflated bases 

reduced from about setiger 65, remaining visible in all but last 

50-60 setigers. Far posterior ventral cirri very long, slender, 

digitiform, about twice as long as neuropodial acicular lobes. 

All notopodial cirri slender, tapering, slightly inflated basally. 

Anterior notopodial cirri with up to 7 indistinct articulations; 

articulations lost by setiger 15. 

Limbate setae marginally serrated. Pectinate setae (Figure 

76c,i) tapering, flat or slightly furled. One marginal tooth very 

much longer than all other teeth; 10-15 teeth present. Shafts of 

compound falcigers tapering, marginally finely serrated in 

anterior setigers (Figure 76b), marginally smooth in posterior 

setigers (Figure 76g). Anterior appendages tapering, bidentate. 

Proximal teeth larger than distal teeth, curved, laterally 

directed. Distal teeth distinctly curved. Median and posterior 

appendages less tapering. Proximal teeth triangular, directed 

laterally, straight. Distal teeth similar to those in anterior 

appendages. Guards bluntly to sharply symmetrically pointed, 

marginally sometimes frayed; mucros absent. Pseudocompound 


(Figure 76f) paired in all but few posterior setigers, with dark 

brown to black cores and clear sheaths, tapering, distinctly 

bent. Curvature of aciculae at right angles to each other in some 

parapodia; cross-sections round. Subacicular hooks (Figure 

76h) with dark brown to black cores and clear sheaths, 

bidentate. Hooks first present from setigers 37 and 39 in 2


FIGURE 76.—Eunice multipectinata (holotype, USNM 16919): a, parapodium 20, anterior view; b, compound 

falciger, parapodium 20; c, pectinate seta, parapodium 20; d, anterior end, lateral view; e, parapodium 60, anterior 

view; f, aciculae, parapodium 60; g, compound falciger, parapodium 60; h, subacicular hook, parapodium 60; i, 

pectinate seta, parapodium 60. Eunice murrayi (holotype, BM(NH) ZK 1885.12.1.201): j, anterior end, lateral 

view. (Scale bars in mm.)


specimens from USNM 16880 and from setiger 45 in holotype, 


replacements). Hooks tapering, bent laterally towards distal; 

heads distinct. Proximal teeth very much larger than distal 

teeth, distally curved, laterally directed. Distal teeth nearly 

erecL 

UNKNOWN MORPHOLOGICAL FEATURES.—Pygidium. 





Characters: 4,6, 15,23,42. 


None. 

REMARKS.—Eunice multipectinata is listed with similar 

species in Tables 27 and 32. It is first and foremost 

characterized by gracile, strongly pectinate branchiae and 

slender, digitiform antennae and notopodial cirri; the subacicular 

hooks start rather late, compared to the two other species 

listed in Table 32 with 10 or more branchial filaments. 

128. Eunice murrayi Mclntosh, 1885 

FIGURE 76j; TABLES 41,45 

Eunice murrayi Mclntosh, 1885:288-289, figs. 48, 49, pi. 39: figs. 7, 8, pi. 

20A: figs. 19, 20. 


1885.12.1.201, Simon's Bay, Cape of Good Hope, dredged, 33 

m, Dec 1873. 


length approximately 37 mm; maximal width 4 mm; length 

through setiger 10, 7 mm. Anal cirri long, articulated. 


peristomium, as deep as 1 /i of peristomium. Prostomial lobes 

frontally rounded, dorsally flattened; median sulcus shallow. 

Eyes not seen. Antennae in a horseshoe, evenly spaced, similar 

in thickness. Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostyles tapering, with moniliform articulations; 

complete left A-II with 8 articulations; incomplete A-III 

with 10 articulations. A-I to setiger 1; A-II to setiger 3; 

incomplete A-III currently to setiger 3; however, apparently 

originally longer than A-II. Peristomium cylindrical. Separation 

between rings distinct dorsally and ventrally; anterior ring 

3 A of total peristomial length. Peristomial cirri to frontal margin 

of peristomium, with 3 articulations of which basal ones more 

than half of total cirral length. 

Maxillary formula 1+1,5+5, 5+0,5+8, and 1+1. Mx III long, 

positioned directly behind left Mx II. 



from setiger 5 to setiger 28. Branchiae present to near posterior 

end, present on less than 55% of total number of setigers. Two 

first and last 5 branchiae single filaments, maximum; 9 

filaments at about setiger 10. Branchial stems strong, stiff. 

Filaments arranged in a neat comb, relatively short. 

All neuropodial acicular lobes distally truncate; aciculae 


transverse folds. Prebranchial ventral cirri digitiform. Between 

setigers 5 and 30 ventral cirri basally moderately inflated. 

Inflated bases ovate; narrow tips digitiform. Posterior ventral 

cirri increasingly digitiform, increasing in length, by setiger 50 

as long and prominent as notopodial cirri. All notopodial cirri 

medially inflated, similar in length. Prebranchial cirri with 3 or 

4 articulations; articulations increasingly indistinct posteriorly; 

most notopodial cirri in branchial region with single articulation; 

postbranchial notopodial cirri without articulations. 

Nearly all setae broken. Shape of limbatc and pectinate setae 

and aciculae currently indeterminable. Mclntosh (1885, pi. 

20A: fig. 19) illustrated shafts of compound falcigcrs as mildly 

inflated. Appendage large with both teeth well developed, 

distinctly curved. Guards blunt; mucros absent. Pseudocompound 

falcigcrs and compound spinigers absent. Aciculae 

yellow, distally pointed; cross-sections round. Separation 



teeth in a crest. Hooks first present from setiger 17. 

UNKNOWN MORPHOLOGICAL FEATURES.—Many setal features; 






Characters: 4,6, 13,14,23, 65-68, 81, 82. 


None. 

REMARKS.—Eunice murrayi has been considered a synonym 

of E. australis, which it resembles in distribution of branchiae 

and in shape of the subacicular hooks. It is listed with similar 

species in Tables 41 and 45. It resembles E. prayensis in having 

the separation between the peristomial rings distinct on all 

sides, but differs from the it in having the notopodial cirri 

articulated in the anterior end; in E. prayensis the notopodial 

cirri lack articulations. 

129. Eunice mutabilis Gravier, 1900 


Eunice mutabilis Gravier, 1900:245-248, figs. 105-109, pi. 13: figs. 71-74. 

MATERIAL EXAMINED.—TWO syntypes, MNHN, Paris, 

Djibouti, coll. Jousseaume and Coutiere, 1897. 


based on the shorter of the two syntypes. 

DESCRIPTION.—Both syntypes complete; one with 196 


through setiger 10, 8 mm. Other syntype with 313 setigers; 

length 117 mm; maximal width 7.5 mm; length through setiger

NUMBER 523 

231 

FIGURE 77.—Eunice mulabilis (syntype, MNHN): a, parapodium 55, anterior view; b, pectinate seta, parapodium 

55; c, compound falciger, parapodium 55; d, anterior end, lateral view; e, subacicular hook, parapodium 55. 

Eunice mutilata (holotype, USNM 4789): f, anterior end, lateral view; g, prebranchial parapodium, anterior view; 

h, compound falciger, prebranchial parapodium; i, parapodium 30-35 (these parapodia all mounted together on 

a slide and not readily distinguishable), anterior view; j, compound falciger, parapodium 30-35; k, subacicular 

hook, parapodium 35; 1, pectinate seta, parapodium 30-35; m, compound falciger, parapodium 30-35; n, 

subacicular hook, parapodium 30-35.


10, 8.5 mm. Anterior body cylindrical, becoming dorsoventrally 

flattened by setiger 50. Abruptly tapering to pygidium. 

Anal cirri short, without articulations. 




sulcus deep. Eyes lateral to bases of A-II, dark. Antennae in a 

horseshoe, evenly spaced, with A-I thicker than other 3. 


Ceratostyles tapering, without articulations. A-I barely to 

posterior peristomial ring; A-II to setiger 1; A-I 11 to setiger 2. 

Peristomium cylindrical; lower lip very muscular, forming 

distinct ventrolateral pad. Separation between rings distinct 

dorsally and ventrally only; anterior ring 4 /s of total peristomial 

length. Peristomial cirri to front edge of peristomium or 

posteriormost part of prostomium, basally inflated, tapering to 

slender tips, without articulations. 

Summary maxillary formula 1+1, 4-544-5, 6fO, 4-6+6-8, 

and 1+1. Left Mx IV short; part of distal arc with Mx III, with 

3 large and varying number of very small, knob-like teeth. 



Branchiae from setiger 17-18 to setiger 196. Branchiae present 

to near posterior end; present on more than 65% of total number 

of setigers. First 2-3 pairs single filaments; all other branchiae 

with 3 or more filaments. Where best developed, at about 

setiger 25, branchiae with 12 slender filaments; more usually 

branchiae with 8 to 9 filaments. Branchial stems slender, 

flexible. Filaments longer than notopodial cirri in all but first 

few and last 15-20 setigers. 


rounded by setiger 10 and conical by setiger 30, retaining that 

shape through rest of body; aciculae emerging above midline. 

All presetal lobes low, transverse folds. Anterior postsetal lobes 

higher than acicular lobes, distally rounded, becoming reduced 

to low, transverse folds by setiger 25. Anterior ventral cirri 

thick, tapering, becoming basally inflated from setiger 10, 

retaining inflated condition through rest of body. Inflated bases 

gradually decreasing in importance posteriorly, forming low 

transverse ridges in posterior half of body; narrow tips tapering. 

All notopodial cirri slightly inflated basally, tapering to 

digitiform tips; those in first 3-4 setigers distinctly longer than 

those in next following setigers. Notopodial cirri decreasing 

slowly in length over remainder of body, becoming about half 

as long in last few setigers as in setiger 10. Notopodial cirri 



77b) in distinct fascicles of 15-20 in median and posterior 

setigers, furled, flared. Both marginal teeth longer than other 

teeth; 1 marginal tooth distinctly longer than other; -20 teeth 

present. Shafts of compound falcigers (Figure 77c) slightly 

inflated, marginally smooth, with distinct internal striations. 

Appendages very large, tapering slightly to large heads, 

bidentate. Proximal teeth slightly larger than distal teeth, 

triangular, directed laterally. Distal teeth gently curved or 

nearly erect. Guards symmetrically rounded, marginally 


spinigers absent. Aciculae projecting, with light to 

medium brown cores and clear sheaths, tapering, usually 

straight, slightly curved dorsally in some parapodia; crosssections 

round. Subacicular hooks (Figure 77e) with light to 

medium brown cores and clear sheaths, bidentate. Hooks first 

present from setiger 22-23, present in all setigers thereafter, 

always single (except for replacements). Hooks tapering 

strongly to very small heads. Proximal teeth very much larger 

than distal teeth. Both teeth directed distally. 

UNKNOWN MORPHOLOGICAL FEATURES.—Pygidium. 






ASSUMED STATES FOR PURPOSE OF PRF.PARING KEY.— 

None. 

REMARKS.—Eunice mutabilis is one of many species in 

group B-4 described from the Red Sea by Gravier, it has been 

considered synonymous with E. afra and is listed with similar 


with more than 10 branchial filaments and with the bases of the 

ventral cirri inflated through posterior setigers. 

130. Eunice mutilata Webster, 1884 

FIGURE 77f-n; TABLES 27,31 

Eunice mutilata Webster, 1884:315-316, pi. 9: figs. 36,36i-d, 37-40. 

MATERIAL EXAMINED.—Holotype, USNM 4789, 1 specimen 

in 3 pieces, plus 5 mounted slides of parapodia, Bermuda, 

coll. G. Brown Goode. 


of an anterior end, a median piece, and a long section with 

a posterior end; however, perhaps as much as 50 middle 

setigers are missing. The three pieces match in all characters 

and are here considered part of the holotype, but because an 

unknown number of setigers are missing, no overall measurements 

can be given. 

DESCRIPTION.—Anterior end with 36 setigers; length 30 

mm; maximal width 3.5 mm wide at setiger 10; length through 

setiger 10, 10 mm. 



lobes frontally rounded, dorsally strongly inflated; median 

sulcus deep. Eyes between bases of A-I and A-II, small, black. 

Antennae in a straight line, evenly spaced, similar in thickness. 


Ceratostyles tapering, with up to 5-6 very indistinct, cylindrical 

articulations. A-I to posterior peristomial ring; A-II to 

setiger 1; A-III to setiger 2. Peristomium cylindrical. Separa-


tion between rings distinct on all sides; anterior ring 3 A of total 

pcristomial length. Pcristomial cirri to middle of anterior 





Branchiae from seiiger 7, missing on last 15-20 setigers. 


65% of total number of setigers. First 3 pairs and last half of 

body with single filaments only; maximum 4 filaments at about 

sctiger 25-30. Branchial stems short, tapering. Filaments in 

pectinate branchiae digitiform, much longer than notopodial 

cirri. Branchial filaments short on posterior half of body, 

digitiform, distinctly shorter than notopodial cirri. 

Anterior neuropodial acicular lobes (Figure 77g) short and 

symmetrically rounded; median (Figure 77i) and posterior 

acicular lobes obliquely truncate with aciculae emerging from 

upper and higher ends. All presetal lobes low, transverse folds. 

Anterior postsetal lobes high narrowly rounded, symmetrical, 

becoming broadly rounded in median setigers, turning into low, 

transverse folds in far posterior setigers. Postsetal lobes 

recognizable as free lobes through at least setiger 35. Anterior 

ventral cirri thick, tapering, becoming basally inflated from 

about setiger 10. Inflated bases transverse welts along ventral 

side of neuropodia; narrow tips digitiform. Inflated bases 

missing in median fragment; posterior ventral cirri short, 

digitiform, increasingly slender. All notopodial cirri digitiform, 

without articulations, becoming increasingly slender posteriorly. 

Limbate setae broadly limbate, marginally smooth. Pectinate 

setae (Figure 771) tapering, furled. One marginal tooth longer 

than all other teeth; 12-15 teeth present Shafts of compound 

falcigers (Figure 77h,j,m) inflated, marginally smooth. All 

appendages tapering, nearly triangular in shape, bidentate. Two 

distinctly different kinds of heads present. In most heads both 

teeth directed distally (Figure 77h,m); proximal teeth smaller 

than distal teeth. Both teeth curved; distal teeth nearly erect In 

other, rarer kind of heads proximal teeth triangular (Figure 77j), 

directed laterally. Distal teeth distinctly bent. Both kinds 

present in same parapodia; relative proportions of both kinds 

varying along length of body, with second kind becoming more 

common posteriorly. Both kinds of compound hooks with 

pointed guards, symmetrically and asymmetrically bluntly 

pointed, respectively; mucros absent. Pseudocompound falcigers 

and compound spinigers absent. Aciculae dark brown to 

nearly black, tapering, straight or gently curved; cross-sections 

round. Subacicular hooks (Figure 77k,n) dark brown, bidentate. 


thereafter, always single (except for replacements). All hooks 

with narrowed, sometimes bent necks and distinct heads. 

Proximal teeth somewhat larger than distal teeth, curved 

distally. Distal teeth nearly erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—Jaw structures; 

features associated with far posterior setigers; pygidium and 

anal cirri. 





Characters: 1,2,4,6,40, 74, 78. 


None. 

REMARKS.—Eunice mutilata is listed with similar species in 

Tables 27 and 31. An apparently characterizing feature is the 

presence of free postsetal lobes through a long anterior section 

of body. 

Eunice mutilata samoae Hartmann-Schrtider, 1965 

Eunice mutilata samoae Hartmann-Schroder, 1965a:134~135. 

REMARKS.—Described as subspecies of E. mutilata Webster, 

this form is here considered separate species and treated as 

such below. 

131. Eunice mutilatoides Augener, 1922 

FIGURE 78a-c; TABLES 25,26 

Eunice mutilatoides Augener, 1922b:45. 

MATERIAL EXAMINED.—Holotype, ZMB 6397, Bird Key, 

Dry Tortugas, coll. Hartmeyer, 1907. 

COMMENTS ON MATERIAL EXAMINED.—Holotype has been 

partially dried out; its condition does not warrant illustrations 

of soft parts. 



5 mm. 

Prostomium distinctly shorter and narrower than peristomium, 

as deep as V2 of peristomium. Median sulcus shallow. 

Eyes posterior to bases of A-I, dark. Antennae in a straight line, 


in all antennae, without articulations. Ceratostyles thick and 

digitiform, without articulations. No antennae reaching beyond 

peristomium; A-I short; A-I I and A-III similar in length. 


dorsally and ventrally; anterior ring 3 A of total peristomial 


ovate, without articulations. 



cirri. Branchiae from setiger 8 to setiger 48. Branchiae 


of total number of setigers. All branchiae short, stubby 

filaments; no more than 2 filaments apparently present in any 

setiger. 

Anterior neuropodial acicular lobes rounded or truncate; 

posterior acicular lobes currently sharply pointed, possibly due 

to condition of type. Shape of pre- and postsetal lobes cannot be

234 


.02 a, b, c, f, g 

d. e 

\ 

FIGURE 78.—Eunice mulilaioides (holotype, ZMB 6397): a, pectinate seta, parapodium 95; b, subacicular hook, 

parapodium 95; c, compound falciger, parapodium 95. Eunice narconi (syntype, BM(NH) ZB 1972.73): d, 

subacicular hook, about parapodium 50; e, acicula, about parapodium 50; f, pectinate seta, about parapodium 50; 

g, compound falciger, about parapodium 50. (Scale bars in mm.) 

determined. First 7 ventral cirri tapering. Ventral cirri long 

ventrolateral ridge in median setigers; narrow tips tapering. 

Posterior ventral cirri tapering. Notopodial cirri apparently 

short, medially inflated. Notopodial cirri without articulations. 


78a) long, slender, tapering, furled. One marginal tooth 

distinctly longer than other teeth; -10 teeth present. Compound 

falcigers (Figure 78c) small. Shafts inflated, marginally 

serrated. Appendages short with relatively large heads, bidentate. 

Proximal teeth tapering to sharp, curved tips. Distal teeth 

abruptly tapering, directed laterally. Guards asymmetrically 

bluntly pointed, following outline of hooks closely; mucros 


absent. Aciculae tan-colored, tapering to blunt tips, straight; 

cross-sections round. Subacicular hooks (Figure 78b) light to 

dark tan color, bidentate. Hooks first present from setiger 19, 


replacements). Hooks with distinct heads. Proximal teeth large, 

directed laterally. Distal teeth about as long as proximal teeth, 

slender, directed obliquely distally. 


some parapodial features; pygidium and anal cirri. 





Characters: 4,6-8,44-50, 74, 78.



None. 

REMARKS.—Eunice mutilatoides was characterized very 

briefly by Augener (1922b) and has remained unreported since 

that time. It is listed with similar species in Tables 25 and 26. 

In addition to E. mutilatoides, the only species in Table 26 with 

poorly developed, palmate branchiae and without articulations 

of the ceratostyles is E. palauensis. The branchiae outreach the 

notopodial cirri in E. palauensis; they are distinctly shorter 

than the notopodial cirri in all setigers in E. mutilatoides. 

132. Eunice narconi Baird, 1869 

Eunice narconi Baird. 1869:350-351. 

FIGURE 78d-g; TABLES 19,21 

MATERIAL EXAMINED.—Three syntypes, BM(NH) ZB 

1972.73, Narcon Island, Antarctic Seas, Erebus and Terror 

Southern Seas Expedition, 1839-1843. 


has been dried out at one time, so illustrating the soft-parts 

would be meaningless. The material consists of three anterior 

ends and some fragments of median and posterior setigers. One 

anterior end is in very bad shape and even the number of 

segments cannot be determined; the description is based on the 

other two anterior fragments. 

DESCRIPTION.—Larger fragment with about 75 setigers; 


6 mm. Shorter fragment with about 50 segments; length 17 

mm; maximal width 1.5 mm; length through setiger 10,5 mm. 

Prostomium distinctly shorter and narrower than peristomium. 

Prostomial lobes frontally rounded, dorsally inflated; 

median sulcus shallow. Shape and length of antennae cannot be 

determined. Peristomium cylindrical. Anterior peristomial ring 

3 A of total peristomial length. Peristomial cirri short, without 



Branchiae present, palmate. Branchiae from setiger 8 to 

setigers 33-35. Branchiae terminating well before posterior 

end, present on less than 55% of total number of setigers. 

Where best developed at around setiger 15 at least 3 filaments 

present. 

Parapodial shape cannot be described in detail; ventral cirri 

with distinctly inflated bases in branchial region. 


setae (Figure 780 tapering, flat. One marginal tooth very much 

longer and stronger than all other teeth; -10 teeth present 

Shafts of compound falcigers (Figure 78g) distinctly inflated, 

marginally serrated. Appendages long, narrow, bidentate. Both 

teeth gently curved. Guards distally bluntly pointed, marginally 

serrated; mucros absent Pseudocompound falcigers and compound 

spinigers absent Aciculae (Figure 78e) paired, yellow, 

tapering to blunt points, gently curved ventrally; cross-sections 



78d) yellow, bidentate. Hooks first present from setiger 35-38. 

Hooks distally curved. Proximal teeth larger than distal teeth. 

Both teeth directed distally. 


and peristomial features; jaw structure; nearly all parapodial 

features; pygidium and anal cirri. 





Characters: 1, 2, 4, 6, 11-23, 25, 32-34, 39, 40, 43-52, 54, 

55,57-64,81,82. 


None. 

REMARKS.—Eunice narconi has been considered incompletely 

known (Augener, 1931:287). Hartman (1964:118) 

synonymized it with some doubt with E. pennata. It agrees with 

E. pennata in that it has yellow, bidentate subacicular hooks 

and branchiae limited to some anterior segments. However, E. 

pennata, at least in European waters, appears to have branchiae 

starting at setigers 3-4, whereas in E. narconi branchiae do not 

start before setiger 8. The two species differ about as much as 

any two species in the groups do, so the synonymy is here 

considered unacceptable. 

Eunice narconi is listed with similar species in Tables 19 and 

21; it is too poorly known to be included in the key. 

133. Eunice nesiotes (Chamberlin, 1919) 


Leodice nesiotes Chamberlin, 1919a:253-256, pi. 57: figs. 6, 7. 

MATERIAL EXAMINED.—Holotype, USNM 19749, Albatross, 

1899-1900 (no station indicated), Marshall Islands, 12 

fathoms, came up on anchor. 


been dried out or exposed to very strong alcohol at one time, so 

most of notopodial cirri and branchiae are badly distorted. 

DESCRIPTION.—Holotype incomplete mature female with 

large eggs in body cavity of 101 setigers; length 90 mm; 

maximal width 6 mm; length through setiger 10,10 mm. 


than peristomium, less than l fr as deep as peristomium. 

Prostomial lobes frontally obliquely truncate, dorsally flattened, 

tilted laterally; median sulcus shallow, continued as 

ridge along prostomial midline. Antennae in a horseshoe, with 

A-I isolated by a gap, similar in thickness. Ceratophores 


tapering, irregularly articulated with up to 5 long, cylindrical 

articulations in A-III. A-I to posterior edge of peristomium; 

A-II to setiger 7; A-III to setiger 10. Peristomium cylindrical, 

somewhat ribbed dorsally, ventrally folded into several shallow 

folds. Separation between rings distinct dorsally, and barely

236 


FIGURE 79.—Eunice nesiotes (holotype, USNM 19749): a, anterior end, lateral view; b, pectinate seta, 

parapodium 35; c, subacicular hook, parapodium 35; d, compound falciger, parapodium 35; e, parapodium 35, 

anterior view. Eunice nicidiqformis (holotype, USNM 5213): f, parapodium 16, anterior view; g, pectinate seta, 

parapodium 16; h, compound falciger, parapodium 16; i, anterior end, lateral view; j, parapodium 47, anterior 

view; k, pectinate seta, parapodium 47; 1, compound falciger, parapodium 47; m, subacicular hook, parapodium 



visible ventrally; anterior ring 4 /s of total peristomial length. 

Peristomial cirri to front edge of peristomium or somewhat 

beyond, slender and tapering, with 2 long articulations. 


forming part of distal arc with left Mx IV. 



Branchiae from setiger 6 to end of fragment. First 2 branchiae 

single, short filaments; maximum 14 filaments present. 

Branchial stems short, tapering. Filaments long, slender. 

Neuropodial acicular lobes distally rounded or truncate; 

aciculae emerging dorsal to midline. Pre- and postsetal lobes 

low, transverse folds. First 5 ventral cirri thick, tapering. 

Ventral cirri basally inflated in branchial region. Inflated bases 

thick, transverse welts; narrow tips very long, distally truncate, 

digitiform. Notopodial cirri very long, tapering from slightly 

inflated bases, less dominant in posterior than in anterior 

setigers. Notopodial cirri without articulations. 

Pectinate setae (Figure 79b) coarse, tapering, flat. One 

marginal tooth slightly longer than other teeth; -10 coarse teeth 

present. Shafts of compound falcigers (Figure 79d) slightly 

inflated, marginally smooth. Appendages slender, bidentate. 

Both teeth directed laterally. Guards asymmetrically bluntly 

pointed, with convexity covering proximal teeth; mucros 


absent. Aciculae paired, black, tapering to bluntly conical tips, 

straight; cross-sections round. Subacicular hooks (Figure 79c) 

black, bidentate. Hooks first present from setiger 31, present in 


Hooks slender. Proximal teeth twice as large as distal teeth, 



with posterior parapodia; pygidium and anal cirri. 





4, 6, 13,14, 36-38; 40, 42,47, 50, 57-60, 63, 74, 78. 


38,1. 

REMARKS.—Eunice nesiotes is listed with similar species 

Tables 27 and 31. The ventral cirri are basally inflated through 

the posterior end in this species; however, the shape cannot be 

determined accurately. Two other species listed in Table 31 

have such distribution of the inflated bases. Eunice borneensis 

has short peristomial cirri; E. guildingi and E. nesiotes have 

peristomial cirri reaching the middle of the prostomium. The 

peristomial cirri lack articulations in E. guildingi; they have 

two long articulations in E. nesiotes. 

134. Eunice nicidioformis Treadwell, 1906 

FIGURE 79f-m; TABLES 11,24,25 

Eunice nicidiaformis Treadwell, 1906:1169, figs. 49-51. 

MATERIAL EXAMINED.—Lectotype, USNM 5213, and 5 

paralectotypes, USNM 142035, Albatross sta 4098, Hawaiian 

Islands, 6.5 miles off Puniawa (= Pauwela) Point, Maui 

(~21°O1X 156°12'W), 23 Jul 1902, 174-278 m, fine coral 

sand and foraminifera, beamtrawl. Four paralectotypes, USNM 

5419, Albatross sta 4077, Hawaiian Islands, 6.1 miles off 

Puniawa (= Pauwela) Point, Maui (-H'Vl'N, 156°15'W), 21 

Jul 1902, 181-194 m, fine coral sand and foraminifera, 

beamtrawl. 

DESCRIPTION.—Holotype complete female with large eggs 

in body cavity with 97 setigers; total length 45 mm; maximal 

width 2 mm; length through setiger 10, 6 mm. Body slender, 

cylindrical with long, slender antennae and long, tapering anal 

cirri. Anal cirri as long as last 15 setigers combined. 


peristomium, less than 1 /2 as deep as peristomium. Prostomial 


rim; median sulcus deep. Eyes near bases of A-II, dark. 

Antennae in a horseshoe, with A-I isolated by a gap, similar in 


articulations. Ceratostyles tapering, with up to 10 distinct 

articulations; articulations cylindrical, drop-shaped distally in 

A-I. A-I to setiger 1; A-II to setiger 7; A-II I to setiger 13. 

Antennae of paratypes generally shorter, but A-I always 

outreaching peristomium and A-III longer than all other 

antennae in all specimens. Peristomium slightly flaring 

anteriorly, lower lip scalloped, muscular. Separation between 

rings distinct dorsally and ventrally; anterior ring ~ 5 /6 of total 


slender and tapering, with 9 articulations. 

Maxillary formula of a paratype from USNM 5213 1+1,7+7, 

8+0, 6+12, and 1+1. Teeth small, slender. Mx III part of distal 

arc with left Mx IV. 

Branchiae (Figure 79f) present, single filaments, distinctly 

shorter than notopodial cirri. Branchiae from setiger 6-8 to 

setigers 40-50. Branchiae terminating well before posterior 

end, present on less than 55% of total number of setigers. First 

20 branchiae digitiform, slightly shorter than notopodial cirri. 

In last several branchial setigers branchiae short spurs on dorsal 

side of notopodial cirri; spurs forming part of vascularized 

sleeves surrounding bases of notopodial cirri (Figure 79j) in 

varying number of median setigers. Spurs decreasing in size 

posteriorly, disappearing at about setigers 40-50. 

All neuropodial acicular lobes symmetrically conical; aciculae 

emerging at midline. Anterior presetal lobes excavate, 

becoming low, transverse folds by setiger 25. All postsetal 

lobes low, transverse folds. First 9 ventral cirri thick, tapering. 

Median ventral cirri basally inflated. Inflated bases very large, 

nearly spherical; narrow tips digitiform. From about setiger 20 

inflated bases rapidly reduced, completely missing by setiger 

30. Posterior ventral cirri triangular, tapering to distinct tips. 

Anterior notopodial cirri very long, nearly filiform, with 6 to 8 

cylindrical articulations. Notopodial cirri reduced to half length 

by setiger 20, without articulations. Posterior notopodial cirri


TABLE 11.—Variable and invariable features of the type lot of Eunice nicidioformis (N = number of individuals 



N 

Max. 

Mm. 

Mean 

SD 


Total length 

Maximal width 


No. of antcnnal articulations 


Branchiae first from setiger no. 





10 

10 

10 

10 

9 

10 

9 

9 

10 

10 

10 

97 

47 

2.2 

6.5 

15 

10 

8 

50 

26 

6 

25 

58 

25 

1 

4.2 

5 

4 

6 

19 

12 

4 

14 

86.70 

36.15 

1.80 

5.21 

10.22 

8.60 

6.56 

32.00 

21.80 

4.30 

21.40 

11.15 

7.39 

0.33 

0.67 

2.77 

1.78 

0.73 

10.32 

3.91 

0.67 

3.03 




Notopodial articulations present 





N=10 

visible donally and ventrally 

1 


brown 

brown 

2 

indistinct 

distinctly digitiform, abruptly tapering distally. Notopodial 

cirri longer than branchiae in all setigers. 

Limbate setae marginally smooth. Anterior pectinate setae 

(Figure 79g) tapering, flat, becoming slightly flaring in median 

and posterior setigers (Figure 79k). One marginal tooth 

distinctly longer than all other teeth; -10 teeth in anterior 

setigers; -15 teeth in median and posterior setigers. Shafts of 

compound falcigers (Figure 79h,l) tapering, marginally serrated. 

Anterior appendages slender, slightly tapering, bidentate. 

Proximal teeth shorter than distal teeth, triangular. Distal teeth 

slender, curved. Median and posterior appendages shorter than 

those in anterior setigers, more distinctly tapering, bidentate. 

Proximal teeth longer than distal teeth, tapering, rather than 

triangular. Distal teeth similar to those in anterior appendages. 

Guards asymmetrically bluntly pointed; mucros absent. Pseudocompound 


paired, brown, tapering with blunt tips, slightly bent 

distally; cross-section round. Subacicular hooks (Figure 79m) 

brown, bidentate. Hooks first present from setiger 25 (setigers 

19-26 in paratypes), present in all setigers thereafter, always 

single (except for replacements). Hooks curved, tapering. 

Proximal teeth larger than distal teeth, triangular, directed 

laterally. Distal teeth erect. 


anal cirri. 




SCORED.—Inappropriate Characters: 33, 34, 39-42, 56, 58, 

59. Unknown Characters: 4,6, 74, 78. 


None. 

REMARKS.—Eunice nicidioformis is characterized by reduced 

branchial equipment, reflected in Treadwell's choice of 

specific name. The species is listed with similar species in 

Tables 24 and 25. The structure of the posterior branchiae is 

very unusual and may be unique in the genus; they form a 

vascularized sleeve around the base of the notopodial cirri in 

this species. In all other species of Eunice posterior single 

branchial filaments are similar to the anterior ones in structure 

in that they form a spur or a filament attached to the dorsal side 

of the notopodial cirrus or emerge directly from the body wall. 

In other features E. nicidioformis resembles E. gravieri as far as 

the features listed in the tables are concerned, but lacks the 

second pair of eyes described for the latter species. 

135. Eunice nigricans Schmarda, 1861 


Eunice nigricans Schmarda, 1861:131, 7 figs. 

Eunice macrobranchia.—Augcner, 1925:27-28 [in part, not Eunice macro - 

branchia Schmarda, 1861]. 

Eunice aphroduois.—HitVman, 1959:311 [not Nereis ophrodilois Pallas. 

1788]. 

MATERIAL EXAMINED.—Holotype, ZMW, Inv. No. 1066, 

South of Port Royal, Jamaica, coll. L. K. Schmarda. 


length 100 mm; maximal width 7 mm at setigers 25-30; length 

through setiger 10, 10 mm; width at setiger 10, 6 mm. Body 

strongly dorsoventrally flattened. 


than peristomium, as deep as x /i of the peristomium. Prostomial 

lobes frontally obliquely truncate, dorsally excavate with 

thickened rim; median sulcus deep. Eyes between bases of A-I


FIGURE 80.—Eunice nigricans (holotype, ZMW, Inv. No. 1066): a, anterior end, lateral view; b, parapodium 48, 

anterior view; c, compound falciger, parapodium 48; d, pectinate seta, parapodium 48; e, aciculae, parapodium 

48; f, subacicular hook, parapodium 48. Eunice northioidea (holotype, USNM 15728): g, anterior end, lateral 

view; h, parapodium 19, anterior view; i, compound falciger, parapodium 19; j, pectinate seta, parapodium 19; k, 

subacicular hook, parapodium 41; 1, pectinate seta, parapodium 41; m, parapodium 41, anterior view. (Scale bars 

in mm.)


and A-II. Antennae in a horseshoe, evenly spaced, with A-III 

thicker than other four. Ceratophores ring-shaped in all 


to 20 short, cylindrical articulations in A-III. A-I to middle of 

anterior peristomial ring; A-II to seuger 3; A-III to setiger 6. 

Peristomium cylindrical. Separation between rings visible only 

dorsally between bases of peristomial cirri; anterior ring 4 /s of 

total peristomial length. Peristomial cirri to middle of prostomium, 

slender, with 7 articulations. 

Jaws missing. Illustration in Schmarda (1861) indicating a 

maxillary formula of 1+1, 4+5, 6+0, +6, and 1+1. Mx III 

forming part of distal arc with left Mx IV. Mx VI possibly 

present. 



Branchiae from setiger 6 on one side and setiger 7 on other side 

to setiger 180. Branchiae present to near posterior end, present 

on more than 65% of total number of setigers. All branchiae 

with 2 or more filaments; maximum 18 filaments present. 

Branchial stems thick, tapering, erect. Filaments distinctly 

shorter than notopodial cirri in all branchial setigers, thick, 

digitiform. 


obliquely truncate by setiger 25, symmetrically conical by 

setiger 100; aciculae emerging dorsal to midline. All pre- and 

postsetal lobes low, transverse folds. First 4 ventral cirri 

tapering. Ventral cirri distinctly inflated between setiger 5 and 

setiger 30. Inflated bases ovate; narrow tips tapering. Inflated 

bases decreasing in importance from about setiger 30; upper 

face of cirri becoming excavated to form shallow scoops 

covering lower edge of neuropodia. Scoops becoming shallower 

in far posterior setigers; last few setigers with slender, 

tapering ventral cirri. All notopodial cirri basally inflated, 

tapering to slender tips, with up to 7 articulations in 

prebranchial setigers; numbers of articulations reduced to 3 by 

last setigers. Notopodial cirri dominant parapodial features in 

most setigers, becoming increasingly dominant in last setigers 

present. 


(Figure 80d) in thick fascicles in median and posterior setigers, 

tapering, flat. Both marginal teeth distinctly longer than other 

teeth; 15 teeth present. Shafts of compound falcigers (Figure 

80c) tapering, marginally smooth. Appendages relatively short, 

bidentate. Both teeth of same size. Proximal teeth triangular, 

directed laterally. Distal teeth directed obliquely distally. 

Guards symmetrically bluntly pointed; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Notopodial 

aciculae paired, brown. Aciculae (Figure 80e) paired in 

most setigers, with brown cores and clear sheaths, tapering to 

slender tips, curved; cross-section round. Subacicular hooks 

(Figure 800 with brown cores and clear sheaths, bidentate. 



slender, tapering. Proximal teeth distinctly larger than distal 

teeth, directed laterally. Distal teeth curved. 


anal cirri. 







None. 

REMARKS.—Eunice nigricans was considered a synonym of 

E. macrobranchia by Augener (1925), which lead Hartman 

(1959:313) to refer it to E. aphroditois, based on the synonymy 

of E. macrobranchia with E. aphroditois proposed by 

Mclntosh (1905). The species belongs to group B-2 as do other 

two taxa named; they arc compared to other tax a in the group 

in Table 27. In contrast to E. aphroditois, which lacks 

articulations of the ceratostyles and notopodial cirri, both £. 

macrobranchia and E. nigricans have articulated ceratostyles 

and, in the case of the latter, also articulated noiopodial cirri. 

Among the species listed in Table 29, peristomial cirri well 

outreach the peristomium in E. kinbergi, E. microprion, E. 

nigricans, E. philocorallia, and E. plicata; in the other species, 

including E. macrobranchia, the peristomial cirri are considerably 

shorter. Of the species with long peristomial cirri, E. 

philocorallia and E. microprion have short branchiae; in the 

other species the branchiae are about as long as the notopodial 

cirri or clearly outreach them. In E. nigricans posterior ventral 

cirri are tapering from the bases, in E. kinbergi posterior ventral 

cirri are scoop-shaped, and in E. plicata bases of posterior 

ventral cirri are inflated and welt-shaped. 

136. Eunice northioidea Moore, 1903 

FIGURE 80g-m; TABLES 27,32 

Eunice northioidea Moore, 1903:433-435, pi. 25: figs. 36-38. 


sta 3718, Japan, 1.2 miles off Ose Zaki, Honshu Island, 

(-35°02X 138°47'E) 11 May 1900, 117 m, volcanic sand, 

shells and rock, beam trawl. 



10, 7 mm. Body cylindrical; only slightly dorsoventrally 

flattened posteriorly. 




sulcus deep. Eyes at bases of A-II, reddish. Antennae in a 

horseshoe, with A-I isolated by a gap, similar in thickness. 


Ceratostyles slender and tapering; A-II and A-III with 

cylindrical articulations; A-I with distally drop-shaped and


proximally cylindrical articulations; maximum 9 articulations 


A-III to setiger 6. Peristomium cylindrical, with a distinct 

muscular lower lip. Separation between rings distinct dorsally 

and visible ventrally; anterior ring ~ 3 A of total peristomial 



Maxillary formula 1+1,6+6,8+0,5+8, and 1+1. Mx HI part 

of distal arc with left Mx IV. 



Branchiae from setiger 3 to end of fragment. First 3 and last 45 

pairs single filaments; maximum 3-4 filaments between 

setigers 10-20. Filaments slender, digitiform, about as long as 


Anterior neuropodial lobes symmetrically bluntly pointed; 

median and posterior acicular lobes (Figure 80m) conical; 



cirri basally inflated from about setiger 5 to about setiger 25. 


cirri tapering, conical. Anterior notopodial cirri slender, 

filiform, with 6 to 7 articulations. Notopodial cirri decreasing 

gradually in length, retaining same shape in all setigers; 

articulations lost in first part of branchial region. 


80j,l) tapering, flat. One marginal tooth much longer than other 

teeth; 7 teeth in anterior setigers; -12 teeth in posterior setigers. 

Shafts of compound falcigers (Figure 80i) slightly inflated, 

marginally smooth. Appendages long, tapering, bidentate. 

Proximal teeth very much shorter than distal teeth, triangular, 

directed laterally. Distal teeth distinctly hooked. Guards 

symmetrically bluntly pointed; mucros absent Pseudocompound 

falcigers and compound spinigers absent Aciculae 

paired in anterior parapodia, single in median and posterior 

parapodia, with brown cores and clear sheaths, tapering, 

bluntly pointed, straight; cross-section round. Subacicular 

hooks (Figure 80k) with dark brown cores and clear sheaths, 



Hooks tapering, slightly bent Proximal teeth larger than distal 

teeth, triangular, directed laterally. Distal teeth short, nearly 

erect 

UNKNOWN MORPHOLOGICAL FEATURES.-A11 features associated 

with posterior setigers and pygidium. 





Characters: 1,2,4,6, 36-40,47, 50, 63. 


38,1. 

REMARKS.—Eunice northioidea is listed with similar species 

in Tables 27 and 32. It is first and foremost characterized 

by having long, slender branchial filaments and long, strongly 

articulated antennae. All species in Table 32, with the 

exception of E. northioidea, have articulated peristomial cirri; 

these cirri are short and completely without articulations in E. 

northioidea. 

137. Eunice norvegica (Linnaeus, 1767) 


Nereis norvegica Linnaeus, 1767:3116.—Gunnerus, 1768, pl.2: fig.7. 

Nereis madreporae pertusae Gunnerus, 1768:45-51, pi. 2: fig. 11. 

Leodice norwegica Lamarck, 1818:323 [in part].—Orsted, 1845:402, 406 [in 

part]. 

Leodice gunneri Storm, 1881:92-95. 

Eunice norwegica.—Cuvier, 1817:525 [ in part].—Audouin and Milne 

Edwards, 1833:219 [in part].—Grube. 1850:292. 

MATERIAL EXAMINED.—Neotype and 1 additional specimen, 

KNVSM, No. 63, ROberg, Statsbygd, Trondheimsfjorden, 

Norway, 2 specimens. 

DESCRIPTION.—Both specimens complete; one with 115 

setigers of which last 15 in regeneration. Neotype with 157 

setigers with last 20 discolored, possibly regenerating; total 


10,13 mm. 




Eyes hidden below overhanging peristomial fold, located 

behind bases of A-II, black. Antennae in a horseshoe, evenly 


antennae, without articulations. Ceratostyles digitiform, with 

up to 16 distinct cylindrical articulations in A-III. A-I to middle 

of peristomium; A-I to setiger 3; A-III to setiger 4. Peristomium 

massive, muscular, cylindrical; frontal margin folded into 

series of short scallops. Separation between rings distinct 

dorsally and ventrally; anterior ring 4 /s of total peristomial 

length. Peristomial cirri to front edge of prostomium, slender, 

with one long and 7 short, bead-like articulations. 

Maxillary formula of neotype 1+1, 7+7, 9+0, 4+9, 1+1, and 

1+1. Mx III long; however, forming a distal arc with very short 

left Mx IV. 

Branchiae (Figure 81e) present, pectinate, about as long as 




of setigers. First and last 3 branchiae single filaments, all others 

pectinate with up to 7 branchial filaments. Filaments short, 

digitiform. 

Anterior neuropodia truncate, becoming rounded in posterior 

setigers; aciculae emerging at midline. All pre- and postsetal 

lobes low, transverse folds. First 6 ventral cirri thick, tapering. 

Ventral cirri basally inflated from about setiger 7 through 

setiger 80. Inflated bases ovate; narrow tips digitiform. Inflated 

bases gradually decreasing posteriorly; last 50 ventral cirri

242 


FIGURE 81.—Eunice norvegica (neotype, KNVSM No. 63): a, anterior end, lateral view; b, compound falciger, 

parapodium 54; c, subacicular hook, parapodium 54; d, pectinate seta, parapodium 54; e, parapodium 54, anterior 

view. Eunice ornata (syntype, USNM 4874): f, pectinate seta, parapodium 15; g, compound falciger, parapodium 

15; h, parapodium 15, anterior view; i, anterior end, lateral view; j, parapodium 40, anterior view; k, aciculae, 

parapodium 40; 1, compound falciger, parapodium 40; m, acicula, parapodium 40; n, pectinate seta, parapodium 

45; o, subacicularhook, parapodium 45. (Scale bars in mm.)


digitiform. Anterior notopodial cirri long, tapering, with 3 

cylindrical articulations. Notopodial cirri retaining same size 

and shape throughout, being especially prominent in posterior 

setigers; articulations lost by beginning of branchial region. 


(Figure 8Id) tapering, flat Both marginal teeth slightly longer 


falcigers (Figure 81b) tapering, marginally smooth. Appendages 

short, tapering to very large heads, bidentate. Proximal 

teeth larger than distal teeth, tapering, directed laterally. Distal 

teeth curved. Guards distally irregularly, symmetrically blunt; 


spinigers absent. Aciculae amber or chestnut colored, tapering, 

straight; cross-section round. Subacicular hooks (Figure 81c) 

from dark amber or chestnut colored, bidentate. Hooks first 

present from setigcr 42 (44 in other specimen), present in all 

setigers thereafter, sometimes paired. Hooks strongly tapering 

to small heads. Proximal teeth very much larger than distal 

teeth. Both teeth directed distally. 


anal cirri. 





Characters: 4,6,42,74, 78. 


None. 

REMARKS.—No type material is available for this species. 

Linnaeus accepted the species as new in a letter to Bishop 

Gunnerus in Trondheim dated 1 December 1766 (Gunnerus, 

1768:51, footnote) and published a brief description next year 

(Linnaeus, 1767). It is not clear from Gunnerus' remarks or 

from the original description if Linnaeus saw any specimens or 

just reacted to Gunnerus' description and appended illustrations, 

which are excellent Incidentally, the species is called 

Nereis norvegica on Gunnerus (1768, pi. 2: fig. 7) even if he 

used a different name in his text. 

Gunnerus (1768) is a description of deep-water corals of 

Norway and their faunal associates; he mentioned several 

localities including Statsbygd (Gunnerus, 1768:38). He specifically 

stated that he had seen fresh material of corals from 

Statsbygd and that he had found his Nereis madreporae 

pertusae on fresh coral material (pp. 44-45). 

The reef at ROberg, which is in Statsbygd, is here considered 

the type locality for the species and one of two specimens listed 

above has been designated as neotype to define and delimit a 

species that has been reported from a variety of localities 

worldwide. 

For most of last century two species, both present on 

ahermatypic reefs, were confounded under the name E. 

norvegica. This synonymy led to the description of Leodice 

gunneri by Storm; this problem is discussed in some detail 

above. 

Eunice norvegica belongs to group B-2 as the groups were 

defined by Hartman (1944) and Fauchald (1970), but aciculae 

and subacicular hooks are relatively light-colored compared to 

the dark-brown to black aciculae and hooks present in most 

species in the group. It is listed with similar species in Tables 

27 and 30. It is the only species in Table 30 to combine short 

branchiae with inflated bases of ventral cirri limited to median 

setigers. 

138. Eunice notata (Treadwell, 1921) 

TABLES 33,37-39 

Uodice notata Treadwell, 1921:52-54, figs. 164-173, pi. 4: fig.12. 

Eunice afra.—Hartman 1956:282 [in part, not Peters, 1854]. 


has been lost. The description by Treadwell, although 

somewhat incomplete, is sufficient to characterize the species. 

DESCRIPTION.—Complete specimen with 95 setigers; total 

length 40 mm; width about 1 mm. 

No antennae outreaching peristomium, without articulations. 

Peristomial cirri short, without articulations. 

Maxillary formula 1+1, 4+4, 5+0, 3+6, and 1+1; Mx III 

illustrated as short; part of distal arc with left Mx IV. 

Branchiae present, not reduced in mid-body region. Branchiae 

from setiger 14 to setiger 70. Branchiae terminating well 


of setigers. Small specimens with single, strap-like filaments 

only; larger specimens with up to 3 filaments. 

Ventral cirri basally inflated from about setiger 10; distribution 

of inflated bases otherwise unknown. Notopodial cirri 

short, tapering, supported by internal aciculae in all setigers. 


Limbate setae marginally smooth. Pectinate setae tapering, 

furled. Both marginal teeth longer than other teeth; -20 teeth 

present. Shafts of compound falcigers inflated, marginally 

serrated. Appendages with nearly parallel sides, bidentate. 


slightly basally. Distal teeth distinctly curved. Guards symmetrically 

bluntly pointed, marginally serrated; mucros absent. 

Pseudocompound falcigers and compound spinigers absent 

Aciculae tapering, distally slightly bent. Aciculae and subacicular 

hooks with light- to dark-brown cores and clear sheaths. 

Subacicular hooks bidentate. Both teeth directed distally; 

proximal teeth somewhat larger than distal teeth. 






SCORED.—Inappropriate Characters: 22, 34, 39, 40 Unknown 

Characters: 4-19, 23-26, 28, 32, 42-50, 52, 54-60, 

80-82. 


None.


REMARKS.—The species was described from Tobago in the 

Windward Islands and is listed with similar species in Tables 

33 and 37-39. It was well characterized by Treadwell; 

however, without access to type material or new material from 

the type locality, it is difficult to relate it to any of many species 

assigned to group B-4. It was omitted from the key. 

Eunice (Marphysa) novaehollandiae (Kinberg, 1865) 

Nauphanta novae Hollandiae Kinberg, 1865:564; 1910:43, pi. 16: figs. 23, 

23B-23G.—Augencr, 1922d:26.—Fauchald, 1987:378-379, fig. 2. 

Eunice (Marphysa) novae Hollandiae.—Grube, 1878b: 165-166. 

REMARKS.—This species has usually been considered a 

synonym of Nauphanta mossambica. It was redescribed by 

Fauchald (1987) as type species of the genus Nauphanta. 

Eunice oerstedii Stimpson, 1854 

Eunice oerstedii Stimpson, 1854:34-35. 

Eunice oerstedii.—Mclntosh, 1885:273-275, pi. 38: figs. 1, 2, pi. 19A: figs. 

14, 15. 

Eunice oerstedii.—Fauvel, 1914:143-145, pi. 10: figs.5-10. 

REMARKS.—No original material is available. The only 

information of any value in Stimpson's description concerns 

the distribution of branchiae, which should be present from 

setiger 4 and be absent posterior to setiger40 with maximum of 

5 filaments. Mclntosh (1885) and Fauvel (1914) discussed two 

very different species under this name. An examination of 

Mclntosh's material shows that his species belongs to group 

A-l, possibly to an undescribed species. Fauvel's species 

belongs to group B-l. The only feature the two species have in 

common is the branchial distribution, which approximates the 

one specified by Stimpson. This kind of branchial distribution 

is found in many different species (e.g., E. pennata and E. 

vittata) and cannot be used by itself to identify any taxon. The 

fauna of the type area in the northwest Atlantic is known to 

include several species. 

Eunice oerstedii is here considered indeterminable, not least 

because two different traditions have developed in identifying 

the species. 

139. Eunice oliga (Chamberlin, 1919) 

TABLES 46,47 

Leodice oliga Chamberlin, 1919a:244-248, pi. 55: fig.11, pi. 56: figs. 2-9. 

COMMENTS ON MATERIAL EXAMINED.—The types are no 

longer available. The following is a summary of Chamberlin's 

description. 

DESCRIPTION.—Type with 82 segments; length 16.5 mm; 

maximal width 2 mm. 


thickened rim; median sulcus deep. Eyes not mentioned. 


Ceratostyles tapering, with up to 30 short, nearly moniliform 

articulations in A-III. A-I to posterior peristomial ring; A-II to 

setiger 7; A-III to setiger 14. Peristomium cylindrical. 

Separation between rings distinct dorsally and ventrally. 

Peristomial cirri to middle of prostomium, slender and 

tapering, with 8 articulations. 

Maxillary formula 1+1, 8+8, 9+0, +10, and presumably 

1+1. 

Branchiae present, palmate, reduced in mid-body region. 



of setigcrs. First and last branchiae single filaments; maximum 

3 filaments at about setiger 12-16. Filaments slender, tapering. 

Branchial stems short. Several median setigers with single 

filaments; numbers increasing to 2 in a short pre-pygidial 

region. 

Neuropodial acicular lobes truncate. Anterior and posterior 

ventral cirri tapering; median ventral cirri basally inflated. 

Notopodial cirri tapering, with 5 to 6 articulations. 

Pectinate setae tapering, flat. One marginal tooth very much 


falcigers inflated, marginally smooth; internal striation present. 

Appendages thick, tapering, with large heads, bidentate. 

Proximal teeth shorter than distal teeth, tapering, directed 

laterally. Distal teeth curved, tapering, directed laterally. 



yellow, tapering to pointed tips; cross-section round. 

Separation of cores and sheaths indistinct in both aciculae and 


teeth in a crest. Hooks tapering, with distinct necks. Proximal 

fang large, directed laterally. Secondary and tertiary fangs 

emerging from joint bases, curved; tertiary fangs very small. 


associated with prostomium; parapodia and setal structures; 






Characters: 4-6, 10-17, 24, 26, 32, 44-52, 54-56, 60, 

80-82. 


None. 

REMARKS.—Eunice oliga was described from a single 

specimen from Makemo Island, Paumotou Islands, in shallow 

water. It is listed with similar species in Tables 46 and 47. 

Eunice oliga is one of two species listed in Table 47 with 

palmate branchiae (i.e., with reduced branchial stem); the other 

species is E. bicirrata. The latter is poorly known; the two 

species may be best compared by matching the descriptions 

directly.


Eunice oliga papeetensis (Chamberlin, 1919) 

leodice oliga papeetensis Chamberlin, 1919a:248-249 

REMARKS.—Originally described as subspecies, this form is 

here considered a distinct species and described as such below. 

140. Eunice ornata Andrews, 1891 

FIGURE 81f-o; TABLES 12, 46,48 

Eunice ornata Andrews, 1891:284-285, pi. 13: figs. 6-13. 

MATERIAL EXAMINED.—Eight syntypes, USNM 4874 and 

4875, Beaufort, North Carolina, 1885. 

COMMENTS ON MATERIAL EXAMINED.—Data for all syntypes 

is summarized in Table 12. Syntype in USNM 4875 is a 

juvenile, apparently of the same species, but is not further 

considered in this description. 

DESCRIPTION.—Syntype described complete with 110 setigers; 

total length 45 mm; maximal width 2 mm wide; length 

through setiger 10, 6 mm. Body cylindrical, slightly dorsoventrally 

flattened posteriorly. Anal cirri slender, as long as last 15 

setigers combined, without articulations. 


peristomium, less than ! /2 as deep as peristomium. Prostomial 

lobes frontally rounded, dorsally strongly inflated; median 

sulcus deep. Palpal regions distinct by frontal grooves. Eyes 

between bases of A-I and A-II, indistinct. Antennae in a 



slender and tapering, with up to 15 moniliform to drop-shaped 

articulations in A-III. A-I to posterior peristomial ring; A-II to 

setiger 1; A-III to setiger 3. A-III always longer than A-II. 

Peristomium cylindrical, with distinct muscular lower lip. 

Separation between rings distinct dorsally and ventrally, 

indistinct only over a very short lateral distance; anterior ring 

3 A of total peristomial length. Peristomial cirri to posterior edge 

of prostomium or front edge of peristomium, slender and 

tapering, with up to 7 long, cylindrical articulations; most 

specimens with 4 to 5 articulations. 

Summary maxillary formula 1+1, 8+8-9, 7-9+0, 6+9-10, 

and 1+1. Mx III long, located behind left Mx II. 


than notopodial cirri, reduced in mid-body region in some 

syntypes, erect. Branchiae from setiger 5 to setiger 110. 


65% of total number of setigers. Only first branchiae single 

filaments; all other branchiae with 3 or more filaments, up to 20 

filaments present. Branchial stems erect, tapering. Filaments 

about as long as notopodial cirri in anterior and median 

setigers, slender and digitiform. In second half of body number 

of filaments reduced to 3 (Figure 81j); filaments decreasing in 

length so in last 

l h of body notopodial cirri longer than 

branchiae. Most specimens with no increase in number of 

filaments towards posterior end; in 1 specimen an increase to 4 

filaments in posterior end was noted 

Anterior neuropodial lobes asymmetrically conical with 

aciculac emerging on dorsal side; farther posteriorly acicular 

lobes become flattened and symmetrically rounded; aciculac 

emerging at midline. All presetal lobes obliquely transverse 

folds. Anterior postsetal lobes forming collars, about as high as 

acicular lobes; by setiger 30 postsetal lobes reduced to low, 

transverse folds. First 9 ventral cirri thick and tapering. Bases 

of ventral cirri inflated, with nearly spherical glandular 

structure from about setiger 10 to setiger 35-40; narrow tips 

digitiform. Inflated bases reduced over next 5-10 setigers. Far 

posterior ventral cirri slender and digitiform, very nearly as 

long as notopodial cirri. Anterior notopodial cirri tapering, with 

3 to 5 articulations. Articulations lost in first branchial setigers; 

all other notopodial cirri slender, tapering, increasingly more 

prominent as branchiae become reduced. 

Limbate setae marginally frayed. Anterior pectinate setae 

(Figure 8 If) tapering, furled. Both marginal teeth of about same 

length; 10 teeth present. Median and posterior pectinate setae 

(Figure 8In) slightly flaring, flat. One marginal tooth distinctly 


falcigers (Figure 81g,l) slightly inflated, marginally serrated. 

Anterior appendages tapering, bidentate. Proximal teeth shorter 

than distal teeth, slender, directed laterally. Distal teeth slender, 

directed obliquely distally. Posterior appendages bidentate with 

proximal teeth longer than distal teeth, triangular, directed 

laterally. Distal teeth tapering, bent. Guards asymmetrically 

bluntly pointed, marginally serrated; mucros absent. Pseudocompound 


yellow; anterior aciculae tapering to blunt cones, bent; 

cross-section round. Median and posterior aciculae (Figure 

81k) flattened in anterior-posterior axis with distal end 

distinctly bidentate, bent dorsally. Proximal teeth larger than 

distal teeth (Figure 81m), directed laterally. Distal teeth erect. 


and subacicular hooks. Subacicular hooks (Figure 8I0) yellow, 

tridentate with teeth in a crest. Hooks first present from setiger 

22-25, present in all setigers thereafter, always single (except 

for replacements). Hooks with large main fang and 2 distal 

fangs emerging from common base. 


anal cirri. 







None. 

REMARKS.—Eunice ornata is listed with similar species in 

Tables 46 and 48. The structure of median and posterior 

aciculae is unusual as is the shift between pectinate setae with 

even marginal teeth to ones with one long marginal tooth from 

anterior to posterior setigers. It has up to 20 branchial 

and


TABLE 12.—Variable and invariable features in the type lot of Eunice ornata (N a number of individuals 




Total length 

Maximal width 



No. of perisiomial cirral articulations 









Notopodial articulations 





N 

3 

3 

5 

5 

6 

5 

7 

6 

7 

7 

Max. 

131 

83 

4 

8 

15 

9 

20 

46 

5 

25 

N=7 

visible dorsally and vent rally 

prostomium 

5 


yellow 

bent; bifid or bidentate 

yellow 

3 

Min. 

110 

45 

0 

1.7 

5 

4 

9 

20 

4 

14 

Mean 

123.33 

67.00 

2.54 

5.74 

11.33 

7.00 

16.43 

36.83 

4.71 

22.57 

SD 

11.59 

19.70 

1.25 

2.41 

3.33 

2.12 

3.87 

9.09 

0.49 

3.91 

filaments; no other species listed in Table 48 has more than 

eight. 

Andrews (1891:285) stated that this species is "not uncommon 

on sponges, etc., in 2 to 3 fathoms in sheltered sounds. The 

young were also found cast up on the Fort beach, after storms, 

upon seaweeds." 

141. Eunice ovalifera Fauvel, 1936 

TABLES 24,27,31 

Eunice ovalifera Fauvel, 1936:67-69, fig. la-h. 

COMMENTS ON MATERIAL EXAMINED.—The type is not 

available. The following represents a summary of information 

in the original description. 

DESCRIPTION.—Type a posteriorly regenerating specimen; 

length 21.5 mm; width 5 mm with parapodia; regenerating 

portion 1.5 mm long. Number of segments 37 in region 

preceding regenerate. 

Prostomium very short, with clavate antennae; A-III to 

setiger 1, with 4 large, slightly barrel-shaped articulations. 


Branchiae present, pectinate. Branchiae from setiger 5, 

present on last segment in non-regenerating part of body. 

Maximum number of filaments 5-6. 

Ventral cirri apparently inflated in most of branchial region. 

Notopodial cirri clavate, without articulations. 

Pectinate setae illustrated with both marginal teeth longer 

than other teeth, narrow, tapering with -10 teeth. Shafts of 

compound falcigers tapering. Appendages short, with large 

heads, bidentate. Proximal teeth larger than distal teeth. Guards 


falcigers and compound spinigers absent. Aciculae and 

subacicular hooks black. Subacicular hooks bidentate. Hooks 

first present from setiger 35. Proximal teeth larger than distal 

teeth, directed laterally. Hooks with distinct subterminal 

inflated region. 




None. 



1, 2, 4-19, 23-29, 32-34, 36-40, 42-56, 58, 59, 65, 

67,74-76,81,82. 


None. 

REMARKS.—This species belongs to group B, but cannot 

safely be assigned to a subgroup, because the branchial 

distribution in intact animals remains unknown; for that reason 

it has been listed both in Tables 24 and 27 as well as in Table 

31. It is too poorly known to be safely identified from the 

original description. The species was reported from southern 

Japan. 

142. Eunice pacifica Kinberg, 1865 

FIGURE 82a-e; TABLES 33-35 

Eunice pacifica Kinberg, 1865:562 [in part]; 1910:41-42, pi. 15: fig. 15 (in 

part].—Hartman, 1948:75-76 (in part]. 

MATERIAL EXAMINED.—Lectotype and 2 paralectotypes,


h. i 

FIGURE 82.—Eunice pacifica (lectotype, RM 310): a, anterior end, lateral view; b, parapodium 34, anterior view; 

c, pectinate seta, parapodium 34; d, subacicular hook, parapodium 34; e, compound falciger, parapodium 34. 

Eunice panamena (holotype, USNM 19354): f, anterior end, lateral view; g, compound falciger, parapodium 10; 

h, parapodium 10, anterior view; i, parapodium 69, anterior view; j, subacicular hook, parapodium 69; k, pectinate 

seta, parapodium 69; 1, acicula, parapodium 69. (Scale bars in mm.)

248 


RM 310, Eimeo (= Moorea), Society Islands, Eugenie 

Expedition 1159. Kinberg (1865:562) gave the locality 

information as "Insula Tahiti et Eimeo mar pacifici, inter 

corallia fundo pedis." 

COMMENTS ON MATERIAL EXAMINED.—The three syntypes 

and two posterior fragments all have been dried out at one time. 

The smaller, complete specimen was selected as lectotype for 

the species. A fourth specimen present belongs to a different 

species and is described above as E. eimeorum. Two pairs of 

maxillae were isolated in a small vial; the two pairs are very 

similar, and are here assumed to belong to this species, rather 

than to E. eimeorum. 

DESCRIPTION.—Paralectotypes incomplete with 121 and 158 

setigers respectively. Lectotype complete with 102 setigers of 

which last 20 in regeneration; total length 33 mm; maximal 

width 3 mm at setiger 10; length through setiger 10,6 mm. 




thickened rim; median sulcus shallow. Eyes posterior to bases 

of A-I. Antennae in a horseshoe; A-I and A-II close together, 

well separated from A-III; A-III slimmer than other 4. 

Ceratophores long in all antennae, without articulations. 

Ceratostyles tapering and slender, with 4 or 5 indistinct long 

articulations. A-I to middle of anterior peristomial ring; A-II to 

posterior peristomial ring; A-III to setiger 1. Peristomium 

massive, cylindrical. Separation between rings distinct dorsally 

and ventrally; anterior ring 4 /s of total peristomial length. 

Peristomial cirri to middle of anterior peristomial ring, 

medially inflated, without articulations. 

Maxillary formula 1+1, 4+4-5, 6+0, 4+8-10, and 1+1. Mx 

III part of distal arc with left Mx IV. Mx VI absent 


cirri, not reduced in mid-body region, flexible. Branchiae 

from setiger 17 to setiger 102 in lectotype and from setigers 

20-21 in paralectotypes. Branchiae present to near posterior 

end, present on more than 65% of total number of setigers. 

Maximum number of filaments 5 from about setiger 40 and 

continued to posterior end. 


acicular lobes rounded (Figure 82b); posterior acicular lobes 

conical; aciculae emerging at midline. Neuropodia carried on 

lateral ridges from about setiger 60 to posterior end. All preand 

postsetal lobes following outline of acicular lobes closely. 


from about setiger 10, retaining inflated glandular bases at least 

through median setigers, forming ventral ridges in most 

parapodia examined; narrow tips digitiform. Far posterior 

ventral cirri with reduced basal inflation, digitiform. Anterior 

notopodial cirri medially inflated, becoming digitiform farther 



(Figure 82c) tapering or slightly flaring, flat. Both marginal 

teeth slightly longer than other teeth; 12 teeth present Shafts of 

compound falcigcrs (Figure 82e) slightly inflated, marginally 

smooth. Appendages short, rather thick, bidentate. Proximal 

teeth larger than distal teeth, triangular, directed laterally. 

Distal teeth slender, curved, directed obliquely laterally. 

Guards symmetrically rounded, marginally serrated; mucros 


absent. Aciculae paired, brown, tapering, blunt, straight; 

cross-section round. Subacicular hooks (Figure 82d) brown, 

bidentate. Hooks first present from setiger 23 in lectotype and 

from setigers 27-28 in paralectotypes, present in all setigers 


narrowing distally to small, distinct heads. Teeth similar in 

size, blunt, directed distally. 


anal cirri. 





Characters: 4, 6, 39-40, 74, 78. 


None. 

REMARKS.—The presence of two species in the type lot 

forced the selection of one specimen as lectotype. The lectotype 

matches both Kinberg's and Hartman's descriptions. Eunice 

pacifica resembles E. afra and related species and is listed with 

similar species in Tables 33, 34, and 35. Four species in Table 

34 have long ceratophores in all antennae; in addition to E. 

pacifica, this includes E. modesta, E. petersi, and E. sonorae. 

Of these species, E. modesta has exclusively single branchial 

filaments; the other three species have more complex branchiae. 

Eunice sonorae has simple, falcate or tapering subacicular 

hooks; the subacicular hooks are bidentate in E. petersi and 

E. pacifica. In E. pacifica both teeth of the subacicular hooks 

are directed distally, in E. petersi, the proximal teeth are 

directed distinctly laterally. 

Due to the poor condition of the specimens it is difficult to 

ascertain where the inflated bases of the ventral cirri ends. 

Kinberg (1865:562) indicated that he had seen specimens 

from Tahiti as well as from Eimeo; all present specimens are 

from Eimeo. 

143. Eunice palauensis Okuda, 1937 

TABLES 24,26,27,28 

Eunice palauensis Okuda, 1937:285-286, figs. 29, 30. 

COMMENTS ON MATERIAL.—This species was described 

from near Guarakabaru Island, Palau Islands; no type material 

is known to have survived. The information given by Okuda is 

summarized below. 

Okuda had four anterior ends to his disposal, the longest of 

these with 42 setigers for 20 mm length. Posterior ends were 

also present, but none of the specimens were complete.


DESCRIPTION.—Body cylindrical and very slender. 

Prostomium distinctly shorter than peristomium, about as 

wide as peristomium. Prostomial lobes frontally rounded, 

dorsally inflated; median sulcus shallow. Presence of eyes not 

indicated. Antennae in a horseshoe, evenly spaced, similar in 


articulations. Ceratostyles slender and tapering, smooth or 

faintly articulated. A-I to posterior peristomial ring; A-II to 

setiger 7; A-III to setiger 18. Separation between peristomial 

rings apparently distinct on all sides; anterior ring - 3 /4 of total 

peristomial length. Slender, tapering peristomial cirri to front 

edge of peristomium or possibly to posterior end of prostomium, 


Jaw apparatus symmetrical according to Okuda. His illustration 

(fig. 30g) indicates a maxillary formula of 1+1, 7+8, 8+0, 

7+9, 1+4, and 1+1 (Okuda's formula: 1+1,6+6-7, 10+8-9). 


cirri. Branchiae from setiger 6-8. Branchiae terminating well 

before posterior end. First 5-9 branchiae single filaments; 

maximum number of filaments 3; last 20 segments abranchiate. 

Acicular lobes symmetrically conical anteriorly, becoming 

rounded in median setigers; aciculae emerging at midline. 

Pre-and postsetal lobes low folds or follow outline of the 

acicular lobes closely. Ventral cirri moderately inflated in 

median setigers. Inflated bases ovate. Posterior ventral cirri 

without basal inflations. Notopodial cirri thick and tapering in 

anterior setigers; articulations absent. 

Limbate setae smooth and rather thick. Pectinate setae 

flaring, flat with 1 marginal tooth longer than other teeth; total 

number of teeth 11-13. Shafts of compound falcigers inflated 

and marginally smooth. Appendages short, tapering with a 

small head, bidentate. Distal tooth nearly erect; proximal tooth 

illustrated as triangular, short and wide-based. Guard symmetrically 


compound spinigers absent. Aciculae paired, dark, pointed; 

cross-section round. Subacicular hooks yellowish brown, 

bidentate. Hooks first present from setigers 19-22. Hooks with 

small heads. Proximal tooth directed laterally; distal tooth 

erect. 




None. 



Characters: 1-6, 12-14, 24, 33, 36, 38, 40, 42, 51, 

52, 55, 59, 60, 63, 74, 78, 81, 82. 


REMARKS.—Eunice palauensis was considered a member of 

group B-2 by Fauchald (1970); it is compared to other species 

in that group in Tables 27 and 28. The possible branchial 

distribution patterns and color of aciculae and subacicular 

hooks could make this species a member of group B-l as well, 

so the species has been included also in Tables 24 and 26. The 

problem cannot be resolved without access to well-preserved, 

complete specimens from the type area. 

The species is too poorly known to be included in the key. 

Some comments can be made on its relations to other species 

listed in the two tables. Of species in Table 24 with less than 

five (but more than single) branchial filaments, three have, at 

least in part, non-articulated ceratostyles. These include, in 

addition to E. palauensis, E. barvicensis and E. mutilatoides. 

Eunice barvicensis has a distinct branchial stem and thus 

pectinate branchiae; the other two have reduced branchial 

stems and palmate branchiae. Separation between E. mutilatoides 

and E. palauensis is difficult based on current descriptions 

and material, but the two species differ minimally in the shape 

and relative size of the appendages of the compound hooks. 

These are barely tapering, with large proximal teeth in the 

former and strongly tapering, with small heads according to 

Okuda (1937, fig. 30g). 

Because the total number of setigers was not mentioned, it is 

unclear what significance the absence of branchiae on 20 

posterior setigers might be. Of the species listed in Table 28 

only four have five or fewer branchial filaments. Of these E. 

polybranchia has peristomial cirri reaching the middle of the 

prostomium; the other three species have peristomial cirri 

reaching at most the middle of the peristomium. Of the 

remaining species, E. unidentata has short antennae, barely 

reaching beyond the prostomium; E. palauensis and E. rullieri 

have A-I 11 distinctly longer than A-II. The notopodial cirri are 

slender in E. palauensis and medially strongly inflated in E. 

rullieri. 

Eunice (Eriphyle) paloloides Moore, 1909 

Eunice (Eriphyle) paloloides Moore, 1909:235-295, ph. 7-9. 



the genus Palola; it was so considered previously by Hartman 

(1938 and 1944). 

144. Eunice panamena (Chamberlin, 1919) 

FIGURE 82f-l; TABLES 46,48 

Leodice panamena Chamberlin, 1919a:256-260, pi. 59: figs. 4-8, pi. 60: figs. 

1-5. 

MATERIAL EXAMINED.—Holotype, USNM 19354, near 

Panama (Pacific Ocean), shore, Albatross, 1891. 



10, 6 mm. Color dark gray with black patches scattered over 

anterior end. 



lobes frontally rounded, slightly dorsally flattened dorsally, 

sloping away from midline; median sulcus deep. Eyes at bases


of A-I. Antennae in a horseshoe, with A-III isolated by a gap, 

with A-III slimmer than other 4. Ceratophores ring-shaped in 

all antennae, without articulations. Ceratostyles medially 

inflated; articulations nearly moniliform; maximum 16 articulations 

in A-II and A-III. A-I to middle of anterior peristomial 

ring; A-II and A-III to setiger 1. Peristomium cylindrical. 


anterior ring ~ 4 /5 of total peristomial length. Peristomial cirri to 

front edge of peristomium, slender and tapering, with 5 


Maxillary formula 1+1, 5+6, 8+0,4+7, and 1+1. 



Branchiae from setiger 5 to end of fragment. First branchia 

single filaments; all others pectinate; maximum number of 

filaments 8 at about setiger 15. Number of filaments decreasing 

to 3 by setiger 35; this number maintained to last segment 

present (Figure 82i). 

All neuropodial acicular lobes distally transverse aciculae 


folds. First 3 ventral cirri thick, tapering. From setiger 4 ventral 

cirri basally inflated. Inflated bases very large, bag-like, 

glandular; narrow tips first digitiform, later short and buttonshaped. 

Notopodial cirri basally inflated, tapering to blunt tips, 

with 2 to 4 distinct, cylindrical articulations. 

Limbate setae with narrow limbation, marginally serrated. 

Pectinate setae (Figure 82k) tapering, flat. Marginal teeth no 

larger than other teeth, -12 teeth present. Shafts of compound 

falcigers (Figure 82g) distally inflated, marginally serrated. 

Appendages short; head small, bidentate. Proximal teeth very 

much smaller than distal teeth, forming a slight triangular 

elevation on side of appendage. Distal teeth tapering, nearly 

erect. Guards asymmetrically bluntly pointed, marginally 

distinctly serrated; mucros absent. Pseudocompound falcigers 

and compound spinigers absent. Most aciculae broken; those 

present yellow, bluntly and weakly hammer-headed (Figure 

821); cross-section round. Separation of cores and sheaths 


hooks (Figure 82j) yellow, tridentate with teeth in a cresL 



teeth decreasing evenly in size from large proximal fang. 


Mx III and left Mx IV; various setal features; pygidium 






1, 2,4, 6, 36-38,40,47, 50, 57-60, 63. 


38,1. 

REMARKS.—Eunice panamena was considered a member of 

group A-5 on the strength of Chamberlin's (1919a:260) 

specific statement that the subacicular hooks should be 

bidentate; they are however tridentate. The species is listed in 

Tables 46 and 48. It has A-II and A-III similar in length as does 

E. stigmatura also listed in Table 48. However, the separation 

between the anterior and posterior peristomial rings is distinct 

dorsally and ventrally only in E. panamena: the separation is 

distinct also laterally in E. stigmatura. 

145. Eunice papeetensis (Chamberlin, 1919) 


Uodice oliga papeetensis Chamberlin. 1919a:248-249. pi. 55: Tigs. 8-10, pi. 

56: fig. 1. 

MATERIAL EXAMINED.—Holotype, USNM 19397, Papeete, 

Tahiti, intertidal, 9 Nov 1899, Albatross. 


length 8.5 mm; maximal width 2 mm; length through setiger 

10, 2.5 mm. 




thickened rim; median sulcus shallow. Eyes behind bases of 

A-I, dark. Antennae in a horseshoe, with A-III isolated by a 

gap, similar in thickness. Ccratophores ring-shaped in all 


to 20 moniliform articulations in A-III. A-I to posterior edge of 


cylindrical. Separation between rings distinct ventrally and 

dorsally. Anterior ring 5 /6 of total peristomial length. Peristomial 

cirri to frontal edge of peristomium, slender, with 6 


Maxillary formula 1+1,6+7,8+0,7+10, and 1+1. Mx III part 


Branchiae present, palmate, not reduced in mid-body region. 

Branchiae from setiger 6 to setiger 30 on one side and 33 on 

other side. Branchiae terminating well before posterior end, 


last 5 pairs single filaments; all other branchiae with 2 or 

maximally 3 filaments. 

Anterior neuropodia distally truncate; median acicular lobes 

rounded; aciculae emerging at midline. Pre- and postsetal lobes 



ovate; narrow tips tapering. Notopodial cirri long, basally 

inflated. Anterior notopodial cirri with about 5 articulations, 

with 3 or 4 articulations in posteriormost segments. 

Pectinate setae (Figure 83a) small, flaring, flat. One marginal 

tooth distinctly longer than other, with -10 teeth present. Shafts 

of compound falcigers (Figure 83d) distally inflated, marginally 

serrated. Appendages rather large; head distinct, bidentate. 

Both teeth well developed, directed laterally. Guards asymmetrically 

bluntly pointed; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculae at least


FIGURE 83.—Eunice papeetensis (holotype, USNM 19397): a, pectinate seta, parapodium 25; b, acicula, 

parapodium 25; c, subacicular hook, parapodium 25; d, compound falciger, parapodium 30; e, acicula, 

parapodium 30; f, anterior end, lateral view. Eunice parasegregata (holotype, ZMH P-14273): g, anterior end, 

lateral view; h, compound falciger, first parapodium of posterior fragment; i, pectinate seta, first parapodium of 

posterior fragment; j, subacicular hook, first parapodium of posterior fragment; k, compound falciger, 

parapodium 30; 1, pectinate seta, parapodium 30; m, first parapodium of posterior fragment, anterior view; n, 

parapodium 30, anterior view. (Scale bars in mm.)


paired, yellow, distally truncate or gently hammer-headed 

(Figure 83b,e); cross-section round. Separation of cores and 

sheaths indistinct in both aciculae and subacicular hooks. 

Subacicular hooks (Figure 83c) yellow, tridentate with teeth in 

a crest. Hooks first present from setiger 21. Hooks with 

well-developed, distinct fangs. 







Characters: 1, 2,4,6, 32,42, 47, 50, 57-60, 63, 81, 82. 


None. 

REMARKS.—Eunice papeetensis was described as a subspecies 

of E. oliga; it differs at the species level from that species 

in that it has branchiae limited to a short anterior region; £. 

oliga has branchiae continued to near the posterior end. 

Eunice papeetensis is listed with similar species in Tables 41 

and 44. It has maximally three branchial filaments, whereas all 

other species in Table 44 have at least eight filaments where the 

branchiae are best developed. 

146. Eunice parasegregata Hartmann-Schroder, 1965 

FIGURE 83g-n; TABLES 19,21-23 

Eunice parasegregata Hartmann-Schroder, 1965b:168-172, figs. 140-144. 

MATERIAL EXAMINED.—Holotype, ZMH P-14273, off 

Punta Lavapie, Chile, 37°08.7'S, 73°38.6'W, 58 m, fine sand 

with rocks and dead algae, dredge and grab, 10 Mar 1960, coll. 

Exp. Mar. Chile I. 


currently in three pieces: an anterior end of 33 setiger, a median 

piece of 10 branchiated setigers, and a mid-posterior abranchiate 

piece of 22 setiger. A number of segments are missing 

between both the anterior end and the short median fragment 

and between the latter and the medioposterior fragment, but 

exactly how many segments are missing cannot currently be 

estimated. 

DESCRIPTION.—Anterior end with 33 setigers; length 22 





shallow. Eyes under peristomial fold, posterior to bases of A-I, 

brown. Antennae in a horseshoe, evenly spaced, similar in 


articulations. Ceratostyles slender and tapering; A-I with 3-4 

drop-shaped articulations; A-I I and A-I 11 with 5 to 6 irregular, 

cylindrical articulations; all articulations on distal half of 

ceratostyles. A-I to posterior peristomial ring; A-II to setiger 5; 

A-I 11 to setiger 6. Peristomium cylindrical. Separation between 

distinct on all sides; anterior ring twice as long as posterior 

ring. Peristomial cirri to middle of prostomium, slender and 


Maxillary formula 1+1, 14+16, 13+0, 10+16, and 1+1. Mx 

III long, located behind left Mx II. Mx VI missing. 

Branchiae (Figure 83n) present, pectinate, distinctly longer 


Branchiae from setiger 3- to end of fragment, present on all 

setigers of median fragment, missing on medioposterior 

fragment. Branchiae terminating well before posterior end. All 

branchiae with 2 or more filaments; maximum 18 filaments at 

setiger 10; branchiae on median fragment with 2 to 3 filaments; 

those of last segments in anterior end with at least 7 filaments. 

All filaments slender, tapering, as long as notopodial cirri 

except in first branchial segment. 

Anterior ncuropodial acicular lobes distally obliquely truncate, 

becoming distinctly truncate in last segments present; 

aciculae emerging dorsal to midline. In medioposterior 

fragment acicular lobes reduced, equally supported by aciculae 

and subacicular hooks (Figure 83m). Anterior pre- and 

postsetal lobes low, transverse folds. Posterior pre- and 

postsetal lobes follow outline of acicular lobes closely. First 5 

ventral cirri thick, tapering, becoming distinctly basally 

inflated in branchiated setigers. Inflated bases nearly spherical; 

narrow tips digitiform. Postbranchial ventral cirri slender and 

nearly digitiform. All notopodial cirri basally slightly inflated, 

tapering to slender tips, without articulations. 


(Figure 83i,l) tapering, flat One marginal tooth much longer 

and heavier than other teeth, -10 teeth present. Shafts of 

compound falcigers (Figure 83hjc) distally inflated, marginally 

serrated. Appendages triangular, bidentate. Proximal teeth 

smaller than distal teeth, triangular, directed laterally. Distal 

teeth gently curved. Guards distally asymmetrically bluntly 

pointed; mucros absent Pseudocompound falcigers and compound 

spinigers absent. Aciculae yellow, distally tapering, 

gently curved; cross-section round. Separation between cores 

and sheaths indistinct in both aciculae and subacicular hooks. 

Subacicular hooks (Figure 83j) yellow, bidentate. Hooks 

present on median and medioposterior fragments, present in all 

setigers, always single (except for replacements). Hooks 

tapering. Proximal teeth larger than distal teeth, directed 

laterally. Distal teeth very short, nearly erect. 

UNKNOWN MORPHOLOGICAL FEATURES.—Parapodial and 

setal features of far posterior setigers; pygidium and anal cirri. 





Characters: 1, 2,4,6, 36, 38, 40, 80. 


REMARKS.—Eunice parasegregata may belong to either 

group A-l or A-2 and has been listed in Tables 19 and 22 as 

well as in Tables 21 and 23. Because of the fragmented state of


the type, there is little in the numerically arranged features that 

appears unique and the setal structures are similar to what is 

present in several species. Six species in Table 21 have 

articulated ceratostyles; of these only E. parasegregata and E. 

tridentata have moniliform or drop-shaped articulations distally 

in the ceratostyles; the other four species have cylindrical 

articulations to the tips of the antennae. In E. parasegregata the 

appendages of the compound falcigers are tapering, not 

particularly small in relation to the shafts of the falcigers; in E. 

tridentata the appendages are very small, slender, and barely 

tapered. Eunice parasegregata is easily separated from other 

species in Table 23, because it has as many as 18 branchial 

filaments; of other species in this table, E. johnsoni comes the 

closest with 10 filaments where the branchiae are best 

developed. 

The dorsal aciculae of setiger 2 were illustrated by 

Hart man n-Schroder as strongly curved; this observation cannot 

be confirmed without damaging the specimen severely, but 

would, if confirmed, represent a wholly unique feature to this 

species. 

The structure of median and posterior parapodia is unusual, 

but the relation between these features and conditions of 

preservation have yet to be elucidated. 

The numbers of maxillary teeth reported here are slightly 

higher than those reported by Hartmann-Schrftder (1965b); the 

basal teeth are very small and easily overlooked. 

Eunice parca Grube, 1878a:99. 

Eunice parca Grube, 1878 

REMARKS.—This species was listed in Grube's tabularly 

formatted paper as having branchiae limited to the anterior and 

median part of the body and distinctly articulated antennae. No 

further description was given, no locality was indicated, and no 

specimens are available in German or Polish collections. The 

species is here considered indeterminable rather than as a 

nomen nudum, because some indication of its features has been 

given by its position in Grube's paper. 

206. Eunice parva Hansen, 1882 

FIGURE 84a-h; TABLES 41,44 

Eunice parva Hansen, 1882:7, pi. 2: figs. 4-7. 

MATERIAL EXAMINED.—Holotype, ZM Leiden, no. 1481, 

Rio de Janeiro, coll. E. v. Beneden. 


now in two pieces and the anterior end has been ventrally 

dissected. 



setiger 10; length through setiger 10, 4 mm. Body anteriorly 

cylindrical; posterior half dorsoventrally flattened, anteriorly 

truncate, posteriorly slowly tapering to the posterior end. 

Pygidium damaged; only short anal cirri present. 


sulcus shallow. Palpal region set off by horizontal frontal 

grooves. Prostomium distinctly narrower than peristomium, 

shorter than peristomium and more than half the depth of the 

peristomium. Eyes behind bases of A-II, faded purple. 


Ceratophores short, ring-shaped. Ceratostyles tapering; maximum 

12 moniliform articulations in A-II I. A-I to posterior edge 

of anterior peristomial ring; A-II to setiger 3; A-III to setiger 5. 

Anterior peristomial ring tapering anteriorly; posterior peristomial 

ring cylindrical. Nuchal fold with lateral notches; lower lip 

with medial notch. Separation between rings distinct dorsally 

and ventrally; anterior ring 2 /3 of total peristomial length. 

Peristomial cirri to posterior edge of prostomium, slender, 

tapering, with 2 distinct articulations. 

Jaws missing; maxillary formula 1+1 4+7, 10+0, 5+7, and 

1+1 according to Hansen (1882, pi. 2: fig. 6). Mx III long, 

straight, located behind left Mx II. Left Mx IV short, curved. 

Mx VI missing. 

Branchiae from setiger 5 to setiger 20; first 2 and last 2 pairs 

single filaments; maximum 4 filaments at setiger 10. Branchiae 

longer than notopodial cirri. Stems slightly shorter than 

notopodial cirri, cylindrical, erect. Filaments slightly shorter 

than notopodial cirri, digitiform, slightly thinner than notopodial 

cirri. 

Anterior neuropodial acicular lobes asymmetrically truncate; 

median and posterior acicular lobes asymmetrically rounded; 

all with aciculae emerging from upper edge. Pre- and postsetal 

lobes low, transverse folds. First 4 ventral cirri tapering, 

thereafter basally inflated; inflated bases large, spherical. Free 

tips tapering; ~ l /i as wide as the inflated bases. Inflated bases 

decreasing from about setiger 20; totally absent from about 

setiger 30. Posterior ventral cirri longer than those in anterior 

setigers, digitiform or slightly tapering, distinctly shorter than 

notopodial cirri. Anterior notopodial cirri tapering with 4 to 5 

distinct articulations; articulations lost in early branchial 

setigers; notopodial cirri gradually elongating in postbranchial 

region, becoming long, digitiform in far posterior setigers. 

Limbate setae about as long as compound hooks, thick, 

marginally serrated. Posterior pectinate setae slender, small; 

shafts cylindrical. Blades flat, slightly flaring; marginal teeth 

no longer or thicker than other teeth; -10 teeth present. Shafts 

of anterior compound hooks distinctly inflated, without distinct 

beaks, marginal serrations or internal striations. Appendages 

short, tapering with distinct heads. Proximal teeth slightly 

shorter than distal teeth, narrowly tapering, directed laterally. 

Distal teeth tapering, slender, directed obliquely distally, gently 

curved. Hoods asymmetrically bluntly pointed, without 

mucros, marginally serrated. Shafts of posterior compound 

hooks distinctly inflated, without distinct distal beaks; marginal 

serrations and internal striations present. Appendages short, 

tapering, chunky with large heads. Proximal teeth larger than 

distal teeth, broadly triangular, directed laterally. Distal teeth

254 


FIGURE 84.—Eunice parva (holotype, Leiden, no. 1481): a, anterior end, lateral view; b, acicula, parapodium 11; 

c, compound falciger, parapodium 11; d, parapodium 11, anterior view; e, parapodium 40, anterior view; f, 

compound falciger, parapodium 40; g, subacicular hook, parapodium 40; h, pectinate seta, parapodium 40. Eunice 

parvibranchis (syntype, ZMH V-804): i, compound falciger, parapodium SO; j, pectinate seta, parapodium SO; k, 

subacicular hook, parapodium 50; 1, anterior end, lateral view. (Scale bars in mm.)


directed laterally, tapering, thick. Hoods symmetrically 

rounded; mucros and marginal serrations absent. Aciculae 

single, yellow, tapering to blunt tips, anteriorly straight with 

narrow hood, posteriorly gently curved, without hoods. 

Subacicular hooks from setiger 24, single, yellow, tridentate. 

Hooks with distinct bent neck; heads large. Fangs decreasing 

evenly in size, evenly spaced. Proximal fangs triangular, 

parrot-beaked. Secondary fangs slender, nearly straight Tertiary 

fangs triangular, tapering, straight. 

UNKNOWN MORPHOLOGICAL FEATURES.—Anal cirri. 







None. 

REMARKS.—A note in Augener's handwriting in the vial 

with the holotype refers this specimen to Eunice rubra as a 

juvenile. The species by definition belongs to group C-l and is 

compared to similar taxa in Tables 41 and 44; it is the only 

species in Table 44 to have hooded tapering aciculae. The setae 

resemble those of species in group C-2 as well. 

147. Eunice parvibranchis Grube, 1870 

FIGURE 84j-m; TABLES 33, 39 

Eunice parvibranchis Grube, 1870b:55. 

MATERIAL EXAMINED.—TWO syntypes, ZMH V-804, 

Upolu, Fiji Islands; ZMB Q 2266, Fiji Islands, coll. Godeffroy. 

COMMENTS ON MATERIAL EXAMINED.—Both syntypes are 

incomplete and are now very soft. A posterior fragment present 

in same vial as the Hamburg syntype appears to belong to a 

species of Palola in that it lacks subacicular hooks and 

pectinate setae. At one time, the Berlin syntype has been dry 

and is now difficult to measure in detail. The description is 

based on the Hamburg specimen, with notes on the Berlin 

specimen where it differs. As indicated the specimens are very 

soft, and so the structure, especially of the parapodia, cannot be 

determined very accurately. 

DESCRIPTION.—Hamburg syntype incomplete with 133 


setiger 10, 10 mm. Berlin syntype incomplete with -145 

setigers; length about 65 mm. 


peristomium, less than x /i as deep as peristomium. Prostomial 


shallow. Eyes posterior to bases of A-I, faded. Antennae in a 

semicircle with A-I well in front of other antennae, evenly 


antennae, without articulations. Ceratostyles thick and digitiform; 

none outreaching peristomium; A-III longest. Peristomium 

cylindrical, with distinct muscular lower lip. Separation 

between rings distinct on all sides; anterior ring 4 /s of total 



Jaws on left hand side all missing in Hamburg syntype; 

lining of oral cavity intact; left-hand jaws possibly shed or 

never developed. Formula for right hand side 1, 6, 10, and 2 

(Mx III missing). Maxillary formula of Berlin syntype 1+1, 

5+5, 7+0, 2+9, 1+1. Mx III and left Mx IV part of distal arc. 



20 to end of fragment (from setiger 19 in Berlin syntype), most 

branchiae with 2 filaments. Where best developed, around 

setiger 40, 3 filaments present. Filaments longer than notopodial 

cirri except in first 3-4 branchial segments, slender and 

digitiform. 

Neuropodial acicular lobes probably rounded or truncate. All 

presetal lobes low, transverse folds. Anterior postsetal lobes 

possibly as high as acicular lobes, becoming low, transverse 

folds in median and posterior setigers. Anterior ventral cirri 

tapering; basally inflated from about setiger 5-6 at least 

through setiger 50. Inflated bases ovate; narrow tips tapering. 

Posterior ventral cirri tapering. Notopodial cirri short and 



(Figure 84j) distally flaring, slightly furled. Both marginal teeth 

slightly longer than other teeth; -16 teeth present Shafts of 

compound falcigers (Figure 84i) distinctly inflated, marginally 

serrated. Appendages short and tapering, bidentate. Proximal 

teeth about as large as distal teeth, narrowly triangular, directed 

laterally. Distal teeth gently curved. Guards symmetrically 

rounded, marginally serrated; mucros absent. Pseudocompound 


brown, lighter in posterior than in anterior setigers, slightly 

tapered, truncate, straight; cross-section round. Subacicular 

hooks (Figure 84k) brown, bidentate. Hooks from setiger 19 

(from setiger 17 in Berlin syntype), present in all setigers 


abruptly tapering distally. Proximal and distal teeth similar in 

size. Proximal teeth thick, short. Distal teeth about one-half as 

thick as proximal teeth. 


Mx III and left Mx IV; parapodial and setal characters of 

far posterior setigers; pygidium and anal cirri. 





Characters: 1, 2, 4, 6, 36-40, 44,47, 50, 57-60, 63, 74, 78. 


38,1. 

REMARKS.—Eunice parvibranchis belongs to group B-4 and 

closely resembles several other species in the group as 

indicated in Tables 33 and 39. Most species listed in Table 39 

have subacicular hooks starting about 10 segments later than


the first start of the branchiae. Eunice denticulata has the 

reverse relationship, with the branchiae starling about 10 

segments later than the subacicular hooks; in three species, E. 

parvibranchis, E. paupera, and E. spongicola, the branchiae 

and the subacicular hooks start within a few segments of each 

other. Among these three species, E. spongicola has hammerheaded 

aciculae; E. parvibranchis and presumably E. paupera 

have pointed aciculae. The latter is poorly known, but has, as 

described, about half the number of teeth in each of the 

maxillae. 

Eunice parvibranchis appears to differ from most other 

species in the group by having brown, rather than black 

aciculae and subacicular hooks and by the fact that the color of 

the aciculae is lighter towards the posterior than in the anterior 

end, a reverse of the customary condition in the genus. 

Eunice paucibranchis Grube, 1866 

FIGURE 85a-c 

Eunice paucibranchis Grube, 1866a:64. 

Eunice austral is.—Grube, 1870a:294-295 [not Eunice australis Quatrefages, 

1866]. 

MATERIAL EXAMINED.—Holotype, ZMH V-805, Samoa, 

ZMB Q 3434, 3 parapodia in 2 vials, Samoa. 

REMARKS.—The specimen from Hamburg is a posterior end 

of a large specimen. It is completely branchiated with 

maximally five very short branchial filaments, with black 

aciculae and subacicular hooks; the latter are bidentate. 

The three parapodia in ZMB Q 3434 have large branchiae 

(Figure 85a) with up to 10 branchial filaments arranged on a 

stiff branchial stem. Ventral cirri are basally inflated with thick, 

tapering tips. Notopodial cirri are short and tapering. Limbatc 

setae are marginally smooth. Pectinate setae (Figure 85c) taper 

and are flat. One marginal tooth is longer than other teeth and 

about 15 teeth are present. Shafts of the compound falcigers 

(Figure 85b) are slightly inflated and lack marginal serrations. 

The appendages are triangular. Proximal teeth are short and 

triangular. The distal teeth are nearly erect. Guards are distally 

asymmetrically bluntly pointed. Aciculae are light brown or 

chestnut colored and taper distally to a blunt, straight tip. 

Subacicular hooks are not present in any of the parapodia. 

The parapodia from Berlin do not belong to the same species 

as the posterior end from Hamburg judging from the color of 

the aciculae. Neither parapodia nor posterior end can belong to 

E. australis as suggested by Grube (1870). Eunice australis has 

yellow, tridentate subacicular hooks (Fauchald, 1986). 

Eunice paucibranchis is here considered indeterminable. 

148. Eunice paupera Grube, 1878 

TABLES 33,39 

Eunice paupera Grube, 1878b:160-161. 

Eunice afra paupera.—Fauvel in Pruvot, 1930:69.—Fauvel, 1930a:537. 

COMMENTS ON MATERIAL.—No type material is available. 

This form has been recorded as a subspecies of E. afra, 

separated from the main form primarily by the reduced 

branchiae (Fauvel, 1930b, 1932; Fauchald, 1970). 

DESCRIPTION.—Type with 127 setigers. 

Ceratostyles digitiform, without articulations, barely longer 

than prostomium. Anterior peristomial ring 4 /s of total 

peristomial length. Peristomial cirri short 

Maxillary formula 1+1,4+3,4+0,2+6, and presumably 1+1. 

Branchiae present, palmate. Branchiae present from setiger 

23; maximum 3 filaments. 

First 3 ventral cirri tapering. Ventral cirri basally inflated in 

mid body, again becoming tapering in second half of body. 

Notopodial cirri long in first 4 setigers, as long as antennae, 

thereafter relatively short through remainder of pa-branchial 

and early branchial setigers, increasing in length in segments 

with bifid and trifld branchiae, decreasing in length in far 

posterior setigers. Notopodial cirri without articulations. 

Compound falcigers with short, bidentate appendages. 

Pscudocompound falcigers and compound spinigcrs absent. 

Aciculae black. Subacicular hooks black, bidentate, first 

present from setiger 23. 


and peristomial features; relationship between Mx III and left 

Mx IV; most parapodial and sctal features; pygidium and anal 

cirri. 


None. 



Characters: 2-19, 24, 25, 28, 29, 32-34, 36-40, 

42-50, 52, 54, 55, 60,65-68, 70, 74-76, 78, 81, 82. 


None. 

REMARKS.—Eunice paupera clearly belongs to group B-4 

and has been rather widely reported (Fauvel, 1930a, 1953). 

Fauvel apparently used the name E. afra paupera as a name for 

all species similar to E. afra with poorly developed branchiae. 

How many of the reports refer to similar taxa (see Table 39) is 

difficult to determine without careful review of material from 

the Philippines. 

Eunice paupera is listed with similar taxa in Tables 33 and 

39; it was discussed above in relation to E. parvibranchis. It is 

too poorly known to be included in the key. 

149. Eunice pauroneurata (Chamberlin, 1919) 

FIGURE 85d-i; TABLES 27,31 

Ltodice pauroneurata Chamberlin, 1919a:249-253, pi. 57: figs. 8-9, pi. 58: 

figs. 1-9, pi. 59: figs. 1-3. 


sta 3401, off Galapagos Islands, 0°59'S, 88°58'3O"W, 722 

m, Globigerina ooze, 28 Mar 1891.


2 

1 

.1 

b, c, », h 

d 

a 

FIGURE 85.—Eunice paucibranchis (slides of type, 21MB Q 3434): a, median parapodium, anterior view; b, 

compound falciger, median parapodium; c, pectinate seta, median parapodium. Eunice pauroneurata (holotype, 

USNM 19756): d, anterior end, lateral view; e, parapodium 90, anterior view; f, compound falciger, parapodium 

90; g, aciculae, parapodium 90; h, pectinate seta, parapodium 90; i, subacicularhook, parapodium 90. (Scale bars 

in mm.)



been deeply dissected frontally; the illustration of the anterior 

end is a partial reconstruction, especially of the outline of the 

lower lip. 

DESCRIPTION.—Holotype incomplete female with 97 setigers; 

length 42 mm; maximal width 5.5 mm; length through 



than peristomium, less than 1 /2 as deep as peristomium. 


sulcus deep. Eyes not observed. Bases of antennae hidden 

below peristomial fold. Antennae in a horseshoe, evenly 

spaced, with A-III considerably heavier than other 4. Ceratophores 


Ceratostyles tapering, A-I and A-II with 3 or 4 long cylindrical 

articulations along distal half. A-I to posterior peristomial ring; 

A-II to setiger 8; A-III incomplete. Peristomium cylindrical. 



middle of anterior peristomial ring, without articulations. 

Maxillary formula 1+1, 7+7, 8+0, +8,1+1. Mx III long and 

located immediately behind left Mx II. Left Mx IV missing. 



Branchiae from setiger 8 to end of fragment. First branchiae 

single filaments; maximum 5 filaments at setiger 15; most 

segments with 3 or 4 filaments; some with 5 even in last 

segments present. Branchial stems short, tapering. Filaments 

digitiform. 

Neuropodial acicular lobes distally rounded or truncate; 



cirri basally inflated from setiger 5. Inflated bases ovate; 

narrow tips digitiform. Inflated bases reduced from about 

setiger 50. By setiger 70-75 ventral cirri short, slender and 

digitiform. All notopodial cirri prominent, basally inflated, 

tapering to blunt tips, without articulations. 

Pectinate setae (Figure 85h) tapering, flat. One marginal 

tooth slightly longer than other teeth; -12 teeth present. Shafts 

of compound falcigers (Figure 85f) tapering, marginally 

smooth. Appendages large, with distinct heads, bidentate. 


laterally. Distal teeth curved. Guards distally rounded, margin 

smooth, mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae (Figure 85g) paired, dark, 

sharply pointed, straight or gently curved; cross-section round. 

Subacicular hooks (Figure 85i) dark, bidentate. Hooks first 


single (except for replacements). Hooks slender, tapering. 

Proximal teeth about twice as large as distal teeth. Both teeth 


UNKNOWN MORPHOLOGICAL FEATURES.—All characters 

associated with far posterior setigers, pygidium, and anal cirri. 





Characters: 1, 2,4,6, 13, 14, 23, 36-38,40, 74, 78. 


38,1. 

REMARKS.—Eunice pauroneurata is listed with similar 

species in Tables 27 and 31. The shape of the appendages of the 

compound hooks resembles the condition in E. collini, but the 

two species can be separated on the structure of the antennae 

and aciculae. Two additional species in Table 31 may have 

short, flexible branchiae. In E. bipapillata the flexibility was 

not noted, so it is here assumed present for the purpose 

of the discussion; in this species, branchiae arc first present 

from setiger 4 with a maximum of seven filaments; in E. 

pulvinopalpata branchiae arc present from sctigcr 3 with a 

maximum of 25 filaments. In E. pauroneurata branchiae are 

first present from scligcr 8 with a maximum of five filaments. 

Chambcrlin (1919a:253) claimed that the unpaired Mx III 

was missing; this is incorrect: Mx III is present, but left Mx IV 

is missing. 

150. Eunice pectinata Grube, 1869 

FIGURE 86«-C; TABLES 19, 20 

Eunice pectinate Gnibe. 1869:492-493. 

MATERIAL EXAMINED.—Holotype, ZMB F 2005, Tor, Red 

Sea, coll. Ehrenberg. 

COMMENTS ON MATERIAL EXAMINED.—A note on the label 

translates as: "according to Grube's original label = E. pennata 

O.F. Muller." 



6 mm. Body cylindrical. 



Prostomial lobes large, frontally rounded; median sulcus deep. 

Eyes posterior to A-I, faded. Antennae in a horseshoe, evenly 



to 12 closely spaced, short, articulations. Articulations cylindrical 

basally, becoming drop-shaped distally. A-I to posterior 


Peristomium cylindrical; lower lip thick, forming ventral 

cushion. Separation between rings distinct on all sides; anterior 

ring ~ 3 /4 of total peristomial length. Peristomial cirri to middle 

of anterior peristomial ring, digitiform, with 6 articulations. 

Maxillary formula 1+1, 5+5, 6+0, 4+8, 1+1. Jaws very soft 

and friable. Mx III form part of distal arc with left Mx IV. 



Branchiae from setiger 5 to end of fragment. First 2 pairs single 

filaments; maximum 10 filaments; even last setigers with 5


RGURE 86.—Eunice pectinata (holotype, ZMB F 2005): a, anterior end, lateral view; b, parapodium 49, anterior 

view; c, subacicular hook, parapodium 49; d, pectinate seta, parapodium 49; e, compound falciger, parapodium 

49. Eunice pelamidis (holotype, MNHN, A.l(R.)-1868-no. 57a): f, subacicular hook, parapodium 110; g, acicula, 

parapodium 110; h, pseudocompound falciger, parapodium 110; i, pseudocompound falciger, parapodium 110; j, 

pectinate seta, parapodium 110; k, compound falciger, parapodium 5; 1, pectinate seta, parapodium 5; m, 

parapodium 5, anterior view.

260 


filaments. Branchial stems long and slender, increasingly 

flexible in posterior setigers. Filaments digitiform, longer than 


Neuropodial acicular lobes distally rounded, becoming 

broadly transverse in last setigers present; aciculae emerging at 

midlinc. Pre- and postsetal lobes low folds following outline of 

acicular lobes closely. First 3 ventral cirri tapering; ventral cirri 

moderately basally inflated in rest of segments. Bases ovate; 

narrow tips digitiform. Notopodial cirri basally inflated, 

articulated. Anterior notopodial cirri with up to 6 closely 

spaced articulations; number of articulations decreasing to 3 in 

last setigers present 

Limbate setae slender, marginally smooth-edged. Pectinate 

setae (Figure 86d) flaring, flat. One marginal tooth slightly 

longer than all other teeth; -15 teeth present. Shafts of 

compound falcigers (Figure 86e) tapering, marginally smooth. 

Appendages slender, bidentate. Proximal teeth shorter than 

distal teeth, tapering, directed distally. Distal teeth distinctly 

thicker than proximal teeth, curved. Guards asymmetrically 

bluntly pointed; mucros absent Pseudocompound falcigers and 

compound spinigers absent. Aciculae yellow, narrow, tapering, 

straight or gently curved; cross-sections round. Separation 

between cores and sheaths indistinct in both aciculae and 

subacicular hooks. Subacicular hooks (Figure 86c) yellow, 



Hooks wide-shafted. Proximal teeth larger than distal teeth, 

directed laterally. Distal teeth thick, truncate. 

UNKNOWN MORPHOLOGICAL FEATURES.—All characters of 






1,2,4,6, 36-38,40,42,47, 50, 57-60,63. 


38,2. 

REMARKS.—Eunice pectinata differs distinctly from E. 

pennata despite Grube's initial impression (see above) and is 

listed with similar species in Tables 19 and 20. Among other 

differences, all notopodial cirri are articulated in E. pectinata 

and are articulated only in anterior setigers in E. pennata. 

Additionally, the guards of the compound falcigers are distally 

blunt in E. pectinata and distally sharply pointed in E. pennata. 

151. Eunice pelamidis Quatrefages, 1866 

FIGURE 86f-m; TABLES 52,53 

Eunice pelamidis Quatrefages, 1866:322-323. 


1868-no. 57a, Payta, Peru, coll. Gaudichaud. 

COMMENTS ON MATERIAL EXAMINED.—The holotype is in 

two pieces and is extremely poorly preserved; total length was 

not measured. It has been dissected for the jaw apparatus and no 

meaningful illustrations of the anterior end is possible. 

DESCRIPTION.—Holotype complete with 221 setigers. Anterior 

end with 113 setigers; length approximately 100 mm; 

maximal width 12 mm; length through setiger 10, 20 mm. 

Prostomial lobes frontally obliquely truncate, dorsally 

flattened, tapering ventrally from posterior edge, median sulcus 

shallow. Eyes behind bases of A-I, dark. Ceratophores 

ring-shaped in all antennae, without articulations. Ccratostyles 

tapering smoothly; up to 21 moniliform articulations in A-III. 

A-I to middle of anterior peristomial ring; A-I I and A-III to 

posterior edge of anterior peristomial ring. Peristomial rings 

distinct ventrally and in part dorsally; anterior ring 3 A of total 


peristomial ring, tapering, with 4 or 5 articulations. 

Maxillary formula 1+1, 5+5, 6+0, 5+8, 2+2. Mx VI absent 


Branchiae (Figure 86m) present, pectinate, distinctly longer 




of setigers. All branchiae strongly pectinate; maximum 24 

long, slender filaments at about setiger 20. 

Anterior neuropodial acicular lobes distally rounded, becoming 

triangular in median and posterior setigcrs; aciculae 

emerging at midlinc. All pre- and postsetal lobes low, 

transverse folds. All ventral cirri basically tapering and thick. 

Ventral cirri very indistinctly inflated basally from about 


tapering. Posterior ventral cirri tapering. All notopodial cirri 

basally slightly inflated, tapering to thick, blunt tips, with 2 to 

4 cylindrical, often indistinct articulations. 

Limbate setae marginally serrated or frayed. Shafts of 

anterior pectinate setae (Figure 861) thick; blade forming 

scoops at end of setae. Both marginal teeth slightly thicker and 

longer than other teeth; -10 teeth present. Shafts of median and 

posterior pectinate setae (Figure 86j) wide and flat; blades flat, 

flaring. Marginal teeth about as long as other teeth; -15 teeth 

present. True compound falcigers (Figure 86k) in anterior 

setigers only; joints increasingly indistinct posteriorly; by 

setiger 50 no truly jointed setae present. Shafts of compound 

falcigers tapering. Appendages very short, stout, bidentate. 

Proximal teeth shorter than distal teeth, directed distally. Distal 

teeth tapering, directed distally. Guards not seen. Pseudocompound 

falcigers (Figure 86h,i) in posterior end distally evenly 

furcate, rather than bifid; reduced joint sometimes visible only 

as slight thickenings along 1 side with some faint lines running 

obliquely across setae, sometimes more distinct with fused 

appendages at distinct angle to shafts. Pseudocompound 

falcigers amber-colored, very nearly as large as subacicular 

hooks in posteriormost setigers. Compound spinigers absent 

Aciculae usually paired, amber-colored to brown, tapering to 

blunt tips (Figure 86g), distally geniculate or distinctly curved; 

cross-section round. Subacicular hooks (Figure 86f) amber-


colored, tridentate with teeth in a crest. Hooks from setiger 63, 

scattered in next 10-15 setigers, in all neuropodia in posterior 

setigers, always single (except for replacements). Hooks 

slender, tapering, with distinct heads. Proximal fangs large, 

curved. Two smaller fangs in crests. 

UNKNOWN MORPHOLOGICAL FEATURES.—Several prostomial 






Characters: 4,6,10-12, 15-17,24, 70, 74, 78. 


None. 

REMARKS.—Eunice pelamidis has remained unreported 

since it was first described. The species is characterized by 

having pseudocompound falcigers in the posterior end as does 

E. afuerensis. In E. afuerensis the articulations of the 

ceratostyles are drop-shaped or moniliform distally only; in E. 

pelamidis the articulations are moniliform throughout. Peristomial 

cirri are articulated in E. pelamidis and lack articulations 

in E. afuerensis. The two species are listed in Tables 52 and 53. 

152. Eunice pellucida Kinberg, 1865 


Eunice pellucida Kinberg, 1865:562. 

Eunice vitlata.—Hartman, 1948:77 [in part, not Nereis vittata Chiaje, 1828]. 

MATERIAL EXAMINED.—Two syntypes, RM Typ 433, St. 

Thomas, West Indies, coll. Werngren. Kinberg (1865:562) 

gave locality information as "St. Thomas, fundo 10-12 

orgyiarum, unde retulit Verngren." 


of two rather poorly preserved anterior ends and a long 

posterior end, which may be part of the longer of the two 

syntypes. The shorter anterior end has been dissected ventrally. 

DESCRIPTION.—One syntype with 35 anterior setigers. Other 

syntype with 58 anterior setigers; length 22 mm; maximal 

width 2 mm wide; length through setiger 10, 3.5 mm. Posterior 

fragment with 60 setigers; length 32 mm. 




shallow. Eyes absent. Antennae in a horseshoe, evenly spaced, 



moniliform articulations in A-III in short syntype; 20 articulations 

in A-III in long syntype. A-I to setiger 2; A-II to setiger 5; 

A-III to setiger 11. Peristomium cylindrical. Separation 

between rings distinct dorsally and ventrally; anterior ring 4 /5 of 

total peristomial length. Peristomial cirri to posterior edge of 

prostomium, with 3 articulations. 


long, located behind left Mx II. All jaws currently very soft. 


region, erect. Branchiae from setiger 5 to setiger 28 on long 

syntype; on other syntype branchiae from setiger 6 and present 

to last segment with 2 very short filaments, corresponding in 

size to branchiae on last branchial segments in other syntype. 

Branchiae terminating well before posterior end, present on less 

than 55% of total number of setigers. Maximally 8 filaments. 

Branchial stems erect. Filaments slender. Posterior fragment 

entirely abranchiate. 

Anterior neuropodial acicular lobes truncate; median and 

posterior acicular lobes (Figure 87f) distally rounded; aciculae 


folds. First 4 ventral cirri thick, tapering. Ventral cirri basally 

inflated from setiger 5 through about setiger 30. Inflated bases 

ovate; narrow tips digitiform. From about setiger 45 ventral 

cirri long, slender and digitiform, increasing in relative length 

posteriorly. Anterior notopodial cirri medially inflated, becoming 

digitiform in posterior setigers, retaining same absolute 

length, thus much more dominant in posterior than in anterior 



87d) tapering, flat Both marginal teeth slightly longer than 


(Figure 87e) distally inflated. Appendages short, slender; heads 

large, bidentate. Proximal teeth triangular, directed laterally. 

Distal teeth curved. Guards asymmetrically bluntly pointed, 


and compound spinigers absent. Aciculae (Figure 87b) paired, 

yellow, distally hammer-headed, nearly bifid in some segments. 

Separation between cores and sheaths indistinct in both 

aciculae and subacicular hooks. Subacicular hooks (Figure 87c) 


setiger 21 or 23, present in all setigers thereafter, always single 

(except for replacements). Hooks with large curved main fangs; 

distal fangs emerging from joint base, decreasing evenly in 

size. 







Characters: 1,2,4, 6, 32, 39,40,42,47, 50,63. 


None. 

REMARKS.—Eunice pellucida was considered as a possible 

synonym of E. vittata by Hartman (1948). It belongs to group 

C-l as does E. vittata, but differs clearly from the latter in that 

it has hammer-headed aciculae and moniliform antennal 

articulations. Eunice vittata has smoothly tapering aciculae and 

long, cylindrical antennal articulations. Eunice pellucida is 

listed with similar species in Tables 41 and 44. Most species 

listed in Table 44 have tapering, pointed aciculae; E.

262 


FIGURE 87.—Eunice pelluclda (syntype, RM Typ 433): a, anterior end, lateral view; b, aciculae, parapodium 54; 

c, subacicular hook, parapodium 54; d, pectinate seta, parapodium 54; e, compound falciger, parapodium 54; f, 

parapodium 54, anterior view. Eunice pennata (USNM 97393): g, anterior end, lateral view; h, parapodium 3, 

anterior view; i, compound falciger, parapodium 3; j, pectinate seta, parapodium 3; k, parapodium 20, anterior 

view; 1, compound falciger, parapodium 20; m, pectinate seta, parapodium 20; n, pectinate seta, parapodium 50; 

o, subacicular hook, parapodium 50; p, parapodium 50, anterior view. (Scale bars in mm.)


papeetensis has expanded and knobbed aciculae and E. 

pellucida has distinctly hammer-headed aciculae. Branchiae are 

distinctly pectinate in E. pellucida and palmate in E. 

papeetensis. 

153. Eunice pennata (Muller, 1776) 

FIGURE 87g-p; TABLES 19, 20 

Nereis pennata Muller, 1776:217; 1779:60-61, pi. 29: figs. 1-3. 

Leodice nonvegica Lamarck, 1818:323.—Savigny, 1820:51.—Audouin and 

Milne Edwards, 1833:219.—Orsted, 1845:402, 406, pi. 2: figs. 13-15.— 

Grube, 1850:202 [in part, not Nereis norvegica Linnaeus, 1767]. 

Eunice pennata.—Fauvel, 1923:400-401, fig.156h-o. 

MATERIAL EXAMINED.—TWO specimens, USNM 97393, 

Storskjaer, Oslofjorden, Norway, 8 June 1982, dredged, coll. 

and id. Inger Winsnes. 

DESCRIPTION.—Both specimens complete, mature females 

with large oocytes in body cavity. Specimen illustrated with 


through setiger 10, 7.5 mm. Other specimen in posterior 

regeneration, with 100 setigers; last 20 in regenerating portion; 

length 57 mm long of which 9 mm in regenerating portion; 

maximal width 5 mm wide; length through setiger 10, 9 mm. 

Body dorsally strongly convex with flattened ventral surface, 

tapering abruptly frontally and slowly towards posterior end. 

Anal cirri slender, without articulations; long ones as long as 

last 3-4 setigers in illustrated specimen. 



Prostomial lobes frontally obliquely truncate, dorsally inflated; 

median sulcus shallow. Eyes lateral to bases of A-I I, purple. 

Antennae in a horseshoe, with A-III isolated by a gap, with A-I 

thicker than other 3. Ceratophores ring-shaped in all antennae, 

without articulations. Ceratostyles slender and tapering to fine 

tips, with long, irregularly spaced articulations; in A-I 

articulations drop-shaped distally. A-III lost or incomplete in 

both specimens. A-I to posterior peristomial ring; A-I I to 

setiger 4 or 5. Peristomium cylindrical. Separation between 

rings distinct on all sides, but especially well marked dorsally 

and ventrally; anterior ring 2 /3 of total peristomial length. 

Peristomial cirri to middle of prostomium, slender, with 3 or 4 

irregular, but relatively long articulations. 

Maxillary formula (examined in one specimen only) 1+1, 

6+7, 9+0, 6+11, and 1+1. Mx III long, located behind left Mx 

II. Teeth of Mx II relatively coarse and triangular; other teeth 

small, even in size and distally blunt. 

Branchiae (Figure 87h,k) present, pectinate, distinctly longer 


Branchiae from setiger 3 to setiger 39 or 41. Branchiae 


of total number of setigers. First 5 and last 2 pairs single 

filaments; maximally 12 filaments at about setiger 15. 

Branchial stems slender, erect, tapering. Filaments shorter than 

notopodial cirri. All filaments flattened, medially expanded, 

with knife-shaped tips. 

Prebranchial and branchial neuropodial acicular lobes 

obliquely truncate with aciculae emerging from upper, higher 

part. Postbranchial acicular lobes (Figure 87p) obliquely 

rounded. All presetal lobes low, slightly excavate, transverse 

folds. Prebranchial and branchial postsetal lobes free, symmetrically 

rounded lobes, visible behind acicular lobes in most 

setigers; postbranchial postsetal lobes following outline of 

acicular lobes closely. Anterior ventral cirri relatively slender, 

tapering, becoming basally inflated from about setiger 6-7, but 

even in first setigers, distal tips set off from remainder of 

ventral cirri by a groove. Inflated bases ovate, rather modest; 

narrow tips very large and tapering. From about setiger 35 basal 

inflation gradually lost, absent from about setiger 40. Posterior 

ventral cirri slender, tapering, nearly conical. Anterior notopodial 

cirri long, digitiform, with 3 to 4 articulations reduced to 1 

to 2 in early branchiated setigers. In branchial region 

notopodial cirri more distinctly tapering, gradually loosing all 

traces of articulations. Postbranchial notopodia with distinct 

ring-shaped bases and slender, tapering notopodial cirrostyles. 


87j,m,n) tapering, flat. One marginal tooth longer than other 

teeth; number of teeth increasing from 8 to 12 from anterior to 

posterior setigers. Shafts of compound falcigers (Figure 87i,l) 

inflated, internally striated, marginally serrated. Appendages 

tapering, bidentate. Proximal teeth smaller than distal teeth, 

broadly triangular, directed laterally. Distal teeth nearly erect or 

gently curved. Guards asymmetrically sharply pointed in 

anterior setigers, becoming symmetrically sharply pointed in 

median and posterior setigers, marginally serrated in median 

and posterior setigers; mucros absent. Pseudocompound falcigers 

and compound spinigers absent. Aciculae usually paired, 

yellow, tapering to slender tips, gently curved or straight; 

cross-section round. Separation of cores and sheaths indistinct 


(Figure 87o) yellow, bidentate. Hooks first present from setiger 

35 or 43, present in all setigers thereafter, sometimes paired. 

Hooks tapering to small heads. Proximal teeth larger than distal 

teeth, triangular, directed laterally. Distal teeth nearly erect. 






Characters: 4,6, 23. 


None. 

REMARKS.—Muller first mentioned this species in a brief 

note (1776) and then expanded the description in 1779; the 

types are lost, but the type locality was given as Storskjaer in 

Christianiafjord (= Oslofjord). In the 1779 publication Muller 

also describes a Nereis pinnata from Madrepora pertusa reefs 

and refers to Nereis noruegica (note spelling) and Nereis 

madreporae pertusae of Gunnerus as synonyms of his Nereis 

pennata.


Three species of Eunice arc associated with ahermatypic 

coral reefs off Norway; all of these are present on the reef at 

Storskjaer in the Oslofjord (Inger Winsnes, pers. comm.). These 

species are readily identified: one has branchiae along most of 

body and black subacicular hooks (£. norvegicd); the two other 

species have branchiae limited to a short anterior region. The 

two latter species are separable on a variety of features, but 

perhaps most easily on the fact that one has yellow (£. 

pennata), the other dark brown or black subacicular hooks and 

aciculae (£. dubitata). 

Some confusion has arisen as to the identity of £. pennata as 

opposed to E. pinnata. Traditionally, E. pennata has been 

assigned to a species in group A-l sensu Fauchald (1970). 

Nothing in Miiller's description contradicts this tradition; for 

that reason, this tradition is here accepted. Nereis pinnata (= 

Eunice pinnata auctores) is treated below. 

As the first described species in group A-l, E. pennata has 

been widely reported and appeared at one time to have a bipolar 

distribution. Records of this species from the southern 

hemisphere (cf. Hartman, 1964:118, 1967:99) have yet to be 

confirmed. 

Perhaps the most unique feature of the species is the presence 

of ring-shaped bases in posterior notopodia; this is a feature 

that has been reported from only one other species of Eunice (£. 

nicidioformis); it resembles the structure of the notopodia 

among onuphids more than in the eunicids. 

Eunice pennata is listed with similar species in Tables 19 

and 20. Of species listed in Table 20, the following have 

branchiae starting on setiger 3: E. biannulata, E. caeca, E. 

kobiensis, E. mexicana, E. pennata, E. segregata, E. valens, 

and E. websteri; the other species have branchiae first present 

from setiger 4 or later. Of the species listed, E. caeca has as 

many as 24 branchial filaments where the branchiae are best 

developed; E. mexicana has 18 filaments; E. biannulata and E. 

kobiensis have 8 filaments; the remaining species have 11-15 

filaments where the branchiae are best developed. Eunice 

pennata and E. websteri have distally moniliform or dropshaped 

articulations in the ceratostyles; E. segregata and E. 

valens have cylindrical articulations. In E. pennata the first five 

branchiae are single filaments; in £. websteri only one anterior 

segment has single filaments. In contrast, at the posterior end of 

the branchiated region, £. pennata has two segments with 

single filaments; £. websteri has 10 segments. Note that £. 

pennata is the only species in Table 20 with distinct 

ring-shaped notopodial bases in posterior notopodia. 

154. Eunice perimensis Gravier, 1900 

FIGURE 88a-c; TABLES 33, 39 

Eunice perimensis Gravier, 1900:239-242, figs. 94-99, pi. 12: figs. 61, 62. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, Red Sea, 

Par, Perimen, 1894, coll. J. Jousseaume. 


was been compressed dorsoventrally to evert pharynx during 

fixation and is distorted as illustrated. 



through setiger 10, 10 mm. Anterior part of body cylindrical; 

becoming strongly dorsoventrally flattened by setiger 10; 

segments short; crowded. 



Prostomial lobes frontally truncate, dorsally somewhat flattened; 

median sulcus deep. Eyes between bases of A-I and A-II. 



Ceratostyles slender and tapering, without articulations. No 

antennae reaching beyond anterior pcristomial ring; A-I 

shortest; A-I 11 longest. Peristomium cylindrical. Separation 

between rings distinct dorsally and ventrally; anterior ring -2/$ 

of total pcristomial length, possibly distorted. Pcristomial cirri 

to middle of anterior pcristomial ring, without articulations. 

Maxillary formula 1 + 1,4+4,7+0,4+7, 1 + 1. Left Mx IV very 

small; part of distal arc with Mx 111 and left Mx V. Jaws heavily 

calcified. 



Branchiae present from setiger 17 to end of fragment. All but 

first branchia with 2 or more filaments; maximum 8 filaments 

from about setiger 30 to end. Branchial stems slender, tapering 

and flexible. Filaments distinctly longer than notopodial cirri 

except in first branchial segments. 

Neuropodial acicular lobes distally truncate to rounded; 


transverse folds. First 9 ventral cirri tapering. Ventral cirri 

distinctly basally inflated by setiger 10. Inflated bases 

moderate, ovate; narrow tips tapering. Ventral cirri remaining 

basally inflated through rest of fragment. Notopodial cirri 

supported by internal aciculae. Prebranchial notopodial cirri 

long, digitiform, not increasing in length through prebranchial 

region, becoming slightly medially inflated in last prebranchial 

segments, becoming reduced in length in branchial region, 


Limbate setae slender, distinctly frayed. Shafts of pectinate 

setae (Figure 88c) slender, blades slightly furled, flared. One 

marginal tooth longer than all other teeth; -20 teeth present. 

Posterior fascicles with up to 10 pectinate setae each. Shafts of 

compound falcigers (Figure 88d) distally inflated, with distinct 

peak, marginally smooth. Appendages short with very large 

distinct heads, bidentate. Proximal teeth slightly longer than 

distal teeth, triangular, directed slightly basally. Distal teeth 

curved. Guards symmetrically rounded, marginally smooth; 


spinigers absent. Aciculae usually single, sometimes paired, 

light to dark brown, thick, tapering, straight, projecting well 

beyond tip of parapodia in posterior setigers; cross-section 

round. Separation of cores and sheaths indistinct in both


FIGURE 88.—Eunice perimtnsis (holotype, MNHN): a, anterior end, lateral view; b, parapodium 52, anterior 

view; c, pectinate seta, parapodium 52; d, compound falciger, parapodium 52; e, subacicular hook, parapodium 

52. Eunice perrieri (holotype, MNHN): f, anterior end, lateral view; g, compound falciger, parapodium 75; h, 

subacicular hook, parapodium 75; i, pectinate seta, parapodium 75; j, parapodium 75, anterior view. (Scale bars 

in mm.) 

aciculae and subacicular hooks. Subacicular hooks (Figure 88e) 

light to dark brown, bidentate. Hooks first present from setiger 


replacements). Hooks tapering towards very small heads. 

Proximal teeth slightly thicker than distal teeth; both teeth 





TURES.—None.




Characters: 1, 2, 6, 36-38,40,47, 50, 57-60,63. 


38,1. 

REMARKS.—Eunice perimensis is listed with similar species 

in Tables 33 and 39. Of species listed in Table 39, four have 

more than three branchial filaments where the branchiae are 

best developed; these include E. complanata, E.flavapunctata, 

E. magnifica, and E. perimensis. The peristomial cirri reach 

beyond the prostomium in E. complanata and do not outreach 

the peristomium in E. flavapunctata and E. perimensis; the 

length of the cirri is unknown for E. magnifica. The separation 

between the two peristomial rings is distinct on all sides in E. 

flavapunctata and is limited to the dorsal and ventral sides in E. 

perimensis; the separation is unknown for E. magnifica. 

Finally, the branchiae are held stiffly erect in E. magnifica and 

are flexed in E. perimensis. 

155. Eunice perrieri Gravier, 1900 


Eunice perrieri Gravier, 1900:232-236, figs. 83-88. pi. 12: figs. 57-59. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, Red Sea, 

Djibouti, coll. Coutiere, no. 29, 1897. 



10, 12 mm. Body cylindrical throughout, with very modest 

flattening of dorsum towards posterior end. 


than peristomium, as deep as 1 /2 of the peristomium. Prostomial 


rim; median sulcus deep. Eyes on lateral sides of ceratophores 

of A-II. Antennae in a horseshoe, evenly spaced, similar in 

thickness. Ceratophores long in all antennae, without articulations. 

All ceratostyles tapering, without articulations. A-I to 

posterior peristomial ring; A-II and A-III to setiger 1. 

Peristomium tapering slightly anteriorly, but basically cylindrical. 

Lower lip thick, inflated, distinctly set off from rest of 

anterior ring. Separation between rings distinct dorsally and 

ventrally; anterior ring - 3 A of total peristomial length. 

Peristomial cirri to anterior third of anterior peristomial ring, 


Maxillary formula 1+1,4+4, 6+0,4+8, and 1+1. Jaws small 

in relation to size of specimen. Teeth on Mx IV small. Left Mx 

IV part of distal arc with Mx III. 



Branchiae from setiger 18 to setiger 170, absent from last 15 

setigers of body. Branchiae present to near posterior end, 

present on more than 65% of total number of setigers. First 2-3 

and last 4-5 pairs single filaments; maximum 10 slender 

filaments where best developed, at about setiger 30. Branchial 

stems slender. 

Anterior neuropodial acicular lobes wide, rather flattened, 

becoming about as high as wide in posterior end, retaining 

smoothly rounded outlines; aciculae emerging at midline. All 

presetal lobes low, transverse folds. Postsetal lobes about twice 

as high as acicular lobes in first 20 setigers; distally rounded. 

Postsetal lobes reduced to following outline of acicular lobes 

by about setiger 30; posterior postsetal lobes low, transverse 

folds. First 9 ventral cirri basally thick, abruptly tapering with 

slender, pointed tips. Ventral cirri basally inflated from about 

setiger 10 through about setiger 100. Inflated bases thick, 

transverse welts; narrow tips tapering. Far posterior ventral cirri 

slender, digitiform, increasing slightly in length, approximating 

length of notopodial cirri. Notopodial cirri basally slightly 

inflated in anterior 2 /3 of body, digitiform in last third of body. 

All notopodial cirri without articulations. 

Limbatc setae slender, marginally smooth. Shafts of pectinate 

setae (Figure 88i) very slender, blades strongly furled and 

flared. One marginal tooth very much longer than other teeth; 

-20 teeth present. Shafts of compound falcigcrs (Figure 88g) 

inflated, marginally serrated. Appendages small wilh nearly 

parallel sides; heads large, distinct, bidentatc. Proximal teeth 

slightly larger than distal teeth, tapering, directed laterally or 

slightly basally. Distal teeth tapering, directed obliquely 

laterally. Gap between teeth somewhat variable. Guards 

symmetrically rounded, marginally serrated; mucros absent. 

Pseudocompound falcigers and compound spinigcrs absent. 

Aciculae usually single, wilh dark copper-colored to dark 

brown cores and clear sheaths, tapering, distally gently curved; 

cross-section round. Subacicular hooks (Figure 88h) with dark 

copper-colored cores and clear sheaths, bidentate. Hooks first 


single (except for replacements). Hook distally strongly 

tapering; heads indistinct. Teeth similar in size, short, 

indistinct, directed distally. 


anal cirri. 







None. 

REMARKS.—Eunice perrieri resembles E. afra closely and 

has been considered a synonym of that species. A non-type 

specimen in the collections of MNHN, Paris, from Djibouti, 

labeled in Fauvel's handwriting, was identified as "£. afra (= E. 

perrieri)." 

Eunice perrieri is listed with similar species in Tables 33 and 

37. It is the only species in Table 37 with as many as 10 

branchial filaments. It has long ceratophores in all antennae, 

feature it shares with only one species in Table 37, E. aciculata.


These two species are very similar; branchiae have single 

filaments in the first 10 setigers in E. aciculata and only in two 

to three setigers in E. perrieri; in addition, the latter has twice 

as many (10) branchial filaments where the branchiae are best 

developed as the former. 

157. Eunice petersi, new name 


Eunice punclala Peters. 1854:611.—Ehlers, 1897:166. 

MATERIAL EXAMINED.—Holotype, ZMB 44, Mozambique, 

2 slides marked type, BM(NH) ZB 1984.71 and 72. 


now in three pieces. 



10, 7 mm. Body cylindrical; segments short and crowded 

throughout. 


about as wide as pcristomium, as deep as l /i of 


inflated; median sulcus deep. Eyes behind bases of A-I, faded, 

covered by pcristomial fold. Antennae in a straight line, evenly 

spaced, similar in thickness. Ceratophores long in all antennae, 

without articulations. Ceratostyles digitiform, tapering slightly, 

with 2 to 3 indistinctly marked cylindrical articulations. A-I 

and A-II to setiger 1; A-III to setiger 3. Peristomium 

cylindrical; lower lip muscular. Separation between rings 

distinct on all sides; anterior ring 3 A of total peristomial length. 

Peristomial cirri to frontal edge of peristomium, digitiform, 





from setiger 13 to setiger 200; occurrence of branchiae sporadic 

posterior to setiger 100. Branchiae present to near posterior 


Where best developed branchiae with 4 short, thick filaments 

on short, truncate branchial stems. Posterior to setiger 70 most 

branchiae single, short, truncate filaments. 

All neuropodial acicular lobes (Figure 89e) wide, rounded; 

aciculae emerging at midline. All presetal lobes low, transverse 

folds. Anterior and median postsetal lobes free, rounded. 

Posterior to setiger 100 postsetal lobes follow outline of 

acicular lobes closely. First 4 ventral cirri thick, tapering. 


thick, ventrolateral ridges between setigers 6 and setiger 75; 

narrow tips short and button-shaped. Posterior ventral cirri 

slender and tapering. All notopodial cirri basally slightly 

inflated, tapering to slender tips. Anterior and median 

notopodial cirri with 2 articulations; posterior notopodial cirri 



(Figure 89d,i) tapering, flat. One marginal tooth longer than 


(Figure 89b,h) distinctly inflated, marginally serrated or 

smooth. Appendages short; heads very large, bidentate. 

Proximal teeth longer than distal teeth, triangular, directed 

laterally. Distal teeth curved, tapering. Guards symmetrically 

rounded, marginally smooth; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Aciculae (Figure 89f) 

single, with brown cores and clear sheaths, tapering, slender, 

straight; cross-section round. Subacicular hooks (Figure 89c,g) 

with brown cores and clear sheaths, bidentate. Hooks first 


single (except for replacements). Hooks tapering, slender, with 

small heads. Proximal teeth larger than distal teeth, tapering, 

laterally directed. Distal teeth slender, directed obliquely 

distally. 







Characters: 4,6, 39,42. 


None. 

REMARKS.—The slides in BM(NH) are numbered 71 and 72. 

Slide 71 contains a median parapodium showing dark brown 

aciculae, compound hooks with both teeth well developed, and 

blunt, rounded guards, and slender limbate setae and groups of 

narrow pectinate setae without distinctly long marginal teeth. 

Number of teeth in each pectinate seta appears to be about 15. 

Subacicular hook tapers towards the tip and the proximal tooth 

is at nearly right angles with the shaft and about twice the size 

of the distal tooth. Aciculae are conical and abruptly tapered. 

Slide 72 contains two very similar neuropodia. Branchiae are 

not visible on either slide. 

Eunice petersi is listed with similar species in Tables 33 and 

35. Perhaps most characteristic is the scattered distribution of 

branchiae over the posterior half of the body. It is one of two 

species in Table 35 with notopodial cirri articulated in the 

anterior end; the other species is E. fauveli. The latter has 

mucronate compound falcigers; E. petersi has distally rounded 

guards on the compound falcigers. 

A new name is necessary, because the specific name 

punctata was pre-occupied in combination Eunice punctata 

(Risso, 1826). 

The name honors the scientist responsible for describing a 

very valuable, early collection from Mozambique. 

158. Eunice philocorallia Buchanan, 1893 

FIGURE 89j-q; TABLES 27,29 

Eunice philocorallia Buchanan, 1893:173-176, pi. 9: figs. 2-6, pi. 10: figs. 

7-9, pi. 11.

268 


FIGURE 89.—Eunice petersi, new name (holotype, ZMB 44 (a-e) slides, BM(NH) ZB 1984.71 and 72 (f-i): a, 

anterior end, lateral view; b, compound falciger, parapodium 63; c, subacicular hook, parapodium 63; d, pectinate 

seta, parapodium 63; e, median parapodium, anterior view; f, acicula, median parapodium; g, subacicular hook, 

median parapodium; h, compound falciger, median parapodium; i, pectinate seta, median parapodium. Eunice 

philocorallia (holotype, BM(NH) 1920.1.5.1): j, anterior end, lateral view; k, compound falciger, parapodium 3; 

1, pectinate seta, parapodium 3; m, parapodium 3, anterior view; n, parapodium 96, anterior view; o, compound 

falciger, parapodium 96; p. pectinate seta, parapodium 96; q, subacicular hook, parapodium 96. (Scale ban in 

mm.)


MATERIAL EXAMINED.—Holotype, BM(NH) 1920.1.5.1,50 

miles off Bolus Head, Kerry, Ireland; dredged, 375 m, in 

parchment-like tubes on Lophophelia prolifera. Royal Dublin 

Society Survey of Western Ireland, 1891. 



through setiger 10, 15 mm. Anterior end cylindrical; middle 

and posterior ends dorsoventrally flattened with prominent 

parapodia. Anal cirri as long as last 15 setigers. 


peristomium, as deep as 1 /2 of peristomium, bent ventrally. 


sulcus shallow. Eyes between bases of A-I and A-II, hidden 

below peristomial fold in illustration. Antennae in transverse 

row, evenly spaced, similar in thickness. Ceratophores ringshaped 


cylindrical, with up to 12 cylindrical, poorly marked articulations 

in A-I 11. A-I and A-II to middle of posterior peristomial 

ring; A-I 11 to setiger 2. Peristomium -3 times as long as 

prostomium, cylindrical. Separation between rings distinct 

dorsally and ventrally; anterior ring ~ 3 /4 of total peristomial 




Branchiae present, pectinate, distinctly shorter than notopodial 


from setiger 6 to setiger 135. Branchiae present to near 

posterior end, present on more than 65% of total number of 

setigers. First branchia single filaments; maximum 4 filaments. 

Stems very short, truncated. Filaments slender, digitiform, 


Anterior neuropodial acicular lobes (Figure 89m) distally 

rounded; posterior acicular lobes increasingly obliquely conical 

with the high side dorsally (Figure 89n); aciculae emerging 

dorsal to midline. Presetal lobes low, oblique folds with high 

side dorsally. Postsetal lobes folds following outline of acicular 

lobes closely. Anterior ventral cirri broadly attached below 

ventral edge of acicular lobes, tapering to blunt tips. Ventral 

cirri strongly inflated basally from about setiger 4 through 

setiger 45; inflated bases completely reduced by setiger 50. 

Inflated bases nearly spherical; narrow tips tapering. Posterior 

ventral cirri short and tapering, blunt, emerging from posterior 

parapodial faces, directed posteriorly and dorsally. Anterior 

notopodial cirri basally slightly inflated, with 6 articulations. 

Notopodial cirri increasing in length posteriorly, becoming 

slightly longer than body width in last setigers, retaining at 

least 2 articulations in all setigers. 


(Figure 891,p) narrow, tapering, flat. One marginal tooth longer 

than other teeth, with -10 teeth. Shafts of anterior compound 

falcigers (Figure 89k) inflated, marginally indistinctly serrated; 

distal beaks present Appendages thick; heads small, bidentate. 

Proximal teeth slightly shorter than distal teeth, thick, 

triangular. Distal teeth slender, curved or bent. Guards 

asymmetrically bluntly pointed; mucros absent Shafts of 

posterior compound falcigers (Figure 89o) tapering, marginally 

smooth; beaks absent Appendages distinctly tapering; heads 

distinct Proximal teeth larger than distal teeth, narrowly 

triangular, directed laterally. Distal teeth short, blunt, nearly 

erect Guards symmetrically bluntly pointed, marginally 

smooth; mucros absent Pseudocompound falcigers and compound 

spinigers absent Aciculae at least paired, with dark 

brown to black cores and clear sheaths, bluntly pointed, straight 

or gently curved; cross-section round. Notopodial aciculae 

slender, straw-colored to light brown. Subacicular hooks 

(Figure 89q) with dark brown to black cores and clear sheaths, 


setigers thereafter, single in most setigers, up to 3 hooks present 

in some setigers. Proximal teeth very much larger than distal 

teeth, directed laterally. Distal teeth nearly erect often nearly 

missing. 




None. 





None. 

REMARKS.—Eunice philocorallia is listed in Table 27 and 

29. Of the species listed in Table 29, four have less than 10 

branchial filaments where the branchiae are best developed; 

these include E. microprion, E. philocorallia, E. plicata, and E. 

tribranchiata. The peristomial cirri reach the prostomium in E. 

microprion and E. philocorallia and do not outreach the 

peristomium in the two other species. Posterior ventral cirri are 

broadly attached, triangular in E. microprion and tapering from 

a small base in E. philocorallia. 

Eunice pinnata (Miiller, 1779) 

Nereis pinnata Miiller, 1779:62-63, pi. 29: figs. 4-7. 

Eunice pinnata.—Cuvier, 1817:525.—Audouin and Milne Edwards. 1833:219. 

REMARKS.—This species, originally reported from an 

ahermatypic coral reef, presumably in the Oslofjord, was not 

characterized well enough by Miiller to allow identification 

with any of the three species present at the Storskjaer reef in the 

Oslofjord (i.e., E. norvegica, E. dubitata, and E. pennata). No 

type specimens exist and the other authors cited limit 

themselves to quoting Miiller's description. The species is here 


159. Eunice plicata Baird, 1869 


Eunice plicata Baird, 1869:348-349.—Fauchald, 1986:253, figs. 51-55. 

MATERIAL 

EXAMINED.—Holotype, BM(NH) ZH

270 


.05 

.1 

1 

r. P 

FIGURE 90.—Eunice plicata (holotypc, BM(NH) ZH 1861.9.20.25): a, anterior end, lateral view; b, parapodium 

46, anterior view; c, compound falciger, parapodium 46; d, pectinate seta, parapodium 46; e, subacicular hook, 


1861.9.20.25, Fremantle, Australia, coll. Bowerbank. 


length 80 mm long; maximal width 5 mm; length through 





thickened rim; median sulcus deep. Antennae in a horseshoe,



in all antennae, without articulations. Ceratostyles very thick, 

digitiform, with 2 or 3 long, cylindrical articulations. A-I to 

posterior edge of anterior peristomial ring; A-I I and A-I 11 to 

setiger 2. Peristomium anteriorly flared; lower lip muscular and 

very distinct. Separation between rings distinct only dorsally; 

anterior ring ~ 9 /io of total peristomial length. Peristomial cirri 

to middle of anterior peristomial ring, without articulations. 

Maxillary formula 1+1,4+4, 6+0, 6+7, and 1+1. 





of setigers. Last 30 branchiae single filaments; maximum 5 

filaments present. Branchial stems very short; branchiae 

appearing nearly palmately branching. Filaments short, thick, 

shorter than notopodial cirri in all but a few median segments. 

Neuropodial acicular lobes triangular; aciculae emerging at 

midline. All presetal lobes low, transverse folds. Anterior 

postsctal lobes free, as long as acicular lobes, following outline 

of acicular lobes by setiger 30, similar to presetal folds in 

posterior setigers. Anterior ventral cirri thick, tapering, as long 

as postsetal lobe, becoming strongly inflated in median 

setigers. Inflated bases thick, transverse welts; narrow tips short 

and button-shaped. By setiger 50 ventral cirrus present only as 

inflated glandular ridges ventral to neuropodia; narrow tips 

absent. All notopodial cirri thick, basally somewhat inflated, 

with 2 or 3 distinct articulations. 

Pectinate setae (Figure 90d) flared, flat. One marginal tooth 

slightly than other teeth; -15 teeth present. Shafts of compound 

falcigers (Figure 90c) tapering, marginally smooth. Appendage 

short, triangular, head distinct, bidentate. Proximal teeth 

shorter than distal teeth, tapering, directed laterally. Distal teeth 

tapering, curved, directed laterally. Guards symmetrically 

bluntly pointed; margins smooth; mucros absent. Pseudocompound 

falcigers and compound spinigers absent. Notopodial 

aciculae black. Neuropodial aciculae up to 3 in a neuropodium, 

black, tapering, bluntly pointed, straight; cross-section round. 

Subacicular hooks (Figure 90e) black, bidentate. Hooks first 


single (except for replacements). Hooks tapering; both teeth of 

about same size, directed distally. 





rv. 



Characters: 4,6, 13, 14,42, 51, 74, 78. 


None. 

REMARKS.—As far as is known, this species has remained 

unreported since it was briefly described by Baird (1869); it 

belongs to group B-2 (Fauchald, 1970). Relations between E. 

plicata and similar taxa are indicated in Tables 27 and 29. It is 

the only species in Table 29 other than E. kinbergi to have the 

ventral cirri basally inflated throughout the body. It can be 

separated from the latter on the structure of the prostomium, 

which is bilobate in E. plicata and quadrilobate in E. kinbergi. 

160. Eunice polybranchia (Verrill, 1880) 

FIGURE 91a-h; TABLES 13,27,28 

Ltodice polybranchia Venill, 1880:358; 1881:323; 1885:428-429. 

Eunice floriaana.—Hartman, 1942:49-50, 52 [in part, not Eunice floridana 

Ehlers, 1887]. 

Eunice norvegica.—Pettibone, 1963:240-242, fig. 63f [in part, not Nereis 

norvegica Linnaeus, 1767]. 


YPM 2731, Fish Hawk sta 871, 40°02'54"N, 70°23'40"W, 210 

m, fine sand with some mud, dredged. 


based on two incomplete specimens; variability of the species 

has been indicated in Table 13. 

DESCRIPTION.—Long syntype incomplete with 85 setigers; 


8 mm. Short syntype incomplete with 42 setigers; length 23 




peristomium. Prostomial lobes thick, frontally rounded, dorsally 

inflated; median sulcus shallow. Eyes between bases of 

A-I and A-I I, dark. Antennae in a horseshoe, with A-I isolated 

by a gap, similar in thickness. Ceratophores ring-shaped in all 

antennae, without articulations. Ceratostyles tapering, without 

articulations, sometimes with contraction rings. A-I to posterior 

peristomial ring or to setiger 1; A-I I to setigers 4-6; A-I 11 to 

setigers 6-11; A-III broken. Peristomium cylindrical, with 

distinctly inflated lower lip. Separation between rings distinct 

dorsally and ventrally; anterior ring 4 /s of total peristomial 

length. Peristomial cirri to posterior part of prostomium, 


Maxillary formula 1+1,6+5,8+0, 5+9, and 1+1. Mx III long, 

located behind left Mx II. Mx VI absent 



Branchiae from setigers 7-8 to last of segments present (85 

setigers). First branchia short, tapering filaments; maximum 5 

filaments reached at about setiger 15 and maintained over next 

10-15 setigers. Filaments thick, stiff, bent posteriorly, so 

branchiae project over next posterior intersegmental furrow. 

All neuropodial acicular lobes conical; anterior acicular 

lobes asymmetrical, becoming more and more symmetrical in 

median setigers (Figure 9In); aciculae emerging at tip of 

acicular lobes. Pre- and postsetal lobes low, transverse folds. 

First 4 ventral cirri thick and tapering. From about setiger 5 

ventral cirri basally inflated; from about setiger 15 through

272 


|NK 

RGURE 91.—Eunice polybranchia (lectotype, YPM 2731): a, anterior end, lateral view; b, appendage of 

compound falciger, parapodium 75; c, subacicular hook, parapodium 75; d, pectinate seta, parapodium 75; e, 

parapodium IS, anterior view; f, pectinate seta, parapodium IS; g, compound falciger, parapodium 15; h, 

parapodium 75, anterior view. Eunice prayensis (syntype, RM 428): i, anterior end, lateral view; j, compound 

falciger, parapodium 25; k, pectinate seta, parapodium 25; 1, subacicular hook, parapodium 25. (Scale bars in 

mm.)


TABLE 13.—Variable and invariable features in the type lot of Eunice poly bronchia (N = number of individuals 



Maximal width 





N 

7 

7 

7 

7 

6 

Max. 

5 

13 

8 

5 

36 

Min. 

2.5 

6 

7 

4 

24 

Mean 

3.71 

8.71 

7.71 

4.86 

29.17 

SD 

0.99 

2.29 

0.49 

0.38 

4.36 




Pcristomial cirri reach 

Perislomial cirral articulations 








N=7 

absent 


prostomium 

absent 

15 

black 

tapering; straight 

black 

2 

distinct 

15 

setiger 50 ventral cirri thick ventrolateral ridges; narrow tips 

short and button-shaped. Inflated bases reduced from about 

setiger 50; ventral cirri gradually displaced to posterior face of 

neuropodia, directed dorsally behind acicular lobes. In last 

setigers present ventral cirri thick, tapering. Notopodial cirri 

digitiform, without articulations. Posterior notopodial cirri 

curved over next posterior intersegmental furrows. 


(Figure 91d,f) flaring, furled. One marginal tooth distinctly 

larger than other teeth; 15 teeth present. Shafts of compound 

falcigers (Figure 91g) inflated, marginally serrated. Appendages 

(Figure 91b,g) tapering; heads distinct, bidentate. Proximal 

and distal teeth similar in size in anterior and median 

setigers; proximal teeth longer than distal teeth in posterior 

setigers. Proximal teeth triangular, directed laterally. Distal 

teeth gently curved. Guards symmetrically bluntly pointed; 


spinigers absent Notopodial aciculae slender, brown. Aciculae 

paired, with dark brown to black cores and clear sheaths, 

tapering, gently curved, considerably more prominent in 

posterior than in anterior setigers; cross-section round. Subacicular 

hooks (Figure 91c) with dark brown to black cores and 

clear sheaths, bidentate. Hooks first present from setiger 24-36 

(29 and 30 in 2 syntypes described), present in all setigers 


tooth twice as large as distal teeth, directed laterally. Distal 

teeth nearly erect. 


anal cirri. 



and 



Characters: 1,2,4, 6, 36-38,40,47, 50, 63. 


38,1. 

REMARKS.—Verrill (1880) discussed material from a series 

of Fish Hawk stations (865-895). The species was described in 

a footnote on page 358. The paragraph to which the footnote is 

attached discusses hard-bottom fauna from a series of stations, 

"especially from stations 865-867," but no particular station is 

mentioned as the type locality for the species. For that reason, 

all material identified by Verrill as Leodice polybranchia from 

Fish Hawk stations 865-895 must be considered syntypes and 

are treated as such here. In order to stabilize the description, a 

lectotype has been designated above. 

Eunice polybranchia resembles E. norvegica as first noted 

by Verrill (1880), and is listed with this and other similar 

species in Table 27. In contrast to E. norvegica it lacks antennal 

articulations and is listed with other species with which it 

shares this features in Table 28. It is the only species in that 

table to combine long peristomial cirri with having A-III 

distinctly longer than A-II. 

161. Eunice prayensis Kinberg, 1865 

FIGURE 91i-l; TABLES 41-45 

Eunice prayensis Kinberg, 1865:563. 

Eunice vittata.—Hartman, 1948:77 [in part, not Nereis viilata Chiaje, 1828]. 

MATERIAL EXAMINED.—Two syntypes, RM 428, Atlantic 

Ocean, Rio de Janeiro, Eugenie Expedition. Kinberg 

(1865:563) gave locality information as "Portus ad Praya 

grande juxta Rio Janerio." 

DESCRIPTION.—One headless fragment of 42 setigers, 

probably starting at setigers 3-5. Other syntype incomplete,


with 61 setigers; length 31 mm; maximal width 2 mm; length 

through setiger 10,4.5 mm. 

Prostomium (Figure 91i) distinctly shorter than peristomium, 

about as wide as peristomium, as deep as l /i of the 


flattened; median sulcus shallow. Eyes between bases of A-I 

and A-I I, black. Antennae in a horseshoe, evenly spaced. 

Ceratophores long in all antennae, without articulations. All 

ceratostyles missing. Peristomium cylindrical. Separation between 

rings distinct on all sides; anterior ring somewhat more 

than 72 of total peristomial length. Peristomial cirri missing. 

Jaws missing. 




posterior end. First branchiae single filaments; all others 

pectinate with up to 9 filaments at about setiger 12. Branchial 

stems slender, erect. Filaments long, slender. 


at midline. Pre- and postsetal lobes low, transverse lobes. 

First 2 ventral cirri tapering. Ventral cirri basally inflated from 

setiger 3 through setiger 35. Inflated bases nearly spherical; 

narrow tips tapering. Posterior ventral cirri thick and tapering. 

Notopodial cirri slightly inflated basally, without articulations. 


Limbate setae marginally finely serrated. Pectinate setae 

(Figure 91k) very slender, tapering, flat. One marginal tooth 

longer than other teeth; 5-7 long, slender teeth present. Shafts 

of compound falcigers (Figure 91j) distally slightly inflated, 

marginally smooth. Appendages long, narrow, bidentate. 

Proximal teeth triangular, directed basally. Distal teeth nearly 

erect in anterior appendages, sharply curved in median 

appendages. Guards tapering to sharp tips, mucronate. Pseudocompound 

falcigers compound spinigers absent. Aciculae 

numbering 2 or 3, yellow, tapering, straight; cross-section 

round. Subacicular hooks (Figure 911) yellow, tridentate with 

teeth in a cresL Hooks first present from setiger 18, present in 

all setigers thereafter, usually single, rarely paired. Main fang 

very large, curved; secondary and tertiary fangs in crests; 

especially tertiary fang very small. 

UNKNOWN MORPHOLOGICAL FEATURES.—Structure of ceratostyles; 

jaw structure; features associated with posterior 






Characters: 1, 2, 4, 6, 17, 20-23, 27-29,38,47, 50, 63. 


REMARKS.—Eunice prayensis was considered synonymous 

with E. vittata by Hartman (1948:77). It belongs to group C-l, 

as does E. vittata, and is listed with similar species in Tables 

41-45. 

Kinberg (1865:563) described the antennae as follows: 

"Appendices lobi cephalici breves nee moniliformes; tentacula 

media et impar segmentum 3 non attingentia, aequalia." 

(prostomial appendages short and not moniliform; A-I I and 

A-III do not reach third segment and are equal). Assuming that 

antennal articulations were cylindrical, the most probable 

comparison is between E. prayensis and species listed in Table 

42. The subacicular hooks are occasionally paired, but are 

certainly not present in a vertical series as in E. vittata and a few 

similar species in Table 45. The only other species with such 

distribution of the subacicular hooks in Table 42 is £. 

flavocuprea. Eunice flavocuprea has maximally four branchial 

filaments and the 10 last pairs of branchiae are single filaments; 

E. prayensis has maximally nine filaments and the branchiae 

have at least two filaments in the last branchiae present 

Kinberg stated that branchiae should be present from setiger 

3; this cannot be confirmed. Note that the notopodial cirri of 

setigers 3-4 are damaged, and branchiae may have been 

present. 

Eunice procera Grubc, 1866 

Eunice procera Grube, 1866b:68; 1878a:100. 

REMARKS.—According to the original description, E. procera 

has branchiae starting on setiger 22 and continued to far 

posterior end with a maximum of 11 filaments at setigers 

59-72. 

Grube (1878a: 100) in addition, by the placement of the 

species in his review, implied that the species should lack 

articulations in the ceratostyles. 

No specimens are available and the information is clearly 

insufficient to characterize the species. It is here considered 

indeterminable. 

162. Eunice prognatha Mclntosh, 1885 

FIGURE 92a-e; TABLES 27,31,33, 36 

Eunice prognatha Mclntosh, 1885:268-270, figs. 29-31, pi. 37: figs. 16, 17, 

pi. 19A:figs. 10, 11. 


1885.12.1.192, off Ascension Island, 7°54'20"S, 14°28'20"W, 

768 m, volcanic sand, Challenger sta 344, 3 Apr 1876. 


currently in two pieces and the anterior end has been deeply 

dissected. The specimen is otherwise in good condition. 



10, 16 mm. 




sulcus shallow. Peristomial fold large, covering bases of 

antennae and eyes at bases of A-I I. Antennae in a horseshoe, 

with A-I isolated by a gap, similar in thickness. Ceratophores


FIGURE 92.—Eunice prognatha (holotype, BM(NH) ZK 1885.12.1.192): a, compound falciger, parapodium 74; 

b, subacicular hook, parapodium 74; c, pectinate seta, parapodium 74; d, anterior end, lateral view; e, parapodium 

74, anterior view. Eunice pruvoti, new name (MNHN, Paris, syntype of E. anceps; h and i of smaller syntype): 

f, anterior end, lateral view; g, subacicular hook, parapodium 58; h, compound falciger, parapodium 40; i, 

subacicular hook, parapodium 40; j, parapodium 58, anterior view; k, pectinate seta, parapodium 58; 1, compound 

falciger, parapodium 58. (Scale bars in mm.)

276 



slender and digitiform; maximum 5-6 long articulations in 

A-II and III. Three median antennae to middle of setiger 1; A-I 

to middle of peristomium. Peristomium with deeply flaring 

pocket on dorsal side, otherwise conically expanding posteriorly. 


anterior ring 5 /s of total peristomial length. Remnants of 

peristomial cirri present. 

Maxillary formula 1+1, 6+6, 8+0, 5+12, 2+1, and 1+1. Mx 

III very short. Unusual features include 2 teeth on left Mx V 

and presence of distinct Mx VI. 



setiger 10 on left side and 13 on right side to setiger 110. 


more than 65% of total number of setigers. Most branchiae 

single, short filaments; some scattered segments with 2 or 3 

short, cylindrical branchial filaments (Figure 92e); last few 

filaments short, nearly button-shaped. 

Neuropodial acicular lobes rounded in anterior setigers, 

slightly more conical in posterior setigers; aciculae emerging 

dorsal to midline. All pre- and postsetal lobes low, transverse 

folds. First 9 ventral cirri digitiform. Ventral cirri distinctly 

basally inflated in setigers 10 through setiger 30. Inflated bases 

ovate; narrow tips tapering. Far posterior ventral cirri digitiform, 

resembling notopodial cirri in shape and length. All 

notopodial cirri basally slightly inflated, distally tapering with 

slender, digitiform tips, without articulations. 

Limbate setae slender, nearly capillary. Ten or more 

pectinate setae in median and posterior setigers. Pectinate setae 

(Figure 92c) flaring, flat. One marginal tooth longer and thicker 


falcigers (Figure 92a) tapering, with poorly defined internal 

striations, without marginal teeth. Appendages short, slender, 

bidentate. Both teeth similar in size. Guards symmetrically 

rounded; mucros absent Pseudocompound falcigers and 

compound spinigers absent. Aciculae single anteriorly, paired 

in median and posterior setigers, black, tapering. Aciculae 

strongly projecting in median setigers; cross-sections knifeedged, 

distally pointed. Subacicular hooks (Figure 92b) black, 



Hooks tapering smoothly to small head. Proximal teeth about 

twice as large as distal teeth; both teeth directed distally. 


anal cirri. 





Characters: 4, 6, 13, 14,27-29, 39,40, 42, 74, 78. 


None. 

REMARKS.—Eunice prognatha belongs to either group B-2 

or to group B-4; the branchial start is exactly at the (rather 

arbitrarily determined) breakoff point between the two groups. 

It is listed with similar species in both groups (Tables 27 and 

31, and Tables 33 and 36). It is unique in Table 31 in that it has 

palmate branchiae with only three filaments; all other species in 

this table have pectinate branchiae (i.e., a distinct, elongated 

branchial stem is present). It is the only species listed in Table 

36 with branchiae distinctly shorter than the notopodial cirri. 

Mclntosh (1885) reported that the peristomial cirri reached 

the frontal margin of the peristomium and that up to five 

branchial filaments could be present in a branchia; neither one 

of these statements can be confirmed. 

9. Eunice pruvoti, new name 

FIGURE 92f-l; TABLES 33.36 

Eunice anceps Pruvol. 1930:69. 

Eunice afra.—Fauvel in Pruvoi, 1930:69 (in ptrt. noi Eunice afra Peten, 

1854|. 

MATERIAL EXAMINED.—One syntype, MNHN, Paris, Kuts, 

He de Pino, Nouvellc Caledonie, 1928, coll. Mmc Pruvot; 1 

syntype, MNHN, Paris, Collection Francais, Nouvelle Caledonie, 

forme jcunc no. 10. 


currently labeled Eunice afra, and the labels are in Pierre 

Fauvel's handwriting; the second specimen also has a label 

naming it as "Eunice jeune no. 10." The small syntype is 

nearly translucent, slender, and narrow and not distinctly 

dorsally flattened; the anterior end has been deeply dissected 

and the jaws are now missing. The description is based on the 

larger of the syntypes. 

DESCRIPTION.—Large syntype complete, of unknown sex, 

with 115 setigers; total length 34 mm long; maximal width 3 

mm; length through setiger 10, 5 mm. Anterior body 

cylindrical, becoming dorsally flattened and rather wide with 

crowded setigers by setiger 10; posterior end abruptly tapering 

and flattened. Anal cirri as long as the last 15 setigers, without 

articulations. Small syntype complete, of unknown sex, with 

about 100 setigers; total length 22 mm; maximal width 1 mm 

wide; length through setiger 10, 3 mm. 

Prostomium (Figure 92f) distinctly shorter than peristomium, 

distinctly narrower than peristomium, less than x li as 

deep as peristomium. Prostomial lobes frontally truncate, 

dorsally somewhat flattened; median sulcus deep. Palpal 

regions set off by horizontal, frontal grooves. Eyes between 

bases of A-I and A-II, large, dark. Antennae in a horseshoe, 


in all antennae, without articulations. Ceratostyles thick, 

sausage-shaped, digitiform, with 1 or 2 cylindrical, distinct 

articulations. Antennae to middle of anterior peristomial ring. 

Peristomium slightly flaring anteriorly, with distinct, muscular 

lower lip. Separation between rings distinct dorsally and 

ventrally; anterior ring ~ A /s of total peristomial length.


Peristomial cirri to posterior end of anterior peristomial ring, 



Branchiae (Figure 92j) single filaments, present from setiger 

15 to setiger 115. All branchiae distinctly longer than 

notopodial cirri, strap-like, proximally flattened, nearly as long 

as half body width in median and posterior setigers. Branchiae 


number of setigers. 

Median acicular lobes distally rounded with aciculae 


folds. Anterior ventral cirri thick, tapering, becoming basally 

modestly inflated by setiger 10, retaining inflated bases through 

most of body. Bases ovate or spherical; narrow tips tapering. 

Far posterior ventral cirri tapering, rather slender. Prebranchial 

notopodial cirri tapering, basally slightly inflated, becoming 

digitiform in branchial region; all notopodial cirri short, rather 

inconspicuous, without articulations. 


(Figure 92k) flaring, flat. One marginal tooth distinctly longer 

than other teeth, with -15 teeth. Anterior setigers with 2-3 

pectinate setae; median and posterior setigers with up to 25 

pectinate setae. Shafts of compound falcigers (Figure 921) 

distally inflated, marginally smooth, with distinct distal beaks. 

Appendages short, tapering, with rather large heads, bidentate. 

Proximal teeth triangular, directed laterally. Distal teeth 

distinctly shorter than proximal teeth, distinctly bent, distally 

blunt Guards symmetrically rounded, marginally serrated; 


spinigers absent. Neuropodial aciculae single, light brown, 

tapering to blunt, straight tips; cross-sections round. Subacicular 

hooks (Figure 92f) light brown, bidentate. Hooks first 


single. Hooks tapering narrow heads. Teeth directed distally; 

both teeth similar in size. 

COMMENTS ON ADDITIONAL SPECIMEN EXAMINED.— 

Antennae of small syntype short, without articulations. 


Shafts of compound falcigers (Figure 92h) inflated, marginally 

smooth. Appendages very short. Guards symmetrically 

rounded. Teeth slender. Proximal teeth longer than distal teeth, 

narrowly tapering, directed laterally. Distal teeth tapering, 

directed laterally. Subacicular hooks (Figure 92i) from about 

setiger 25. Hooks with narrowed necks and distinct heads. Both 

teeth directed distally and similar in size. 

UNKNOWN MORPHOLOGICAL FEATURES.—Location of 

maximum width and width at setiger 10. Condition of dorsal 

edge of pygidium. Jaw structure. 


Mx III forming arc with left Mx IV; Mx VI absent. 





None. 

REMARKS.—Eunice anceps was first used in a list of 

synonyms of E. afra with "Pruvot M.S." (Fauvel in Pruvot, 

1930:69) as author. The name has never been used as a valid 

name for the taxon and is thus not available as a name 

according to the Code. The new name honors the person 

responsible for recognizing the distinct status of the species, G. 

Pruvot. 

The larger of the two specimens belong to the same group of 

species as E. afra and appears to be distinguishable from that 

species. The features observable indicate that the smaller 

specimen belongs to the same species as the larger one, but 

some features crucial for species identification are missing and 

the association between the two specimens is by no means 

certain. The species, as defined by the large syntype, is 

compared to other species in group B-4 in Tables 33 and 36. 

Eunice pruvoti is the only species listed in Table 36 with 

exclusively single branchial filaments. 

163. Eunice pulvinopalpata Fauchald, 1982 


Eunice pulvinopalpata Fauchald, 1982b:781-785, fig. la-f. 

MATERIAL EXAMINED.—Holotype and paratype, USNM 

74304 and 74305, Pacific Ocean off western Mexico, 20°50'N, 

109°06'W, 2633 m, Alvin dive 1214, sample #7, 20 Apr 1982, 

at base of white smoker. (Other paratypes in BM(NH) and 

AHF). 


length 450 mm long; maximal width 11 mm; length through 

setiger 10, 23 mm. Paratype examined complete with 353 

setigers; total length 320 mm with posterior end in regeneration; 

maximal width 11 mm; length through setiger 10,18 mm. 

Body anteriorly cylindrical, becoming ventrally flattened by 

setiger 100, retaining strongly convex dorsum in all setigers. 

Body tapering abruptly anteriorly and slowly posteriorly; even 

in last setigers body 6 mm wide. 




rim; median sulcus deep. Peristomial fold deep. Eyes not 

observed. Antennae in a horseshoe, evenly spaced, similar in 


articulations. Ceratostyles slender and tapering, with up to 19 

irregularly cylindrical articulations in A-III. A-I to posterior 

peristomial ring; A-I I and A-III to setiger 2. Peristomium 

tapering anteriorly, with a distinct muscular lower lip. 



middle or front edge of prostomium, slender and tapering, with 

-12 irregular, cylindrical articulations. 

Maxillary formula (examined in USNM 74305) 1+1, 17+17,

278 


2 

1 

.02 

.1 

a 

e 

b, c 

FIGURE 93.—Eunice pulvinopalpata (holotype, USNM 74304): a, anterior end, lateral view; b, compound 

fakiger, parapodium 105; c, pectinate seta, parapodium 105; d, subacicular hook, parapodium 105; e, parapodium 

105, anterior view. (Scale bars in mm.)


18+0, 12+12, and 1+1. Mx III behind left Mx II. Both Mx IV 

with same number of teeth. All teeth very small and even in 

size. 



Branchiae from setiger 3 to end of body. Branchiae present to 


of setigers. First pair single filaments, all other branchiae with 

at least 4 filaments; maximally 25 filaments from setiger 15 to 

setiger 275; thereafter number slowly reduced; 15 filaments 

present in last setigers. Branchial stem thick, folded over in 

most parapodia. Filaments filiform. 

Neuropodial acicular lobes asymmetrically rounded with 

aciculae emerging on dorsal side. All presetal lobes obliquely 

truncate. All postsetal lobes low, transverse folds. First 5 or 6 

ventral cirri thick, digiliform. Ventral cirri basally inflated by 

setigers 6-7; inflated bases retained through about setiger 250. 

Inflated bases nearly spherical; narrow tips tapering. Far 

posterior ventral cirri tapering. Notopodial cirri very long, 

especially in anterior part of body. Where best developed, at 

about setiger 100, longer than width of body, slender, tapering, 



93c) tapering, flat. One marginal tooth very much longer than 


(Figure 93b) slightly inflated, marginally smooth. Appendages 

tapering, bidentate. Both teeth of same size, slender, tapering. 

Proximal teeth directed distally; distal teeth gently curved. 


falcigers and compound spinigers absent. Most 

aciculae in triplets, dark brown to black, with tapering slender 

tips, straight; cross-section round. Separation between cores 

and sheaths indistinct in both aciculae and subacicular hooks. 

Subacicular hooks (Figure 93d) dark brown to black, bidentate. 

Hooks first present from setiger 68 in holotype (from setigers 

62-72 in paratypes), present in all setigers thereafter, always 

single (except for replacements). Hooks slender, tapering to a 

small head. Proximal teeth slightly larger than distal teeth; both 

teeth directed distally. 


anal cirri. 





Characters: 1, 2,4, 6, 13, 14, 36. 


None. 

REMARKS.—Eunice pulvinopalpata is a member of group 

B-2; however, it also resembles some of the deep-water 

members of group A-1 and should be compared to these species 

as well (Tables 19-21). It is branchiated to near the posterior 

end, in contrast to most other deep-water species. It is listed 

with similar species in group B-2 in Tables 27 and 31. The only 

and 

other species in Table 31 with more than 20 branchial filaments 

is E. franklini. The branchial filaments in the latter are very 

short and stubby; those in E. pulvinopalpata are slender, 

digitiform and certainly not unusually short. 

Eunice punctata (Risso, 1826) 

Ltodice punclata Risso, 1826:421-422. 

Eunice punctata.—Quatrefages, 1866:331. 

REMARKS.—This species, of which no authoritative material 

exists, was considered either indeterminable or doubtfully as a 

member of Marphysafallax by Fauvel (1923:451). The species 

described by Heider (1925) under this name cannot be a 

Marphysa because it has peristomial cirri; it is here considered 

as E. harassii. Eunice punctata (Risso) is indeterminable. 

Eunice punclata Peters, 1854:611. 

Eunice punctata Peters, 1854 

REMARKS.—The specific name is preoccupied in the 

combination E. punctata (Risso, 1826). Peters' species is 

redescribed above as E. petersi. 

Eunice punctata Grube, 1856 

Eunice punctata Grube, 1856:59-60. 

Eunice binominata Quatrefages, 1866:327. 

REMARKS.—Quatrefages (1866:327) pointed out that 

Grube's specific name was pre-occupied by Risso (1826); 

Quatrefages apparently had overlooked Peters' use of the name, 

issued two years before Grube *s paper, so the name was in fact 

doubly pre-occupied by the time Quatrefages reviewed the 

matter. Grube's species is here discussed as E. binominata. 

164. Eunice purpurea Grube, 1866 


Eunice purpurea Grube, 1866b:68. 

Eunice vlolacea.—Grube, 1861:60-61 [not Eunice violacea Grube, 1856]. 

MATERIAL EXAMINED.—Two syntypes, ZMB F 2012, 

Portor6, Lesina, Adriatic Sea, coll. Grube. 


based wholly on the large specimen; some notes on the small 

specimen, which is a juvenile, have been appended. The 

peristomium of the large specimen has been ventrally dissected 

and the jaw apparatus torn partially loose; the illustration shows 

the condition with the jaws tucked back into position, but is in 

part a reconstruction where tissue had been removed. 

DESCRIPTION.—Large syntype complete with 106 setigers; 

total length 53 mm; maximal width 4 mm; length through 

setiger 10, 9.5 mm. Other syntype with 40 setigers; length 7 

mm long. 

Prostomium (Figure 94a) distinctly shorter and narrower

280 


J 

.02 

.1 

1 

a 

r. e 

d 

FIGURE 94.—Eunice purpurea (syntype, ZMB F 2012) : a. anterior end, lateral view; b, parapodium 32, anterior 

view; c, pectinate seta, parapodium 32; d, subacicular Hook, parapodium 32; e, compound falciger, parapodium 



than peristomium, as deep as x l2 of the peristomium. Prostomial 


Peristomial fold completely everted making region posterior to 

antennal bases appear very long. Eyes between bases of A-I and 

A-II. Antennae in a horseshoe, evenly spaced, similar in 



articulations in A-III. A-I to posterior peristomial ring; A-II and 

A-I 11 to setiger 2. Peristomium slightly flaring anteriorly, with 

distinct muscular lower lip. Separation between rings visible 

ventrally, slightly better marked dorsally; anterior ring 4 /s of 


peristomial ring, digitiform, with 3 cylindrical rings. 

Maxillary formula 1+1,4+5, 7+0, 2+7, and 1+1. Mx III part 

of distal arc with left Mx IV. Left Mx IV continued as edentate 

plate behind Mx III. Mx VI absent. 





of setigers. First 3 and last 10 pairs single filaments; maximum 

14 filaments at about setiger 20. First 2 pairs short, nearly 

button-shaped. Branchial stems long, erect, often curved in half 

corkscrew posteriorly. Filaments short, slender. 

Neuropodial acicular lobes distally rounded, very low; 

aciculac emerging at midlinc. All pre- and postsetal lobes 

follow outline of acicular lobes closely. First 4 ventral cirri 

tapering. Ventral cirri with inflated bases between setigers 5 

and 50. Inflated bases thick, transverse welts; narrow tips short 

and button-shaped. Posterior to setiger 50 ventral cirri 

becoming tapering, increasing in relative length, becoming 

about as long as notopodial cirri in last setigers present First 50 

notopodial cirri basally distinctly inflated, with decreasing 

numbers of articulations, up to 4 articulations; last articulated 

notopodial cirrus in setiger 55. Posterior notopodial cirri 

increasingly slender, decreasing in length to roughly 2 /3 of 

length in anterior setigers. 

Limbate setae marginally smooth. Median pectinate setae 

(Figure 94c) flaring, flat. One marginal tooth distinctly longer 

than other teeth; -15 teeth present. Compound falcigers (Figure 

94e) very large. Shafts tapering, marginally smooth. Appendages 

tapering, with large heads, bidentate. Proximal teeth 

longer than distal teeth, triangular, directed laterally. Distal 

teeth tapering relatively abruptly, distinctly bent. Guards 

symmetrically rounded, marginally serrated; mucros absent. 


Aciculae paired, with brown cores and clear sheaths, tapering; 

cross-sections round. Subacicular hooks (Figure 94d) with 

brown cores and clear sheaths, bidentate. Hooks first present 


(except for replacements). Hooks very slender, tapering to 

small head. Proximal teeth very much larger than distal teeth, 

triangular, directed laterally. Distal teeth slender, curved. 


anal cirri. 







None. 

REMARKS.—Eunice purpurea is compared to similar species 

in Tables 27 and 30. Of species listed in Table 30 with 10 or 

more branchial filaments where the branchiae are best 

developed, E.flavopicta and E. reducta lack articulations in the 

peristomial cirri. The other heavily branchiated species have 

such articulations. Posterior ventral cirri are basally inflated in 

E. coccinea, widely triangular, nearly scoop-shaped in E. 

laticeps and E. violaceomaculata, tapering from narrow bases 

in E. purpurea, and unknown for E. macrobranchia. Eunice 

purpurea appears to have the branchiae distinctly outreaching 

the notopodial cirri; in E. macrobranchia the notopodial cirri 

are of about the same length as the branchiae. 

The juvenile has lost all antennae; branchiae are present from 

setiger 5 and continued to near the posterior end. Where best 

developed at setiger 15, each branchia with two relatively long 

filaments. Aciculae and subacicular hooks brown; the latter are 

present from setiger 15. The juvenile very probably belongs to 

the same species as the larger specimen, but this cannot be 

confirmed presently. 

165. Eunice pyenobranchiata Mclntosh, 1885 

FIGURE 95a-f; TABLES 14,27,32 

Eunice pyenobranchiata Mclntosh, 1885:294-297, figs. 54, 55, pi. 39: figs. 

13-15, pi. 21 A: figs. 4,5.—Fauchald, 1986:253-255, figs. 56-61. 

Eunice tentaculata.—Fauvel, 1917:269 [in part, not Eunice tentaculata 

Kinberg, 1865, nor Eunice tentaculata Quatrefages, 1866]. 

MATERIAL EXAMINED.—Three syntypes, BM(NH) ZK 

1921.5.1.1997, Challenger sta 162, off East Moncoeur land, 

Bass Strait, 39°10'30"S, 146°37'E, 69 m, dredged, sand and 

shells, 2 Apr 1874. One syntype, BM(NH) ZK 1921.5.1.1998, 

Challenger sta 163A, Twofold Bay, between Melbourne and 

Sydney, 36°59'S, 150°20 / E, 274 m, trawled, green mud. 

COMMENTS ON MATERIAL EXAMINED.—The syntype from 

Twofold Bay was the one examined in detail by Mclntosh; 

most of the description is based on this specimen. The 

specimen has been frontally dissected and the jaws are now 

missing. 

DESCRIPTION.—Syntype described incomplete with 63 setigers; 

length 50 mm; maximal width 8 mm; length through 

setiger 10, 12 mm. Other syntype incomplete with 76 setigers; 


10, 10 mm. 

Prostomium (Figure 95a,b) distinctly shorter and narrower 

than peristomium, less than x fa as deep as peristomium. 


sulcus deep. Antennae in a horseshoe, with A-III isolated by a

282 


FIGURE 95.—Eunice pycnobranchiala (syntype, BM(NH) ZK 1921.5.1.1997 (b); BM(NH) ZK 1921.5.1.1998 (a 

and c-f): a, anterior end, lateral view; b, anterior end, lateral view; c, parapodium 32, anterior view; d, subacicular 

hook, parapodium 32; e, compound falciger, parapodium 32; f, pectinate seu, parapodium 32. Eunice quinquefida 

(holotype, USNM 15714): g, anterior end, lateral view; h, parapodium 19, anterior view; i, compound falciger, 

parapodium 32; j, pectinate seta, parapodium 19; k, parapodium 35, anterior view; 1, pectinate seta, parapodium 

35; m, subacicular hook, parapodium 35; n, aciculae, parapodium 35. (Scale bars in mm.)


TABLE 14.—Variable and invariable features in the type lot of Eunice pycnobranchiata (N = number of 

individuals examined; SD = standard deviation; measurements in mm; * = SD not calculated). 



Total length 

Maximal width 








N 

2 

2 

4 

4 

4 

4 

4 

4 

4 

4 

Max. 

140 

105 

7 

13 

17 

5 

8 

5 

50 

31 

Min. 

129 

70 

2 

5 

8 

3 

5 

1 

32 

23 

Mean 

134.5 

87.5 

5 

9.5 

11.75 

4 

6 

3.5 

40.75 

27.5 

SD 

* 

* 

2.16 

3.32 

3.86 

1.15 

1.41 

1.91 

7.37 

3.42 


Separation between rings distinct 


Aciculae distally 



N=4 

dorsally and ventrally 

brown 


brown 

2 

gap, with A-I slimmer than other 3. Ceratophores ring-shaped 


with up to IS irregular, cylindrical articulations. Antennae 

reaching posterior peristomial ring; A-I I and A-I 11 similar in 

length; A-I somewhat shorter. Peristomium cylindrical with 

distinct muscular lower lip. Separation between rings distinct 

dorsally and ventrally; anterior ring about 

5 /« of total 


peristomial ring, with 3 articulations. 

Maxillary formula according to Mclntosh (1885, fig. 54) 

1+1, 5+5,6+0, 6+8, 1+1. Mx III part of distal arc with left Mx 

IV and V. 



5-6 to end of fragments. Where best developed branchiae with 

4 short thick filaments. 

All neuropodial acicular lobes (Figure 95c) rounded; 

aciculae emerging dorsal to midline. Presetal lobes low, 

transverse folds. Anterior postsetal lobes forming collars 

around dorsal edge of acicular lobes. Median postsetal lobes 

low, transverse folds. Anterior ventral cirri digitiform. Median 

ventral cirri with barely visible inflated bases. Inflated bases 

ovate; narrow tips digitiform. Anterior notopodial cirri digitiform, 

with 2 to 4 irregular articulations. Median notopodial 

cirri basally somewhat inflated, without articulations. 

Limbate setae slender. Pectinate setae (Figure 950 flat, 

flaring. Both marginal teeth slightly longer than other teeth; 

-15 teeth present. Shafts of compound falcigers (Figure 95e) 

tapering, marginally smooth with internal striations. Appendages 

short, bidentate. Proximal teeth shorter than distal teeth, 

triangular, directed laterally. Distal teeth tapering, nearly erect. 

Guards distally asymmetrically bluntly pointed; mucros absent. 


Aciculae single in first few setigers, thereafter paired, black, 

tapering, straight; cross-section round. Subacicular hooks 

(Figure 95d) black, bidentate. Hooks first present from setiger 

28 or 29, present in all setigers thereafter, always single (except 

for replacements). Proximal teeth much larger than distal teeth; 

both teeth directed obliquely distally. 






IV. 



Characters: 1, 2,4, 6, 13, 14, 34, 36-40,42, 47, 50, 51, 60, 

63,74. 


38,1. 

REMARKS.—The specimen from Bass Strait differs slightly 

from the specimen from Twofold Bay in length of antennae and 

in development of branchiae. Branchiae more prominent in 

Bass Strait specimens than in the other specimen. These 

differences are not considered to be of value as specific 

characters. Mclntosh commented on the variability in the 

number of teeth in the jaws. Such variability was not observed 

in the current study. 

Eunice pycnobranchiata is compared to similar species in 

Tables 27 and 32. It is the only species in Table 32 in which the 

branchiae and the notopodial cirri are similar in length; in two 

species, the branchiae are distinctly shorter and in the others, 

they are distinctly longer than the notopodial cirri. The length 

differences are very obvious in all cases. 

The species has been considered a junior synonym of E. afra 

and E. laticeps. It differs clearly from E. afra by having 

branchiae from setigers 5-6 rather than from setiger 16. Eunice

284 


laticeps has distally cylindrical antennal articulations and 

maximally about 18 branchial filaments; E. pycnobranchiata 

has distally drop-shaped antennal articulations and maximally 

four branchial filaments. The proposed synonyms cannot be 

accepted. 

Eunice quadrioculata Grube, 1856 

Eunice quadrioculata Gnibe, 1856:60. 

Marphysa quadrioculata.—Quatrefages, 1866:337.—Grube, 1878a: 101 -102. 

REMARKS.—This species, of which Grube actually only saw 

an illustration made by A.S. Orsted, has been considered a 

member of the genus Marphysa because Quatrefages first 

referred it to that genus. 

166. Eunice quinquefida Moore, 1903 

FIGURE 95g-n; TABLES 27,32 

Eunice quinquefida Moore. 1903:435-437, pi. 25: figs. 39-41. 


sta 3698, Japan, Sagami Bay, 4.5 miles off Manazuru Zaki 

(approximate position 35 o l(rN, 139°10'E),5 May 1900,280 m, 

green mud, volcanic ash and sand, beamtrawl. 



10, 8 mm. Body anteriorly cylindrical, becoming slightly 

dorsoventrally flattened by setiger 25-30. 



Prostomial lobes frontally rounded, dorsally strongly inflated; 

median sulcus deep. Eyes between bases of A-I and A-II, dark. 

Antennae in a horseshoe, with A-I isolated by a gap, similar in 


articulations. Ceratostyles slender and tapering, separated 

irregularly into articulations; maximum 10 articulations present 

in A-II. Articulations cylindrical basally drop-shaped distally, 

especially in A-I. A-I to posterior peristomial ring; A-II to 

setiger 4; A-III to setiger 3 (probably incomplete). Peristomium 

slightly flaring anteriorly; lower lip distinct, muscular. Separation 

between rings very distinct dorsally and ventrally, 

indistinct only for a short distance laterally; anterior ring 5 /6 of 

total peristomial length. Peristomial cirri to front edge of 

peristomium, slender and tapering, with 3 cylindrical articulations. 


of distal arc with left Mx IV. Teeth on Mx II slender and 

sharply pointed; other teeth short and delicate. 

Branchiae (Figure 95h,k) present, pectinate, distinctly longer 


Branchiae from setiger 7 to end of fragment. All branchiae with 

at least 2 filaments; maximum 5 filaments at setigers 15-20. 

Branchial stems slender. Filaments slender, nearly filiform. 

Filaments as long as notopodial cirri. 

All neuropodial acicular lobes symmetrically conical; 

aciculae emerging at midline. Presetal lobes obliquely truncate. 

Postsctal lobes low, transverse fold. First 4 or 5 ventral cirri 

thick, tapering. Ventral cirri strongly inflated basally from 

setiger 6-7. Inflated bases nearly spherical in setigers 15-25, 

changing to thick transverse welts from about setiger 26, 

becoming gradually reduced from about setiger 30, missing 

posterior to setiger 40. Narrow lips tapering in all ventral cirri 

with inflated bases. Far posterior ventral cirri slender, tapering. 

Notopodial cirri with distinct thickened cirrophorcs; cirrostyles 

thick, tapering, without articulations. 

Limbate setae marginally frayed. Anterior pectinate setae 

(Figure 95j) tapering, fiat. One marginal tooth distinctly longer 

than other teeth; 8-9 teeth present. Median and posterior 

pectinate setae (Figure 951) distinctly furled, retaining same 

shape and number of teeth. Shafts of compound falcigers 

(Figure 95i) inflated, marginally finely serrated. Appendages 

tapering, bidentatc. Proximal and distal teeth similar in size. 

Proximal teeth short, truncate, directed laterally. Distal teeth 

curved. Guards symmetrically sharply pointed, without 

mucros. Pscudocompound falcigers and compound spinigers 

absent. Aciculae single in anterior parapodia, paired elsewhere 

(Figure 95n), with brown cores and clear sheaths, tapering, 

sharply bent, sometimes in double curves; cross-section round. 

Subacicular hooks (Figure 95m) with brown cores and clear 

sheaths, bidentatc. Hooks first present from setiger 30, present 

in all setigers thereafter, paired in most setigers. Hooks slender, 

tapering, with small head. Proximal teeth larger than distal 

teeth, triangular, laterally directed. Distal teeth short, erect 







Characters: 1, 2,4, 6, 23, 36-40,47, 50, 63. 


38,1. 

REMARKS.—Because of the incomplete condition of the type 

of E. quinquefida, the species was listed in group B-5 by 

Fauchald (1970). It resembles species in group B-2, and is here 

compared to species in Tables 27 and 32. It is one of two 

species in 32 without articulated notopodial cirri. The other 

species, E. rubella, has very short antennae barely outreaching 

the prostomium. Although A-III is probably incomplete in the 

type of E. quinquefida, A-II reaches setiger 4. 

Eunice quoya Quatrefages, 1866 

Eunice quoya Quatrefages. 1866:318.—Fauchald. 1986:255. 

REMARKS.—The original diagnosis of this species is 

incomplete and no material is available. It was characterized as 

indeterminable by Fauchald (1986) and no new evidence has 

been come to light to change that opinion.


167. Eunice reducta Fauchald, 1970 

FIOURE 96a-h; TABLES 27, 30 

Eunice reducta Fauchald. 1970:39-43, pi. 5: figj. a-i. 

MATERIAL EXAMINED.—Holotype, AHF Poly 0338, Gulf of 

California, Mexico, entrance of Bahia Agua Verde, from 

25°31'36"N, lll°03'15"W to 25°31'40"N, lll o 04'13"W, 42- 

48 m, dredge, sand, mollusks, crabs, 17 Mar 1949, Velero IV, 

sta 1743-49. 



10, 15 mm. Body anteriorly truncate, cylindrical, tapering 

slowly posteriorly to slender posterior end with crowded short 

segments; most body segments only slightly wider than long; 

parapodia strictly lateral. Paired anal cirri barely longer than 

pygidium, without articulations; dorsal edge of pygidium 

smooth. 


than peristomium, as deep as 

x li of the peristomium. 

Peristomial lobes frontally obliquely truncate, dorsally excavate 

with a thickened rim; median sulcus shallow, but wide. 

Eyes posterior to bases of A-I, black. Antennae in a horseshoe, 



with up to 3 cylindrical, indistinct articulations. A-I to middle 

of anterior peristomial ring; A-I I and A-I II to posterior 

peristomial ring; A-I 11 slightly longer than A-I I. Peristomium 

cylindrical; lower lip distinct muscular. Separation between 

rings distinct dorsally and ventrally; anterior peristomial ring 

4 /5 of total peristomial length. Peristomial cirri to middle of 

anterior peristomial ring, tapering, without articulations. 

Maxillary formula 1+1, 444, 8+0, 4+10, and 1+1 according 

to Fauchald (1970:42). 




well before posterior end, present on less than 55% of the total 

number of setigers. Last 6 pairs single filaments, all others with 

at least 2 filaments; maximum number of filaments 21 in a trim 

series. Branchial stems long, erect, tapering. Filaments short, 

digitiform. Some filaments furcate. Filaments and notopodial 

cirri similar in length. 

Anterior neuropodial acicular lobes distally truncate with 

aciculae emerging at midline. Posterior neuropodial acicular 

lobes conical with aciculae emerging at tip. All pre- and 

postsetal lobes low folds. First 4 ventral cirri thick, tapering. 

Ventral cirri basally inflated from about setiger 5 through about 

setiger 50. Inflated ventral cirri large, irregularly rounded; 

narrow tips absent. Ventral cirri increasingly digitiform in far 

posterior setigers, but always shorter than notopodial cirri. 

Anterior notopodial cirri thick, medially inflated, decreasing 

rapidly in width posteriorly. Postbranchial notopodial cirri half 

as long as anterior notopodial cirri, slender, digitiform. 

Pre-branchial and branchial notopodial cirri with 2 or 3 

articulations; postbranchial notopodial cirri without articulations. 

Limbate setae tapering, marginally pilose. Pectinate setae 

(Figure 96d,f) with flattened shafts; blades flat, flaring. One 

marginal tooth very much longer than other teeth; -20 teeth in 

median and posterior setae; -10 in anterior setae. Shafts of 

compound falcigers (Figure 96b,e,h) thick, tapering, marginally 

smooth. Appendages short, triangular, bidentate. Proximal 

teeth very much smaller than distal teeth; anterior proximal 

teeth small, distally directed spur, posterior proximal teeth low, 

triangular elevations. Distal teeth very large, blunt, erect. 



paired in anterior setigers, single in posterior setigers, dark 

brown to black, tapering, blunt-tipped; cross-section round. 

Subacicular hooks (Figure 96c) dark brown to black, bidentate. 



tapering gently to small head. Proximal teeth about half as large 

as distal teeth; both teeth directed distally. 


Mx III and left Mx IV. 


TURES.—Mx III forming part of a distal arc with left Mx IV. 



Characters: 4,6,42, 74,78. 


None. 

REMARKS.—Fauchald (1970) indicated that subacicular 

hooks could be present from setigers 34-49 in non-type 

specimens of this species from western Mexico. Eunice reducta 

belongs to group B-2, but differs from most species in the 

group by having a long abranchiate posterior end. It is 

compared to similar species in Tables 27 and 30. The two other 

species in Table 30 with abranchiate posterior end, E. coccinea 

and E. fimbriata, have 10 and five branchial filaments 

respectively; E. reducta has 21 filaments where the branchiae 

are best developed. 

Eunice rissoi Quatrefages, 1866 

Eunice rissoi Quatrefages, 1866:315-316, pi. 10: fig.5. 

Eunice vittata.—Gmbc, 1870a:295 [in part, not Nereis vittata Chiaje, 1829]. 

REMARKS.—No type specimens are available. 

In the original description A-I 11 is mentioned as being very 

much longer than the other antennae; peristomial cirri were 

described as long; branchia present from setiger 3 (ring 4) with 

up to 5 filaments. 

Grube in his review of material in the Paris Museum 

indicated that branchiae began on setiger 4 and were absent 

from the last 10 of a total of 105 segments present in a complete 

specimen. A-I 11 reached setiger 4 and had a few long

286 


FIGURE 96.—Eunice reducta (holotype, AHF Poly 0338): a, anterior end, lateral view; b, compound falciger, 

parapodium 275; c, subacicular hook, parapodium 275; d, pectinate seta, parapodium 275; e, compound falciger, 

parapodium 3; f, pectinate seta, parapodium 3; g, parapodium 10, anterior view; h, compound falciger, 

parapodium 10. Eunice rosaurae (syntype, BM(NH) ZK 1941.1.1.217-221): i, anterior end, lateral view; j, 

parapodium 25, anterior view; k, parapodium 68, anterior view; 1, pectinate seta, parapodium 25; m, compound 

falciger, parapodium 25; n, acicula, parapodium 68; o, pectinate seta, parapodium 68; p, subacicular hook, 



articulations. Grube referred the species, incorrectly, to E. 

vittata. The latter lacks branchiae on the posterior half of the 

body. 

Grube suggested that E. rissoi might be the same as E. 

rubrocincta Ehlers; the branchial distribution certainly is closer 

to the latter than it is to E. vittata. However, Grube's suggestion 

is certainly not adequate to propose a synonymy. 

The information available is insufficient to characterize E. 

rissoi and it is here considered indeterminable. 

168. Eunice rosaurae Monro, 1939 

FIGURE 96i-p; TABLES 24,25.52,53 

Eunice rosaurae Monro. 1939:351-352, fig. 282 a-f [in part]. 

MATERIAL EXAMINED.—Two syntypcs, ZK 1941.1.1.217- 

221, off St. George, Grenada, 12°05'N, 61°49'W, 720-800 m, 

27 Nov 1937, trawled, Rosaura Expedition, Atlantic, 1937-38, 

sta34. 

COMMENTS ON MATERIAL EXAMINED.—Two of the three 

original syntypes fit the species as described by Monro; the 

third syntype belongs to E. collini and is treated above. Of the 

two syntypes, one had been dissected and the jaws were 

separated; it agrees with the specimen described in detail by 

Monro and forms the base of the description given below. 

DESCRIPTION.—One syntype complete with 142 setigers; 

total length 152 mm; maximal width 4 mm; length through 

setiger 10, 18 mm. Other syntype nearly complete with 138 




peristomium, as deep as ! /2 of peristomium. Prostomial lobes 

frontally obliquely truncate, dorsally somewhat flattened; 

median sulcus deep. Eyes between bases of A-I and A-II, black. 


Ccratophores ring-shaped in all antennae, without articulations. 

Ceratostyles medially inflated; inner half without articulations; 

outer half of ceratostyles increasingly articulated; outer 

articulations distinctly moniliform; maximum 13 articulations 

in A-III. A-I to posterior margin of peristomium; A-II to setiger 

3; A-III to setiger 3, but slightly longer than A-II. Peristomium 

cylindrical. Separation between rings distinct dorsally only; 


front edge of posterior peristomial ring, medially inflated, 



Branchiae (Figure 96j) present, single filaments, distinctly 

shorter than notopodial cirri, not reduced in mid-body region. 



number of setigers. First and last 10 branchiae small tubercles. 

All other branchiae single, slender filaments. 

Anterior neuropodial acicular lobes distally truncate with 

aciculae emerging near upper corner of lobes. Posterior acicular 

lobes distally triangularly rounded with both aciculae emerging 

centrally. Anterior and median pre- and postsetal lobes follow 

outline of acicular lobes closely. Posterior pre- and postsetal 

lobes low, transverse folds. First 5 ventral cirri basically 

digitiform, slightly tapering. Ventral cirri basally inflated from 

setiger 6 through branchial region. Inflated bases ovate; narrow 

tips digitiform. Posterior ventral cirri retaining short, inflated 

bases as sausage-shaped thickenings along ventral edge of 

parapodia; narrow tips digitiform. Prebranchial notopodial cirri 

slender, gently inflated basally; branchial notopodial cirri 

somewhat more inflated. Postbranchial notopodial cirri (Figure 

96k) slender with distinct separation between cirrophores and 

cirrostyles. 

Limbate setae slender, marginally obscurely serrated. Pectinate 

setae (Figure 961,o) tapering, flaL One marginal tooth 

distinctly longer than other teeth; -10 teeth present. Shafts of 

compound falcigers (Figure 96m) distally tapering, marginally 

serrated. Appendages short, bidentate. Proximal teeth smaller 

than distal teeth, very short, tapering, curved laterally. Distal 

teeth gently curved, slender. Guards bluntly pointed, marginally 

serrated; mucros absent. Pseudocompound falcigers and 

compound spinigers absent. Aciculae mostly paired, single in 

anterior setigers, with black cores and clear sheaths, distally 

tapering, gently curved (Figure 96n). Subacicular hooks 

(Figure 96p) with black cores and clear sheaths, sometimes 

including all teeth; some hooks distally tridentate with 2 large 

distal teeth in tandem and small, nearly clear proximal teeth. In 

other hooks distal teeth fused, so hooks apparently bidentate. 

Bidentate hooks in full lateral view identical in appearance to 

tridentate ones, with both teeth pointing distally. Hooks first 

present from setiger 38 (32 in the other specimen), present in all 

setigers, always single (except for replacements). 

be- 

UNKNOWN MORPHOLOGICAL FEATURES.—Relationship 

tween Mx III and left Mx IV; pygidium and anal cirri. 







None. 

REMARKS.—Eunice rosaurae is listed with similar species in 

Tables 24 and 25. It resembles two additional species in Table 

25 in having short branchiae consisting of single strap-like 

filaments. These two species, E. gravieri and E. nicidioformis, 

have exclusively bidentate subacicular hooks. Tridentate 

subacicular hooks with the teeth in tandem present in some 

neuropodia have not been found in any other species in group 

Bl. A total of four species are known to have the distal teeth in 

tandem in the genus; these are listed in Tables 52 and 53. Two 

of these have light yellow aciculae and subacicular hooks. The 

only other species with dark aciculae and subacicular hooks is 

E. cirrobranchiata. All aciculae are pointed in E. rosaurae: the 

aciculae are in part hammer-headed in E. cirrobranchiata.

288 


169. Eunice roussaei Quatrefages, 1866 


Eunice roussaei Quatrefages, 1866:309-311, pi. 10: fig. 1-4.—

NUMBER 523 

289 

FIGURE 97.—Eunice roussaei (MNHN, slides from Santander): a, anterior parapodium, anterior view; b, 

compound falciger, anterior parapodium; c, pectinate seta, anterior parapodium; d, compound falciger, 

medioposterior parapodium; e, pectinate seta, medioposterior parapodium; f, medioposterior parapodium, 

anterior view; g, subacicular hook, medioposterior parapodium; h, compound falciger, median parapodium; i, 

pectinate seta, median parapodium; j, median parapodium, anterior view; k, compound falciger, posterior 

parapodium; 1, pectinate seta, posterior parapodium; m, subacicular hook, posterior parapodium; n, posterior 

parapodium, anterior view. (Scale bars in mm.)

290 


.4 

n 

s 

FIGURE 98.—Eunice rubella (holotype, Canterbury Museum): a, anterior end, lateral view; b, pectinate seta, 

parapodium 45; c, subacicular hook, parapodium 45; d, compound falciger, parapodium 45; e, parapodium 45, 

anterior view. Eunice rubra (holotype, ZMC): f, anterior end, lateral view; g, pectinate seta, parapodium 20; h, 

compound falciger, parapodium 20; i, parapodium 20, anterior view; j, parapodium 49, anterior view; k, 

compound falciger, parapodium 49; 1, pectinate seta, parapodium 49; m, subacicular hook, parapodium 49; n, 

aciculae, parapodium 49; o, pectinate seta, parapodium 111; p, subacicular hook, parapodium 111; q, compound 

falciger, parapodium 111; r, aciculae, parapodium 111; s, parapodium 111, anterior view.


antennae, without articulations. Ceratostyles relatively stout 

and digitiform, with up to 7 moniliform articulations in A-II; 

innermost articulations half length of each antenna. Antennae 

reaching middle of peristomium. Peristomium cylindrical, 

more than twice as long as prostomium. Separation between 

rings distinct dorsally and laterally, indistinct ventrally; 


about middle of peristomium, tapering, with 3 articulations. 


and 1+1 according to Knox. 



to end of fragment. Where best developed, between sctigers 7 

and 29, branchiae with 2 short, thick filaments; otherwise all 

branchiae with single filaments (Figure 98e). 

Anterior ncuropodial acicular lobes very short, distally 

truncate; aciculae emerging at midline. Pre- and postsetal lobes 


Ventral cirri basally from setiger 8 through setiger 27. Inflated 

bases ovate; narrow tips tapering. Posterior ventral cirri large; 

open scoops covering emergent parts of subacicular hooks. 

Anterior notopodial cirri thick and medially inflated, with 3 

articulations. Notopodial cirri retaining same size in all 

segments present, becoming digitiform by setiger 20, without 


Pectinate setae (Figure 98b) tapering, flat. Both marginal 

teeth slightly longer than other teeth; -12 teeth present. Shafts 

of compound falcigers (Figure 98d) tapering, marginally finely 

dentate. Appendage short, rather wide, bidentate. Both teeth 

similar in size, short and blunt Guards bluntly truncate, 


and compound spinigers absent. Aciculae paired in median and 

posterior parapodia, black, very thick, tapering, bent dorsally; 

cross-sections round. Subacicular hooks (Figure 98c) black, 



Hooks curved, with large head. Proximal teeth larger than distal 

teeth, triangular, directed laterally. Distal teeth directed 

dorsally. 


Mx III and left Mx IV; some features associated with 






Characters: 1, 2,4, 6, 36-38,40, 74, 78. 


38,1. 

REMARKS.—Eunice rubella differs from most other species 

with black, bidentate subacicular hooks by having very 

strongly beaded ceratostyles and by the poor development of 

branchiae. It is listed with similar species in Tables 27 and 32; 

it is the only species in Table 32 with the separation between 

peristomial rings visible only dorsally; in all other species the 

separation is visible also ventrally. 

Branchiae were present to setiger 80 according to Knox; the 

distribution on the fragments present agrees with this pattern. 

Eunice rubra Grube, 1856:59. 

171. Eunice rubra Grube, 1856 

FIGURE 98f-s; TABLES 46,47 

MATERIAL EXAMINED.—Holotype, ZMC, St. Thomas, West 

Indies, coll. A.S. Orsted and H. Krdyer. 

DESCRIPTION.—Holotype nearly complete with 118 setigers; 


10, 7.5 mm. 

Prostomium (Figure 980 about as long as peristomium, 

distinctly narrower than peristomium, as deep as x h- of the 

peristomium. Prostomial lobes forming blunt 3-sided pyramid, 

frontally obliquely truncate, dorsally flattened; median sulcus 

shallow. Palpal region marked off by shallow horizontal 

groove. Eyes between bases of A-I and A-II. Antennae in a 



digitiform, with up to 23 moniliform articulations in (apparently 

incomplete) A-III. A-I to middle of anterior peristomial 

ring. Longest A-II to setiger 1; incomplete A-III to setiger 3. 




slender and tapering, with 10 cylindrical articulations. 


Branchiae (Figure 98i,j,s) present, pectinate, distinctly 

longer than notopodial cirri, not reduced in mid-body region, 

erect. Branchiae from setiger 4 to end of fragment (-15 setigers 

missing). First branchia with 2 filaments; all other branchiae 

strongly pectinate with at least 6 filaments. Maximum 

filaments 21 at about setiger 15. Number of filaments reduced 

to 6 by setiger 40; this number retained in rest of specimen. No 

increase in number of filaments towards posterior end, but a 

slight increase in length of filaments present. 

All neuropodial acicular lobes distally rounded; aciculae 


transverse folds. Presetal lobes distinctly shorter than acicular 

lobes in anterior and median setigers, following outline of 

acicular lobes closely in posterior setigers. First 3 ventral cirri 

thick and tapering. Ventral cirri basally inflated from about 


tapering. Posterior ventral cirri increasingly digitiform, about 

half as long as notopodial cirri in median and posterior setigers. 

All notopodial cirri tapering from slightly inflated bases. 

Anterior notopodial cirri with up to 6 articulations; median and 

posterior notopodial cirri with 3 to 4 articulations. 


(Figure 98g,l,o) tapering, flat. One marginal tooth distinctly

292 


longer than other teeth; -10 teeth present. Shafts of anterior and 

median compound falcigers (Figure 98hjc) inflated, marginally 

serrated; those of posterior falcigers tapering and smooth 

(Figure 98q). Appendages thick with very nearly parallel sides, 


directed laterally. Distal teeth nearly straight, directed obliquely 

distally. Guards asymmetrically bluntly pointed; mucros 


absent. Aciculae paired, yellow. Anterior aciculae tapering to 

blunt tips, gently curved or straight. In median setigers inferior 

aciculae (Figure 98n) blunt-tipped, distinctly sharply bent. 

Posteriorly inferior aciculae (Figure 98r) indistinctly bifid, 

slender, superior aciculae distinctly bifid with teeth of different 

sizes, thick; cross-sections round. Separation between core and 


Subacicular hooks (Figure 98m,p) yellow, tridcntate with teeth 

in a crest. Hooks first present from setiger 27, present in all 

setigers thereafter, paired in some setigers. Two distal fangs 

grouped on common bases; tertiary fangs very small in median 

setigers, becoming more distinct in posterior setigers. 







Characters: 1,2,4,6,23, 36-38,40. 


38,1. 

REMARKS.—Eunice rubra is listed with similar species in 

Tables 46 and 47. It has both pointed and weakly bidentate 

aciculae. Other than E. elseyi, it has a far higher number of 

branchial filaments than any other species with modified 

aciculae listed in Table 47. Eunice rubra has crested tridentate 

subacicular hooks; E. elseyi has hooks with the distal pair of 

teeth in tandem. 

172. Eunice rubrivittata (Treadwell, 1921) 


Leodice rubrivitlata Treadwell, 1921:34-36, figs. 85-94, pi. 1: fig. 18. 

Eunice viitata.—Augener, 1927:40.—Hartman, 1956:252 [in part, not Nereis 

vittata Chiaje, 1829]. 

MATERIAL EXAMINED.—Holotype, AMNH 1263, Buccoo 

Bay, Tobago land, 8 Apr 1918. 



10, 5.1 mm. 

Prostomium and peristomium about equal in length (Figure 

99a), distinctly narrower than peristomium, less than V2 as deep 

as peristomium. Prostomial lobes frontally rounded, dorsally 

slightly flattened; median sulcus deep. Eyes not observed. 



Ceratostyles distally blunt, digiliform, with 2 or 3 irregularly 

arranged, cylindrical articulations. A-I to middle of peristomium; 

A-II and A-III to setiger 1. Peristomium cylindrical. 

Separation between rings distinct on all sides of body; anterior 

ring slightly longer than posterior ring. Anterior ring with 

frontal, yoke-shaped thickening covering at pcristomial fold. 

Peristomial cirri incomplete, probably slender and digitiform. 

Jaws not available. 

Branchiae (Figure 99c) present, pectinate, distinctly shorter 




number of setigers. First 6 branchiae single, long filaments 

which increase somewhat in length posteriorly. From setiger 9 

another filament added; maximum 5 filaments from setiger 25 

to 40. Number of filaments reduced to 1 by setiger 45. 

Anterior ncuropodial acicular lobes distally obliquely transverse 

with distinct, rounded lobes dorsal to emergence of 

aciculae; posterior acicular lobes increasingly withdrawn into 

body wall with aciculae emerging from tips; subacicular hooks 

supporting a secondary, flattened conical structure along 

ventral edge of parapodium (Figure 99b). All prc- and postsetal 

lobes follow outline of acicular lobes closely. First 5 or 6 

ventral cirri digitiform. Ventral cirri basal I y inflated from 

setiger 6 through setiger 50. Inflated bases nearly spherical; 

narrow tips digitiform. Ventral cirri digitiform from setiger 60, 

emerging well distal to bases of parapodia. Anterior notopodial 

cirri tapering, with 2 or 3 indistinct cylindrical articulations. 

Posterior notopodial cirri long, digitiform, without articulations. 

Limbate and pectinate setae not observed. Shafts of 

compound falcigers (Figure 99c) inflated, marginally smooth. 

Appendages bidentate. Proximal teeth much larger than distal 

teeth. Distal teeth directed obliquely distally. Guards with very 

short distal mucros. Pseudocompound falcigers and compound 

spinigers absent. Aciculae single, yellow, tapering, straight; 

cross-sections round. Notopodial aciculae distinct in anterior 

and median setigers. Separation between core and sheath 


hooks (Figure 99d) yellow, tridentate with teeth in a crest. 



very large, curved main fangs; fangs decreasing in size to very 

short tertiary fang. 


pectinate setae; pygidium and anal cirri. 





Characters: 1,2,4,6, 13,14, 27-29,65-68. 


None.

NUMBER 523 

293 

FIGURE 99.—Eunice rubrivUtata (holotype, AMNH 1263): a, anterior end, lateral view; b, parapodium 70, 

anterior view; c, compound falciger, parapodium 29; d, subacicular hook, parapodium 29; e, parapodium 29, 

anterior view. Eunice rullieri (syntype, Zool. Mus., Tel-Aviv University, 7563): f, anterior end, lateral view; g, 

pectinate seta, parapodium 27; h, subacicular hook, parapodium 27; i, compound falciger, parapodium 27; j, 

parapodium 27, anterior view. (Scale bars in mm.).


REMARKS.—Eunice rubrivittata is listed with similar species 

in Tables 41 and 42. The only other species with notopodial 

articulations limited to anterior setigers in Table 42 is E. 

medicina. In the latter posterior branchiae have at least paired 

filaments; in the former the last 10-11 pairs of branchiae are 

single filaments. 

173. Eunice rubrocincta Ehlers, 1868 

TABLES 19,21 

Eunice rubrocincta Ehlers, 1868:344-347, pi. 15: figs. 4-14. 

COMMENTS ON MATERIAL EXAMINED.—No material is 

available, the notes below are based on the original description 

and illustrations. 

DESCRIPTION.—A live specimen with 100 segments 68 mm 

long and 3 mm wide; a preserved specimen with 106 segments 

45 mm long and 3.5 mm wide. 

Ceratostyles digitiform, without articulations. A-III to 

setiger 1, A-I and A-II to middle of peristomium. Pcristomial 

rings distinct on all sides. Peristomial cirri to posterior part of 

prostomium, without articulations. Antennae and pcristomial 

cirri slender, nearly filiform. 

Maxillary formula 1+1, 6+7,7+0, 3+8,1+1, and 1+1. Mx III 

part of a distal arc with left Mx IV. Mx VI very small, but 

distinct. 


cirri, erect. Branchiae from setiger 3 to setiger 80. 


more than 65% of total number of setigers. Maximum number 

of filaments 8; posterior half of body with very small, single 

branchial filaments. Branchiae of a median segment illustrated 

as having strong, tapering branchial stems and digitiform 

filaments. 

Ventral cirri distinctly inflated basally in median region, 

tapering in anterior and posterior setigers. Notopodial cirri 


Pectinate setae illustrated as flared and furled with both 

marginal teeth longer than other teeth and a total of -15 teeth. 

Shafts of compound falcigers may have been inflated with 

internal striations. Appendages illustrated as strongly tapering 

with very short heads, bidentate. Both teeth directed laterally 

and similar in size. Guards asymmetrically bluntly pointed; 


spinigers absent. Aciculae yellow, tapering; cross-sections 


aciculae and subacicular hooks. Subacicular hooks yellow, 

bidentate. Hooks with large, laterally directed proximal teeth 

and smaller, obliquely directed distal teeth. 


and peristomial features; jaw structure; details of parapodial 

structures. 


None. 



Characters: 4-19, 24, 26, 33, 39, 42-51, 54-56, 60, 80-82. 


None. 

REMARKS.—Described from Kvamcr Gulf in the Adriatic 

Sea, this species has been considered the same as E. harassii 

(Fauvel, 1923:399). Ehlers was aware of the similarities and 

gave a differential description, demonstrating at what points he 

believed his species differed from E. harassii. Eunice rubrocincta 

has branchiae ending well before the end of the body; E. 

harassii has (apparently) branchiae continued to the far 

posterior end. Eunice rubrocincta has light yellow aciculae and 

subacicular hooks; /•.'. harassii has light brown aciculae and 


Eunice rubrocincta belongs to group A-1 rather than to B-2 

as suggested by Fauchald (1970) and is listed with similar 

species in Tables 19 and 21. Of species without articulated 

ceratostyles listed in Table 21, only two have less than 10 

branchial filaments where the branchiae arc best developed. Of 

these two species, E. manihine and E. rubrocincta, the former 

has branchiae present from setiger 8, the latter has branchiae 

from setiger 3. 

174. Eunice rullieri, new name 

FIGURE 99f-j; TABLES 27.28 

Eunice aphroditois punctata Fishelson and Rullicr, 1969:74-76, fig. 2a-c. 

MATERIAL EXAMINED.—Two syntypes. Zoological Museum, 

Tel-Aviv University, 7134 and 7563, Cundabilu, Red 

Sea. 

DESCRIPTION.—Large syntype (7563) complete with 72 


through setiger 10, 5 mm. Other syntype with 49 setigers; 


4 mm. Remaining color pattern a grayish brown base color, 

with 3 rows of large white blotches dorsally. Body rather wide 

anteriorly, tapering to slender posterior end, basically conical 

with both dorsal and ventral side convex. Anal cirri long, 

subdistally inflated. 


than peristomium, as deep as x h of peristomium. Prostomial 


Eyes at bases of A-II, small. Antennae in a horseshoe, evenly 


antennae, without articulations. Ceratostyles thick and digitiform, 

without articulations. Three median antennae to posterior 

peristomial ring; A-I to middle of anterior peristomial ring. 

Peristomium cylindrical with thickened yoke forming nuchal 

fold; lower lip thickened and slightly set off from rest of 

peristomium. Separation between rings distinct only dorsally; 


posterior edge of anterior peristomial ring, ovate, without 

articulations.





from setiger 5 to setiger 70. Branchiae present to near posterior 

end, present on more than 65% of total number of setigers. All 

branchiae, except last 4-5 pairs, with 2 or more filaments; 

maximum 5 filaments at about setiger 10. Branchial stems 

tapering, flexible. Filaments digitiform, thick, shorter than 


Neuropodial acicular lobes symmetrically rounded, becoming 

narrower posteriorly, with elevated portions where superiormost 

aciculae emerge. All presetal lobes low, transverse 

folds. Anterior postsetal lobes rounded, outreaching acicular 

lobes, becoming low transverse folds from about setiger 15. 


from setiger 5 to about setiger 30. Inflated bases ovate; narrow 

tips tapering. Posterior ventral cirri broadly attached, tapering. 

All notopodial cirri prominent, thick, spindle-shaped, strongly 

inflated medially (Figure 99j), without articulations, very much 

longer than branchiae in posterior half of body. 

Limbale setae marginally serrated. Pectinate setae (Figure 

99g) tapering, flat. Both marginal teeth slightly longer than 

other teeth; -10 teeth present. Number of pectinate setae 

increasing posteriorly, becoming dominant setal element in 

dorsal fascicle. Shafts of compound falcigers (Figure 99i) 

tapering, marginally serrated, with distinct internal striations. 

Appendages triangular, tapering to large heads, bidentate. 

Proximal teeth larger than distal teeth, slender, tapering, 

directed laterally. Distal teeth distinctly bent. Guards angularly 

bluntly and symmetrically rounded, strongly serrated towards 

base; mucros absent. Pseudocompound falcigers and compound 

spinigers absent. Aciculae paired, with dark brown cores 

and clear sheaths, tapering, distally straight, projecting well 

beyond tip of acicular lobes; cross-sections round. Subacicular 

hooks (Figure 99h) with dark brown cores and clear sheaths, 


setigers thereafter, paired in most setigers. Hooks slender, 

tapering to small heads. Proximal teeth larger than distal teeth, 

curved, directed laterally. Distal teeth narrow, obliquely 

curved. 

UNKNOWN MORPHOLOGICAL FEATURES.—Jaw structure. 







None. 

REMARKS.—Eunice rullieri was originally described as a 

subspecies of E. aphroditois. The name applied to the 

subspecies, punctata, is preoccupied in several different 

combinations (see above) and because the material examined is 

of a distinct species, a new name had to be applied. 

The current specimens are small and differ from other 

species in group B-2 (Tables 27 and 28) by several major 

features, including distribution and shape of subacicular hooks. 

Some of these features are rather juvenile in appearance; 

however, the overall appearance of the specimens is not that of 

a juvenile. Eunice rullieri is one of three species listed in Table 

28 with routinely more than single subacicular hooks in every 

segment with such hooks; the other two species are E. roussaei 

and E. sebastiani. The relationship between E. roussaei and E. 

sebastiani was discussed above. Eunice sebastiani has about 40 

branchial filaments where the branchiae are best developed, E. 

rullieri has five. 

ETYMOLOGY.—It is a pleasure to name this species for the 

well-known polychaete expert, the late Dr. F. Rullier. 

175. Eunice samoae Hartmann-Schroder, 1965 

FIGURE 100a-h; TABLES 24, 25 

Eunice mulilata samoae Hartmann-Schroder, 1965a:134-135. 

MATERIAL EXAMINED.—Holotype ZMH P 14272, Samoa, 

Tutuila, Matu'u Bay, 21-23 Oct 1959, algae on edge of 

fringing reef, coll. G. Hartmann and G. Hartmann-SchrOder. 






lobes frontally obliquely truncate; dorsal surface flattened; 

median sulcus deep. Eyes not seen. Antennae in a horseshoe, 


in all antennae, without articulations. Ceratostyles medially 

inflated, thick and digitiform, with up to 5 cylindrical, 

indistinctly marked articulations in A-III. A-I to posterior 


Peristomium flaring anteriorly. Separation between rings 

distinct on all sides; anterior ring ~ 3 A of total peristomial 





Branchiae (Figure lOOd) present, pectinate, distinctly longer 




number of setigers. First 2 and last 10 pairs single filaments; 

other branchiae pectinate with maximum 6 filaments at setiger 

15. Filaments thick with abruptly tapering, narrow tips, about 

as long as notopodial cirri. 

Anterior neuropodial acicular lobes rounded, becoming 

triangular in far posterior setigers (Figure lOOe); aciculae 

emerging above midline. Presetal lobes low, transverse folds. 

Postsetal lobes free, rounded lobes in first 30-35 setigers, 

outreaching acicular lobes in first 30 setigers. Median postsetal 

lobes follow outline of acicular lobes closely. Posterior

296 


FIGURE 100.—Eunice samoae (holotype, ZMH P 14272): a, anterior end, lateral view; b, compound falciger, 

parapodium 20; c, pectinate seu, parapodium 20; d, parapodium 20, anterior view; e, parapodium 100, anterior 

view; f, pectinate seu, parapodium 100; g, compound falciger, parapodium 100; h, subacicularhook, parapodium 

100. Eunice schizobranchia (MNHN): i, parapodium 14, anterior view; j, compound falciger, parapodium 14; k, 

compound falciger, parapodium 14; 1, anterior end, lateral view; m, pectinate seu, parapodium 683; n, subacicular 

hook, parapodium 683; o, acicula, parapodium 683; p, compound falciger, parapodium 683; q, parapodium 683, 

anterior view. (Scale bars in mm.)


posiscial lobes low, transverse folds. First 5 ventral cirri thick 

and tapering. Ventral cirri basally inflated from setiger 6 

through setiger 40. Inflated base large, transverse welts; narrow 

tips tapering. Posterior ventral cirri thick and tapering and 

about as long as notopodial cirri, but distinctly thicker. All 

notopodial cirri thick, basally inflated, without articulations. 

Limbatc setae slender, marginally smooth. Anterior pectinate 

setae (Figure 100c) furled, slightly flaring. One marginal 

tooth distinctly longer than other teeth; -15 teeth present 

Posterior pectinate setae (Figure 1000 slightly less furled, 

distinctly more flared. One marginal tooth longer than other 


100b,g) distinctly inflated, marginally serrated; beak distinct. 

All appendages similar in size, triangular with distinct heads, 

bidentate. Proximal teeth narrow, slender, directed or curved 

slightly distally. Distal teeth distinctly larger than proximal 

teeth, gently curved. All guards symmetrically bluntly pointed, 

nearly rounded; mucros absent. Pseudocompound falcigers and 

compound spinigers absent. Aciculae single or paired, dark 

brown, tapering, nearly straight, blunt-tipped; cross-sections 

round. Separation between core and sheath indistinct in 

aciculae. Subacicular hooks (Figure lOOh) with dark brown 



(except for replacements). Hooks tapering, slightly bent. 

Proximal teeth larger than distal teeth; both teeth directed 

distally. 


anal cirri. 





Characters: 1,2,6,13,14. 


None. 

REMARKS.—Eunice samoae was originally described as a 

subspecies of E. mutilata. The branchial distribution differ 

markedly in the two forms. The former has branchiae limited to 

a short anterior part of the body, the latter has branchiae 

distributed over most of the body. The shape of the branchiae 

and the compound falcigers are also different. 

Eunice samoae is listed with similar species in Tables 24 and 

25. Of the four species in Table 25 that have cylindrical 

articulations in the ceratostyles, two species, E. coccinea and E. 

samoae, have short peristomial cirri; in the other two species 

the peristomial cirri reach at least the peristomium. Eunice 

samoae differs from E. coccinea in that it has the ventral cirri 

inflated only in the median region; in the latter the ventral cirri 

are basally inflated to near the posterior end. 

The ceratostyles are very thick, nearly sausage-like, and the 

branchial filaments are thick and distally abruptly tapering with 

a slender tip; both features are missing in other species listed in 

Table 24. 

and 

176. Eunice savignyi Grube, 1878 

TABLES 19.20 

Eunice savignyi Gnibc, 1878b:150-151. 

DESCRIPTION.—Grube had available a single specimen, 

which apparently now has been lost; the notes below are based 

on his description. 

Type specimen incomplete with 105 segments (nearly 

complete according to Grube); length 63 mm. 

Ceratostyles with long, cylindrical articulations. A-III to 

setiger 16 with 14 articulations. Peristomial cirri reaching 

beyond peristomium, with 5 articulations. 


Branchiae pectinate. Branchiae from setiger 4 to setiger 45. 

Branchiae terminating well before posterior end, present on less 

than 55% of total number of setigers. Maximum 12 filaments. 

Limbate and pectinate setae and compound falcigers 

unknown. Aciculae and subacicular hooks yellow. Subacicular 

hooks bidentate. 


and peristomial features; jaw structure; all parapodial features; 

limbate and pectinate setae; compound falcigers. 


None 



Characters: 1-20, 23-26, 28, 32-34, 39, 40, 42-72, 

74-76, 78, 80-82. 


71,2; 72,2. 

REMARKS.—The species was well characterized by Grube; 

but some crucially important features were not mentioned by 

Grube. It has been included in Tables 19 and 20 with similar 

species, but has been excluded from the key. 

Eunice savignyi cannot be distinguished from similar species 

without examination of fresh material. The original specimen 

came from an unnamed locality in the Philippines. 

Eunice schemacephala Schmarda, 1861 

Eunice schemacephala Schmarda, 1861:132, 7 figs., pi. 32: fig. 260.—Ebbs, 

1966:527-534, fig. 9a-j. 

Staurocephalus gregaricus Mayer, 1900:1. 

IMayeria gregaricus.—Merrill, 1900:650. 

REMARKS.—Schmarda's original description and illustrations 

identify the species as a member of the genus; the only 

relatively unique feature appears to be the short, unidentate 

appendages of the compound falcigers. Ebbs identified the 

species with the Atlantic palolo. However, the shape of the 

appendages of the compound falcigers is not unique to this 

species, nor does the color pattern described by Ebbs agree with 

that illustrated by Schmarda. 

Mayer described the Atlantic palolo as Staurocephalus 

gregaricus; Verrill (1900) moved this species into a new genus,

298 


Mayeria, on the grounds that Mayer's eunicid species could not 

be referred to a dorvilleid genus. I am not convinced that the 

Atlantic palolo can be identified with E. schemacephala and 

prefer to use the name E.fucata Ehlers for the species described 

in great detail by Ebbs; presumably the same seen by Mayer. 

The names associated with Mayer's species are listed also 

under E.fucata. 

Eunice schemacephala is indeterminable. 

177. Eunice schizobranchia Claparede, 1870 

FIGURE lOOi-q; TABLES 22,23 

Eunice schizobranchia Claparede, 1870:394, pi. 2: fig. 6.—Fauvel, 1923:407- 

408, fig. 160 a-k. 

MATERIAL EXAMINED.—One specimen, MNHN, Paris, Gulf 

of Naples, Oct. 1899, identified by P. Fauvel. 

DESCRIPTION.—Specimen complete with 731 setigers plus 

short, regenerating posterior end with pygidium; total length 

655 mm; maximal width 5 mm at setiger 10; length through 

setiger 10, 10 mm. Body cylindrical through first half, 

thereafter tapering very nearly imperceptibly towards posterior 

end. All segments of about the same length, with relatively 

short laterally situated parapodia. 

Prostomium (Figure 1001) distinctly shorter and narrower 


Prostomial lobes frontally obliquely truncate, dorsally flattened, 

sloping from posteromedial region obliquely forwards; 

median sulcus distinct ventrally, very short dorsally. Eyes not 

observed. Antennae in a horseshoe with A-I and A-II emerging 

close together, well lateral to midline; A-III isolated on small 

medial elevation, similar in thickness. Ceratophores ringshaped 


slender and tapering, without articulations. A-I and A-III to 

middle of anterior peristomial ring; A-II to setiger 1. 

Peristomium flaring anteriorly; lower lip large and muscular. 



barely reaching posterior 73 of anterior peristomial ring, 


Maxillary formula 1+1,4+4, 6+0, 2+6, and 1+1. Mx III part 

of arc with left Mx IV. Mx V reduced with barely distinct teeth. 

Mx VI absent. 

Branchiae (Figure lOOq) present, pectinate, distinctly longer 




of setigers. First 100 branchiae single filaments; maximum 

seven filaments. Number of filaments retained through rest of 

body; even in last branchiated setigers 6 filaments may be 

present. Branchial stems short, strongly tapering. Filaments 

long and digitiform, outreaching notopodial cirri in all but few 

first setigers. 

Anterior neuropodial acicular lobes (Figure lOOi) rounded 

with aciculae emerging above midline, above emergence of 

aciculae distinct, tapering cirri present. Median and posterior 

neuropodial acicular lobes triangular, retaining small superior 

tab even in last setigers. All presetal lobes low, transverse folds. 

Anterior postsetal lobes higher than acicular lobes and rounded, 

following outline of acicular lobes closely from about setiger 

30. First 9 ventral cirri tapering to digitiform tips. Ventral cirri 

basally inflated from setiger 10 through rest of body. Inflated 

bases nearly spherical; narrow tips tapering. Bases of posterior 

ventral cirri thick, transverse welts; narrow tips short and 

button-shaped. All notopodial cirri basally slightly inflated, 

retaining same length throughout, decreasing in thickness 



100m), apparently missing in first 20 setigers, slender, 

tapering, furled with thickened margins. One marginal tooth 

very long; -7 rather coarse teeth present. Compound falcigers 

in thick, double fascicles in anterior setigers, decreasing in 

numbers posteriorly, reduced to single anterior fascicle by 

setiger 25. Shafts of compound falcigers (Figure lOOj.k.p) 

slightly inflated, marginally serrated; internal striations and 

beaks distinct. Appendages of hooks in anterior fascicles of 

anterior setigers (Figure 100k) long and slender, tapering, 

bidentate. Both teeth distinct and slender. Proximal teeth 

slightly smaller than distal teeth, directed laterally. Distal teeth 

slender and nearly erect. In posterior fascicles of anterior 

setigers, especially towards upper ends of fascicles appendages 

relatively longer and teeth reduced (Figure 100J). Proximal 

teeth small knobs and distal teeth short, erect knobs. In 

posterior setigers appendages long, slender and tapering 

(Figure lOOp); head large. Proximal teeth large, laterally 

directed, triangular. Distal teeth curved, very much smaller than 

proximal teeth. Guards longer than appendages in all hooks, 

symmetrically rounded and marginally strongly serrated; 


spinigers absent. All aciculae (Figure lOOo) tapering, distally 

straight; cross-sections round. In anterior setigers up to 4 

aciculae in oblique series, in posterior setigers all aciculae 

single. Anterior aciculae black, becoming lighter posteriorly, 

light brown near posterior end. Subacicular hooks (Figure 

lOOn) clear and translucent, rather than yellow, bidentate. 

Hooks first present from setiger 60, first irregularly occurring, 

by setiger 300 present in all setigers, always single (except for 

replacements). All hooks emerging nearly at right angles with 

aciculae, projecting well beyond ventral cirrus in all setigers. 

Hooks distally tapering. Proximal teeth larger than distal teeth; 

both teeth directed distally. 


anal cirri. 





Characters: 13, 14,42, 74, 78. 

and



None. 

REMARKS.—Claparede did not leave any type material; the 

specimen examined is one of the specimens on which Fauvel 

based his definition of the species; it agrees with Claparede's 

description and confirms the description of the species given by 

Fauvel and used in all later publications. The species is listed 

with similar species in Tables 22 and 23. Eunice schizobranchia 

is unique in having translucent, nearly clear subacicular 

hooks and dark aciculae. Branchiae start very late and are 

continued to the far posterior end. 

208. Eunice scombrinis Quatrefages, 1866 


Eunice scombrinis Quatrefages, 1866:319-320.—Grubc, 1870a:296-297. 

MATERIAL EXAMINED.—MNHN, Paris A.l(R.)-l868-no. 

54a, Guayaquil, Ecuador, coll. Eydoux and Souleyet. 


130 setigers. Total length 47 mm; maximal width 4 mm at 

setiger 10. Length through setiger 10, 7 mm. Anterior half of 

body cylindrical; posterior half strongly dorsoventrally flattened. 

All segments short and crowded. 

Prostomium frontally rounded, dorsally inflated; median 

sulcus deep. Prostomium distinctly shorter than peristomium, 


peristomium. Both prostomial lobes separated into lobes by 

transverse frontal grooves. Eyes not observed. Antennae in a 

deep horseshoe, evenly spaced, similar in thickness. Ceratophores 

short and ring-shaped in all antennae, without articulations. 

Ceratostyles tapering, without articulations. A-I to 

second peristomial ring; A-II and A-III to setiger 3; A-III 

slightly longer than A-II. Peristomium cylindrical with distinct 

muscular, scalloped lower lip. Separation between peristomial 

rings distinct dorsally and ventrally; anterior ring 9 /io of total 

peristomial length. Peristomial cirri barely outreach posterior 

peristomial ring. Peristomial cirri tapering, without articulations. 

Maxillary formula 1+1, 9+, 8+0, 7+9, 1+1, 1+1. Right Mx 

II damaged. Mx III long; located left Mx II. All teeth long and 

delicate. 


cirri, not reduced in mid-body region. Branchial stems 

flexible. Branchiae present from setiger 7 to posterior end. 


65% of total number of setigers. First and last 5 pairs of 

branchiae single filaments. Maximum 17 branchial filaments 

present. Branchial filaments slender, shorter than notopodial 

cirri, digitiform. 

Neuropodial acicular lobe distally truncate; aciculae emerging 

dorsal to midline. All presetal lobes low transverse folds. 

Anterior postsetal lobes project as free, rounded lobes; median 

and posterior postsetal lobes follow outline of acicular lobes 

closely. Anterior ventral cirri thick and tapering. Ventral cirri 

basally inflated from about setiger 6; inflated bases continued 

through rest of body. Inflated bases thick, transverse welts. 

Ventral cirri with short, button-shaped free tip in all setigers. 

Upper, free edge of ventral cirri scoop-shaped in median and 

posterior setigers. Notopodia supported by internal aciculae; all 

notopodial cirri basally strongly inflated, strongly tapered. 

Inferior lower edge produced as small, free lobe in all except 

first 2 parapodia. Notopodial cirri without articulations. 

Limbate setae smooth, slender. Pectinate setae numbering 

2-3 in median and posterior parapodia. Pectinate setae 

tapering, flat. One marginal tooth slightly longer than other 

teeth; total number of teeth -15. Shafts of compound falcigers 

tapering; distal beak; marginal serrations and internal striations 

absent. Appendages short, strongly tapering to large head, 

bidentate. Proximal tooth conical, shorter than distal teeth, 

directed laterally. Distal teeth directed obliquely distally, 

distinctly curved. Guards asymmetrically bluntly pointed, 

marginally smooth, without mucro. Pseudocompound and 

falcigers and compound spinigers absent. Aciculae paired, dark 

brown, tapering, distally pointed, straight; cross-section round. 

Separation between sheath and core distinct in both aciculae 

and subacicular hooks. Subacicular hooks light brown anteriorly, 

dark brown posteriorly, bidentate, tapering to small heads. 

Subacicular hooks first present from setiger 29, present in all 

setigers thereafter, usually 3 in a vertical series with slimmest 

hook dorsalmost and coarsest hook ventralmost. Both teeth 

directed obliquely distally; distal tooth distinctly larger than 

proximal teeth. Guards distally smoothly rounded. 


anal cirri. 





Characters: 13, 14, 65,66. 


None. 

REMARKS.—Eunice scombrinis is compared to similar 

species in Tables 27 and 28. In addition to E. scombrinis, the 

only species listed in Table 28 to have multiple subacicular 

hooks in most parapodia is E. sebastiani. In E. sebastiani, the 

appendages of the compound falcigers are very slender and are 

supported by distinctly inflated shafts; in E. scombrinis, the 

appendages are short and tapering and the shafts are distally 

tapering. The subacicular hooks are bidentate in E. scombrinis 

and distally simple and falcate in E. sebastiani. 

178. Eunice sebastiani Nonato, 1965 

FIGURE 102; TABLES 27,28,50 

Eunice sebastiani Nonato, 1965:133-139, figs. 1-4. 

MATERIAL EXAMINED.—Praia do Araca, Sa Sebastiao,

300 


FIGURE 101.—Eunice scombrinis (holotype, MNHN, Paris, A.l(R.)-1868-no. 54a): a, anterior end, lateral view; 

b, parapodium 5, anterior view; c, compound falciger, parapodium 5; d, parapodium 59; e, pectinate seta, 

parapodium 59; f, subacicular hook, parapodium 59; g, compound falciger, parapodium 59. (Scale bars in mm.)

NUMBER 523 

301 

FIGURE 102.—Eunice sebastiani (Praia do Baraquecaba, Sao Sebasliao, Brazil): a, anterior end, lateral view; b, 

compound falciger, parapodium 3; c, pectinate seta, parapodium 3; d, parapodium 3, anterior view; e, parapodium 

681, anterior view; f, compound falciger, parapodium 681; g, subacicular hook, parapodium 681; h, pectinate seta, 

parapodium 681; i, compound falciger, parapodium 75; j, pectinate seta, parapodium 75; k, parapodium 75, 

anterior view. (Scale bars in mm.)


Brazil, 23°49'S, 45°25'30"W, muddy beach intertidal zone, 21 

June 1986, coll. A.E. Migotto and M.A. da Mota, 1 specimen. 

Praia do Baraquec,aba, Sao Sebastiao, Brazil, 23°50'S, 

45°26'W, fine sand, intertidal, 23 Jun 1986, coll. Moists A. da 

Mota, 1 specimen. 

COMMENTS ON MATERIAL EXAMINED.—The types are not 

available; the two specimens examined come from the type 

locality; they were made available by A.E. Migotto through P. 

da Cunha Lana. The specimen illustrated and described is from 

Praia do Baraquecaba. 

DESCRIPTION.—Specimen described in detail complete with 



10, 21 mm. Other specimen complete with 689 setigers; total 


through setiger 10, 30 mm. Anterior part of body through 

setiger 100 cylindrical with small parapodia; both dorsum and 

ventrum convex. Body dorsoventrally flattened posteriorly, 

tapering slowly over the last 250 setigers to posterior end, 

which is ~5 mm wide. Anterior end abruptly tapering. Anal 

cirri missing in both specimens. Both specimens dark-bronze 

colored as preserved; anterior appendages and notopodial cirri 

light colored with darker cross-bands or spots. 



lobes frontally obliquely, dorsally inflated; median sulcus 

shallow, nearly vertically directed, narrow. Each prostomial 

lobe divided longitudinally in large lateral part and smaller 

medial part Lateral parts ovate, frontally and dorsally flattened, 

sloping from high edge at junction with medial parts laterally; 

medial parts inflated, frontally rounded. Antennae in a 

horseshoe, with A-I directly in front of A-II; with A-III isolated 

by a gap, similar in thickness. Bases of A-I and A-II located in 

continuation of lines separating lateral from medial portions of 

each half prostomial half; base of A-III at head of median 

prostomial sulcus. Ceratophores long in A-I and ring-shaped in 

other 3, without articulations. Ceratostyles slender and tapering, 

none reaching beyond anterior peristomial ring, without 

articulations. A-II thicker and somewhat longer than A-III. 

Peristomium cylindrical, massive; lower lip muscular. Separation 

between rings distinct dorsally and barely visible ventrally; 


barely reaching beyond front edge of posterior peristomial ring, 

basally inflated, without articulations. 

Maxillary formula 1+1, 9+5, 6+0, 3+8, 1+1, and 1+1. All 

teeth, except Mx VI slender, tapering and circular in 

cross-section and strongly opalized with dark, nearly black tips. 

Mx VI low, knife-edged ridges. Mx III long and located behind 

left Mx II. 

Branchiae (Figure 102e,k) present, pectinate, distinctly 

longer than notopodial cirri; length of branchiae ~ 2 h width of 

body where best developed, not reduced in mid-body region, 

erecL Branchiae from setiger 7-14 to setiger 686. Branchiae 


number of setigers. All except for last 3 pairs pectinate; 

maximum 40 filaments from about setiger 35. Number of 

filaments slowly reduced posteriorly; in last 200 segments less 

than 20 filaments present in each branchia. Branchial stems 

long, slender and tapering. All filaments longer than notopodial 

cirri except in last 100 branchiated setigers. 

Anterior neuropodial acicular lobes (Figure 102d) bilobed, 

with aciculae emerging from depressions between halves; 

neuropodial acicular lobes low, rounded to truncate from early 

branchial setigers through rest of body. Presetal lobes low 

transverse folds. Postsetal lobes follow outline of acicular lobes 

closely. Anterior ventral cirri short, nearly conical with pointed 

tips. Ventral cirri basally inflated from first branchial setigers; 

inflated bases retained in all remaining setigers. From about 

setiger 40 through setiger 500 ventral cirri thick, flattened 

ridges supported on thick folds of tissue; distal lips entirely 

absent. Far posterior ventral cirri with thick tapering tips. All 

notopodial cirri with ventrally directed expansion attached near 

bases; especially in anterior branchial region, this accessory 

notopodial lobe pendant and nearly drop-shaped. Distal end of 

anterior notopodial cirri conical, tapering to slender lips. Distal 

end of median notopodial cirri tapering to slender tips, 

becoming rather thick and nearly triangular in posterior 


Limbale setae short, tapering and marginally pilose. Anterior 

pectinate setae (Figure 102c,j) tapering, furled, becoming 

nearly fiat in posterior setigers (Figure 102h). One marginal 

tooth longer than other distinctly short, trim teeth; -12 teeth 

present. Shafts of compound falcigers (Figure 102b, f.i) inflated 

and marginally smooth; inner ends of shafts dark brown or 

black in most setigers. Appendages long, slender, with nearly 

parallel sides and a very small head, bidentate. Relative length 

of appendages decreasing posteriorly. Proximal teeth triangular, 

directed laterally. Distal teeth nearly erect in most hooks, 

gently curved in some median setigers. Guards symmetrically 

bluntly to sharply pointed, marginally smooth; mucros absent. 


Aciculae usually numbering 3-4, black, tapering, blunt with 

straight tips; cross-sections round. Separation of acicular cores 

and sheaths indistinct. Subacicular hooks (Figure 102g) with 

dark brown to black core and clear sheath, tapering to slender, 

slightly curved, simple tips. Hooks first present from setiger 51 

or 56, present in all setigers thereafter; multiple hooks in most 

setigers, up to 4 hooks present. 

UNKNOWN MORPHOLOGICAL FEATURES.—Anal cirri. 







None. 

REMARKS.—This species was originally described with 

bidentate subacicular hooks, which appear to be present in


smaller specimens. Most species that become as large as this 

species does tend to have modified occurrence and structure of 

the subacicular hooks, and the fact that the hooks in these large 

specimens are unidentate is not here considered to be of great 

importance. The species is compared to similar species in 

Tables 27 and 28 and with other species with simple, spine-like 

subacicular hooks in Table 50. 

Of the four species in Table 28 with very short antennae, E. 

aphroditois and E. suviensis have bidentate subacicular hooks, 

as far as known at all times; E. unidentata always has simple, 

spine-like subacicular hooks and E. sebastiani may have both. 

Eunice unidentata was described with long, articulated 

ceratophores; the ceratophores of E. sebastiani are long in A-I; 

they are short and ring-shaped in A-I I and A-I II; they all lack 

articulations. Eunice sebastiani also resembles E. scombrinis 

and is discussed above in relation to that species. 

179. Eunice segregata (Chamberlin, 1919) 

FIOURE 103; TABLES 19. 20 

Ltodice segregata Chamberlin, 1919a:237-240, pi. 54: figs. 1-4 [in pan]. 

Eunice segregata.—Fauchald, 1969:6-8. fig. 3a-g. 

MATERIAL EXAMINED.—USNM 19153, Albatross sta 

D5695, off southern California 33°33'N, 120°17'30"W, 26 Apr 

1911, 977 m, green sand, Globigerina, beam trawl. 

COMMENTS ON MATERIAL EXAMINED.—As indicated by 

Fauchald (1969), none of the original specimens remaining fit 

the description given by Chamberlin (1919a); these specimens 

were described as a new species by Fauchald (1969). The 

specimen here described is the one Fauchald (1969) used in the 

re-description of this species. 

DESCRIPTION.—Specimen incomplete female with 92 setigers; 


through setiger 10,5 mm; width at setiger 10,3.5 mm. Anterior 

body dorsally convex with flattened ventrum, becoming 

cylindrical by about setiger 25. 




sulcus shallow. Palpal region marked by horizontal grooves. 

Eyes between bases of A-I and A-I I, hidden by peristomial fold, 

faded purple. Antennae in a nearly straight line, with A-I only 

slightly in front of other antennae, with A-I isolated by a gap, 


without articulations. Ceratostyles slender, slightly tapering, 

but basically digitiform, with up to 10 indistinct, cylindrical 

articulations in A-I 11. A-I to setiger 1; A-II to setiger 5; A-III to 

setiger 8. Peristomium cylindrical with slightly inflated 

posterior ring, ~ x h wider than prostomium; lower lip distinct, 

muscular. Separation between rings distinct on all sides; 


middle of anterior peristomial ring, slender, tapering to slender 

tips, with 5 to 6 long, cylindrical articulations. 


long, straight and located behind left Mx II. Mx VI absent. 

Jaws very badly decalcified; very soft. 





number of setigers. First 2 pairs single filaments; all other 

branchiae with at least 2 filaments; maximum 15 filaments 

present between seugers 25 and 30. Filaments slender, tapering 

to fine tips, about as long as notopodial cirri. 

Anterior neuropodial acicular lobes (Figure 103b) symmetrically 

conical, retaining same shape through branchial region, 

becoming lower and more flattened posteriorly (Figure 103h); 

aciculae emerging at midline. Postbranchial neuropodial 

acicular lobes flattened conical with tip associated with inferior 

aciculae. Presetal lobes low, transverse folds. Pre-branchial and 

branchial postsetal lobes high, rounded free lobes; postbranchial 

postsetal lobes follow outline of acicular lobes closely. 

First 7 ventral cirri tapering. Ventral cirri with inflated bases 

from about setiger 8, gradually lost from about setiger 30. 

Inflated bases nearly spherical; narrow tips digitiform. 

Postbranchial ventral cirri digitiform. Anterior notopodial cirri 

long, slender, tapering to fine tips, about as long as peristomial 

cirri in first few setigers. Notopodial cirri less prominent in 

branchial region, retaining about same shape. Postbranchial 

notopodial cirri long, basally slightly inflated and distally 

digitiform. Anterior notopodial cirri with up to 5 cylindrical, 

indistinct articulations; articulations gradually lost in branchial 

region. Postbranchial notopodial cirri without articulations. 

Limbate setae slender, tapering with narrow, marginally 

smooth limbation, longer than compound falcigers in all 

setigers. Anterior pectinate setae (Figure 103d) slender, 

tapering, furled. One marginal tooth large, rather thick; ~5 teeth 

present. Median and posterior pectinate setae (Figure 103g,l) 

flat, distally slightly flaring. One marginal tooth distinctly 

longer and thicker than other teeth; -10 teeth present. Shafts of 

anterior and posterior compound falcigers (Figure 103c j) 

tapering; distal beaks present, marginally smooth. Shafts of 

median falcigers (Figure 1030 inflated; distal beaks present, 

marginally indistinctly serrated. Appendages similar in all 

setigers, long, tapering to small heads, bidentate. Proximal 

teeth smaller than distal teeth, reduced triangular, directed 

laterally. Distal teeth nearly erect, tapering. Guards sharply and 

asymmetrically pointed, without mucros. Pseudocompound 

falcigers and compound spinigers absent. Aciculae (Figure 

103j) paired, yellow, tapering, pointed. Most aciculae distally 

straight with both aciculae similar, in posterior setigers superior 

aciculae with distinct core and sheath construction and thicker 

than inferior acicula, gently curved; cross-sections round. 

Subacicular hooks (Figure 103k) yellow, bidentate. Separation 

between cores and sheaths indistinct in all subacicular hooks. 

Hooks first present from setiger 36, present in all setigers

304 


FIGURE 103.—Eunice segregata (USNM 19153): a, anterior end, lateral view; b, parapodium 3, anterior view; c, 

compound falciger, parapodium 3; d, pectinate seta, parapodium 3; e, parapodium 21, anterior view; f, compound 

falciger, parapodium 21; g, pectinate seta, parapodium 21; h, parapodium 75, anterior view; i, compound falciger, 

parapodium 75; j, aciculae, parapodium 75; k, subacicular hook, parapodium 75; 1, pectinate seta, parapodium 75. 

(Scale bars in mm.)



tapering towards small heads. Proximal teeth nearly twice as 

large as distal teeth, directed laterally. Distal teeth nearly erect. 


anal cirri. 


TURES.—None 



Characters: 1, 2,47, 50,63. 


None. 

REMARKS.—Eunice segregata is listed with similar species 

in Tables 19 and 20. Of species in Table 20 with branchiae 

starting from setiger 3, two species, E. biannulata and E. 

kobiensis, have eight branchial filaments where the branchiae 

are best developed; one species, E. caeca has 24 filaments; the 

other species have from 11 to 18 filaments. Of the remaining 

species, E. mexicana, E. pennata, and E. websteri have distally 

drop-shaped to moniliform articulations of the ceratostyles; E. 

segregata and E. valens have cylindrical articulations throughout. 

The two peristomial rings are distinct on all sides in E. 

segregata: the separation are distinct only dorsally and ventrally 

in E. valens. The relationship between this species and similar 

species was also explored in Fauchald (1969). 

180. Eunice semisegregata Fauchald, 1969 

FIGURE 104a-d; TABLES 19, 21 

Eunice semisegregata Fauchald, 1969:8-10, fig. 4a-e. 

Leodice segregata Chamberlin. 1919a:237-240, pi. 54: figs. 1-4 [in part]. 

MATERIAL EXAMINED.—Paratype, USNM 19398, Albatross 

sta 3417, off southwestern Mexico, 16 o 32'00"N, 99°48'(XTW, 

11 Apr 1891, 902 m, green mud, small beam trawl. 

DESCRIPTION.—Paratype incomplete with 84 setigers; length 


Body cylindrical, slightly flattened posteriorly, essentially 

truncate anteriorly. 




sulcus deep. Eyes posterior to bases of A-I. Antennae in a 



slender and tapering, with up to 8 cylindrical articulations in 

A-III. A-I to middle of anterior peristomial ring; A-I I to setiger 

4; A-III to setiger 5. Peristomium inflated, barrel-shaped. 

Separation between rings distinct on all sides; anterior ring 2 /3 


anterior peristomial ring, slender and digitiform, without 



long, located behind left Mx II. Mx VI absent. 



Branchiae from setiger 3 to setiger 65 in paratype and 69 in 

holotype. Branchiae terminating well before posterior end, 

present on less than 55% of total number of setigers. All 

branchiae pectinate; maximum 38 filaments from about setiger 

15. Branchial stems slender, tapering, erect. Filaments slender, 

nearly filiform. 

Neuropodial acicular lobes symmetrically rounded, rather 

broadly in anterior setigers, becoming nearly conical in median 

and posterior setigers (Figure 104a); aciculae emerging at 


ventral cirri thick and tapering. Ventral cirri basally inflated 

from about setiger 5 through setiger 40; inflated bases totally 

missing by setiger 50. Inflated bases modest, ovate, narrow tips 

tapering. Posterior ventral cirri slender and tapering. Anterior 

notopodial cirri about as thick as antennae, tapering. In 

branchial region notopodial cirri slender and tapering from 

slightly inflated bases. Postbranchial notopodial cirri slender, 

nearly filiform. Notopodial cirri without articulations. 


104c) tapering, flat. One marginal tooth longer and thicker than 


(Figure 104c) distally inflated, marginally serrated. Appendages 

long, tapering towards small heads, bidentate. Proximal 

teeth smaller than distal teeth, short, broadly triangular, 

directed laterally. Distal teeth gently curved, tapering. Guards 

symmetrically sharply pointed, marginally serrated; mucros 


absent. Aciculae paired in most parapodia, yellow, tapering to 

blunt tips, distally bent; cross-sections round. Separation 


subacicular hooks. Subacicular hooks yellow, bidentate. Hooks 

first present from setiger 51 in holotype and from setiger 55 in 

paratype, present in all setigers thereafter always single (except 

for replacements). Hooks tapering to small heads. Teeth similar 

in size and directed distally. 


with posterior parapodia; pygidium and anal cirri. 





Characters: 1,2,4, 6,47, 50, 63, 65-68. 


None. 

REMARKS.—Eunice semisegregata was separated from type 

material of E. segregata on a series of distributional features 

detailed in part in Tables 19 and 21. Eunice semisegregata has 

branchiae extending farther back or with many more branchial 

filaments, or both, than other species in listed in Table 19, with 

the exception of E. megabranchia and E. validobranchiata. 

Subacicular hooks are first present from setiger 35 in E. 

megabranchia and from setigers 37-40 in E. validobranchiata.

306 


FIGURE 104.—Eunice semisegregata (paratype, USNM 68947): a, parapodium 40, anterior view; b, compound 

falciger, parapodium 4; c, pectinate seta, parapodium 18; d, anterior end, lateral view. Eunice sonorae (hoiotype, 

AHF Poly 0283): e, anterior end, lateral view; f, parapodium 6, anterior view; g, compound falciger, parapodium 

6; h, parapodium 380, anterior view; i, pectinate seta, parapodium 380; j, compound falciger, parapodium 380; k, 

subacicular hook, parapodium 380. (Scale bars in mm.)


The ceratostyles lack articulations in E. megabranchia; they 

have cylindrical articulations to the tip in E. semisegregata and 

have distally drop-shaped articulations in E. validobranchiata. 

Eunice siciliensis Grube, 1840:83. 

Eunice siciliensis Grube, 1840 



the genus Palola. 

Eunice simplex Peters. 1854: 611. 

Eunice simplex Peters, 1854 

REMARKS.—Grube (1878a: 101) referred this species to 

Palola (= Eunice) siciliensis under some doubt. There is 

nothing in Peters' original description to contradict such a 

conclusion. The issue will be further discussed in a review of 

the genus Palola. 

181. Eunice sonorae Fauchald, 1970 

FIGURE 104e-k; TABLES 33, 35, 50.51 

Eunice sonorae Fauchald, 1970:45-48, pi. 6: figs. a-g. 

MATERIAL EXAMINED.—Holotype, AHF Poly 0283, Puerto 

Penasco, Sonora, Mexico, rocky intertidal, 8 Apr 1967, coll. P. 

Pickens. 



10, 14 mm. Anterior body cylindrical, tapering slowly, with 

numerous, very crowded segments posteriorly. Anal cirri long, 



than peristomium, as deep as 1 /2 of peristomium. Prostomial 


sulcus deep. Eyes behind bases of A-I, black. Antennae in a 


long, confluent with prostomium, without articulations. Ceratostyles 

slightly tapering, with a maximum of 4 irregular, 

cylindrical articulations in A-III. A-I to anterior peristomial 

ring; A-II to posterior edge of peristomium; A-III to setiger 1. 

Peristomium with very voluminous, muscular lower lip, 

otherwise cylindrical. Separation between rings distinct on all 

sides; anterior ring 2 h of total peristomial length. Peristomial 

cirri to middle of anterior peristomial ring, without articulations. 





well before posterior end, present on more than 65% of total 

number of setigers. First branchia with 2 filaments; maximum 

6 filaments from about setiger 65; from about setiger 400 

number of filaments decreasing and length of filaments 

decreasing rapidly. From setiger 475 branchiae very short, 

tapering single filaments decreasing in size posteriorly. 

Branchial stem very slender. Filaments very long, slender, 

nearly filiform. 

Anterior neuropodial acicular lobes (Figure 104f) obliquely 

truncate with aciculae emerging near midline. Posterior 

acicular lobes symmetrically rounded with aciculae emerging 

slightly superior to middle. All presetal lobes low, transverse 

folds. Anterior postsetal lobes twice as high as acicular lobes, 

symmetrically rounded. Postsetal lobes decreasing in size 

posteriorly, retained as short rounded free lobes superior to and 

behind acicular lobe in all setigers. First 4 ventral cirri thick and 

tapering. Ventral cirri basally inflated from about setiger 5 

through rest of body. Inflated bases thick, elongated welts, less 

conspicuous posterior to setiger 450; narrow tips tapering. 

Anterior notopodial cirri medially inflated; posterior notopodial 

cirri tapering from a wide base, retaining about same length 

in all setigers. Notopodial cirri without articulations. 

Anterior setigers with thick supra-acicular and posterior 

fascicles of limbate setae and masses of compound falcigers in 

subacicular and anterior positions; pectinate setae absent. 

Median and posterior setigers with thinner fascicles of limbate 

setae and compound falcigers; pectinate setae present Limbate 

setae slender, marginally smooth. Posterior pectinate setae in 

large numbers; some pectinate setae (Figure 104i) unusually 

large; shafts flattened; blade flat, widely flaring. One marginal 


anterior compound falcigers (Figure 104g) tapering, with 

distinct internal striations, marginally smooth, without distal 

beak. Appendages slender, slightly tapering, bidentate. Proximal 

teeth much smaller than distal teeth, tapering, directed 

distally. Distal teeth nearly erect, thick, blunt. Shafts of 

posterior compound falcigers (Figure 104j) inflated, without 

internal striation, marginally finely serrated; distal beak 

present. Appendages shorter than in anterior setigers; heads 

distinct, rather large, bidentate. Proximal teeth larger than distal 

teeth, triangular, laterally directed. Distal teeth tapering, 

directed laterally. Guards symmetrically rounded in all setigers, 

smooth-edged in anterior setigers, marginally serrated in 

posterior setigers; mucros absent Pseudocompound falcigers 

and compound spinigers absent. Notopodial aciculae honeycolored 

and especially prominent in posterior setigers. Aciculae 

up to 4 in anterior setigers, becoming reduced to 1 in posterior 

setigers, with dark brown to black cores and clear sheaths, 

tapering to slender, straight tips; cross-sections round. Subacicular 

hooks (Figure 104k) with dark brown to black cores and 

clear sheaths, tapering to slender single tips. Hooks first present 


(except for replacements). Hooks slender, distally abruptly 

tapering to simple, blunt tips. 



TURES.—Mx III short; forming part of distal arc with left Mx 

IV.






None. 

REMARKS.—Eunice sonorae belongs to the small group of 

species with dark, simple spine-like subacicular hooks and was 

compared to other species in group D4 by Fauchald (1970). It 

is listed with similar species in Tables 33,35, 50, and 51. It has 

multiple aciculae in anterior setigers and subacicular hooks first 

present from setiger 54; other species in Table 35 have single 

aciculae and the subacicular hooks are present much farther 

forward; in addition, the subacicular hooks are simple and 

spine-like in E. sonorae and are bidentate in the other species in 

this table. 

Eunice splendida Grube, 1856:58. 

Eunice splendida Grube, 1856 

REMARKS.—No material is available of this species. The 

original description indicates that A-III should reach sctigcr 5 

and have 26 moniliform articulations. The peristomial cirri 

have cylindrical articulations and reach the prostomium. The 

notopodial cirri lack articulations. Branchiae from setiger 5 to 

posterior end of the complete specimen; maximum 15 

filaments; number of filaments decrease posteriorly. 

The specimen, which was collected at Valparaiso, Chile, 

resembles E. lucei according to Grube. 

The features mentioned are consistent with species in group 

C-2, but without information about setal structures the species 

cannot be assigned to a group. 

The information is inadequate to characterize the species and 

it is here considered indeterminable. 

182. Eunice spongicola (Treadwell, 1921) 


Leodice spongicola Treadwell, 1921:25-27, figs. 53a-j. 

Eunice filamentosa.—Hartman, 1956:283 [in part, not Eunice filamentosa 

Grube, 1856]. 

MATERIAL EXAMINED.—Holotype, AMNH 1383, Montego 

Bay, Jamaica, Jul 1921. 



through setiger 10, 6.0 mm. Body tapering evenly towards 

posterior end from widest point at about setiger 10. 



Prostomial lobes frontally truncate, dorsoventrally flattened; 

median sulcus deep. Eyes behind bases of A-I, black. Antennae 

in a horseshoe, evenly spaced, similar in thickness. Ceratophores 


Ceratostyles tapering, without articulations. All antennae 

outreaching tip of prostomium, none outrcaching peristomium. 

Peristomium nearly twice as long as prostomium, nearly twice 

as wide, cylindrical. Separation between rings distinct only 


cirri to posterior edge of anterior peristomial ring, 


Jaws half everted. Maxillary formula 1+1, +, 7+0,3+5, and 

1+1; dentition of Mx II not dcterminable without destroying 

specimen. 




near posterior end, present on more than 65% of tolal number 

of setigers. First 5 branchiae with single filaments; maximum 

filaments 3 reached at midbody and maintained to near 

posterior end. Branchial stems very short, tapering. AH 

filaments very long, digiliform, longer than notopodial cirri in 

most setigers. 

Anterior ncuropodial acicular lobes distally truncate; posterior 

acicular lobes conical; aciculae emerging at midlinc. All 

prcsctal lobes low, transverse folds. Anterior postscial lobes 

distinct free lobes, about as high as acicular lobes; posterior 


digitiform. Ventral cirri basally inflated between sciigcrs 4 and 

50. Inflated bases large, ovate; narrow tips digitiform. Inflated 

bases rapidly reduced from setiger 50; posterior ventral cirri 

digitiform. Anterior notopodial cirri digitiform, about twice as 

long as ventral cirri, increasing in length posteriorly, nearly as 

long as branchial filaments in last setigers. Notopodial cirri 


Limbate setae long, slender, marginally finely dentate. 

Pectinate setae (Figure 105c) flat, widely flaring. Both 

marginal teeth slightly longer than other teeth; -15 teeth 

present. Shafts of compound falcigers (Figure 105e) distally 

inflated, marginally serrated, without distinct beak. Appendages 

with distinct heads, slightly tapering, bidentate. Proximal 

and distal teeth similar in size. Proximal teeth tapering, directed 

slightly basally. Distal teeth strongly tapered, directed laterally. 

Guards symmetrically rounded, marginally smooth; mucros 


absent. Aciculae (Figure 1050 single, amber-colored, darker 

near base than at tip, darker in posterior than in anterior 

setigers, distally expanded dorsoventrally into hammer-headed 

structures; shafts straight; cross-sections round. Subacicular 

hooks (Figure 105d) amber colored, darker near base than at 

tip, darker in posterior than in anterior setigers, bidentate. 



narrow necks; large heads. Proximal teeth larger than distal 

teeth, parrot-beaked, directed basally. Distal teeth nearly 

digitiform, distally blunt. 



EXPECTED STATES OF UNKNOWN MORPHOLOGICAL FEA-

NUMBER 523 

309 

FIGURE 105.—Eunice spongicola (holotype, AMNH 1383): a, anterior end, lateral view; b, parapodium 66, 

anterior view; c, pectinate seta, parapodium 66; d, subacicular hook, parapodium 66; e, compound falciger, 

parapodium 66; f, acicula, parapodium 66. Eunice Stanleyi, new species (holotype from MCZ 810, one syntype 

of Eunice antillensis): g, anterior end, lateral view; h, parapodium 10, anterior view; i, compound falciger, 

parapodium 70; j, subacicular hook, parapodium 70; k, pectinate seta, parapodium 10; 1, compound falciger, 


310 


rv. 



Unknown Characters: 42, 74,78. 


None. 

REMARKS.—Eunice spongicola was considered a synonym 

olE.filamentosa by Hartman (1956); the former has very short 

antennae and the inflated bases of the ventral cirri are ovate; in 

E. filamentosa the antennae reach well beyond the tip of the 

antennae and the inflated bases form thick transverse ridges. 

Eunice spongicola is listed with similar species in Tables 33 

and 39. It is one of two species with hammer-headed aciculae 

listed in Table 39, the other being E. denticulata. The 

ceratophores are long in all antennae in E. denticulata; they are 

short and ring-shaped in E. spongicola. In E. denticulata the 

subacicular hooks start -10 segments before the start of the 

branchiae; in E. spongicola the start of branchiae and 

subacicular hooks are separated only by two segments. 

183. Eunice stanleyi, new species 

FIGURE 105g-l; TABLES 24, 25 

Eunice antillensis Ehlers, 1887:84-85 [in pan]. 

MATERIAL EXAMINED.—Holotype, MCZ 810,1 syntype of 

Eunice antillensis Ehlers, Blake sta 45,25°33'N, 84°21'W, 185 

m. 

COMMENTS ON MATERIAL EXAMINED.—As indicated above, 

the material of E. antillensis was composed of two species, one 

of which appears to have been overlooked previously. 


130 setigers; total length 58 mm; maximal width 4 mm wide; 

length through setiger 10, 7 mm. Body cylindrical; parapodia 

very short; anterior end truncate. 


than peristomium, as deep as x h of peristomium. Prostomial 


Eyes posterior to bases of A-I, black, hidden below peristomial 

fold. Antennae in shallow horseshoe; A-I separated from A-I I 

and III by distinct gap, similar in thickness. Ceratophores 


digitiform, with up to 11 cylindrical articulations in A-III. A-I 

to posterior edge of peristomium; A-II to setiger 3; A-III to 

setiger 6. Peristomium cylindrical, somewhat longer ventrally 

than dorsally. Separation between rings distinct dorsally and 

barely noticeable ventrally; anterior ring 3 /4 of total peristomial 

length. Peristomial cirri to middle of peristomium, tapering, 





to setiger 33. Branchiae terminating well before posterior end, 



last 8 branchiae single filaments (Figure 105h); maximum 2 

filaments, found scattered from setiger 12 through setiger 25. 

Filaments short, thick, longer than notopodial cirri in most 

setigers. 


rounded to triangular posteriorly; aciculae emerging at 


ventral cirri thick, tapering, rather prominent. In branchial 

region ventral cirri with indistinctly inflated bases; postbranchial 

ventral cirri tapering. Inflated bases ovate; narrow tips 

tapering. Notopodial cirri basally slightly inflated, tapering to 

digitiform tips, with 3 distinct articulations. 

Limbate setae slender, with narrow limbations. Pectinate 

setae (Figure 105k) short, flaring, flat. One marginal tooth 

thicker and longer than other teeth, with -15 teeth. Shafts of 

anterior compound falcigcrs (Figure 1051) slightly inflated, 

marginally serrated. Shafts of posterior falcigcrs (Figure 105i) 

tapering, marginally smooth. Anterior appendages with very 

low, triangular, laterally directed proximal teeth and gently 

curved distal teeth, bidentatc. Posterior appendages with long, 

tapering proximal teeth and distal teeth more distinctly bent 

than in anterior appendages. Guards asymmetrically bluntly 

pointed; anterior guards marginally serrated; posterior guards 

marginally smooth; mucros absent. Pscudocompound falcigcrs 

and compound spinigers absent. Aciculae paired, dark brown, 

tapering, straight; cross-sections round. Subacicular hooks 

(Figure 105j) dark brown to black, bidentatc. Hooks first 


single (except for replacements). Hooks tapering. Proximal 

teeth much larger than distal teeth, triangular, directed laterally. 

Distal teeth narrow, tapering, erect. 




Mx III forming distal arc with left Mx IV; Mx VI absent. 



Characters: 4, 6, 74, 78. 


None. 

REMARKS.—Eunice stanleyi is listed with similar species in 

Tables 24 and 25. It was discussed in relation to the rather 

similar E. dubitata above. In addition to the distributional 

differences mentioned there, the two species differ in the 

structure of the compound hooks. The appendages of these 

hooks have nearly erect, distal teeth in E. dubitata and have 

distinctly bent distal teeth in E. stanleyi. In the subacicular 

hooks of E. dubitata, the proximal teeth are narrower than the 

distal teeth; in E. stanleyi, the proximal teeth are very much 

thicker than the distal teeth. 

ETYMOLOGY.—The species is named for Mr. Stanley S. 

Hirsch, who at a very important time gave the author much 

needed assistance.


184. Eunice stigmatura (Verrill, 1900) 

FIGURES 106,22,23, 46, 48 

Ltodice stigmatura Verrill, 1900:641-643.—Treadwell, 1921:20-22, figs. 

31-40, pi. 1: figs. 10-13. 

Eunice villata.—Hartman, 1942:9 [in part, nol Nereis vittata Chiaje, 1829]. 

MATERIAL EXAMINED.—TWO syntypes, YPM 1232, Bermuda 

Islands, low water, 1898, A.E. Verrill; YPM 1041 and 

2095, Bermuda Islands, low water, 1898, coll. A.E. Verrill, 3 

specimens; YPM 2094, Castle Harbor, Bermuda Islands, 1898, 

coll. A.E. Verrill, 3 specimens; YPM 12%, Bermuda Islands, 

low water, 1901, coll. A.E. Verrill. 

DESCRIPTION.—Large syntype incomplete mature female 


through setiger 10, 4 mm. One specimen from YPM 2095 

(presumably part of original material, but not designated as 

type) complete with 151 setigers, length 60 mm. 




sulcus shallow. Eyes hidden beneath peristomial fold, posterior 

to bases of A-I. Antennae in a horseshoe, evenly spaced, similar 

in thickness. Ceratophores ring-shaped in all antennae, without 

articulations. Ceratostyles slender and digitiform, with up to 15 

short, cylindrical articulations in A-I 11 of short syntype; A-I 11 

lost in large syntype. A-I to setiger 1, A-I I to setiger 8; A-I 11 (in 

short syntype) to setiger 10. Peristomium cylindrical, with 

somewhat expanded lower lip. Separation between rings very 

distinct on all sides; anterior ring 2 h of total peristomial length. 

Peristomial cirri to tip of prostomium, slender and digitiform, 

with 8 to 9 short, cylindrical articulations. 

Maxillary formula of specimen from YPM 2095 1+1, 6+6, 

8+0, 6+8, and 1+1. Max III long, located behind left Max II. 

Max VI absent. Jaws poorly sclerotinized. 

Branchiae (Figure 106a) present, pectinate, about as long as 


Branchiae from setiger 3 to setiger 110. Branchiae on setigers 

70-80 very short and easily overlooked (Figure 106e); 

filaments increase in length in last 10-15 branchiated setigers. 

Branchiae present to near posterior end, more than 65% of total 

number of setigers. First 4 and roughly last 40 branchiae single 

filaments; maximum 5 filaments at about setiger 15. Branchial 

stems short, slender. Most filaments long and slender. 

Anterior neuropodial acicular lobes obliquely rounded with 

aciculae emerging near upper edge; median and posterior 

acicular lobes symmetrically rounded with aciculae emerging 

above distal point of neuropodia. All pre- and postsetal lobes 

low, transverse folds. First 6 ventral cirri long and tapering. 

Ventral cirri basally somewhat inflated from about setiger 7 

through setiger 45. Inflated bases ovate; narrow tips tapering. 

Posterior ventral cirri digitiform with rounded tips, increasingly 

prominent posteriorly. All notopodial cirri prominent, increasing 

in length from setiger 1 through setiger 7-8, thereafter 

decreasing slowly in length, but distinctly longer than 

branchiae in all branchiated setigers. Anterior notopodial cirri 

with up to 13 distinct articulations; articulations become less 

distinct and fewer posteriorly; last 25 notopodial cirri without 

obvious articulations. 


setae (Figure 106fJc) very narrow, tapering, flat. One marginal 

tooth about twice as thick and long as other teeth; 5-6 teeth 

present in pectinate setae from all parts of body. Shafts of 

compound falcigers (Figure 106c,g) distally inflated, marginally 

finely serrated; distinct beak present. Appendages decreasing 

in length from anterior to posterior setigers, bidentate. 

Anterior appendages with proximal teeth smaller than distal 

teeth, short, tapering, blunt-tipped, directed distally. Distal 

teeth very slender, directed obliquely distally. Posterior 

appendages with proximal teeth about as long as distal teeth, 

triangular, directed basally. Distal teeth tapering, directed 

obliquely distally. All guards asymmetrically bluntly pointed, 

becoming distinctly more symmetrical in posterior setigers, 


and compound spinigers absent. Aciculae (Figure 106b) paired, 

yellow, distally abruptly tapering. Superior aciculae always 

thinner than inferior aciculae; cross-sections round. Separation 


subacicular hooks. Subacicular hooks (Figure 106h,i, j) yellow, 

bi- or tridentate. Hooks first present from setigers 29 and 31 in 

syntypes (from setigers 27-35 in other material), present in all 

setigers thereafter, always single (except for replacements). All 

hooks with large, laterally directed proximal fang. Distal ends 

with thick; broad shafts crowned by either 2 fangs or terminated 

bluntly as a single, thick, angular fang. Tertiary fang small in 

most specimens. Hooks not obviously worn; newly forming 

hooks with 2 or 3 distinct teeth present. 


anal cirri. 







None. 

REMARKS.—Specimens that are not listed as types above are 

assumed to be part of the original material. 

Treadwell (1921) gave no evidence of having seen the types, 

but did examine material from the type locality. 

Eunice stigmatura has been considered synonymous with E. 

vittata, but the branchial distribution is different in that the 

former has branchiae continued to near the posterior end; the 

latter has branchiae terminating well before the posterior end. 

The two syntypes have exclusively single, thick, truncate distal 

teeth, rather than paired fangs in the subacicular hooks; the 

other specimens of the original material have three teeth. Only 

a single specimen, from YPM 2094, was observed to have 

some subacicular hooks with single large teeth and others with 

three fangs.

312 


J 

.02 

1 

,5 

e 

b, c, 

d 

f. g. h, i. j. k 

a 

\ 

FIGURE 106.—Eunice stigmatura (syntype, YPM 1232): a, parapodium 13, anterior view; b, acicula, parapodium 

13; c, compound falciger, parapodium 13; d, anterior end, lateral view; e, parapodium 79, anterior view; f, 

pectinate seta, parapodium 79; g, compound seta, parapodium 79; h, subacicular hook, parapodium 79; i, 

subacicularhook, parapodium 85; j, subacicularhook, parapodium 102; k, pectinate seu, parapodium 102. (Scale 

bars in mm.)


Eunice stigmalura is listed with similar species in Tables 46 

and 48. It is also listed in Tables 22 and 23 among species with 

bidentate subacicular hooks. The thick, blunt distal teeth in the 

subacicular hooks present in part of the material make the 

species by definition a member both of groups A-2 and C-2, 

even if all other characters suggest that it to be a member of C-2 

only. 

The separation between anterior and posterior rings of the 

pcristomium is distinct on all sides in E. stigmatura and E. 

havaica. In E. havaica A-111 is distinctly longer than A-II; in E. 

stigmatura they are similar in length. Of the three species in 

Table 23 with long, cylindrical articulations in the ceratostyles, 

E. johnsoni has short peristomial cirri; in E. gracilicirrata and 

E. stigmatura the peristomial cirri reach at least the middle of 

the prostomium. In E. gracilicirrata the branchiae clearly 

outreach the notopodial cirri; in E. stigmatura they are of about 

the same length. 

Eunice stragulum Grube, 1878 

Eunice stragulum Grube, 1878b:163-164. 

Marpkysa stragulum.—Crossland, 1903:136. 

REMARKS.—This species was described in detail and listed 

by Grube in what he recognized informally as section 

Marphysa within the genus Eunice. It was formally transferred 

to Marphysa by Crossland and will be treated in a forthcoming 

review of that genus. 

Eunice subdepressa Grube, 1866 

Eunice subdepressa Gnibe, 1866b:68; 1878a:101. 

REMARKS.—Described from Puerto Cabello, Venezuela, this 

species was characterized as having branchiae starting on 

setiger 24 and continued to end of the fragmented specimen 

(203 setigers). Maximum number of branchial filaments is four 

and A-II I does not reach beyond the peristomium. The listing in 

Grube (1878a: 101) implies that the antennae lack articulations. 

The information is clearly inadequate to characterize the 

species; no specimen is available and the species is here 


185. Eunice suviensis (Treadwell, 1922) 

TABLES 27,28 

Leodice suviensis Treadwell, 1922:138-139, figs. 12-16, pi. 2: figs. 8-13. 


COMMENTS ON MATERIAL EXAMINED.—The type is no 

longer available in AMNH. The following notes are based on 

Treadwell's description. 

DESCRIPTION.—Total length 370 mm; maximum width 9 

mm. 

Eyes dark, lateral to bases of A-II. Antennae in a horseshoe. 


Ceratostyles without articulations. A-III to setiger 1; other 

antennae slightly shorter. Peristomial cirri to middle of anterior 

peristomial ring, without articulations. 

Maxillary formula 1+1, 5+5, 7+0, 6+8, and 1+1; Mx III 

short; forming an arc with left Mx IV. 



from setiger 9 through 160 segments (however, as total number 

of setigers was not mentioned, the fraction of branchiated 

setigers cannot be estimated). Maximum number of filaments 

9. Branchial stems long and slender. Filaments slender and 

digitiform, very much longer than short, tapering notopodial 

cirri. 

Ventral cirri inflated from about setiger 10 at least through 

setiger 50. Posterior ventral cirri without basal inflation. 

Notopodial cirri short and tapering in all setigers, without 


Pectinate setae slightly flaring with 1 long marginal tooth, 

with 16 teeth. Shafts of compound falcigers inflated. Appendages 

short with both teeth well developed and directed 

obliquely distally. Guards symmetrically rounded; mucros 


absent. Aciculae and subacicular hooks "very black." Aciculae 

tapering and distally blunt, presumably straight. Subacicular 

hooks tapering with large, laterally directed proximal teeth and 

erect, smaller distal teeth. 


and peristomial features; parapodial features. 


None. 



Characters: 1, 4-12, 16, 17, 20, 24-26, 28, 29, 37-40, 

42-52, 54-56, 60, 65,67, 74, 78, 80-82. 


38,1. 

REMARKS.—The shape of the subacicular hooks, branchial 

distribution, and development of branchiae indicate that the 

synonymy proposed by Hartman (1956) cannot be correct. The 

species was well characterized, but not sufficiently to be 

included in the key; it is compared to similar species in Tables 

27 and 28. Compared to other species with short antennae and 

bidentate subacicular hooks listed in Table 28, E. suviensis has 

slender, tapering notopodial cirri, rather than medially or 

basally strongly inflated ones. New material from Rat Passage, 

Suva Harbor, from which it was originally described, should be 

examined to establish features that cannot be determined from 

original description. 

Eunice taenia Claparede, 1864 

Eunice taenia Claparede. 1864:580-581, pi. 4: fig. 11.—Quatrefages, 

1866:653-654. 

Eunice siciliensis.-GniYx, 1866b:68.—Ehlers, 1868:353-358.

314 

REMARKS.—No material is available; nothing in the description 

or illustrations argues against the synonymy proposed first 

by Grube (1866b). 

Eunice tahitana Kinberg, 1865 

Eunice tahitana Kinberg, 1865:562; 1910:42, pi. 16: fig.17. 

Eunice antennata.—Hartman, 1948:78-79 [in part, not Leodice antennata 

Lamarck, 1818]. 

REMARKS.—No material is available in Riksmusect, Stockholm 

(R. OlerOd, in litt.); when Hartman examined the 

material, only fragments without an anterior end were left of 

this species. The species was characterized as having antennae 

with moniliform articulations and articulated peristomial and 

notopodial cirri, and branchiae from setiger 4 with up to six 

filaments. Hartman (1948:79) mentioned that the subacicular 

hooks were yellow and tridentate; that the aciculae were 

yellow, tapering and distally slightly curved; and that the 

compound falcigers were distally bidentate. 

Eunice tahitana belongs to group C-2, but cannot be 

adequately characterized without access to fresh material. The 

species is here considered indeterminable. 

186. Eunice tentaculata Kinberg, 1865 


Eunice tentaculata Kinberg, 1865:562; 1910:41, pi. 15: fig.13. 

Eunice valenciennesii Grube, 1878b:55.—Hartman, 1948:76-77. 

MATERIAL EXAMINED.—TWO syntypes, RM 434, 2°30'S, 

107°30'E, 88 m. 


in poor condition and the soft parts cannot be illustrated 

meaningfully. Most of the description is based on the larger of 

the syntypes. A set of maxillae are present in the vial; it is 

approximately of the size to be expected for the longer of the 

two syntypes and is here considered belonging to this species. 

DESCRIPTION.—One syntype with 9 setigers only, frontally 

dissected; jaws missing. Other syntype nearly complete with 

about 75 setigers; length 50 mm. 

Prostomium apparently frontally rounded; median sulcus 

shallow. Eyes present. Ceratophores ring-shaped in all antennae, 

without articulations. Ceratostyles with up to 8 long, 

cylindrical articulations in A-III. A-I to middle of peristomium; 

A-II to setiger 3 and A-III to setiger 10. Peristomium 


anterior ring 3 A of total peristomial length. Peristomial cirri 

nearly to frontal margin of prostomium, tapering, with 4 


Maxillary formula 1+1,5+6,7+0,8+9, and 1+1. Mx III long, 

located behind left Mx II. Mx V very soft, possibly due to poor 

preservation. 


region, erecL Branchiae from setiger 6 to setiger 28. Branchiae 



of total number of setigers. First branchiae single filaments; all 

other branchiae pectinate, with possible exception of last few. 

Maximum number of filaments 9 according to Kinberg (1865). 

Shape of neuropodial acicular lobes indeterminable. Ventral 

cirri inflated in some median setigers. Notopodial cirri medially 

inflated at least in branchial region. 

Limbate setae marginally smooth. Pectinate setae with both 

marginal teeth slightly longer than other teeth; -15 teeth 

present. Shafts of anterior compound falcigers (Figure 107a) 

distally inflated, marginally smooth. Shafts of posterior 

compound falcigers (Figure 107b) barely inflated, marginally 

distinctly serrated. Anterior appendages tapering, with distinct 

head, bidentate. Proximal teeth smaller than distal teeth, 

triangular, directed laterally. Distal teeth sharply tapering, 

curved. Anterior guards asymmetrically bluntly pointed, 

marginally smooth; mucros absent. Posterior appendages 

tapering, with distinct head, tridentate. Proximal teeth smaller 

than distal teeth, narrowly triangular, nearly tapering, directed 

laterally. Distal teeth replaced by large, erect shaft terminating 

in 2 small blunt teeth. Guards symmetrically sharply pointed, 


and compound spinigers absent. Aciculae (Figure 107d) 

yellow, tapering, distinctly bent distally; cross-sections round. 


and subacicular hooks. Subacicular hooks (Figure 107c) 


setiger 23, present in all setigers thereafter, single in setiger 23; 

most setigers with 4 or 5 hooks in dorsoventral row. Hooks 

tapering to narrow necks and distinct heads. Main fang large, 

triangular, directed slightly basally; secondary and tertiary 

fangs on same bases; fangs decreasing evenly in size from main 

to tertiary fangs. Some hooks with paired protuberances on 

both sides of main fang appearing as laterally placed additional 

teeth under certain light conditions. 


and peristomial features; parapodial features; all features 

associated with posterior setigers. 


TURES.—MX III long; located behind left Mx II. 



Characters: 1-6, 8, 10-12, 14-17, 20, 32, 42-52, 54-56, 

60,61,63,64. 


None. 

REMARKS.—The name E. tentaculata was proposed by 

Kinberg (1865) for this taxon; a year later Quatrefages (1866) 

used the same name for a completely different species (for a 

detailing of problem see Fauchald, 1986); the latter species was 

renamed E. laticeps by Ehlers (1868). Grube (1878b:55) 

renamed Kinberg's species, in the mistaken belief that 

Kinberg's paper was issued after Quatrefages' massive tomes. 

Hartman (1948) repeated Grube's mistake.

NUMBER 523 

31! 

.1 

.5 

a, b, c t n h d 

FIGURE 107.—Eunice tentaculata (syntype, RM 434): a, compound falciger, parapodium 9; b, compound 

falciger, parapodium 45; c, subacicularhook, parapodium 45; d, acicula, parapodium 45. Eunice tenuis (holotype, 

AMNH 1913-1271): e, parapodium 103, anterior view; f, compound falciger, parapodium 103; g, pectinate seta, 

parapodium 103; h, subacicularhook, parapodium 103. (Scale bars in mm.)

316 


Kinberg (1865:562) gave the original locality as "Mare 

pacificum, long, orient. 107°10\ lat. austr. 2°30', in fundo 48 

orgyiarum. VERNGREN." Kinberg (1910) repeated the same 

information and the label in the vial is the same. Hartman 

(1948:76) indicates that specimens should have been found off 

Clipperton Island in the Pacific Ocean, i.e., at 107° WEST rather 

than EAST. It is here assumed that label and Kinbcrg's 

information is correct and that the specimens were found in the 

strait between Sumatra and Borneo. 

Eunice tentaculata is listed with similar species in Tables 41 

and 42. The maximum number of branchial filaments indicated 

by Kinberg appears to be correct, but cannot be absolutely 

confirmed, due to the poor state of preservation of the material. 

Eunice tentaculata has, at least in part, tridentate appendages in 

the compound falcigers; furthermore the guards of these setae 

are blunt, rather than mucronate. The relationship with other 

species with multiple subacicular hooks was discussed in the 

remarks on E. indica. 

Eunice tentaculata Quatrefages, 1866 

Eunice tentaculata Quatrefages, 1866:317-318. 

Eunice lat ice ps Ehlers, 1868:312.—Fauchald, 1986:251-252. figs. 46-50. 

REMARKS.—As detailed by Fauchald (1986) the valid name 

of this species is E. laticeps and it has been treated as such 

above. 

Eunice tenuicirrata (Verrill, 1900) 

Leodice tenuicirrata Verrill, 1900:643-644.—Hartman, 1942:9. 

Leodice rtigma/wa.—Treadwell,1921:20-22, figs. 31-40, pi. 1: figs.10-13 

[not Leodice stigmatura Verrill, 1900]. 

REMARKS.—The type is not present in the Peabody 

Museum, Yale University, and has been missing at least for 40 

years (Hartman, 1942; W. Hartman, in litt.). Treadwell did not 

justify referring this species to E. stigmatura and did not 

mention examining the types of either species. There is 

insufficient information in the original description to allow any 

identification of the species; it is here considered indeterminable. 

187. Eunice tenuis (Treadwell, 1921) 

FIGURE 107e-h; TABLES 33,38 

Leodice tenuis Treadwell, 1921:51-52, figs. 154-163, pi. 4: fig. 11. 

Eunice tenuis.—Hartman, 1956:284. 

MATERIAL EXAMINED.—Holotype, AMNH 1913-1271, Dry 

Tortugas, Florida. 

COMMENTS ON MATERIAL EXAMINED.—The anterior end of 

the holotype has been sliced off frontally, so only a remnant of 

the peristomial rings is present on the dorsal side. The specimen 

is complete ventrally only from setiger 5-6; thus no 

illustrations could be made of the anterior end. 



10, -4.5 mm. 

Prostomium according to Treadwell (1921:51) conspicuously 

bilobed. Ceratophores ring-shaped in all antennae, 

without articulations. Ccratostylcs slender and digitiform, 

without articulations. Outreaching prostomium, with A-III 

longest. Peristomial rings distinct on all sides; relauve length of 

rings indeterminable. Pcristomial cirri short, ovate, without 


Jaws missing; maxillary formula 1+1, 7+5, 7+0, 3+6, and 

1+1 (rewritten after statement in Treadwell, 1921:52). 

Branchiae (Figure 107e) present, palmate, distinctly longer 


Branchiae from setiger 81 to posterior end. Branchiae present 


of setigers. First 10 branchiae single filaments; number of 

filaments increasing to 3 by setiger 150, finally to 4 by setiger 

400. Numbers of filaments decreasing to 3 in last 100 

segments; last few segments with reduced branchiae. 

All ncuropodial acicular lobes distally rounded; aciculae 

emerging at midlinc. AH prc- and postsctal lobes low, 

transverse folds. First 200 ventral cirri digitiform. Ventral cirri 

basally inflated in middle 200 segments. Inflated bases ovate; 

narrow tip tapering. Posterior ventral cirri short, nearly 

tubercular, without inflated bases. All notopodial cirri digitiform. 


Limbate setae missing in scattered setigers, gently curved, 

marginally frayed. Pectinate setae (Figure 107g) long and 

prominent in posterior setigers, narrow, tapering, flat. Marginal 

teeth no longer than other teeth; less than 10 teeth present. 

Shafts of compound falcigers (Figure 1070 distally inflated, 

marginally serrated. Appendages short, stout, bidentate. Proximal 

teeth very much larger than distal teeth, tapering, directed 

laterally. Distal teeth tapering, sharply bent. Guards symmetrically 

rounded, marginally frayed; mucros absent. Pseudocompound 


single, light amber-colored, tapering to blunt tips, straight; 

cross-sections round. Subacicular hooks (Figure 107h) light 

amber-colored, bidentate. Hooks first present from setiger 124, 

present in all setigers thereafter, paired in most setigers. 

Proximal teeth very much larger than distal teeth, directed 

laterally. Distal teeth directed distally. 





IV. 



Unknown Characters: 4, 6-8, 10-17, 24, 26, 40, 74, 78. 


None. 

REMARKS.—Eunice tenuis was compared to E. cariboea by


Hartman (1956, as E. (Nicidion) cariboea) after a reexamination 

of the types of the former. The latter lacks 

branchiae and has black subacicular hooks and aciculae; the 

former has branchiae and amber-colored subacicular hooks and 

aciculae. Eunice tenuis is listed with similar species in Tables 

33 and 38; it differs from all other species in that group by the 

very late appearance of branchiae, from setiger 81 in E. tenuis 

and from setiger 28 in E. tentaculata, the species with the next 

highest number. It is the only species listed in Table 38 to have 

paired subacicular hooks in most sctigers; other species have 

single hooks. It has light amber-colored aciculae and subacicular 

hooks, rather than dark brown to black as usual in most other 

species. 

Eunice teretiuscula Schmarda, 1861 

Eunice teretiuscula Schmarda, 1861:129, 6 figs., pi. 32: fig. 259. 

Marphysa teretiuscula.—Quatrefages, 1866:337.—Ehlcrs, 1868:359. 

REMARKS.—This species was originally described as lacking 

peristomial cirri, a feature characteristic of the genus 

Marphysa. The species will be treated in a forthcoming review 

of that genus. 

188. Eunice thomasiana Augener, 1922 

FIGURE 108a-i; TABLES 24, 25 

Eunice thomasiana Augener, 1922b:45. 

MATERIAL EXAMINED.—Two syntypes, ZMH V-9763 and 

ZMB 6398, St. Thomas, West Indies, Kukenthal and Hartmeyer 

coll. 

COMMENTS ON MATERIAL EXAMINED.—The Hamburg syntype 

is complete and in good condition; the Berlin syntype is 

incomplete and has been partially dehydrated. The description 

and illustrations are based on the Hamburg syntype with 

comments where the Berlin syntype differs. 

DESCRIPTION.—Hamburg syntype complete female with 

small eggs, with 99 setigers; total length 45 mm; maximal 

width 3 mm; length through setiger 10, 6 mm. Body with 

roughly circular cross-section, tapering evenly towards posterior 

end. Berlin syntype incomplete with 97 setigers; length 

about 40 mm; maximal width 3 mm. 




median sulcus shallow. Eyes between bases of A-I and 

A-II. Antennae in a horseshoe, with A-I isolated by a gap, 



up to 11 long, distally well-separated, ovate articulations in 

A-I 11, basally articulations cylindrical. A-I to setiger 3; A-II to 

setiger 5; A-I 11 to setiger 6. Peristomium contracted in fixation, 

cylindrical. Separation between rings distinct dorsally only; 

anterior ring 4 /5 of total peristomial length; anterior ring 

dorsally folded into several longitudinal folds. Peristomial cirri 

to front edge of peristomium, digitiform, with 5 articulations. 




setiger 4 to setiger 79, from setiger 4 to last setiger present in 

incomplete syntype. Branchiae terminating well before posterior 


Most branchiae single filaments (Figure 108e); maximum 

number of filaments 3 in a few setigers around setiger 15. 


First few neuropodial acicular lobes symmetrically rounded 

(Figure 108d); all other acicular lobes distally conical; aciculae 


transverse folds. First 4 ventral cirri thick and tapering. Ventral 

cirri basally inflated from about setiger 5 through setiger 40. 

Inflated bases thick, transverse welts; narrow tips tapering. 

Posterior ventral cirri tapering, without basal inflations. All 

notopodial cirri digitiform, longer than branchiae, with up to 4 

articulations anteriorly; posterior setigers usually with 2 

articulations; distalmost articulation ovate. 

Limbate setae slender, marginally finely serrated in anterior 

setigers, marginally smooth in posterior setigers. Anterior 

pectinate setae (Figure 108b) furled, flaring. One marginal 

tooth distinctly longer than other teeth; -15 teeth present 

Posterior pectinate setae (Figure 108f) tapering, otherwise 

similar to anterior pectinate setae. Shafts of compound falcigers 

distally tapering, marginally smooth, without distal beak. 

Anterior appendages (Figure 108c) tapering, with distinct head, 


directed laterally. Distal teeth nearly erect, tapering. Posterior 

appendages (Figure 108h) less distinctly tapering, bidentate. 

Proximal teeth larger than distal teeth, narrow, directed 

laterally. Distal teeth distinctly bent than in anterior appendages. 

Guards asymmetrically bluntly pointed, marginally 


spinigers absent. Aciculae (Figure 108i) paired, brown, 

curved slightly dorsally, flattened, knife-edged, tapering to 

blunt tips. Subacicular hooks (Figure 108g) brown, bidentate. 

Hooks first present from setiger 22 (from setiger 24 in Berlin 

syntype), present in all setigers thereafter, mostly single, rarely 

paired. Hooks tapering to narrow necks; heads large. Proximal 

teeth larger than distal teeth, tapering, directed laterally, but 

distally slightly upturned. Distal teeth erect, tapering. 





IV. 



Characters: 4, 6, 39, 40, 74, 78. 

ASSUMED STATES FOR PURPOSE OF PREPARING 

None. 

KEY.—

318 


e~ 

FIGURE 108.—Eunice thomasiana (holotype, ZMH V-9763): a, anterior end, lateral view; b, pectinate seta, 

parapodium 10; c, compound falciger, parapodium 10; d, parapodium 10, anterior view; e, parapodium 58, 

anterior view; f, pectinate seta, parapodium 58; g, subacicular hook, parapodium 58; h, compound falciger, 

parapodium 58; i, aciculae, parapodium 58. Eunice tibiana (lectotype, MCZ): j, parapodium 3, anterior view; k, 

compound falciger, parapodium 3; 1 pectinate seta, parapodium 3; m, anterior end, lateral view; n, parapodium 35, 

anterior view; o, subacicular hook, parapodium 35; p, compound falciger, parapodium 35; q, pectinate seta, 

parapodium 35; r, subacicular hook, parapodium 35. (Scale bars in mm.)


REMARKS.—Eunice thomasiana has not been reported since 

its original description. It is listed with similar species in Tables 

24 and 25. It can be separated from all other species in these 

tables by the presence of knife-edged aciculae, a feature more 

common in species in Table 27. 

189. Eunice tibiana (Pourtales, 1867) 

FIGURE 108j-r, TABLES 27,31 

Marphysa tibiana Pourtales, 1867:108. 

Eunice tibiana.—Ehlers, 1887:90-91, pi 27: figs. 1-13. 

MATERIAL EXAMINED.—Lectotype, MZC, Caribbean Sea, 

22°09'30"N, 82°21'30"W, 443 m, Sigsbee sta 11. 



through setiger 10, 5.5 mm. Body roughly circular in 

cross-section, tapering abruptly anteriorly, tapering gently 

towards posterior end. Anal cirri as long as last 3 setigers, 

basally inflated. 

Prostomium (Figure 108m) distinctly shorter and narrower 



median sulcus rather shallow. Eyes posterior to bases of 

A-I, chestnut-colored. Antennae in transverse row, evenly 



tapering, with up to 20 irregular, indistinct cylindrical 

articulations in A-I 11. A-I to setiger 1; A-I I to setiger 11; A-I 11 

to setiger 13. Peristomium cylindrical. Separation between 

rings distinct ventrally and dorsally; anterior ring 4 /5 of total 

peristomial length. Peristomial cirri reaching beyond tip of 

prostomium, slender and tapering, with 7 or 8 indistinct, 

irregular cylindrical articulations. 

Maxillary formula 1+1, 7+8, 8+0, 8+12, and 1+1. Mx HI 

long, located behind left Mx II. 

Branchiae (Figure 108n) present, pectinate, distinctly shorter 



well before posterior end; present on more than 65% of total 

number of setigers. First and last 10-15 pairs single filaments; 

maximum 5 filaments. Branchial stems short, tapering. 

Filaments digitiform, shorter than notopodial cirri. 

Anterior neuropodial acicular lobes (Figure 108j) distally 

conical, becoming obliquely triangular in median and posterior 

setigers; aciculae emerging either at midline or dorsal to 

midline. Presetal lobes low, transverse folds. Anterior postsetal 

lobes nearly as high as acicular lobes, transverse, becoming 

reduced to low folds by setiger 30. Ventral cirri in first setiger 

long, tapering. Next 3 ventral cirri thick, much shorter than in 

setiger 1. Ventral cirri basally inflated from setiger 5 through 

setiger 45, thereafter inflated bases rapidly reduced over next 

10 setigers. Inflated bases thick, transverse welts; narrow tips 

tapering. Posterior ventral cirri short, triangular, located behind 

subacicular hooks. Notopodial cirri basally inflated, without 

articulations. Notopodial cirri of setiger 1 longer than those of 

next following setigers and considerably less inflated basally. 

Limbate setae relatively short, marginally smooth, sharply 

tapering. Pectinate setae (Figure 1081,p) flat, flaring. Marginal 

teeth no longer than other teeth, with 15 teeth. Number of 

compound falcigers high in first third of body; in most 

remaining setigers number reduced to 1 or 2 per parapodium. 

Shafts of compound falcigers (Figure 108k,p) inflated, marginally 

serrated. Appendages tapering, rather thick, especially in 

median and posterior setigers, bidentate. Proximal teeth thicker 

than distal teeth, triangular. Distal teeth gently curved, nearly 

erect Guards asymmetrically sharply pointed, but not mucronate. 

Pseudocompound falcigers and compound spinigers 

absent. Aciculae paired, honey-colored, tapering, pointed, 

straight; aciculae especially heavy in second third of body; 

cross-sections round. Subacicular hooks (Figure 108o,r) honeycolored, 

bidentate. Hooks first present from setiger 26, present 

in all setigers; up to 5 hooks in vertical row present in each 

setiger. Hooks very thick, with small, abruptly tapering heads. 

Proximal teeth slightly larger than distal teeth, directed 

laterally. Distal teeth erect. Ventralmost hook often worn, may 

appear unidentate. 



None. 



Characters: 4, 6, 74,78. 


None. 

REMARKS.—The specimen selected as lectotype is one of the 

specimens examined and described by Ehlers (1887). Pourtales 

(1867) described only the tube, as being horn-colored with 

alternating openings. Ehlers added the first description of the 

animals inhabiting these tubes. No material is left from 

Pourtales study, thus the concept of this species has always 

been based on Ehlers' description. 

Eunice tibiana is compared to similar species in Tables 27 

and 31. Multiple subacicular hooks are present in two species in 

Table 31; of these two, E. borneensis has very short peristomial 

cirri. The peristomial cirri in E. tibiana outreach the prostomium. 

Other differences can be found by comparing entries in 

Table 31, illustrations, and descriptions. 

190. Eunice torquata Quatrefages, 1866 


Eunice torquata Quatrefages, 1866:312-313, pi. 10. 

MATERIAL EXAMINED.—Holotype, MNHN, Paris, A.l(R.)- 

1868-no. 47a, SL-Jean-de-Luz, coll. Quatrefages. 

DESCRIPTION.—Specimen incomplete with 120 setigers; 

length 97 mm; maximal width 4 mm; length through setiger 10,

320 


FIGURE 109.—Eunice torquata (Tholotype, MNHN A.l(R.)-1868-no. 47a): a, anterior end, lateral view; b, 

parapodium 47, anterior view; c, compound falciger, parapodium 47; d, pectinate seta, parapodium 47; e, 

subacicular hook, parapodium 47; f, aciculae, parapodium 47. Eunice torresiensis (syntype, BM(NH) 

ZK. 1885.12.1.193): g, anterior end, lateral view; h, compound falciger, parapodium 34; i, subacicular hook, 

parapodium 34; j, parapodium 34, anterior view. (Scale bars in mm.) 

10.5 mm. Body cylindrical throughout with well-defined 

segments. 




Eyes between bases of A-I and A-II, faded, hidden below 

peristomial fold. Antennae in a horseshoe, evenly spaced, 

similar in thickness. Ceratophores ring-shaped in all antennae,



moniliform articulations in A-III. First articulation of ceratostyle 

long and cylindrical. A-I to posterior peristomial ring; 

A-II to setiger 1; A-III to setiger 2. Peristomium cylindrical. 



middle of anterior peristomial ring, tapering, with 4 long, 

cylindrical or drop-shaped articulations. 




from setiger 3 through end of fragment. All branchiae with 2 or 

more filaments; maximum number of filaments 7 at about 

setiger 15. Posterior to setiger 30 most branchiae with 2 or 3 

filaments (Figure 109b). Branchial stems short, tapering. 



and posterior acicular lobes distinctly more rounded; aciculae 


folds. First 4 ventral cirri thick, tapering; ventral cirri with 

indistinctly inflated bases between setigers S and 30. Inflated 

bases ovate; narrow tips tapering. Posterior ventral cirri thick 

and tapering shape. Notopodial cirri basally slightly inflated, 

tapering. Anterior notopodial cirri with up to 5 distinct, long 

articulations; posterior notopodial cirri with 3 articulations. 


108d) narrow, tapering, flat. One marginal tooth longer than 

other teeth; -12 teeth present. Shafts of compound falcigers 

(Figure 109c) tapering, marginally indistinctly serrated. Appendages 

short, nearly triangular, bidentate. Proximal teeth 

smaller than distal teeth, reduced triangular, sharp-tipped, 

directed laterally. Distal teeth thick, blunt, curved. Guards 

asymmetrically bluntly pointed, marginally smooth; mucros 


absent. Aciculae (Figure 109f) paired, chestnut-brown, tapering, 

slightly curved dorsally; cross-sections round. Separation 


subacicular hooks. Subacicular hooks (Figure 109e) with 

chestnut-brown shafts up through neck and clear heads, 


setigers thereafter, paired in some setigers. Hooks tapering to 

narrow necks; heads small. Proximal teeth larger than distal 

teeth, directed obliquely distally. Distal tooth blunt, directed 

obliquely distally. Guards blunt. 





IV. 



Characters: 1, 2,4,6, 36-38,40. 


38,1. 

REMARKS.—Eunice torquata is compared to other species in 

group B-2 in Table 27; in addition to E. torquata, only one 

additional species in this Table, E. annulicornis, has strongly 

moniliform articulations in the ceratostyles. The two species 

have been compared above in the discussion of E. annulicornis. 

Eunice torquata has been widely reported from the Mediterranean 

Sea and adjacent areas. 

191. Eunice torresiensis Mclntosh, 1885 

FIGURE 109g-j; TABLES 15,46,47 

Eunice torresiensis Mclntosh, 1885:270-272, figs. 32, 33, pi. 37: figs. 18-21, 

pi. 19A: figs. 12, 13 [in part].—Fauchald, 1986:256-258. figs. 67-70. 

MATERIAL EXAMINED.—Four of the original 6 syntypes, 

British Museum (Natural History) ZK 1885.1.12.193, Challenger 

sta 186, Torres Strait, 10°30'S, 142°18'E, 15 m, coral 

sand. 

COMMENTS ON MATERIAL EXAMINED.—TWO additional 

syntypes were assigned to E. tribranchiata by Fauchald (1986, 

see below). The remaining material consists of a large and a 

small specimen; the large specimen is described in detail 

below; the small specimen, illustrated by Mclntosh, consists of 

16 setigers and has been cut, presumably from a much longer 

specimen; the remainder is now missing. 

DESCRIPTION.—Large syntype complete with 72 setigers; 

total length 47 mm; maximal width 3 mm at setiger 10; length 

through setiger 10,6.5 mm. Anterior body with highly convex 

dorsum and flattened ventrum, becoming circular in crosssection 

in mid body and dorsoventrally flattened posteriorly, 

abruptly tapering anteriorly, slowly tapering posteriorly. 


than peristomium, as deep as l /i of peristomium. Prostomial 

lobes frontally rounded, dorsally inflated, well separated 

medially. Reddish eyes between bases of A-I and A-II. 

Antennae in a straight line, with A-I isolated by a gap, similar 

in thickness. Ceratophores long in all antennae, without 

articulations. Ceratostyles tapering, strongly articulated; distal 

articulations distinctly moniliform; maximal number of articulations 

about 30 in A-III. A-I to setiger 1; A-II to setiger 7; A-III 

to setiger 9. Peristomium about as wide as anterior part of body. 

Separation between rings very indistinct, noticeable only 

ventrally and at cirral bases; anterior ring nearly 3 /A of total 

peristomial length. Peristomial cirri to front edge of the 

peristomium, slender, with -12 articulations. 

Maxillary formula (rewritten from Mclntosh, 1885:271, fig. 

32) 1+1, 4+3, 6+0, 5-6+8, 1+1. 


cirri, distinctly reduced in mid-body region (Figure 109j), 

flexible. Branchiae from setiger 5 to setiger 70. Branchiae 


number of setigers. First branchiae single filaments; all other 

branchiae with at least 2 filaments; maximum 7 filaments by 

setiger 15; thereafter number decreasing to 3; this number

322 


TABLE 15.—Variable and invariable features in the type lot of Eunice torresiensis (N = number of individuals 

examined; SD = standard deviation; measurements in mm; * = SD not calculated). 



Total length 

Maximal width 









N 

2 

2 

4 

4 

4 

4 

4 

4 

4 

4 

3 

Max. 

99 

51 

5 

9 

40 

9 

6 

7 

30 

10 

28 

Min. 

81 

32 

2.5 

6 

34 

4 

5 

6 

28 

4 

19 

Mean 

90.00 

41.50 

3.50 

7.05 

35.75 

7.00 

5.25 

6.50 

29.50 

6.75 

24.67 

SD 

• 

» 

1.08 

1.37 

2.87 

2.16 

0.50 

0.58 

1.00 

3.20 

4.93 



No. of pectinate teeth 






N=4 


10 

tapering 

yellow 

pointed or blunt 

yellow 

3 

maintained in middle region of body; towards posterior end 

number of filaments increasing to 7; this number maintained in 

all but last few segments. 


at midline. All pre- and postsetal lobes low, transverse 

folds. First 4 ventral cirri digitiform. Ventral cirri basally 

inflated from setiger 5 through setiger 30. Inflated bases ovate; 

narrow tips digitiform. Posterior ventral cirri digitiform without 

inflated bases. Notopodial cirri long and digitiform; prebranchial 

cirri with up to 6 moniliform articulations; farther back 

articulations long, slightly drop-shaped; most cirri with 4 


Limbate and pectinate setae not observed. Shafts of 

compound falcigers (Figure 109h) tapering without ornamentation. 

Appendages short, with large heads, bidentate. Proximal 

and distal teeth similar in size. Proximal teeth triangular, 

directed laterally. Distal teeth tapering, sharply bent. Guards 

asymmetrically bluntly pointed; mucros absent. Pseudocompound 


paired, yellow, distally bent tapering to sharp tips; crosssections 



(Figure 109i) yellow, tridentate with teeth in a crest. Hooks first 


single (except for replacements). Hooks with slender main 

fangs; secondary teeth nearly as large as main fang and small 

tertiary teeth closely appressed to the secondary teeth. 




TURES.—MX III long, located behind left Mx IV; anal cirri with 

moniliform articulations. 



Characters: 65-68. 


None. 

REMARKS.—A clarification of the status of the various types 

of this species was issued by Fauchald (1986:257) and the 

species is here accepted as defined on that occasion. 

Eunice torresiensis is listed with similar species in Tables 46 

and 47. Of the species with reduced number of branchial 

filaments in a mid-body region, two, E. margariticaea and E. 

oliga, have long peristomial cirri reaching at least the middle of 

the prostomium; in the other three species, the peristomial cirri 

do not outreach the peristomium. Eunice interrupta has 

maximally three branchial filaments where the branchiae are 

best developed; the two remaining species, E. antennata and E. 

torresiensis, have seven filaments. Eunice antennata has stiff, 

erect branchiae; in E. torresiensis the branchiae are flexible. 

Eunice triantennata (Risso, 1826) 

Leodice irianlennata Risso, 1826:422.—Fauvel, 1923:451. 

Eunice irtantennala.—Grube, 1850:292. 

REMARKS.—Grube (1850:292) referred this species to the 

section he called Leodicae Marphysae; Fauvel (1923:451) 

called the species possibly a member either of Lysidice or 

possibly as Marphysa fallax. No material exist of Risso's 

species; it is here considered a Marphysa and will be further 

discussed in a review of that genus.


FIGURE 110.—Eunice tribranchiata (syntype erf Eunice torresiensis, BM(NH)ZK.1885.12.1.193): a, parapodium 

35, anterior view; b, compound falciger, parapodium 35; c, pectinate seta, parapodium 35; d, subacicular hook, 

parapodium 35; e, anterior end, lateral view. Eunice tridentata (holotype, ZMH PE699): f, anterior end, lateral 

view; g, acicula, parapodium 20; h, compound falciger, parapodium 20; i, parapodium 20, anterior view; j, 

parapodium 162, anterior view; k, pectinate seta, parapodium 162; 1, subacicular hook, parapodium 162. (Scale 

bars in mm.) 

192. Eunice tribranchiata Mclntosh, 1885 


Eunice tribranchiata Mclntosh, 1885:297, pi. 21A: figs. 6, 7.—Fauchald, 

1986:258, figs. 71-75. 

Eunice torresiensis Mclntosh, 1885:270-272, figs. 32, 33, pi. 37: figs. 18-21, 

pi. 19A:figs. 12,13 [in part]. 

MATERIAL EXAMINED.—Holotype and paratype, BM(NH) 

ZK.1885.12.1.205,2 syntypes of Eunice torresiensis, BM(NH) 

ZK.1885.12.1.193, Challenger sta 186, Torres Strait, l 

142°18'E, 15 m, coral sand. 

COMMENTS ON MATERIAL EXAMINED.—An account of the 

material of this species can be found in Fauchald (1986:258); 

the description below is based on one of the two syntypes 

removed from the material of E. torresiensis. 

DESCRIPTION.—Holotype a posterior fragment of 90 median 

and posterior setigers. Specimen described and illustrated 

incomplete with 35 setigers, 31 mm long; maximal width 7 mm 

at setiger 10; length through setiger 10,13 mm. Other specimen 

with 18 setigers; length 15 mm; maximum width 8 mm; length 

through setiger 10,11 mm. Body cylindrical, abruptly tapering 

anteriorly.



than pcristomium, less than V2 as deep as peristomium. 


sulcus deep. Antennae in a horseshoe, with A-I isolated by a 



to 6 cylindrical articulations of varying lengths in A-III. A-I to 

posterior peristomial ring; A-I I and A-III to setiger 1. 

Peristomium as wide as anterior part of body, cylindrical, with 

distinct muscular lower lip. Separation between rings well 

marked dorsally and less distinct ventrally; anterior ring - 4 /5 of 


peristomial ring, slightly inflated basally, with 5 articulations. 

Maxillary formula (illustrated in Mclntosh, 1885, plate 37: 

fig. 20) 1+1,4+4, +0, 6+6, 1+1; Mx III absent in illustration, 

but here assumed present in specimen. 



Branchiae from setiger 4 or 5 to end of fragments. All branchiae 

with 2 or more filaments; maximum 8 filaments. Branchial 

stems tapering, flexible; filaments longer than stems, digitiform. 

Neuropodial acicular lobes symmetrically triangular; aciculae 

emerging at midline. Pre- and postsetal lobes low, 

transverse folds. First 5 ventral cirri very large, digitiform. 

Ventral cirri basally inflated from about setiger 6. Inflated 

bases large, scoop-shaped (subacicular hooks emerging at 

bases of scoops when they start); narrow tips tapering. 

Notopodial cirri basally slightly inflated with 2 or 3 long, 

indistinct articulations. 


(Figure 110c) flat, flaring with thick shafts. All teeth similar in 

length and thickness; -20 teeth present. Shafts of compound 

falcigers (Figure 110b) long, essentially un-tapering, marginally 

smooth. Appendages slender, tapering, bidentate. Proximal 

teeth larger than distal teeth, triangular and directed laterally or 

slightly distally. Distal teeth tapering, very nearly erect. Guards 

asymmetrically bluntly pointed; mucros absent. Pseudocompound 


paired, black, distally tapering, straight; cross-sections round. 

Subacicular hooks (Figure HOd) black, bidentate. Hooks first 


single (except for replacements). Heads small; teeth similar in 

size. Proximal teeth directed laterally, curved. Distal teeth 

triangular, directed distally. Guards short and truncate. 

UNKNOWN MORPHOLOGICAL FEATURES.—Structure of Mx 

III and its relationship to left Mx IV; pygidium and anal cirri. 


TURES.—MX III short, forming part of distal arc with left Mx II. 



1, 2,4, 6, 13, 14, 36-38,40,42, 57-60, 63, 74, 78. 


38,1. 

REMARKS.—The relationship between E. tribranchiata and 

similar species in Australia was discussed by Fauchald (1986). 

It is listed with similar species in Tables 27 and 29. Of the four 

species with less than 10 branchial filaments where the 

branchiae are fully developed listed in Table 29, two, E. 

microprion and E. philocorallia, have peristomial cirri reaching 

the middle of the prostomium; the two remaining species have 

short peristomial cirri. The peristomial cirri arc articulated in E. 

tribranchiata and lack articulations in E. plicata. 

193. Eunice tridentata Ehlers, 1905 

FIGURE 110f-l; TABLES 19, 21 

Eunice tridenlala Ehlers. 1905:288-290. pi. 9: figi. 3-10— F«uch«ld, 

1986:258-259. figs. 76-82. 

MATERIAL EXAMINKD.—Holotypc, ZMH PE699. New Zealand. 

DESCRIPTION.—Holotypc complete with 185 sctigers; total 


through setiger 10, 12 mm. Body circular in cross-section, 

abruptly tapering anteriorly and slowly tapering posteriorly. 


than pcristomium, as deep as V2 of peristomium. Prostomial 


Antennae in a straight line, evenly spaced, similar in thickness. 


Ceratostyles slender and digitiform, with up to 8 cylindrical, 

distinct articulations in A-III. A-I to posterior margin of 

peristomium; A-I I to setiger 2; A-III to setiger 3. Peristomium 

a thick, cylindrical collar around prostomium; lower lip 

distinctly scalloped, muscular. Separation between rings very 

distinct both dorsally and ventrally, appearing nearly as a 

separate segment, but indistinct laterally; anterior ring up 4 fc of 

total peristomial length. Peristomial cirri nearly to tip of 


Jaw apparatus missing. 

Branchiae (Figure HOi) present, pectinate, distinctly longer 




number of setigers. First and last 15 pairs single filaments; all 

other branchiae pectinate with maximum 14 filaments at setiger 

30. Branchial stems soft, pliable. Filaments long and slender, 

very much longer than notopodial cirri where best developed. 

Anterior neuropodial acicular lobes asymmetrically rounded 

with aciculae emerging above midline, near midline in 

posterior setigers (Figure HOj). Posterior neuropodial acicular 

lobes symmetrically conical. Anterior presetal lobes obliquely 

transverse, sloping from a high point at dorsal margins towards 

bases of acicular lobes at ventral margin. Anterior postsetal 

lobes transverse low folds. Posterior pre- and postsetal lobes 

follow outline of acicular lobes closely. First 4 ventral cirri 

thick and sharply tapering. Ventral cirri with inflated bases


from setiger 5 through setiger 25. Inflated bases nearly 

spherical; narrow tips tapering, broadly conical. Postbranchial 

ventral cirri increasingly digitiform, gradually loosing basal 

inflation. Prebranchial and early branchial notopodial cirri 

medially inflated, about half as long as peristomial cirri, 

becoming basally distinctly inflated, abruptly tapering with 

long, digitiform tips in most of branchial region. Postbranchial 

notopodial cirri medially inflated, similar in shape to prebranchial 

cirri but shorter. Notopodial cirri without articulations. 

Limbate setae slender, nearly capillary. Pectinate setae 

(Figure 110k) flat, flaring, with thick shafts. One marginal 


compound falcigers (Figure HOh) have thick, smoothly 

tapering, without marginal dentition. Appendages very small, 

slender, with indistinct head, bidentate. Proximal teeth triangular, 

directed laterally. Distal teeth much smaller than proximal 

teeth, nearly erect Guards symmetrically bluntly pointed; 


spinigers absent. Aciculae (Figure HOg) paired, yellow, 

distally bent ventrally, abruptly tapering; cross-sections round. 


and subacicular hooks. Subacicular hooks (Figure 1101) 

yellow, bidentate. Hooks first present from setiger 51, irregular 

in occurrence, always single (except for replacements). Hooks 

distally abruptly tapered, with small head; both teeth directed 

obliquely distally; proximal and distal teeth similar in length; 

proximal teeth slender and tapering; distal teeth massive and 

blunt. 









None. 

REMARKS.—Eunice tridentata is compared to similar species 

in Tables 19 and 21. Perhaps the most unusual feature of 

this species is that the branchiae first reach their maximum 

number of filaments at about setiger 30; in all other species in 

Table 21, the maximum number of filaments is reached at 

setiger 15, with the exception of E. heterochaeta, in which it is 

reached at setiger 25. The notopodial cirri are articulated in the 

anterior end in E. heterochaeta and wholly lack articulations in 

E. tridentata. 

Eunice tristriata Grube, 1870 

Eunice tristriata Grube, 1870b:55; 1878a: 100. 

REMARKS.—No specimens are available. The prostomial 

antennae are very indistinctly articulated; the median one 

reaches setiger 5. Branchiae are present from setiger 5 and are 

absent on the last 22 setigers; the maximum number of 

filaments is 10. The branchiae do not outreach the "remarkable 

thick" notopodial cirrus. 

The species must be considered indeterminable. 

194. Eunice tubicola (Treadwell, 1922) 


Leodice tubicola Treadwell. 1922:139-142, figs. 17-23, pi. 3: figs. 1-6. 

Eunice tubifex.—Hartman, 1956:282-283 [in part, not Eunice tubifex 

Crossland, 1904]. 

MATERIAL EXAMINED.—Holotype, AMNH 1540, Pago 

Pago, Samoa, 1920. 


had been dissected and the jaws are now missing. The lower lip 

had been sliced off, so the illustration is partly a reconstruction 

(below the dashed line indicated in the illustration). Remnants 

of a stiff horny tube was also present in sample, as were two 

fragments of another specimen, apparently of the same species. 


length 26.5 mm; maximal width 1.2 mm at about setiger 10; 

length through setiger 10,4.0 mm. 

Prostomium (Figure Ilia) distinctly shorter and narrower 



groove shallow. Eyes not observed. Antennae in a horseshoe, 



without articulations. All antennae similar in length, none 

reaching beyond anterior peristomial ring. Peristomium cylindrical. 


anterior ring - 4 /s of total peristomial length. Peristomial cirri 

digitiform, without articulations, barely reaching middle of 

anterior peristomial ring. 

Jaws missing. 

Branchiae (Figure llle) present; single filaments, about as 

long as notopodial cirri. Branchiae from setiger 22 on one side 

and 24 on other side to setiger 69. Branchiae terminating well 


of setigers. All branchiae digitiform filaments, about as long as 


Anterior neuropodial acicular lobes (Figure 11 lb) obliquely 

truncate with small, rounded tabs above aciculae; aciculae 

emerging at midline. Median and posterior neuropodial 

acicular lobes nearly triangular with straight dorsal and distal 

edges. Pre- and postsetal lobes low, transverse folds. First 5 

ventral cirri digitiform, thereafter basally strongly inflated, 

through rest of body. Inflated bases thick, transverse welts; 

narrow tips short and button-shaped. Anterior notopodial cirri 

medially inflated, becoming digitiform in posterior setigers, but 

retaining same length as in anterior setigers. Notopodial cirri 


Limbate setae marginally serrated. Pectinate setae small

326 


FIGURE 111.—Eunice tubicola (holotype, AMNH 1540): a, anterior end, lateral view; b, parapodium 8, anterior 

view; c, compound spiniger, parapodium 8; d, appendage of compound falciger, parapodium 8; e, parapodium 26, 

anterior view; f, compound falciger, parapodium 26; g, pectinate seta, parapodium 26; h, subacicular hook, 


(Figure lllg), delicate, tapering, flat. About 12 teeth present. 

Shafts of compound falcigers (Figure 11 If) strongly inflated. 

Appendages short and slender, bidentate. Proximal and distal 

teeth well developed, slender, tapering, curved and nearly 

parallel in median setigers, diverging in anterior setigers 

(Figure Hid). Guards blunt with marginal teeth; mucros 

absent. Pseudocompound falcigers absent. Compound spinigers 

(Figure 111c) in first 12 setigers. Shafts strongly inflated;


appendages long, knife-like and marginally serrated. Aciculae 

single, yellow, smoothly tapering, straight; cross-sections 

round. Subacicular hooks (Figure lllh) dark yellow (honeycolored) 

or amber, bidentate. Hooks first present from setiger 


replacements). Hooks about as thick as aciculae, gently curved 

with distinct narrow necks and distinct heads. Proximal teeth 

very much larger than distal teeth, strongly curved. Distal teeth 

nearly erect. 




TURES.—Mx III long and straight, located behind left Mx II. 


SCORED.—Inappropriate Characters: 22, 33, 34, 39-42, 56, 

60. Unknown Characters: 13, 14, 74, 78. 


KEY.— 

None. 

REMARKS.—Eunice tubicola was considered a synonym of 

E. tubifex by Hartman (1956). It is listed in Tables 52 and 53 

with similar species. The only three species in Eunice with 

compound spinigers are E. impexa, E. tubicola, and E. tubifex. 

The first has distinctly articulated ceratostyles; the other two 

have non-articulated ceratostyles. In E. tubicola the separation 

between the peristomial rings is distinct both dorsally and 

ventrally; in E. tubifex it is distinct dorsally only. Eunice 

tubicola has very short antennae, barely outreaching the 

prostomium; in E. tubifex the antennae are distinctly longer 

than the prostomium with the three median antennae similar in 

size. 

195. Eunice tubifex Crossland, 1904 


Eunice tubifex Crossland, 1904:303-310, figs. 52-55, pi. 21: figs. 1-8. 


BM(NH) ZK 1924.3.1.90/91, Wasin Harbour, S Kenya coast, 

dredged, average depth 18 m, extensive Telesto cover. 

Crossland Zanzibar and British East Africa collections, 

1901-1902. 

COMMENTS ON MATERIAL EXAMINED.—The specimens 

examined are the three smaller specimens mentioned by 

Crossland (1904:308) as differing in structure from the adult 

specimens. The description is based on the two larger 

specimens; some comments are made on the third small 

specimen. Lectotype with jaws partially everted; paralectotype 

with jaws fully everted. 

DESCRIPTION.—Lectotype incomplete with 51 setigers; 


through setiger 10,7 mm. Large paralectotype incomplete with 

52 setigers; length 36.5 mm; maximal width 2 mm; length 

through setiger 10,7 mm. Anterior body inflated; median body 

dorsoventrally flattened. 

Prostomium (Figure 112a,i) distinctly shorter than peristomium, 

about as wide as peristomium, as deep as V2 of the 

peristomium or deeper. Prostomial lobes frontally rounded, 

dorsally excavate with a flattened rim; median sulcus deep. 

Black eyes between bases of A-I and A-II. Antennae in a 

horseshoe; A-I 11 separated from A-I and A-II by distinct gap, 


without articulations. Ceratostyles slender and digitiform 


A-II to setiger 4; A-III incomplete in lectotype, to setiger 4 in 

paralectotype. Peristomium inflated anteriorly, massive, with 

muscular lower lip. Separation between rings distinct dorsally; 

2 transverse furrows present ventrally, separating ventrum in 3 

equal parts. Anterior ring 4 /s of total peristomial length. 

Peristomial cirri thick, to middle of anterior peristomial ring, 

basally inflated, without articulations. 

Maxillary formula of paralectotype 1+1,4+3, 7+0, 2+7, and 

1+1; partial formula of lectotype 1+1,4+4,7+0,2+, and 1+1. 

Mx VI absent; Mx III long, straight, located behind left Mx II. 

Distal arc absent. 



Branchiae from setiger 17 as single filaments; maximum 4 

filaments by setiger 30; maximum number retained to posterior 

end of fragments. Branchial stems shorter than notopodial cirri, 

tapering, flexible. Filaments longer than notopodial cirri, 

slender, filiform. 

Neuropodial acicular lobes (Figure 112c,m) obliquely 

truncate with distinct superior tab; aciculae supporting superior 

tab. Presetal lobes low, transverse folds. Anterior postsetal 

lobes free, projecting as far as acicular lobes, rounded. Median 



through rest of fragments. Inflated bases thick, transverse 

welts; narrow tips short and button-shaped. Prebranchial 

notopodial cirri medially inflated, increasing in length through 

setiger 10, thereafter retaining the same length through rest of 

prebranchial region. Notopodial cirri of branchial region 

increasingly shorter, becoming tapering. Notopodial cirri 


Limbate setae longer than all other setae, narrow, marginally 

smooth. Anterior and median pectinate setae (Figure 112fJ,r) 

similar, furled, tapering. Shafts slender, cylindrical. Both 

marginal teeth as long as other teeth, distinctly heavier. About 

10 teeth present. Anterior parapodia with dense masses of 

compound spinigers. Shafts of spinigers (Figure 112b,j) 

slightly inflated, smooth, with distinct distal beaks. Appendages 

slender, tapering, marginally smooth. Spinigers replaced 

by compound falcigers from early branchial setigers. Shafts of 

falcigers (Figure 112e) inflated, marginally smooth, without 

distal beaks. Appendages longer than inflated end of shaft, 

tapering. Heads distinct, bidentate. Proximal teeth longer than 

distal teeth, narrowly tapering, directed laterally. Distal teeth 

slender, tapering, distinctly bent Guards wide, distally 

rounded, amucronate; margins smooth. Pseudocompound 

falcigers absent Aciculae paired, thick, inferior aciculae black;

328 


FIGURE 112.—Eunice tubifex (synv/ês, BM(NH) ZK 1924.3.1.90/91): a, anterior end, lateral view; b, compound 

spiniger, parapodium 7; c, parapodium 7, anterior view; d, parapodium 42, anterior view; e, compound falcigcr, 

parapodium 42; f, pectinate seta, parapodium 42; g, subacicularhook, parapodium 42; h, acicula, parapodium 42. 

(i-r of smaller syntype): i, anterior end, lateral view; j, compound spiniger, parapodium 7; k, compound falciger, 

parapodium 7; 1, pectinate seta, parapodium 7; m, parapodium 7, anterior view; n, parapodium 33, anterior view; 

o, subacicular hook, parapodium 33; p, appendage of compound falciger, parapodium 33; q, acicula, parapodium 

33; r, pectinate seu, parapodium 33.


superior aciculae light brown; all aciculae distally geniculate 

(Figure 112h,o), expanded into small, flattened tabs; crosssections 

of shafts round. Subacicular hooks (Figure 112g) 

slender, bidentate, with black cores and clear sheaths. Hooks 

first present from setiger 29 (32 in paralectotype), present in all 


Hooks tapering, straight. Proximal teeth tapering, directed 

laterally; distal teeth smaller than proximal teeth, slender, 

tapering, directed obliquely laterally. 


anal cirri. 

EXPECTED STATES OF SELECTED MORPHOLOGICAL FEA- 




Characters: 1,2,4,6, 36-40,47, 50, 63, 74, 78. 


38,1. 

REMARKS.—The smaller specimen differs from the specimens 

described above in the following features: 

Branchiae (Figure 112n) single filaments, rather than 

pectinate with up to 4 filaments. 

Distal end of inflated ventral cirri relatively much larger than 

in larger specimens. 

Pectinate setae (Figure 112o) with one marginal tooth longer 

than other in small specimen. Median pectinate setae distinctly 

furled in small specimen. Shafts of both compound spinigers 

and falcigers (Figure 112j,k) with very large distal beaks; shafts 

of falcigers (Figure 112k) serrated as is margin of guards. 

Falcigers present already from about setiger 5 in small 

specimens and not until about setiger 15 in larger specimens. 

Subacicular hooks (Figure 112o) distinctly beaked with large 

head and parrot-beaked teeth. 

These differences, and the differences Crossland claimed for 

the specimens described above and the adult specimens he 

described in the first pages of the description, are the kind of 

differences that may be related to the size of the specimen, but 

may also be of specific value under certain circumstances. The 

species is clearly valid, but without access to large adults from 

the type areas, preferably from Zanzibar, it is going to be 

difficult to assess the relations among the three different forms 

Crossland included in his description. 

The relationship between E. tubifex and the closely similar E. 

tubicola is explored above; the two described forms may in fact 

belong to a single widespread species, but without knowledge 

of variability with increasing size, this problem cannot be 

solved. 

196. Eunice unidentata Rioja, 1962 

TABLES 27,28, 50 

Eunice unidentata Rioja, 1962:175-178, figs. 77-83. 

COMMENTS ON MATERIAL.—Rioja 

did not leave any 

material of this or any of the other species he described from 

Mexico (Maria Elena Caso and Vivianne Solis Weiss, in litt.). 

The following description summarizes what is known about 

this species based on his description and illustrations. 

DESCRIPTION.—Specimens 80-100 mm long and 2-3 mm 

wide. 

Prostomium distinctly shorter than peristomium, about as 

wide as peristomium. Prostomial lobes frontally rounded; 

median groove shallow. Small eyes hidden below peristomial 

fold. Antennae in a horseshoe, evenly spaced, similar in 

thickness. Ceratophores long in all antennae, with 4 to 5 

distinct, short articulations. Ceratostyles slender and tapering, 

not articulated. All antennae short; not outreaching peristomium; 

A-III longest, A-II slightly shorter and A-I shortest. 

Peristomium cylindrical. Separation between rings distinct at 

least dorsally. Peristomial cirri tapering, short, without articulations. 




from setiger 23 absent on last x h or 7* of body. Branchiae 


of total number of setigers. Maximum 5 filaments reached at 

about setiger 35 and lasting through setiger 70. Filaments long, 

digitiform. 

Parapodial features not described in detail by Rioja. Median 

parapodium (Rioja, 1962, fig. 78) with truncate, possibly 

slightly excavate neuropodial acicular lobes; aciculae emerging 

at midline. Presetal and possibly postsetal lobes low, transverse 

folds. Ventral cirri basally inflated with large, thick, tapering 

tips in median parapodia. Notopodial cirri tapering, without 


Pectinate setae flat, flaring. One marginal tooth longer than 


inflated, marginally smooth. Appendages short and triangular 

with very large heads, bidentate. Proximal teeth triangular and 

directed laterally. Distal teeth of about same size as proximal 

teeth, curved. Guards symmetrically rounded, mucros absent. 


Aciculae with dark (black) cores and clear sheaths, distally 

straight, bluntly pointed; cross-sections round. Subacicular 

hooks with dark cores and clear sheaths, distally tapering, 

unidentate. Hooks first present from setigers 20-25. 

UNKNOWN MORPHOLOGICAL FEATURES.—Many prostomial 

and peristomial features; jaw structure; many parapodial 



None. 



Characters: 1, 4-6, 8, 12, 14, 24-26, 36, 39, 40, 45, 47, 48, 

50-52,57-61,63,64,81,82. 


None. 

KEY.—


REMARKS.—Eunice unidentata was collected near Acapulco, 

Mexico. It is first and foremost characterized by the 

structure of the subacicular hooks and the very characteristic 

bases of the antennae. The basal articulations appear to be 

unique to the whole genus. The species is here considered valid 

and is listed with similar species in Tables 27 and 28. It is listed 

with other species with simple, spine-like subacicular hooks in 

Table 50. 

197. Eunice unifrons (Verrill, 1900) 

FIGURE 113a-j; TABLES 41,42 

Uodice unifrons Verrill, 1900:644.—Tread well, 1921:17-20, figs. 21-30, pi. 

1: figs. 5-9. 

Eunice vittata.—Hartman, 1942:9 [in part, not Nereis vittata Chiaje, 1929]. 

MATERIAL EXAMINED.—Holotype, YPM 1241, Bermuda, 

1898, A.E. Verrill and company coll.; YPM 1321, Platts 

Village, Bermuda, beach to 10 feet, A.E. Verrill and company, 

1898,4 specimens. 

COMMENTS ON MATERIAL EXAMINED.—The holotype material 

now consists of an anterior fragment of 31 anterior setigers 

and a median fragment of about 60 setigers; the specimen has 

been completely dried out at one time (Treadwell, 1921:20 

reported the type dry); the only information gained from the 

specimen concerns the structure of the subacicular hooks and 

other setae and some details of branchial distribution. 

The four specimens from YPM 1321 were part of the original 

material, but were not designated as types; they are in better 

shape than the holotype even if they are rather soft and have 

lost most of the antennae. 

DESCRIPTION.—All specimens from YPM 1321 incomplete 

with up to 90 setigers; specimen described and illustrated with 






distinct ventrally and at frontal edge, but not dorsally. Eyes not 

seen. Antennae in a horseshoe, with A-I isolated by a gap, 


without articulations. Ceratostyles digitiform, with cylindrical 

articulations; innermost articulation (other than ceratophoral 

ring) - V2 of total antennal length; maximum 4 articulations in 

A-I I and A-I 11. A-I to posterior peristomial ring; A-I I and A-I 11 

to setiger 2. Peristomium cylindrical. Separation between rings 

distinct on all sides; anterior ring less than 2 /3 of total 

peristomial length. Peristomial fold covering base of prostomium 

is unfolded on all 4 specimens. Peristomial cirri to 

middle of anterior peristomial ring, slender, digitiform, with 5 


Maxillary formula of 1 specimen 1+1, 8+8, 5+0, 8+10, and 

1+1. Mx VI missing. Mx III behind left Mx II, but relatively 

short. 



Branchiae from setiger 3 through setigers 45-54 (of more than 

70 setigers, no specimens complete). First 4 and last 10 pairs 

single filaments; maximum 5 filaments at about setiger 15-30. 

Branchial stems slender. Filaments slender. 

Anterior neuropodial acicular lobes rounded with a distinct, 

short nipple-like projection near middle of lobe and posterior to 

emergence of aciculae. Median and posterior neuropodial 

acicular lobes conical; aciculae emerging at midline. Pre- and 


conical. Ventral cirri basally distinctly inflated from about 

setiger 6. Inflated bases ovate; narrow tips digitiform. Posterior 

ventral cirri increasingly tapering with inflated portion less 

noticeable. Anterior notopodial cirri basally inflated, becoming 

slender and digitiform in branchial region. All notopodial cirri 

with 2 or 3 cylindrical articulations. 

Limbate setae slender, finely serrated. Pectinate setae 

(Figure 113c,f,i) tapering, flat. One marginal tooth very much 

longer and heavier than all other teeth; 7-10 teeth present. 

Shafts of compound falcigers (Figure 113b,g) slightly inflated 

and marginally serrated. Anterior and median appendages long 

and narrow with nearly parallel sides, bidentatc. Teeth very 

similar in size, both sharply pointed. Proximal teeth directed 

laterally. Distal teeth distinctly bent Posterior appendages 

(Figure 113j) very much shorter with slender teeth; proximal 

teeth shorter than distal teeth. Guards distally asymmetrically 

bluntly pointed and marginally serrated; mucros absent 


Aciculae single; clear yellow, distally tapering, straight; 

cross-sections round. Separation between cores and sheaths 


hooks (Figure 113e,h) yellow, tridentate in a crest. Hooks first 

present from setiger 25 in holotype and from setigers 26-31 in 

other specimens, present in all setigers thereafter, always single 

(except for replacements). Hooks with large curved main fangs 

and very small tertiary fangs; 2 distal fangs forming group 

separated from main fang. 


anal cirri. 




SCORED—Inappropriate Characters: 56, 58, 59. Unknown 

Characters: 1, 2,13, 14, 38, 63. 


REMARKS.—Eunice unifrons is listed with similar species in 

Tables 41 and 42. It is one of five species in Table 42 with blunt 

guards on the compound falcigers. The other species include E. 

aucklandica, E. multicylindri, E. tentaculata, and E. vittata. Of 

these species, only E. multicylindri and E. unifrons have single 

subacicular hooks throughout. The other species listed have at 

least paired subacicular hooks in most setigers. The peristomial 

cirri are articulated in E. multicylindri and lack articulations in 

E. unifrons.


b. c, e, f, g, h. i, j. n, o 

FIGURE 113.—Eunice unifrons (a-g of nontype, original material, YPM 1321; h-j of holotype, YPM 1241): a, 

anterior end, lateral view; b, compound falciger, parapodium 23; c, pectinate seta, parapodium 23; d, parapodium 

23, anterior view; e, subacicular hook, parapodium 37; f, pectinate seu, parapodium 37; g, compound falciger, 

parapodium 37; h, subacicularhook, parapodium SO; i pectinate seu, parapodium SO; j, appendage of compound 

falciger, parapodium SO. Eunice valens (holotype, MCZ 120): k, anterior end, lateral view; 1, parapodium 17, 

anterior view; m, compound falciger, parapodium 17; n, pectinate seu, parapodium 17; o, appendage of 

compound falciger, posterior parapodium; p, subacicular hook, posterior parapodium; q, aciculae, posterior 


332 

Eunice valenciennesii Grube, 1878 

Eunice valenciennesii Grube, 1878a:99.—Hartman, 1948:76-77. 

Eunice tentaculata Kinberg, 1865:562. 

REMARKS.—As noted elsewhere (Fauchald, 1986:252) and 

above, this name was proposed as a replacement name for 

Eunice tentaculata Kinberg in the mistaken belief that the 

species name proposed by Kinberg was preoccupied. The name 

is here considered an objective junior synonym of E. 

tentaculata Kinberg, because that name is valid and the 

replacement name was proposed (by inference) for the material 

treated by Kinberg, and no additional material has ever been 

named as types or otherwise. 

198. Eunice valens (Chamberlin, 1919) 

FIGURE 113k-q; TABLES 19,20 

Leodice valens Chamberlin, 1919c:257-258,pl. 1: figs. 6-8. 

Eunice valens.—Fauchald, 1969:10-12. fig. 5a-d. 

MATERIAL EXAMINED.—Holotype, MCZ 120, Mendocino, 

California, coll. A. Agassiz. 



through setiger 10, 12.5 mm; width at setiger 10, 8.5 mm. 

Anterior part of body circular in cross-section, becoming 

ventrally flattened with convex dorsal side posteriorly. Anterior 

end truncate; body tapering slowly posteriorly to narrow 

pygidium. Anal cirri as long as the last 5 to 6 setigers, with 5 or 

6 cylindrical articulations. 

Prostomium (Figure 113k) distinctly shorter and narrower 



median groove shallow. Transverse grooves separating 

palpal region present. Dark purple eyes lateral to bases of A-II. 

Antennae in a horseshoe, with A-I and A-II emerging close 

together, separated from A-III by distinct gaps, similar in 



articulations in A-III. A-I to front of posterior peristomial ring; 

A-II to back of posterior peristomial ring; A-III to setiger 1. 


dorsally and ventrally and indistinct only for short distance 

laterally; anterior ring 3 A of total peristomial length. Peristomial 

cirri to middle of anterior ring, slender and tapering, with 

three indistinct rings. 

Maxillary formula 1+1, 6+8, 9+0, 6+8 according to 

Chamberlin (1919c), presumably also 1+1. Mx III long and 

located behind left Mx II. 





number of setigers. First 2 and last 10 pairs single filaments; all 


other branchiae with at least 2 filaments; maximum 11 

filaments. Branchial stems tapering and erect. Filaments 

digitiform, relatively thick; filaments about as long as 


Anterior neuropodial acicular lobes symmetrically rounded 

with aciculae emerging dorsal to midline. Posterior acicular 

lobes nearly conical. All presetal lobes low folds. Anterior 

postsetal lobes free, triangular lobes about as high as acicular 

lobes, becoming low folds following outline of acicular lobes 

closely from about setiger 20. First 9 ventral cirri thick and 


through about setiger 60. Inflated bases thick transverse welts; 

narrow tips tapering. Posterior ventral cirri thick, tapering and 

reflexed behind acicular lobes in far posterior setigers. Anterior 

notopodial cirri basally inflated, with 3 to 4 indistinct 

articulations. Articulations lost in middle branchial setigers; 

postbranchial notopodial cirri slender, tapering, without articulations. 

All limbate setae outrcaching compound hooks, slender and 

marginally finely serrated. All pectinate setae (Figure 113n) 

with cylindrical, slender shafts, distally tapering, flat. One 

marginal tooth longer than other teeth; -10 teeth present. Shafts 

of all compound falcigcrs (Figure 113m) very coarse, nearly as 

thick as aciculae, tapering from wide proximal part of hinge. 

Beaks absent; shafts with distinct, dark-colored cores and 

dark-yellowish sheaths. Appendages short, narrow and tapering 

to small head, bidentate. Proximal teeth short, triangular and 

directed laterally, more prominent in posterior than in anterior 

setigers. Distal teeth nearly erect in all setigers, sharply pointed 

in anterior setigers, tapering and blunt in posterior setigers. All 

guards asymmetrical, sharply pointed, but not mucronate in 

anterior setigers, bluntly pointed in posterior setigers. Pseudocompound 

falcigers and compound spinigers absent. All 

aciculae and subacicular hooks with core and sheath structure. 

Aciculae (Figure 113q) paired in most setigers, but up to 4 

aciculae in some setigers, yellow, tapering to blunt tips and 

gently curved; cross-section round. Subacicular hooks (Figure 

113p) yellow, bidentate. Hooks first present from setiger 43, 

present in all setigers thereafter, single in most, paired in some 

setigers. Hooks about as thick as aciculae, tapering to distinct, 

bent necks and distinct heads. Both teeth directed distally; 

proximal teeth larger than distal teeth. 








None. 

REMARKS.—Eunice valens belongs to group Al and is 

compared to similar taxa in Tables 19 and 20. The holotype was 

not dissected for the dentition; thus Chamberlin must have had 

available at least one paratype. When examined in 1967, the


paratype consisted of several unidentifiable fragments. 

The relationship between this and similar species was 

discussed in detail by Fauchald (1969); no new information is 

available. It is discussed in relation to E. segregata above. 

Eunice valida Gravier, 1900 

Eunice valida Gravier, 1900:264-267, figs. 134-136, pi. 13: figs. 80-82. 

Eunice siciliensis.—Grassland, 1904:323-326, pi. 22: figs. 8, 9. 

REMARKS.—This species was described with branchiae 

starting at setiger 112 and continuing to the end, with 

maximum a single branchial filament and as lacking both 

subacicular hooks and pectinate setae. The species was 

considered a synonym of Eunice siciliensis by Crossland; it 

certainly belongs to the genus Palola as does siciliensis. It will 

be further considered in a review of Palola. 

Eunice validissima Grube, 1866a:67. 

Eunice validissima Grube, 1866 

REMARKS.—Grube applied this name to Mediterranean and 

Adriatic specimens resembling E. gigantea. Specimens of E. 

gigantea from the Red Sea and Fiji had branchiae from setiger 

6; E. validissima had branchiae from setigers 9-11. Grube 

must have had large specimens of a species in group B-2. His 

description is inadequate to identify the species; no material is 

available. The species is here considered indeterminable. 

199. Eunice validobranchiate Monro, 1937 

FIGURE 114a-g; TABLES 16,19,21 

Eunice validobranchiata Monro, 1937:288-289, fig. 13a-f. 

MATERIAL EXAMINED.—Three syntypes, BM(NH) ZK. 

1937.9.2.312-6, Arabian Sea, South of Oman, 21°50WN, 

59°52'00"E, 1046 meters, green mud, trawled, 3 Nov 1933, 

John Murray Expedition sta 54. 

COMMENTS ON MATERIAL EXAMINED.—All three syntypes 

are incomplete; one with 103 setigers is deeply dissected and 

probably formed the base of Monro's description; another has 

111 setigers and forms the base of the description and 

illustrations given here. The third type consists of 52 setigers. 

DESCRIPTION.—All 3 syntypes incomplete. One syntype 


through setiger 10, 10 mm. Another syntype 111 setigers; 


through setiger 10, 7 mm. Third syntype with 52 setigers; 


10, 6 mm. 




sulcus deep. Antennae in a horseshoe, with A-I isolated by a 


antennae, without articulations. Ceratostyles tapering, without 

articulations. A-I to setiger 1; A-II to setiger 4; A-III to setiger 


dorsally, indistinct ventraily and indistinguishable laterally; 

anterior ring 3 A of total peristomial length. Peristomial cirri 

outreaching prostomium, slender and tapering, without articulations. 

Maxillary formula 1+1,9+9,11+0, 8+11, and 1+1 according 

to Monro. Mx III long. Left and right Mx IV apparently 

matching. Distal teeth on left Mx II illustrated as being very 

small. 





number of setigers. All branchiae except last 2 or 3 pairs 

pectinate; maximum filaments 33 in largest syntype, 17 and 13 

in 2 other syntypes; maximum filaments reached at setigers 

15-20. Branchial stems slender, erect Filaments long and 

slender. 

Anterior and median neuropodial acicular lobes truncate, 

becoming irregularly rounded in posterior setigers (Figure 

114c); aciculae emerging at midline. Pre- and postsetal lobes 

low transverse folds. First 3 ventral cirri thick and tapering. 

Ventral cirri modestly basally inflated in median setigers. 


cirri not basally inflated, digitiform. Notopodial cirri long, 

slender and tapering, with distinct internal aciculae, without 


Limbate setae slender. Shafts of pectinate setae (Figure 

114e) slender, blade flaring, flat One marginal tooth longer 


falcigers (Figure 114d) tapering, dentate along cutting margins. 

Appendages nearly triangular in outline, bidentate. Proximal 

teeth low, triangular and directed laterally. Distal teeth delicate, 

tapering, nearly erect Guards symmetrically pointed, but not 

mucronate, marginally serrated. Pseudocompound falcigers 

and compound spinigers absent Aciculae single anteriorly, 

paired posteriorly, yellow, slender, tapering, straight; crosssection 

round. Separation of cores and sheaths indistinct in both 


114f,g) yellow, bidentate. Hooks first present from setiger 37, 

30, and 28 in 3 syntypes, present in all setigers thereafter, 

always single (except for replacements). Hooks slender; 

proximal teeth larger than distal teeth, directed distinctly 

laterally. 


anal cirri. 





Characters: 1, 2, 4, 6, 13, 14, 63. 


None. 

KEY.—


FIGURE 114.—Eunice validobranchiata (syntype, BM(NH) ZK 1937.9.2.312-6): a, anterior end, lateral view; b, 

parapodium 17, anterior view; c, parapodium 64, anterior view; d, compound falciger, parapodium 17; e, pectinate 

seta, parapodium 64; f, subacicular hook, parapodium 64; g, subacicular hook, parapodium 64. Eunice 

violaceomaculata (syntype, MCZ 163): h, anterior end, lateral view; i, parapodium 3, anterior view; j, compound 

falciger, parapodium 3; k, pectinate seta, parapodium 3; 1, parapodium 45, anterior view; m, pectinate seta, 

parapodium 45; n, compound falciger, parapodium 45; o, subacicular hook, parapodium 45. (Scale bars in mm.)


TABLE 16.—Variable and invariable features in the type lot of Eunice validobranchiata (N = number of 



N 

Max. 

Min. 

Mean 

SD 

Maximal width 





No. of teeth of pectinates 


3 

3 

3 

3 

3 

3 

3 

5 

12 

52 

28 

45 

9 

42 

4 

10 

41 

20 

40 

8 

28 

4.33 

10.67 

47.67 

24.33 

43.33 

8.33 

37.00 

0.58 

1.15 

5.86 

4.04 

2.89 

0.58 

7.81 

LNVARIABLE FEATURES 




Perislomial cirral articulations 


Articulations in notopodial cirri 








N=3 

absent 



absent 

3 

absent 


inflated 

yellow 


yellow 

2 

indistinct 

REMARKS.—Eunice validobranchiata is a member of group 

Al and resembles several of the other deep-water species from 

the same group closely. These species are listed in Tables 19 

and 21. The detailed relation between this species, E. 

hawaiensis and E. megabranchia, is discussed in the "Remarks" 

section of E. hawaiensis. 

Eunice violacea Grube, 1856:55-56. 

Eunice violacea Grube, 1856 

REMARKS.—This species was described from Puntarenas, 

Costa Rica; no specimens are currently available. 

The information given in the original description indicates 

that A-III reaches setiger 7 and has 5 articulations; A-II and A-I 

are similar in length; each barely reaching beyond the 

peristomium and with two articulations. The peristomial cirri 

are two-thirds the length of the peristomium. Notopodial cirri 

lack articulations. Branchiae are present from setiger 6 and 

continued to the end of the incomplete specimen (68 setigers) 

and with maximally 28 filaments. Setal structures were poorly 

characterized by Grube; he did mention that the aciculae and 

subacicular hooks were dark. 

The species is not sufficiently characterized to identify it 

even to group; it may be a member of group B-2 and related to 

a long series of rather similar species, but cannot be sufficiently 

identified to allow a safe assignment of synonymy. It is here 


200. Eunice violaceomaculata Ehlers, 1887 

FIGURE 114h-o; TABLES 17,27,30 

Eunice violaceomaculata Ehlers, 1887:86-87, pi. 24: figs. 11, 12, pi. 25: figs. 

1-7. 

MATERIAL EXAMINED.—4 syntypes, MCZ 163, Tortugas, 

Florida, shallow water. 


based on a complete syntype with 215 setigers. The material 

includes in addition a very large anterior fragment of 132 

setigers, a rather small complete specimen with 114 setigers, 

and finally an incomplete specimen, consisting of 165 setigers. 

The three smaller specimens clearly belong to the same species 

and the concept of the species is here wholly based on these 

specimens. 

DESCRIPTION.—One syntype complete with 215 setigers; 

total length 170 mm; maximal width 7 mm at setiger 10; length 

through setiger 10, 12 mm. Second specimen complete with 

114 setigers; length 62 mm; maximal width 5 mm at setiger 10. 

Third specimen incomplete with 165 setigers; length 225 mm; 

maximal width 9 mm at setiger 10. Fourth specimen 

incomplete with 132 setigers; length 340 mm long; maximal 

width 22 mm at setiger 10. 


than peristomium, as deep as x /i of peristomium. Prostomial 

lobes widely diverging and frontally obliquely truncate, 

dorsally somewhat flattened; median sulcus deep. Dark eyes 

between bases of A-I and A-II. Antennae in a horseshoe, evenly


TABLE 17.—Variable and invariable features in the type lot of Eunice violaceomaculata (N = number of 

individuals examined; SD = standard deviation; measurements in mm; * = SD not calculated). 



Total length 

Maximum width 



No. of pcristomial cirral articulations 






N 

2 

2 

4 

4 

3 

3 

4 

4 

2 

3 

4 

Max. 

215 

170 

22 

40 

18 

9 

7 

30 

70 

10 

77 

Min. 

114 

62 

5 

7 

7 

5 

6 

12 

50 

9 

25 

Mean 

164.50 

116.00 

10.75 

18.25 

13.33 

6.67 

6.25 

20.25 

60.00 

9.67 

44.25 

SD 

* 

* 

7.68 

14.80 

5.69 

2.08 

0.50 

7.41 

• 

0.58 

22.65 




No. of notopodial cirral articulations 

Notopodial cirri articulated 



Guards of compound falcigers 






N=4 



6 

throughout body 

flat; tapering 

tapering 

symmetrically rounded 

dark brown 

tapering 

brown 

2 

distinct 



tapering, with up to 10 cylindrical articulations in A-III. A-I to 

posterior peristomial ring; A-II to setiger 4; A-III to setiger 7 

(complete only in a single specimen). Peristomium flaring 

anteriorly, more strongly so when distorted by everted jaws, 



peristomial length. Peristomial cirri to front margin of 

peristomium or to middle of prostomium, slender and tapering, 


Maxillary formula (summarized for 2 specimens) 1+1, 5+5, 

6-7+0,4-6+9, and 1+1. Mx III short, forming part of distal arc 

with left Mx IV. Mx VI missing. 



Branchiae from setiger 6-7 to setiger 215. Branchiae present to 


of setigers. Maximum 20 filaments at about setigers 15-20. 

Branchial stems thick, relatively pliable, doubled over in 

several segments. Filaments digitiform, distinctly shorter than 


Neuropodial acicular lobes obliquely truncate with aciculae 

emerging from elevated knobs near superior end of lobes. All 

presetal lobes low transverse folds. Anterior and median 

postsetal lobes following outline of acicular lobes closely, 

becoming free, inferiorly elevated folds in posterior setigers! 

First 4 ventral cirri thick and tapering, nearly triangular in 

outline. Ventral cirri basally inflated from about setiger 5 and 

retaining inflated base through first half of body. Inflated bases 

thick transverse welts, relatively low in setigers 25-35, 

thereafter becoming more distinctly rounded cushions in next 

20 or so setigers. Far posterior ventral cirri triangular in outline 

and tapering to blunt tips, forming shallow scoops around 

lower edge of parapodia. All notopodial cirri tapering, with up 

to 6 long, cylindrical articulations in anteriormost setigers 

(Figure 114i). Notopodial cirri of larger specimens rather thick 

basally, retaining tapered tips in all setigers. Notopodial cirri 

shorter than branchiae except in anteriormost and posteriormost 

branchiated setigers. 


114k,m) flat, tapering, with rather wide, delicate teeth. 

Marginal teeth no longer than other teeth; -7-8 teeth present. 

Shafts of compound falcigers (Figure 114j,n) marginally 

smooth, tapering. Appendages tapering and relatively short, 

bidentate. In 1 specimen proximal teeth directed laterally, in 

other specimens directed obliquely distally. Distal teeth slender 

and obliquely curved in all specimens. Guards marginally 

smooth, symmetrically rounded; mucros absent. Pseudocompound 


single, dark brown to nearly black, tapering to slender tips, 

distally curved or straight; cross-sections round. Subacicular 

hooks (Figure 114o) dark brown, bidentate. Hooks first present 

from setiger 25, 36, and 39, present in all setigers thereafter,



tapering to relatively small head. Proximal teeth curved and 

directed laterally in 2 specimens and triangular and directed 

laterally in smallest specimen. Distal teeth smaller than 

proximal teeth, curved in all specimens. 


anal cirri. 





Characters: 39,40, 74, 78. 


None. 

REMARKS.—Part of the variability in the material may be 

due to the wide range of sizes of the type specimens. The very 

large specimen has very short antennae, not reaching beyond 

the prostomium and short peristomial cirri, and the subacicular 

hooks are not present before setiger 77; it may belong to the 

same species as the other three specimens, but without adequate 

intermediate specimens it is difficult to assign it to the same 

species. Similarly, there are some morphological differences 

between the two intermediate-sized specimens and the smallest 

one: the small specimen has nearly symmetrical neuropodial 

acicular lobes and laterally directed proximal teeth on the 

subacicular hooks. Furthermore, the ventral cirri are tapering 

and relatively narrow and do not form a scoop in posterior 

setigers. It is thus possible that the type lot of E. violaceomaculata 

consists of three different species, but this cannot be 

explored without access to sufficient and well-preserved 

material. 

The species are listed with similar species in Tables 27 and 

30. Of the species with more than 10 branchial filaments where 

the branchiae are best developed, two, E. flavopicta and E. 

reducta, have articulated peristomial cirri, the other five species 

lack articulations in the peristomial cirri. Of these five, E. 

macrobranchia has branchiae about as long as the notopodial 

cirri; the remaining species have the branchiae outreaching the 

notopodial cirri. In E. coccinea, the ventral cirri remain inflated 

through the posterior end, in the remaining three species, E. 

laticeps, E. purpurea, and E. violaceomaculata, posterior 

ventral cirri lack inflations; they are tapering in E. laticeps and 

E. purpurea and widely triangular, nearly scoop-shaped in E. 

violaceomaculata. 

Palola viridis Gray, 1847:18. 

Eunice viridis auctores 

REMARKS.— This species is the type species of the genus 

Palola. It is often considered as a Eunice and is for that reason 

listed also in this context. It is being considered in a 

forthcoming review of Palola. 

and 

201. Eunice vittata (Chiaje, 1829) 

Nereis vittata Chiaje, 1829:195. 

Eunice vittata.—Grube, 1850:293. 

Eunice minuta.—Grube, 1866b:68. 

FIGURE 115a-i; TABLES 18.41,42 

MATERIAL EXAMINED.—Gulf of Salerno, coll. A. Giangrande, 

4 specimens. 

COMMENTS ON MATERIAL EXAMINED.—The types are no 

longer in existence. The material examined was collected close 

to the type locality in the Gulf of Naples. 

DESCRIPTION.—Specimen described and illustrated complete 

adult female with 73 setigers; total length 23.5 mm; 


mm. Body circular in cross-section with relatively prominent 

parapodia throughout; posterior end tapering relatively 

abruptly towards pygidium. Longest anal cirri less than last 5 

setigers combined. 



lobes frontally rounded, dorsally mildly inflated; median sulcus 

distinct ventrally, completely indistinct dorsally. Palpal region 

distinctly marked as paired, ovate anteroventrally directed 

fields on ventral surface of prostomium. Eyes hidden below 

translucent peristomial fold, black, behind bases of A-I. 

Antennae in a horseshoe with A-I well in front of other 

antennae, evenly spaced, similar in thickness. Peristomial fold 

covering bases of A-II and A-III. Ceratophores ring-shaped in 


digitiform, tapering abruptly to short, blunt tips, with up to 8 

very poorly marked, cylindrical articulations in A-III. A-I to 


cylindrical. Separation between rings very distinct on all sides; 

each ring slightly inflated. Peristomial cirri to middle or nearly 

the front edge of prostomium; slender and tapering, with 3 

long, cylindrical articulations. 

Summary maxillary formula for 2 specimens 1+1, 9-10+9- 

10, 8-9+0, 6+8-12, and 1+1. Jaws not calcified, with slender, 

tapering teeth. Mx III long, straight and located behind left Mx 

11. Mx VI missing. 

Branchiae (Figure 115d) present, pectinate, about as long as 




number of setigers. Last branchia with single filaments, 

maximum number of filaments 12 reached at about setiger 12. 

Branchial stems tapering, held erect over body in most setigers. 

Filaments slender, tapering to very fine tips, about as long as 

notopodial cirri in best-developed branchiae. 

Anterior neuropodial acicular lobes symmetrically rounded; 

median and posterior neuropodial acicular lobes slightly more 

conical and slightly asymmetrical with aciculae emerging 

above midline (Figure 115e). All presetal lobes are low, 

transverse folds. Anterior postsetal lobes with high sections

338 


J 

1 

.5 

1 

.5 

t>. c. t, g, h, i, k. n, o. p. q. < 

a 

d 

. m, s 

FIGURE 115.—Eunice vittata (Gulf of Salemo): a, anterior end, lateral view; b, pectinate seta, parapodium 8; c, 

compound falciger, parapodium 8; d, parapodium 8, anterior view; e, parapodium 49, anterior view; f, subacicular 

hooks, parapodium 49; g, compound falciger, parapodium 49; h, pectinate seta, parapodium 49; i, aciculae, 

parapodium 49. Eunice vittalopsis (holotype, AHF Poly 0337): j, anterior end, lateral view; k, compound falciger, 

parapodium 2; 1, parapodium 2, anterior view; m, parapodium 25, anterior view; n, pectinate seta, parapodium 25; 

o, compound falciger, parapodium 25; p, subacicular hook, parapodium 81; q, pectinate seta, parapodium 81; r, 

compound falciger, parapodium 81; s, parapodium 81, anterior view. (Scale bars in mm.)


TABLE 18.—Variable and invariable features in the material examined of Eunice vittata (N = number of 




Total length 

Maximal width 







N 

4 

4 

4 

4 

4 

4 

4 

4 

4 

Max. 

78 

24 

2 

4 

8 

27 

12 

25 

22 

Min. 

62 

9 

0.75 

1.75 

4 

20 

6 

19 

20 

Mean 

69.75 

19.13 

1.69 

3.44 

7.00 

23.75 

9.00 

23.50 

20.75 

SD 

7.14 

6.98 

0.63 

1.13 

2.00 

2.99 

2.45 

3.00 

0.96 



Perislomial cirri reach 



Notopodial articulations 








N=4 



3 

3 

absent 


5 

inflated 

yellow 

yellow 

3 

indistinct 

superiorly; these sections wrapped around superior edge of 

acicular lobes and terminating on front of acicular lobes 

approximately at midline, covering emerging aciculae. Remainder 

of postsetal lobes following outline of acicular lobes 

closely. High sections located inside presetal lobes on anterior 

face of parapodia, reaching well beyond acicular lobes. First 4 

ventral cirri tapering and blunt-tipped. Ventral cirri basally 

inflated from about setiger 5 through setiger 25. Inflated base 

ovate and very distinctly set off from large, tapering narrow 

tips. Bases of ventral cirri merge into body wall posterior to 

setiger 25, in posterior setigers only tapering, blunt tips of 

ventral cirri retained. Anterior notopodial cirri basally slightly 

inflated, tapering distally to slender, digitiform tips. Posterior 

notopodial cirri reduced in length to ~ x li of length in anterior 

setigers, tapering from base. Notopodial cirri without articulations. 

Limbate setae nearly capillary with very indistinct limbation, 

relatively thick with short, tapering tips, marginally very finely 

pilose. All pectinate setae (Figure 115c,h) very small, tapering, 

flat One marginal tooth very thick and long; ~5 teeth present. 

Anterior pectinate setae slightly larger than posterior ones. 

Shafts of compound falcigers (Figure 115b,g) inflated with 

distinct distal beak, marginally coarsely serrated. Anterior 

appendages slightly longer than posterior ones, and slightly 

less tapering, bidentate. Proximal teeth narrowly triangular and 

directed slightly basally. Distal teeth distinctly shorter than 

proximal teeth, bent and directed laterally, thick and relatively 

blunt in posterior setigers, slender and tapering in anterior 

setigers. Anterior guards tapering to sharp, symmetrical tips, 

but not mucronate. Posterior guards symmetrically rounded, all 

guards marginally finely serrated; mucros absent. Pseudocompound 


(Figure 115i) paired, tapering, clear yellow; tips distinctly bent 

ventrally; cross-sections round. Dorsal aciculae in posterior 

setigers with short, pointed guards. Separation between core 


Subacicular hooks (Figure 115f) yellow, tridentate with teeth in 

a crest. Hooks first present from setiger 20-22, present in all 

setigers thereafter; multiple hooks in most setigers. 








None. 

REMARKS.—Eunice vittata is listed with similar species in 

Tables 41 and 42. The relationship between this species and 

other species with multiple subacicular hooks and articulated 

ceratostyles was indicated in the remarks to E. indica. 

Eunice vittata has been reported from all over the world. 

Most of these reports probably are of related species, including 

the species listed in Table 42.

340 

202. Eunice vittatopsis Fauchald, 1970 

FIGURE 115j-s; TABLES 41, 43 

Eunice vittatopsis Fauchald, 1970:50-52, pi. 7: figs. a-d. 

MATERIAL EXAMINED.—Holotype, AHF Poly 0337, Velero 

III sta 739-37, Mexico, Gulf of California, Ensenada de San 

Francisco, 27°57'05"N, lll°03'20"W, 30 Mar 1937, intertidal, 

shingle, hand collected. 


length 38 mm; maximal width 2.5 mm at setiger 10; 

length through setiger 10, 6 mm. Body with ovate crosssection; 

each segment more than twice as wide as long, tapering 

slightly towards anterior end. 

Prostomium (Figure 115j) distinctly shorter and narrower 


Prostomial lobes frontally obliquely truncate; dorsal surface 

pustulate; median sulcus shallow. Eyes between bases of A-I 

and A-I I, dark purple. Antennae in a horseshoe, evenly spaced, 


without articulations. Ceratostyles slender and tapering; 3 

medial antennae with cylindrical articulations; A-I with distally 

drop-shaped articulations; maximum number of articulations 

six in A-I and A-III. A-I to setiger 2; A-II and A-III to setiger 

7. A-II slightly longer and distinctly heavier than A-III. 

Peristomium distorted by protruded jaws, apparently cylindrical. 

Separation between rings distinct on all sides; anterior ring 

3 A of total peristomial length. Peristomial cirri reaching well 

beyond tip of prostomium, slender, tapering, with 4 long 



long and located behind left Mx II. Mx VI absent. 

Branchiae (Figure 115m,s) present, pectinate, distinctly 

longer than notopodial cirri, not reduced in mid-body region, 

erect. Branchiae from setiger 3 to end of fragment. First 2 pairs 

single filaments; all other branchiae with 3 or more filaments; 

maximum 10 filaments present. Branchial stems slender, 

pliable, tapering. Filaments very long and slender, as long as 

notopodial cirri in all setigers. 

Anterior neuropodial acicular lobes distally rounded (Figure 

1141); median acicular lobes symmetrically rounded; aciculae 

emerging at midline. Posterior acicular lobes obliquely truncate 

with aciculae emerging from upper end. Pre- and postsetal 

lobes low folds. First 4 ventral cirri tapering and slightly 

inflated. Ventral cirri distinctly basally inflated by setiger 5 

through about setiger 55-60. Inflated base ovate; narrow tips 

digitiform. In last setigers present ventral cirri slender, 

digitiform, longer than those in anterior setigers. Notopodial 

cirri with 3 to 4 long, cylindrical articulations. Anterior 

notopodial cirri medially, thick, increasing in length through 

first 5 setigers, with long, slender digitiform tips in anterior 

branchial setigers, tapering and slender from about setiger 20. 

Limbate setae rather wide, marginally serrated. Pectinate 

setae (Figure 115n,q) very narrow, with cylindrical shafts; 


blades fiat and tapering. One marginal tooth very large; 6-7 

teeth present. Shafts of compound falcigers tapering in anterior 

and median setigers (Figure 115k,o), becoming slightly inflated 

in posterior setigers (Figure 115r). Anteriorly shafts marginally 

smooth, becoming marginally serrated in last setigers present. 

Appendages rather similar in all setigers, tapering, with rather 

large heads, bidentatc. Proximal teeth narrowly triangular, 

directed laterally. Distal teeth about as large as proximal teeth, 

curved. Guards asymmetrically bluntly pointed; marginal 

serrations absent in anterior, present in posterior setigers; 


spinigers absent. Aciculae paired in most sctigers, yellow, 

slender, tapering to sharp tips; most straight, but some sharply 

bent; cross-sections round. Separation between core and sheath 


hooks (Figure 115p) yellow, tridentate with teeth in a crest. 

Hooks first present from seliger 39, present in all setigers 

thereafter, always single (except for replacements). Main fangs 

large; secondary fang small and tertiary fang very small, 

appended to back of secondary fang. 

UNKNOWN MORPHOLOGICAL FEATURES.—Posterior distribution 

of branchiae; pygidium and anal cirri. 





Characters: 1, 2, 36-38,40,42, 63. 


38,2. 

REMARKS.—Eunice vittatopsis belongs to group C-l; Fauchald 

(1970) reported non-type specimens with a limited 

numbers of pairs of branchiae. It is compared to similar species 

in Tables 41 and 43. It is the only species in Table 43 in which 

the peristomial cirri project clearly beyond the prostomium. 

Eunice vivida Stimpson, 1854 

Eunice vivida Stimpson, 1854:35, fig.26. 

REMARKS.—This species, of which no specimens are 

available, was described from Grand Manan in the Bay of 

Fundy. The description and illustration showed a species with 

long, slender, tapering peristomial cirri and branchiae present 

from setiger 3 through setiger 42, and branchiae pectinate with 

a maximum of nearly 20 filaments. Nothing is mentioned about 

the setal distribution or structure, nor are there any additional 

details that allow identification. Stimpson may have had a 

specimen of E. pennata (see above), but the information 

present in the original description is insufficient to allow 

identification. Eunice vivida is here considered indeterminable. 

203. Eunice wasinensis, new name 

FIGURE 116a-d; TABLES 33, 40 

Nicidion gracilis Crossland, 1904:327-329, figs. 65,66 pi. 22: figs. 10, 11.


1 

.05 

I 

1 

h 

a 

FIGURE 116.—-Eunice wasinensis (syntype, BM(NH) ZK 1928.4.11.50): a, anterior end, lateral view; b, 

parapodium 60, anterior view; c, pectinate seta, parapodium 60; d, subacicular hook, parapodium 60. Eunice 

websleri (holotype of Eunice loncicirrala Webster, USNM 4792): e, anterior end, lateral view; f, pectinate seta, 

anteromedian parapodium; g, compound falciger, anteromedian parapodium; h, anteromedian parapodium, 

anterior view; i, posterior parapodium, anterior view; j, pectinate seta, posterior parapodium; k, compound 

falciger, posterior parapodium; 1, subacicular hooks, posterior parapodium. (Scale bars in mm.)

342 

MATERIAL EXAMINED.—TWO syntypes, BM(NH) ZK 

1928.4.11.50, Wasin Harbour, near border of Kenya and 

Tanganyika, dredged, Crossland Zanzibar and British East 

Africa collection, 1901-1902, average depth 10 fathoms, 

extensive cover of Telesto. 

COMMENTS ON MATERIAL EXAMINED.—One of the two 

syntypes is complete; the other is an anterior fragment. 

DESCRIPTION.—Complete syntype of unknown sex; with 84 

setigers; posterior regenerate including last 15 setigers; total 


through setiger 10, 6 mm. Anterior fragment with 77 setigers; 

length 14 mm; width 2.0 mm. 

Prostomium (Figure 116a) about as long and as wide as 

peristomium, as deep as V2 of the peristomium. Prostomial 

lobes large, frontally rounded, dorsally inflated; median sulcus 

deep. Eyes between bases of A-I and A-II; black. Antennae in 

a shallow horseshoe, evenly spaced, similar in thickness. 


Ceratostyles sausage-shaped, without articulations; none projecting 

clearly beyond tip of prostomium or beyond posterior 

peristomial rings. Peristomium cylindrical, slightly flaring 

ventrally. Separation between rings distinct dorsally and 

ventrally, less distinct, but visible laterally; anterior ring ~ 2 /3 of 

total peristomial length. Peristomial cirri barely to front edge of 

posterior ring, tapering, without articulations. 

Maxillary formula 1+1, 5+5, 4+0, 3+8, and 1+1 according 

to Crossland (1904, fig. 65). Mx III part of distal arc with left 

MxIV. 


Anterior neuropodial acicular lobes rounded; acicular lobes 

conical in median and sharply pointed in far posterior setigers 

(Figure 116b); aciculae emerging at midlinc. Presetal lobes 

low, transverse folds. First 20 postsetal lobes distinct, rounded, 

shorter than acicular lobes, following outline of acicular lobes 

in median and posterior setigers. First 4 ventral cirri thick, 

tapering. Ventral cirri basally indistinctly inflated in median 

setigers. Inflated bases transverse welts; narrow tips absent. 

Posterior ventral cirri without inflated bases, short, nearly 

tubercular. Notopodial cirri short, without articulations; median 

and posterior notopodial cirri increasingly filiform, retaining 

same length as in anterior setigers. Notopodial cirri without 


Limbate setae not observed. Pectinate setae (Figure 116c) 

short, flaring, furled. Both marginal teeth longer than other 

teeth; -10 coarse teeth present. Shafts of compound falcigers 

inflated, marginally smooth. Appendages bidentate. Guards 


falcigers and compound spinigers absent Aciculae single, 

black, pointed, straight; cross-sections round. Subacicular 

hooks (Figure 116d) black, bidentate. Hooks first present from 

setiger 31 in 1 syntype and from setiger 27 in other syntype, 


replacements). Hooks tapering, with small head. Proximal teeth 


smaller than distal teeth, directed laterally, triangular. Distal 

teeth tapering, erect 









None. 

REMARKS.—A new name is needed because the name E. 

gracilis is preoccupied in the combination E. gracilis Grube, 

1867. 

The species is listed with similar species in Tables 33 and 40. 

Three species in Table 40 combine a lack of articulations in the 

ceratostyles with inflated bases of the ventral cirri limited to 

median setigers. In addition to E. wasinensis, the two other 

species are E. carihoea and E. goodei. In E. cariboea the 

inflated bases are transverse ridges; in the two other species, 

they are ovate and more or less free from the ventral edge of the 

ncuropodia. In E. wasinensis the inflated ventral cirri lack a free 

tip; such tips are present in E. goodei. 

204. Eunice websteri Fauchald, 1969 

FIGURE 116e-l; TABLES 19. 20 

Eunice websteri Fauchald, 1969:12-14, fig. 6. 

Eunice longicirrala Webster, 1884:318-319, pi. 12: figs. 75-80. 

Leodice margantacea Vcrrill, 1900:644-645. 

Leodice longicirrata.—Tie&dv/cM, 1921:11-14, figs. 3-12, pl.l: figs. 1-4 [not 

Eunice longicirrala Kin berg, 1865]. 

Eunice longicirrala.—Hartman, 1942:9 [not Eunice longicirrala Kinberg, 

1865]. 

MATERIAL EXAMINED.—Holotype, 1 specimen and 8 

mounted slides, USNM 4792, Bermuda, coll. G. Brown Goode. 



through setiger 10, 8 mm. Body slightly dorsoventrally 

flattened, but basically convex dorsally and more flattened 

ventrally. Anal cirri as long as last 15 setigers, slender, with 15 



than peristomium, as deep as l /i of peristomium. Prostomial 

lobes frontally truncate, dorsally flattened; median sulcus very 

shallow. Eyes lateral to bases of A-II, dark. Antennae in a 



slender and digitiform, with mostly long, cylindrical articulations. 

Distal articulations in A-I short, very well set off from 

each other and nearly moniliform; maximum number of 

articulations 8 in A-I 11. A-I to posterior peristomial ring; A-II to 

setiger 6; A-I 11 to setiger 8. Peristomium cylindrical. Separation 

between rings distinct dorsally and ventrally; anterior ring


3 A of total peristomial length. Peristomial cirri to middle of 

prostomium, very slender, with 6 long, cylindrical articulations. 






number of setigers. First and last 10 pairs single filaments; 

maximum 15 filaments. Branchial stems slender, tapering. 

Filaments slender, longer than notopodial cirri in median 

branchial setigers Anterior neuropodial acicular lobes 

broadly and nearly symmetrically rounded, becoming conical 

in posterior setigers (Figure 116i); aciculae emerging at 

midline. All presetal lobes low, transverse folds. Anterior 

postsetal lobes low, rounded, free folds, becoming low, 

transverse folds by end of branchial region. Postsetal lobes 

shorter than acicular lobes in all setigers. First 6 ventral cirri 

thick, nearly triangular and tapering. Ventral cirri basally 

inflated from about setiger 7 through about setiger 55. Inflated 

bases modest, ovate inflated; narrow tips digitiform. Postbranchial 

ventral cirri digitiform. All notopodial cirri with 2 or more 

long, cylindrical articulations. Anterior notopodial cirri tapering, 

long and distally truncate; median and posterior notopodial 

cirri digitiform. 

All limbate setae slender and marginally smooth. Pectinate 

setae (Figure 116f, j) narrow, tapering, flat. One marginal tooth 

longer than other teeth; 10 teeth present. Shafts of compound 

falcigers (Figure 116g,k) tapering and internally striated, but 

without marginal teeth. Appendages short, tapering with 

distinct heads, bidentate. Proximal teeth triangular, directed 

laterally or slightly basally. Distal teeth obliquely curved. 

Guards distally round, marginally smooth; mucros absent 


Two or 3 aciculae in a parapodium, yellow. Anterior aciculae 

bluntly tapering or nearly truncate; median aciculae distally 

expanded to distinct rounded tabs; posterior aciculae becoming 

slender, tapering to fine tips, sometimes distinctly bent 

dorsally; cross-sections round. Separation between core and 


Subacicular hooks (Figure 1161) yellow, bidentate. Hooks first 

present from setiger 31, present in all setigers thereafter, paired 

in most setigers. Hooks tapering, with large, laterally directed 

proximal teeth. Distal teeth smaller than proximal teeth, nearly 

erect 








None. 

REMARKS.—As noted by Fauchald (1969), the name 

originally applied by Webster to this taxon was preoccupied 

and had to be replaced. Eunice websteri is a member of group 

A-l and is compared to similar taxa in Tables 19 and 20. 

Median aciculae are distally expanded into rounded tabs, a 

feature present only in one additional species listed in Table 20, 

E. antillensis. Eunice websteri has about twice the number of 

branchial filaments as E. antillensis; furthermore, in E. 

websteri, the articulations of the ceratostyles are distally 

drop-shaped; in E. antillensis, they are cylindrical along the 

whole length of the antennae. 

205. Eunice woodwardi Baird, 1869 


Eunice woodwardi Baird, 1869:347-348. 

MATERIAL EXAMINED.—Holotype, BM(NH) ZH 

1863.8.19.13, La Corufta, Spain. H. Woodward, Esq. coll. 


poorly fixed and is now very soft. The illustration of the 

anterior end is somewhat idealized to avoid showing wrinkles 

caused by collapse of internal structure. 



through setiger 10, 7 mm. Body shape indeterminable. 




flattened; median sulcus deep. Eyes not observed. Antennae in 

a horseshoe, with A-I isolated by a gap, similar in thickness. 


Ceratostyles slender and digitiform, with up to 4 or 5 long, 

rather indistinct articulations in A-III. A-I to posterior 

peristomial ring; A-II to setiger 3; A-III to setiger 9 or 10. 


dorsally and ventrally; anterior ring about 

2 /3 of total 

peristomial length. Peristomial cirri to frontal edge of peristomium, 



Branchiae present pectinate, distinctly longer than notopodial 




setigers. All branchiae, except last 4 or 5 with 2 or more 

filaments maximum number of filaments 12 at about setiger 15. 

Branchial stems slender. Filaments longer than notopodial cirri 

in median branchiferous setigers. 

Neuropodial acicular lobes truncate; aciculae emerging at 


ventral cirri tapering and digitiform. Ventral cirri basally 

inflated from about setiger 4 through setiger 40, Inflated bases 

ovate; narrow tips tapering. Postbranchial ventral cirri slender 

and tapering. Prebranchial notopodial cirri medially inflated, 

becoming increasingly slender posteriorly and digitiform in

344 


•02 b. c, d 

5 

FIGURE 117.—Eunice woodwardi (holotype, BM(NH) ZH 1863.8.19.13): a, anterior end, lateral view; b, 

compound falciger, parapodium 45; c, acicula, parapodium 45; d, subacicular hook, parapodium 45. (Scale bars 

in mm.) 

postbranchial region. Notopodial cirri without articulations. 

Limbate setae rather wide and marginally finely serrated. 

Pectinate setae not observed. Shafts of compound falcigers 

(Figure 117b) tapering, marginally smooth. Appendages large 

with distinct heads, bidentate. Proximal teeth larger than distal 

teeth, triangular, directed slightly basally. Distal teeth curved. 

Guards symmetrically rounded with a distinct mucro, marginally 

serrated. Pseudocompound falcigers and compound spinigers 

absent. Aciculae (Figure 117c) paired, yellow, tapering 

and bent dorsally; cross-sections round. Separation between 


hooks. Subacicular hooks (Figure 117d) yellow, tridentate with 

teeth in a crest Hooks first present from setiger 31, present in 

all setigers thereafter, most postbranchial setigers with 4 to 5 

hooks in each segment. Hooks with large main fangs; distal pair 

of fangs in a crest. 






SCORED.—Inappropriate Characters: 56, 58, 59. Unknown


Characters: 1, 2, 13, 14, 63,65-68. 


None. 

REMARKS.—Eunice woodwardi was considered indeterminable 

by Hartman (1959); it is listed with similar species in 

Tables 41 and 42. It is the only species in Table 42 to combine 

the presence of multiple subacicular hooks with incomplete 

separation between the anterior and posterior peristomial rings. 

Eunice zonata Chiaje, 1841 

Eunice zonata Chiaje, 1841:94.—Quatrefages, 1866:330.—Ehlers, 

1868:312.—Fauvel, 1923:451. 

REMARKS.—Quatrefages (1866:330) listed this species 

without any comments; Ehlers and Fauvel both considered it 

incertae sedis. As far as known, no material exists and it is here 


TABLE 19.—Comparison of species in group A-l. Column numbers are the same as character numbers used to 

prepare the key (see explanation below; fora complete listing of characters and character states, see Appendix A). 

11 

13 

15 

19 

25 

26 

36 

55 

85 

86 

94 

98 

114 

120 

121 

132 

146 

150 

153 

173 

176 

179 

180 

193 

198 

199 

204 

Species 

E. antarctica 

E. antillensis 

E. arcturi 

E. arliculata 

E. benedicti 

E. biannulata 

E. caeca 

E. edwardsi 

E. hawaiensis 

E. heterochaeta 

E. japonica 

E. kobiensis 

E. manihine 

E. megabranchia 

E. mtxicana 

E. narconi 

E. parasegregala 

E. pectinata 

E. pennata 

E. rubrocincta 

E. savignyi 

E. segregata 

E. semisegregata 

E. tridentata 

E. valens 

E. validobranchiala 

E. websleri 

1 

122 

U 

110 

U 

105 

139 

90 

118 

U 

105 

110 

U 

127 

U 

106 

U 

u 

u 

114 

100 

U 

U 

U 

185 

179 

U 

150 

2 

102 

U 

80 

U 

38 

105 

103 

70 

U 

27 

50 

U 

75 

U 

87 

U 

U 

U 

73 

68 

U 

U 

U 

170 

140 

U 

120 

3 

3.5 

3 

U 

3 

2.5 

3 

6 

3.5 

7 

3.5 

2.5 

3.5 

4 

7 

3.5 

2 

3 

2 

3 

3 

U 

4 

6 

7 

9 

4 

4 

5 

11, 

7 

U 

U 

5 

12 

14 

8 

13 

4 

7 

6.5 

9 

12 

10.5 

6 

8 

6 

7.5 

U 

U 

5 

9 

12 

12.5 

4 

8 

35 

3 

4 

6 

4 

3 

3 

3 

3 

3 

3 

4 

3 

8 

3 

3 

8 

3 

5 

3 

3 

4 

3 

3 

3 

3 

3 

3 

36 

39-44 

36 

42 

55 

39 

50 

48 

34 

54 

85 

46 

41 

43 

54 

40 

33-35 

U 

U 

39-41 

80 

45 

39 

65/69 

80 

76 

37-40 

53 

80 

31-44 

33 

36 

40 

21 

38 

42 

28 

30 

22 

34 

30 

29 

35 

32 

35-38 

U 

41 

35-43 

U 

U 

36 

51/55 

51 

43 

28-37 

31 

U = feature is unknown. Numbers separated by a dash indicate a range; numbers separated by a slash indicate 

paired alternative observations. Explanation of column heads: 

1 = Total number of setigers (complete specimens only). 

2 = Total length in mm (complete specimens only). 

3 = Maximum width in mm. 

5 = Length through setiger 10 in mm. 

35 = Branchiae first present from setiger number. 

36 = Branchiae present through setiger number. 

80 = Subacicular hooks first present from setiger number.

346 


TABLE 20.—Comparison of Eunice species in group A-l with articulated pcristomial cirri. Stub (Q gives 

character numbers; column heads are species numbers. Characters and character states are listed in Appendix A. 

Species are in order by number, corresponding species names are listed at the end of the table. 

c 

7 

8 

9 

11 

12 

14 

15 

16 

17 

20 

22 

24 

IJ 

26 

27 

28 

32 

34 

39 

40 

41 

42 

43 

44 

45 

48 

51 

54 

55 

60 

61 

64 

13 

1 

2 

2 

2 

2 

1 

1 

3 

1 

1 

1 

2 

J 

4 

3 

2 

3 

1 

2 

2 

6 

U 

1 

3 

2 

2 

3 

1 

2 

1 

4 

1 

15 

U 

U 

U 

U 

U 

U 

U 

U 

U 

U 

U 

U 

| T 

U 

u uuuuuu10uuuuuuuuuuu 

u 

u2 

1 

19 

1 

2 

3 

2 

1 

1 

3 

1 

6 

1 

2 

2 

1 

1 

3 

3 

2 

1 

1 

1 

2-3 

6 

15 

U 

U 

U 

Uuuu2 

4 

2 

26 36 94 98 121 150 

SOFT BODY CHARACTER STATES 

1 

2 

3 

1 

2 

2 

1 

1 

1 

1 

2 

1 

3 

2 

3 

2 

1 

1 

1 

2 

1 

3 

1 

1 

3 

2 

2 

2 

1 

3 

1 

1 

1 

1 

2 

2 

3 

2 

3 

2 

1 

U 

1 

1 

1 

1 

1 

2 

2 

1 

2 

2 

2 

2 

1 

1 

1 

1 

2 

1 

3 

1 

4 

1 

1 

1 

2-3 

8 

10 

1-2 

3 

2 

2 

5 

1 

3 

2 

2 

2 

3 

2 

2 

1 

1 

1 

0 

24 

15 

2 

3 

2 

3 

3 

1 

2 

1 

4 

2 

3 

2 

1 

1 

1 

2 

2-3 

11 

U 

3 

2 

2 

2 

3 

1 

2 

2 

1 

2 

2 

1 

1 

1 

1 

3 

2 

8 

U 

3 

2 

2 

2 

3 

1 

2 

1 

2 

2 

3 

2 

1 

1 

1 

0 

0-3 

IK 

15 

2 

3 

2 

2 

9 

1 

3 

2 

3 

2 

3 

1 

3 

2 

1 

2 

1 

1 

1 

1 

2 

1 

3 

1 

2 

1 

2 

2 

U 

10 

u 

1-2 

2 

1 

1 

3 

1 

3 

U 

2 

1 

153 

2 

1 

2 

2 

1 

1 

1 

3 

4 

1 

2 

2 

2 

1 

2 

1 

1 

5 

2 

12 

15 

1 

3 

2 

3 

5-6 

1 

2 

1 

4 

2 

176 

179 

U 1 

u 

2 

2 

uuuuuuu1 

2 

1 

3 

3 

2 

1 

3 

1 

u 1 

11 V.' 

u 2uuuuu12uuuuuuuuuuu 

u uuuuu 

3 

1 

1 

1 

1 

2 

0 

15 

25 

3 

2 

2 

3 

7 

1 

3 

2 

3 

2 

198 

2 

2 

2 

2 

:» 

2 

.J 

3 

1 

1 

1 

1 

2 

10 

11 

U 

2-3 

3 

2 

3 

9 

2 

2 

1 

1 

2 

204 

1 

2 

2 

2 

2 

2 

1 

1 

1 

3 

2 

2 

.» 

3 

2 

2 

1 

1 

1 

10 

15 

U 

2 

2 

2 

3 

6 

1 

3 

2 

3 

1 

65 

66 

74 

76 

78 

82 

2 

1 

2 

2 

2 

2 

2 

u 

2 

1 

2 

1 

2 

1 

1 

1 

2 

1 

2 

2 

SETAL CHARACTER STATES 

2 U 2 1 

1 U 1 1 

2 2 2 1 

1 

2 

1 

1 

2 

2 

1 

2 

1 

1 

1 

5 

U = feature is unknown; dash (-) = character inappropriate. Numbers separated by a dash indicate a range; 

numbers separated by a slash indicate paired alternative observations. 

Character states of characters not included in the comparisons: 10,1; 13,1; 18,3; 19,2; 21,1; 23,2; 29,1; 30,1; 

31,1; 33,2; 37,2; 38,2; 52,1; 53,1; 56,-; 57,2; 58,-; 59,-; 69,1; 70,2; 71,2; 72,2; 73,1; 75.1; 77,1; 79,1; 81.1. 

Species included: 13 E. anlillensis; 15 E. arcturi; 19 E. articulata; 26 E. biannulaia; 36 E. caeca; 94 E. 

japonica; 98 E. kobiensis; 121 E. mexicana; 150 E. pectinala; 153 E. pennata; 176 E. savignyi; 179 E. segregata; 

198 E. valens; 204 E. websteri. 

2 

2 

2 

1 

2 

1 

2 

1 

2 

1 

2 

2 

1 

1 

1/2 

1 

2 

1 

2 

1 

2 

1 

2 

2 

2 

1 

2 

1/3 

2 

3


TABLE 21.—Comparison of Eunice species in group A-1 without articulated peristomial drri. Stub (C) gives 

character numbers; column heads are spedes numbers. Characters and character states are listed in Appendix A. 

Species are in order by number, corresponding spedes names are listed at the end of the table. 

c 

7 

8 

9 

11 

12 

13 

14 

15 

16 

17 

20 

21 

22 

23 

24 

25 

26 

27 

28 

31 

32 

34 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

53 

54 

55 

57 

58 

59 

60 

61 

64 

11 

3 

1 

2 

1 

1 

U 

U 

1 

1 

1 

3 

1 

1 

2 

2 

3 

4 

2 

2 

1 

3 

1 

2 

6-7 

2 

5 

15 

1 

2 

1 

1 

3 

3 

1 

1 

3 

2 

_ 

_ 

2 

2 

1 

15 

25 

U 3 

U 2 

U 2 

U 

U 

U 

U 

uuuuuuuuuuuuuuu2uu10uuuuuuuuuuuuuuuu 

u uu2 

1 

uu 

2 

1 

1 

2 

1 

1 

1 

1 

2 

_ 

2 

2 

4 

2 

2 

2 

1 

1 

2 

2 

3 

0 

12 

15 

3-4 

3 

2 

2 

4 

1 

1 

2 

3 

1 

2 

2 

_ 

2 

3 

55 85 86 114 120 132 

3 

2 

2 

2 

1 

1 

2 

3 

3 

1 

3 

1 

1 

2 

2 

4 

2 

2 

2 

1 

2 

1 

2 

5 

2 

6 

10 

2-3 

2 

2 

2 

5 

1 

2 

1 

2 

2 

- 

- 

1 

1 

3 


3 1 2 3 

1 2 2 1 

3 2 3 2 

2 2 2 2 

2 1 1 2 

1 2 2 1 

3 _ _ 2 

1 1 1 1 

1 1 1 1 

1 2 U 1 

1 1 1 1 

2 1 2 2 

_ 1 - _ 

2 2 U 2 

U 1 1 2 

u2 3 3 4 

2 2 3 

1 1 2 3 

1 1 2 1 

1 1 2 1 

1 1 1 1 

1 1 - 1 

2 1 2 2 

0 3 - 0 

4 15 - 7 

30 7 3 47 

U 25 U 15 

2 1-2 1 1 

2 2 2 3 

2 2 2 2 

2 2 2 2 

3 4 4 3 

1 3 1 1 

1 1 1 1 

1 2 2 1 

2 2 1 2 

2 2 2 2 

- - - - 

_ _ - - 

1 2 1 1 

1 1 1 3 

3 2 3 2 

3 

1 

2 

2 

U 

u uu 

u 

u2u3 

1 

5 

2 

U 

U 

2 

U 

U 

3 

15 

U 

U 

U 

U 

u u1uuuuuuuu 

2 

146 

3 

1 

2 

2 

2 

1 

2 

2 

2 

2 

4 

2 

2 

2 

1 

1 

1 

2 

0 

U 

18 

10 

1 

3 

2 

2 

5 

1 

1 

1 

3 

2 

- 

_ 

2 

1 

2 

173 

U 

U 

U 

u uuuu3 

2 

_ 

4 

U 

4 

U 

2 

2 

1 

1 

1 

1 

U 

11 

8 

U 

Uu 

u 

u 

u1 

1 

uu2 

- 

- 

u 2 

3 

180 

3 

1 

3 

2 

1 

1 

2 

1 

1 

1 

1 

1 

1 

2 

2 

4 

2 

1 

2 

1 

1 

1 

2 

0 

0 

38 

15 

2 

2 

2 

2 

4 

1 

1 

1 

2 

2 

- 

- 

1 

3 

3 

193 

3 

1 

3 

2 

2 

U 

U 

3 

1 

1 

3 

1 

2 

2 

1 

3 

4 

2 

1 

1 

1 

1 

2 

1 

15 

14 

30 

2-3 

2-3 

2 

2 

4 

1 

1 

1 

2 

2 

- 

- 

2 

2 

3 

199 

3 

1 

3 

2 

1 

U 

u 

1 

2 

1 

1 

2 

_ 

2 

2 

3 

3 

3 

1 

1 

1 

1 

2 

0 

2-3 

33 

15 

1 

2 

2 

2 

3 

1 

1 

1 

3 

2 

- 

- 

2 

3 

3 

65 

66 

70 

74 

77 

81 

82 

2 

1 

2 

2 

1 

1 

1 

2 

1 

1 

2 

1 

1 

2 

U 

U 

2 

2 

1 

U 

u 


2 

1 

1 

2 

1 

1 

1 

2 

1 

2 

2 

1 

1 

1 

2 

2 

2 

2 

1 

1 

5 

1 

2 

1 

U 

2 

1 

1 



Character states of characters not induded in the comparison: 10,1; 18,3; 19,2; 29,2; 30,1; 33,2; 37,2; 56,-; 

69.1; 71.2; 72,2; 73.1; 75.1; 76.1; 78.2; 79,2. 

Species included: 11 E. antarclica; 15 E. arcturi; 25 E. benedicti; 55 E. edwardsi; 85 E. hawaiensis; 86 E. 

heterochaeta; 114 E. manihine; 120E. megabranchia; 132 E. narconi; 146 E. parasegregata; 173 E. rubrocincta; 

180 E. semisegregata; 193 E. iridentata; 199 E. validobranchiata. 

1 

2 

2 

1 

U 

u 

2 

1 

2 

2 

1 

1 

1 

1 

2 

2 

2 

1 

U 

U 

U 

U 

2 

4 

1 

1 

1 

2 

2 

2 

1 

1 

2 

1 

2 

2 

2 

2 

1 

1 

1

348 


TABLE 22.—Comparison of Eunice species in groups A-2 *nd A-4. Column heads are the same as character 

numbers used to prepare the key (see explanation below; for » complete listing of characters and character states, 

see Appendix A). 

18 

76 

95 

102 

146 

177 

184 

Species 

E. armillata 

E. gracilicirrala 

E.johnsoni 

E. leptocirris 

E. parasegregala 

E. schizobranchia 

E. stigmatura 

1 

114 

323 

160 

U 

U 

731 

U 

2 

46 

214 

75 

U 

u 

655 

U 

3 

2 

2.5 

3 

3 

3 

5 

2 

5 

6 

9 

7 

9 

8 

10 

4 

35 

6 

3 

24 

3 

3 

67 

3 

36 

109 

181 

130 

U 

u 

730 

110-126 

80 

34 

63 

31 

51 

U 

60 

27-35 

U = feature is unknown. Numbers separated by a dash indite a range. 

Explanation of column heads numbers: 







80 = Subacicular hooks first present from setiger number. 

TABLE 23.—Comparison of Eunice species in group A-2. Stub (C) gives 

character numbers; column heads give species numbers. Characters and 

character states are listed in Appendix A. Species are in order by number, 

corresponding species names are listed at the end of the table. 

TABLE 23.—Continued. 

c 

18 

76 95 102 146 

177 

184 

C 

18 76 95 102 146 

177 

184 

7 

8 

9 

12 

13 

14 

15 

16 

17 

18 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

32 

37 

38 

39 

40 

41 

42 

43 

44 

46 

47 

48 

49 

3 

1 

3 

1 

U 

u 

3 

1 

1 

1 

1 

1 

3 

2 

2 

4 

4 

1 

2 

2 

1 

1 

1 

1 

U 

3 

U 

1 

3 

2 

2 

3 

2 


3 

1 

3 

2 

U 

U 

1 

3 

3 

3 

1 

1 

1 

2 

2 

4 

4 

2 

2 

1 

1 

2 

2 

2 

36 

7 

20 

1 

2 

2 

2 

3 

2 

3 

1 

3 

1 

1 

3 

1 

1 

1 

3 

4 

1 

1 

2 

2 

4 

3 

1 

2 

2 

1 

2 

1 

3 

20 

10 

U 

2 

2 

2 

2 

2 

2 

2 

2 

3 

2 

U 

U 

1 

3 

1 

1 

3 

2 

- 

2 

2 

4 

2 

3 

1 

2 

3 

1 

1 

U 

U 

6 

15 

2-3 

3 

2 

2 

2 

2 

3 

1 

2 

2 

1 

2 

1 

1 

1 

3 

1 

1 

2 

2 

2 

4 

2 

2 

2 

2 

1 

2 

2 

0 

U 

18 

10 

1 

3 

1 

1 

2 

1 

2 

2 

2 

1 

U 

U 

1 

3 

1 

3 

1 

2 

_ 

4 

1 

3 

5 

1 

2 

2 

1 

1 

1 

100 

0 

7 

U 

3 

2 

2 

2 

3 

1 

3 

1 

2 

1 

1 

2 

1 

1 

1 

3 

3 

1 

1 

1 

2 

4 

2 

2 

2 

1 

2 

1 

1 

4 

40 

4 

15 

1 

3 

2 

2 

2 

2 

51 

52 

53 

5^ 

55 

56 

51 

5 

5 

60 

61 

64 

65 

66 

73 

78 

79 

81 

SOFT BODY CHARACTER STATES—Continued 

5 

3 

1 

1 

2 

- 

2 

- 

- 

2 

4 

2 

10 

3 

1 

1 

3 

- 

2 

- 

- 

2 

3 

2 

- 

1 

2 

- 

- 

1 

2 

- 

- 

1 

1 

3 

4 

1 

1 

1 

2 

_ 

2 

- 

- 

2 

1 

2 

5 

1 

1 

1 

3 

_ 

2 

_ 

_ 

2 

1 

2 

2 

2 

1 

2 

2 

1 


2 

1 

1 

2 

2 

1 

1 

1 

1 

2 

2 

1 

2 

1 

1 

2 

2 

1 

2 

1 

1 

2 

2 

1 

9 

1 

1 

1 

2 

- 

1 

2 

1 

_ 

1 

3 

1 

1 

2 

U 

2 

2 

6 

1 

1 

1 

2 

- 

2 

- 

- 

2 

3 

2 

2 

1 

1 

2 

2-3 

1 

U = feature is unknown; dash (-) = character inappropriate. Numbers 

sepai^ted by a dash indicate a range. 

Character states of characters not included in the comparisons: 10,1; 11,2; 

19,2; 30,1; 31,1; 33,2; 69,1; 70,2; 71,2; 72.2; 74,2; 75,1; 76,1; 77,1; 78,2; 82,1. 

Species included: 18 E. armillata; 76 E. gracilicirrata; 95 E. johnsoni; 102 

E. Uptocirris; 146 E. parasegregala; 177 E. schizobranchia; 184 E. 

stigmatura.


TABLE 24.—Comparison of Eunice species in group B-1. The column heads are the same as the character 

numben used to prepare the key (see explanation below; for a complete listing of characters and character states 

see Appendix A). 

7 

23 

41 

42 

54 

58 

78 

99 

120 

131 

134 

143 

168 

175 

183 

188 

Species 

E. amoureuxi 

E. barvicensis 

E. coccinea 

E. coccinoides 

E. dubitata 

E. elegans 

E. gravieri 

E. langi 

E. megabranchia 

E. muxilaloides 

E. nicidioformis 

E. palauensis 

E. rosaurae 

E. samoae 

E. Stanleyi 

E. thomasiana 

1 

U 

U 

u 

133 

U 

71 

U 

110 

97 

U 

142 

U 

130 

99 

2 

13 

U 

U 

u u 

75 

28 

42 

U 

32 

45 

U 

152 

U 

58 

45 

3 

U 

3.5 

5 

5 

7 

3 

2 

2 

7 

1 

2 

U 

4 

4 

4 

3 

5 

U 

6 

10 

8 

16 

8 

U 

5.5 

12 

5 

6 

U 

18 

5 

7 

6 

35 

9 

6 

6 

7-8 

3-5 

3 

5 

3 

3 

8 

6-8 

6-8 

3 

6 

4 

4 

36 

24 

65 

U 

78 

38-48 

33 

20/26 

39 

54 

48 

40-50 

U 

48-55 

51 

33 

79 

80 

17 

19 

38-39 

28/30 

35-38 

30 

25 

30 

35 

19 

19-26 

19-22 

32-38 

26 

29 

22/24 

U = feature is unknown. Numben separated by a dash indicates a range; numben separated by a slash indicated 


1 = Total number of seiigers (complete specimens only). 



5 - Length through setiger 10 in mm. 

35 = Branchiae fint present from setiger number. 

36 • Branchiae present through setiger number. 

80 = Subacicular hooks fint present from setiger number.

350 


TABLE 25.—Comparison of Eunice species in group B-l with antennal 

articulations. Stub (C) gives character numbers; column heads are species 

numbers. Characters and character states are listed in Appendix A. Species are 

in order by number, corresponding species names are listed at the end of the 

table. 

c 

7 

8 

9 

11 

12 

14 

15 

16 

17 

18 

20 

22 

23 

24 

25 

26 

27 

28 

29 

31 

32 

33 

34 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

57 

58 

59 

60 

61 

64 

65 

66 

74 

75 

76 

78 

79 

82 

41 

3 

1 

3 

2 

2 

U 

3 

1 

1 

3 

3 

1 

2 

1 

3 

5 

1 

2 

1 

1 

1 

2 

1 

2 

0 

U 

10 

30 

1 

3 

2 

2 

4-5 

1 

1 

4 

1 

4 

2 

- 

3 

2 

2 

2 

U 

1 

1 

U 

2 

1 

54 

3 

2 

2 

2 

2 

2 

1 

1 

4 

3 

1 

2 

2 

2 

3 

5 

2 

2 

1 

2 

3 

2 

- 

2 

5 

10-15 

3 

10 

1-2 

2 

2 

2 

10 

3 

1 

2 

2 

- 

- 

2 

1 

2 

2 

1 

1 

1 

1 

1 

2 

1 

58 78 134 143 168 


2 

2 

2 

2 

2 

2 

1 

1 

U 

1 

3 

1 

U 

2 

4 

3 

2 

1 

1 

1 

1 

2 

1 

3 

U 

U 

U 

U 

3 

U 

U 

U 

3 

3 

1 

1 

U 

5 

- 

3 

2 

1 

3 

3 

- 

- 

3 

3 

3 

2 

1 

4 

1 

2 

1 

3 

1 

2 

1 

1 

3 

5 

2 

1 

1 

3 

3 

- 

_ 

3 

2 

1 

1 

U 

U 

1 

1 

1 

3 

1 

- 

2 

U 

4 

3 

1 

1 

2 

2 

1 

U 

_ 

2 2 2 u 

1 - - 5-9 

1 

10 

10 

2 

- 

- 

- 

U 

_ 

1 

- 

3 

U 

3 

U 

2 

3 u 2 2 

2 uuu1 2 1-2 

2 

2 1-2 

4 

9 U 

1 

1 U 

1 u 1 1 

2 u2 3 U 

1 

2 2 

4 _ _ _ 

2 

- 

1 

2 

_ 

1 

4 

2 

_ 

1 

4 

2 

_ 

U 

1 

3 


U 2 2 

1 1 2 

U U U 

1 1 1 

1 1 1 

u u U 

2 2 2 

U 1 U 

2 

1 

2 

1 

1/3 

2 

2 

5 

2 

2 

3 

2 

2 

3 

1 

1 

1 

3 

2 

2 

1 

2 

1 

3 

1 

3 

2 

3 

3 

2 

_ 

2 

- 

_ 

- 

- 

1-2 

1-2 

1 

1 

5 

3 

1 

2 

1 

2 

3 

_ 

1 

3 

2 

1 

1 

1 

1 

1 

2/4 

1 

175 

2 

2 

3 

2 

2 

U 

1 

1 

1 

3 

2 

1 

4 

2 

4 

3 

1 

3 

2 

1 

1 

2 

1 

2 

2 

10 

6 

15 

2 

3 

2 

3 

5 

1 

2 

2 

2 

_ 

_ 

1 

1 

3 

1 

2 

2 

1 

1 

1 

2 

1 

183 

3 

1 

3 

2 

2 

2 

1 

2 

1 

3 

3 

2 

2 

2 

3 

3 

1 

1 

1 

2 

1 

2 

_ 

2 

8 

8 

2 

12 

1-2 

2 

2 

2 

3 

1 

1 

2 

2 

_ 

_ 


separated by a dash indicate a range; numbers separated by a slash indicate 

paired alternative observations. 

Character states of characters not included in the comparisons: 10,1; 13,1- 

19,2; 21,1; 30,1; 37,2; 53,1; 56,-; 69,1; 70,2; 71,2; 72,2; 73,2; 77,2; 81,1. 

Species included: 41 E. coccinea; 54 E. dubitata; 58 E. elegans; 78 E. 

gravieri; 134 E. nicidioformis; 143 E. palauensis; 168 E. rosaurae 175 E. 

1 

1 

1 

2 

2 

U 

1 

1 

u 

2 

1 

188 

2 

2 

2 

2 

1 

3 

1 

2 

1 

3 

1 

2 

2 

2 

1 

4 

1 

2 

1 

2 

3 

2 

_ 

1 

U 

U 

3 

15 

2 

2 

2 

2 

4 

1 

2 

2 

2 

_ 

_ 

1 

4 

1 

1 

2 

U 

2 

1 

U 

2 

2 

TABLE 26.—Comparison of Eunice species in group B-l without antennal 

articulations. Stub (C) gives character numbers; column heads ire species 

numbers. Characters and character stales are listed in Appendix A. Species are 

in order by number, corresponding species names are listed at the end of the 

table. 

c 

7 

8 

9 

11 

12 

14 

15 

18 

20 

23 

25 

27 

28 

29 

31 

32 

34 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

60 

61 

64 

65 

66 

70 

73 

82 

7 

3 

u 

U 

u uuu3 

3 

1 

U 

1 

u 

2 

3 

1 

U 

U 

u 

uuuuuuuuuuuuu 

u 

u1 

2 

u 

23 42 99 120 


3 

2 

3 

1 

1 

3 

1 

1 

1 

2 

1 

1 

1 

2 

1 

2 

I 

3 

10 

4 

25 

2-3 

2 

2 

2 

4 

3 

1 

3 

2 

4 

3 

3 

1 

3 

1 

1 

5 

1 

2 

3 

2 

3 

1 

3 

2 

1 

1 

1 

U 

U 

9 

U 

2 

3 

2 

3 

4 

1 

2 

2 

1 

1 

3 

2 

2 

2 

U 

U 

u 3 

u 

i; 

3 

3 

2 

1 

I 

1 

1 

i: 

u 

H 

15 

1 

2 

2 

2 

4 

3 

1 

2 

3 

4 

3 


2 

2 

2 

2 

2 

2 

2 

2 

2 

1 

2 

2 

2 

2 

1 

\ 

1 

2 

2 

2 

2 

1 

1 

2 

A 

3 

1 

2 

1 

1 

1 

0 

7 

47 

15 

1 

3 

2 

2 

3 

1 

1 

2 

1 

3 

2 

1 

2 

1 

1 

1 

131 

U 

U 

2 

2 

2 

2 

3 

3 

3 

4 

3 

1 

5 

2 

2 

3 

- 

u 

U 

2 

u 

1-2 

U 

I) 

u 

7 

1 

2 

2 

1 

2 

3 

1 

1 

2 

2 

1 

143 

3 

2 

1 

1 

U 

u 

1 

3 

1 

2 

4 

1 

1 

2 

2 

1 

- 

5-y 

U 

u 

2 

2 

1-2 

1-2 

U 

U 

1 

u 

u 

1 

3 


separated by a dash indicate a range. 


16.1; 17,1; 19.2; 21,2; 22,-; 24,2; 26,3; 30,1; 33,2; 37.2; 38.2; 53.1; 56.-; 57,2; 

58,-; 59.-; 69,1; 71,2; 72,2; 74,1; 75,1; 76,1; 77,2; 78,1; 79,2; 81.1. 

Species included: 7 E. amoureuxi; 23 E. barvicensis; 42 E. coccinoides; 99 

E. langi; 120 E. megabranchia; 131 E. mutilatoides; 143 E. palauensis. 

2 

2 

2 

2 

U


TABLE 27.—Comparison of Eunice species in group B-2. The column numbers are the same as the character 

numbers used to prepare the key (see explanation below; for a complete listing of all characters and character 

states see Appendix A). 

Species 

1 

2 

3 

5 

35 

36 

80 

8 

10 

14 

17 

28 

29 

30 

31 

33 

41 

48 

53 

57 

64 

68 

69 

70 

71 

72 

79 

81 

82 

83 

92 

97 

100 

101 

107 

108 

110 

111 

122 

123 

127 

130 

133 

135 

136 

137 

141 

143 

149 

158 

159 

160 

162 

163 

164 

165 

166 

167 

169 

170 

174 

208 

178 

185 

189 

/: amphiheliae 

E. annulicornis 

E. aphroditois 

E. argentinensis 

E. biformicirrata 

E. bilobata 

E. bipapillata 

E. borneensis 

E. bowerbanki 

E. coccinea 

E. contingens 

E. djiboutiensis 

E. eimeorum 

E. funbriata 

E. flavopicta 

E floridana 

E.franklini 

E. frauenfeldi 

E. fucata 

E. grubei 

E. guildingi 

E. guttala 

E. harassii 

E. investigator is 

E. kinbergi 

E. laliceps 

E. laurillardi 

E. longicirris 

E. longisetis 

E. macrobranchia 

E. magellanica 

E. microprion 

E. mindanavensis 

E. mullipectinata 

E. mutilata 

E. nesiotes 

E. nigricans 

E. northioidea 

E. norvegica 

E. ovalifera 

E. palauensis 

E. pauroneurala 

E. philocorallia 

E. plicata 

E. poly bronchia 

E. prognatha 

E. pulvinopalpata 

E. purpurea 

E. pycnobranchiata 

E. quinquefuia 

E. reducta 

E. roussaei 

E. rubella 

E. rullieri 

E. scombrinis 

E. sebastiani 

E. suviensis 

E. tibiana 

40-150 

145 

U 

u 

137 

U 

u 

120 

142 

U 

u uu 

362 

166 

111 

U 

U 

250 

U 

258 

115 

U 

U 

380 

U 

189 

U 

108 

150 

130 

127 

63 

196 

U 

U 

180 

U 

157 

U 

U 

U 

136 

126 

U 

122 

U 

106 

U 

u 

298 

U 

u 

72 

130 

706 

U 

165 

180-200 

135 

U 

u 

217 

U 

U 

42 

95 

U 

U 

u 

230 

215 

80 

U 

U 

195 

U 

138 

52 

U 

110 

398 

U 

150 

U 

40 

200 

140 

82 

40 

172 

U 

u 

100 

u 

200 

u 

u 

u 

115 

80 

U 

80 

U 

53 

U 

U 

318 

U 

U 

24 

47 

1455 

370 

73 

7-10 

4 

22 

5 

7 

6 

U 

6 

7 

5 

U 

10 

3.5 

7 

6 

6 

7 

5 

5 

6 

6 

3 

2.5 

7 

15 

10 

5 

5 

3.5 

U 

7 

5 

4 

7 

3.5 

6 

7 

2.5 

9 

5 

U 

5.5 

6 

4 

3 

7.5 

11 

4 

8 

2.5 

6 

U 

2 

3 

4 

23 

9 

3 

U 

12.5 

55 

12 

17 

7 

U 

7 

11 

10 

U 

20 

7.5 

11 

U 

10.5 

15 

7 

10 

12 

7 

8 

5 

U 

16 

21 

9 

16 

6.4 

U 

17 

10 

7 

16 

10 

10 

10 

7 

13 

U 

U 

6 

6 

8 

8 

16 

23 

9.5 

12 

8 

15 

U 

8 

5 

7 

35 

U 

5 

4 

3 

6 

3 

3 

7 

4 

6 

5 

6 

6 

5 

5 

9 

5 

8-10 

6 

6 

5 

3 

5 

6 

4 

6 

8-9 

4 

3 

3 

6 

7-8 

7-8 

7-8 

6 

7-8 

7 

6 

6-7 

3 

7 

5 

6-8 

8 

6 

7 

7-8 

10/13 

3 

5 

5-6 

7 

4 

6-8 

5 

5 

7 

7-14 

9 

6 

140-150 

134 

U 

u 

132 

U 

u 

120 

136 

U 

U 

u u 

122 

166 

111 

U 

u 

170-180 

U 

236 

105 

U 

U 

365 

U 

184 

U 

104 

U 

120 

127 

63 

1% 

U 

U 

180 

U 

155 

U 

U 

u 

135 

115 

U 

110 

U 

100 

U 

U 

131 

U 

u 

70 

125 

686 

160 

135 

U 

35 

200 

19-22 

27 

25 

U 

30-35 

31 

38-39 

30 

81 

38 

26 

U 

29-40 

34 

27 

U 

24 

24 

32 

28 

44-45 

123 

38 

32-33 

30 

18 

U 

26/30 

38/41 

21 

37-45 

27 

31 

30 

25 

42 

35 

19-22 

27 

34 

25 

24-36 

35 

62-72 

19 

28/29 

30 

45 

U 

28 

19 

29 

56/56 

U 

26



190 

192 

196 

200 

Species 

E. torquata 

E. tribranchiala 

E. unidenlata 

E. violaceomaculata 

1 

U 

U 

U 

215 

2 

U 

U 

80-100 

170 

3 

U 

7 

2-3 

7 

5 

U 

13 

U 

12 

35 

3 

4-5 

23 

6-7 

36 

U 

U 

U 

215 

80 

32 

30 

20-25 

25-39 


paired alternative observations. Explanation of heads: 









TABLE 28.—Comparison of the Eunice species from group B-2 without antennal articulations. Stub (Q gives 

character number, column heads give species number. Characters and character states are listed in Appendix A. 

Species are in order by number; corresponding species names are listed at the end of the table. 

c 

14 

48 

53 

72 

82 84 143 160 169 

174 

178 

185 

196 

208 

7 

8 

9 

11 

12 

14 

15 

16 

18 

19 

20 

23 

24 

25 

26 

27 

28 

31 

32 

33 

34 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

57 

58 

59 

60 

61 

3 

1 

2 

2 

1 

U 

1 

1 

3 

2 

3 

4 

2 

3 

4 

1 

4 

2 

0 

U 

30 

30 

1-2 

2 

2 

1 

250 

2 

3 

2 

1 

2 

2 

_ 

1 

u 

u 

u 

u 

u 

u 

3 

U 

u 

u2 

U 

u 2 

U 

1 

3 

2 

1 

1 

1 

1 

U 

18-19 

15 

1-2 

2 

2 

2 

U 

2 

U 

2 

2 

_ 

_ 

2 

1 

3 

2 

3 

2 

2 

1 

3 

1 

3 

2 

3 

4 

2 

3 

8 

1 

4 

2 

0 

U 

25 

U 

1-2 

3 

2 

2 

5 

1 

1 

3 

2 

_ 

_ 

4 

1 

U 

u 

u 

u 

u 

u 

u 

u 

u 

u 

u 

1 

u 

u 

3 

2 

1 

1 

1 

2 

1 

2 

1 

U 

30 

12 

U 

1 

3 

2 

2 

U 

U 

U 

U 

2 

_ 

_ 

2 

1 


U 3 

U 2 

U 1 

U 1 

U U 

U U 

U 1 

u 1 

3 3 

2 2 

1 1 

2 2 

U U 

U 4 

u 3 

u 1 

u 1 

2 

1 

U 

- 

2 

1 

3 

2 

2 

U 

1 

1 

3 

2 

1 

2 

2 

1 

4 

1 

1 

1 

1 

2 

1 

1 

1 

U 

u 

10 

u 

3 

2 

2 

3 

19 

1 

1 

1 

1 

1 

1 

- 

1 

1 

u 

u 

18-20 

14 

u 

u 

u 

u 

u 

1 

u 

2 

2 

_ 

_ 

2 

4 

2 

U 

5-9 

U 

3 

U 

2 

2 

1-2 

1-2 

U 

U 

1 

U 

2 

_ 

- 

U 

1 

3 

3 

2 

1 

2 

3 

1 

2 

3 

2 

1 

2 

1 

3 

4 

2 

2 

1 

1 

2 

1 

1 

1 

1 

U 

5 

15 

3 

2 

2 

2 

4 

1 

1 

1 

2 

- 

- 

1 

4 

U 

U 

u 

u 

uuuuuuu1 

u 

uu 

1 

u 

1 

1 

u 

1 

1 

1 

u 

u 

47 

u 5 

2 

2 

3 

U 

u 

2 

1 

U 

u 

u 

u 

1 

3 

1 

3 

2 

2 

4 

1 

1 

3 

2 

3 

1 

1 

1 

5 

1 

5 

1 

1 

2 

2 

1 

1 

0 

4-5 

5 

10 

2 

3 

2 

3 

4 

1 

1 

2 

2 

- 

- 

4 

2 

2 

1 

2 

2 

2 

U 

1 

3 

2 

2 

1 

4 

1 

3 

5 

1 

4 

1 

1 

2 

1 

1 

1 

0 

3 

40 

35 

15 

2 

2 

1 

6-13 

1 

2 

4 

1 

2 

4 

- 

1 

U 

U 

U 

U 

U 

6 

1 

U 

3 

2 

U 

4 

U 

u 

u 

1 

u 

1 

1 

2 

1 

1 

1 

u 

u9 

U 

U 

U 

U 

U 

U 

U 

U 

u 

2 

- 

- 

65 

66 

70 

74 

79 

81 

82 

1 

1 

2 

1 

2 

3 

1 

U 

U 

2 

U 

2 

U 

u 

2 

2 

2 

1 

2 

3 

1 

U 

U 

2 

1 

1 

U 

U 


2 U 2 1 

1 u 2 2 

1 2 2 2 

U U U 1 

2 

1 

2 

2 

1 

1 

2 

U 

U 

2 

1 

1 

2 

1 

2 

1 

2 

U 

3 

2 

1 

2 

1 

2 

1 

3 

1 

1 

2 

2 

1 

1 

5 

u 

3 

U 

2 

2 

U 

2 

U 

U 

3 

U 

2 

1 

U 

u 

1 

1 

1 

1 

1 

4 

U 

u 

u 

1 

1 

1 

1 

2 

1 

2 

2 

U 

U 

5 

35 

1 

2 

3 

U 

U 

U 

1 

2 

U 

U 

U 

U 

U 

3 

1 

3 

1 

1 

U 

1 

1 

2 

2 

1 

2 

1 

3 

9 

1 

1 

1 

1 

2 

2 

1 

1 

5 

5 

17 

U 

1 

3 

2 

3 

5 

1 

2 

1 

1 

2 

1 

- 

1 

U = feature is unknown; dash (-) = character inappropriate. Numbers separated by a dash indicate a range. 

Character states of characters not include in the comparisons: 10,1; 12,1; 13,1; 17,1; 21,2; 22,- 29,2; 30,1; 

53.1; 56,-; 64.3; 69,1; 71A 72.2; 73,2; 75.1; 76,1; 77,2; 78,1. 

Species included: 14 E. aphroditois; 48 E. contingens; 53 E. djiboutiensis; 72 E.fucata; 82 E. guttata; 84 E. 

investigatoris; 143 E. palauensis; 160 E. polybranchia; 169 E. roussaei; 174 E. rullieri; 178 E. sebastiani; 185 

E. suviensis; 196 E. unidentata; 208 E. scombrinis. 

2 

2 

2 

1 

1 

U 

U 

- 

- 

2 

1 

2 

1 

5

354 


TABLE 29.—Comparison of the Eunice species from Group B-2 with cylindrical


TABLE 30.—Comparison of Eunice species from group B-2 with cylindrical antennal articulations and notopodial 

articulations limited to anterior setigers. Stub (C) gives character number, column heads give species number. 

Characters and character states are listed in Appendix A. Species are in order by number, corresponding species 

names are listed at the end of the table. 

c 

41 

64 

68 100 108 110 123 

137 

164 

167 

200 

7 

8 

9 

12 

14 

15 

17 

20 

23 

24 

25 

26 

27 

28 

29 

32 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

54 

55 

57 

58 

59 

60 

61 

3 

1 

3 

2 

U 

3 

1 

3 

2 

1 

3 

5 

1 

2 

1 

1 

1 

6 

U 

2 

2 

0 

U 

10 

30 

1 

3 

2 

2 

4-5 

1 

4 

1 

4 

2 

_ 

3 

2 

3 

2 

2 

U 

3 

1 

3 

4 

1 

4 

4 

1 

2 

1 

1 

2 

9 

122 

2 

2 

0 

10-12 

5 

15 

2 

3 

2 

3 

4-5 

2 

1 

2 

- 

_ 

3 

4 


U 

u 

u 

u 

1 

u 

uu2 

u uu1 

u 

2 

1 

1 

5 

166 

1 

1 

0 

0 

18-19 

14 

U 

u uu 

u 

u 

u 

u 

u 

u 

u 

u 

3 

1 

3 

1 

U 

3 

1 

1 

1 

2 

3 

3 

1 

2 

1 

1 

1 

4 

U 

1 

1 

0 

u 

18 

20 

1 

3 

2 

2 

3 

1 

2 

2 

- 

_ 

1 

4 

3 

2 

2 

2 

3 

1 

4 

3 

U 

2 

3 

5 

1 

2 

1 

1 

1 

6 

104 

1 

1 

1 

15 

8 

U 

3 

3 

2 

3 

9 

2 

3 

2 

- 

- 

2 

4 

u 

u 

u 

u 

u 

uuuuuuu1 

u 

u2 

1 

7-8 

U 

1 

1 

5 

U 

15-17 

U 

U 

u 

u 

u 

u 

u 

u 

u 

u 

u 

u 

u 

3 

2 

3 

1 

U 

U 

1 

1 

2 

2 

3 

6 

2 

1 

1 

3 

1 

6 

63 

1 

1 

0 

5 

3 

15 

1-2 

3 

2 

1 

5 

3 

2 

1 

4 

2 

_ 

2 

1 

3 

3 

2 

4 

1 

1 

3 

2 

2 

3 

4 

2 

1 

1 

3 

1 

7 

155 

1 

1 

1 

3 

7 

U 

1-2 

2 

2 

2 

6 

1 

3 

2 

- 

- 

2 

3 

3 

1 

3 

2 

3 

1 

1 

1 

1 

1 

3 

4 

1 

2 

1 

1 

1 

5 

100 

1 

1 

2 

0 

14 

20 

2 

2 

1 

1 

4 

2 

1 

2 

- 

- 

1 

1 

2 

3 

2 

2 

2 

1 

1 

3 

1 

1 

3 

4 

1 

1 

2 

1 

1 

4 

131 

2 

2 

0 

6 

21 

U 

1 

2 

2 

2 

4 

1 

4 

2 

- 

- 

2 

2 

2 

2 

3 

2 

3 

1 

1 

1 

1 

1 

3 

5 

1-2 

1 

1 

1 

1 

6-7 

215 

1 

1 

U 

U 

20 

15 

1 

3 

2 

1 

4 

2 

2 

2 

- 

- 

4 

3 

66 

78 

82 

2 

U 

1 

2 

2 

1 

1 

U 

U 


1 1 1 

U 1 1 

2 1 


Character states of characters not included in the comparison: 10,1; 11,2; 13,1; 16,1; 18,3; 19,2; 21,1; 22,1; 

30,1; 31.1; 33,2; 52,1; 53,1; 56,-; 64,2; 65.2; 69.1; 70,2; 71A 72.2; 73,2; 74,1; 75,1; 76,1; 77,2; 79,2; 81,1. 

Species included: 41 E. coccinea; 64 E. fimbriata; 68 E. flavopicta; 100E. laticeps; 108 E. longisetis; WOE. 

macrobranchia; 123 E. mindanavensis; 137 E. norvegica; 164 E. purpurea; 167 E. reducla; 200 E. 

violaceomaculata. 

u 

2 

1 

1 

1 

U 

2 

2 

1 

1 

2 

U 

1 

1 

U 

1


c 

7 

8 

9 

12 

13 

14 

15 

16 

17 

18 

20 

23 

24 

25 

26 

27 

28 

29 

31 

32 

34 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

57 

58 

59 

60 

61 

65 

66 

70 

74 

75 

78 

81 

82 

TABLE 31.—Comparison of Eunice species in group B-2 with cylindrical antennal articulations, but without 

notopodial articulations. Stub (C) gives character number, column heads give species number. Characters and 

character sutes are listed in Appendix A. Species are in order by number, corresponding species names are listed 

at the end of the table. 

8 30 

1 u 

2 uuuuuuuuu 

3 

U 

2 

_ 

3 

U 

1 

3 

1 3 

4 4 

2 U 

4 U 

5 U 

u 

u 

u 

1 

3 

U 

1 

1 

u 

4 

8 

1 

3 

2 

2 

U 

U 

1 

3 

2 

- 

- 

2 

1 

U 

1 

u 

u 

u 

uu 

U 

U 

7 

U 

2-3 

2 

2 

2 

U 

1 

U 

U 

u 

u 

u3 

U 

2 

2 

2 

U 

1 

U 

u 

u 

31 

1 

2 

3 

1 

U 

U 

1 

1 

1 

3 

1 

1 

2 

3 

6 

1 

1 

1 

1 

1 

2 

1 

1 

0 

10 

8 

15 

1 

3 

2 

3 

4 

1 

2 

1 

1 

2 

1 

- 

1 

2 

2 

2 

U 

1 

U 

1 

4 

57 

3 

2 

3 

1 

1 

2 

1 

1 

1 

3 

1 

4 

2 

3 

5 

1 

3 

2 

U 

U 

5 

20 

1-3 

2 

2 

2 

U 

1 

1 

3 

1 

U 

u 

-2 

2 

1 

2 

U 

1 

U 

u 

u 

70 81 83 110 130 


2 

u 

2 

3 

3 

1 

3 

1 

U 

U 

3 

1 

1 

3 

3 

2 

2 

3 

5 

2 

3 

1 

0 

U 

25 

U 

3 

2 

3 

3 

U 

1 

1 

2 

U 

1 

3 

1 

U 

u 

1 

1 

1 

3 

3 

2 

1 

4 

3 

2 

2 

2 

1 

1 

1 

2 

1 

0 

20 

11 

15 

2 

3 

2 

1 

4 

1 

1 

2 

1 

1 

2 

- 

3 

u u 

u 

4 

3 

1 

3 

1 

2 

2 

4 

4 

1 

1 

2 

1 

1 

1 

1 

1 

3 

U 

10 

15 

1/3 

2 

2 

2 

5 

1 

1 

2 

2 

_ 

- 

4 

1 

u 

uuuuuuuuuuuuu1 

u 

u1 

2 

1 

1 

1 

5 

u 

15-17 

U 

U 

U 

U 

u 

1 

3 

2 

1 

3 

3 

1 

1 

3 

1 

2 

2 

4 

3 

1 

2 

2 

1 

1 

1 

1 

1 

3 

U 

4 

25 

3 

2 

2 

3 

u 

uuuuuuuu 

3 

9 

1 

2 

3 

2 

_ 

- 

2 

4 


2 2 2 u 1 

2 2 1 1 1 

1 2 2 2 2 

U U 2 u U 

1 1 1 u1 1 

u U 2 

U 

1 1 1 u 1 

1 1 1 u 1 

133 

2 

2 

2 

1 

U 

u 

1 

2 

1 

3 

1 

2 

2 

3 

4 

2 

2 

U 

14 

U 

1-2 

3 

2 

2 

5 

1 

2 

3 

U 

U 

U 

U 

1 

2 

1 

2 

U 

1 

U 

1 

1 

141 

U 

U 

u 

u uuuu4 

U 

u 

u uuU 

L 1 

1 

u 

uuuuu 

5-6 

U 

U 

u 

u 

u 

u 

uu 

1 

uu_5 

u 

1 

2 

U 

u 

2 

U 

u 

149 

3 

1 

3 

1 

U 

u 1 

1 

6 

3 

1 

u 2 

3 

3 

1 

2 

2 

1 

3 

2 

1 

1 

1 

U 

5 

15 

1-2 

2 

2 

2 

4 

1 

1 

3 

2 

_ 

_ 

2 

1 

2 

1 

2 

U 

1 

U 

1 

1 

162 

3 

1 

2 

1 

U 

U 

1 

2 

1 

3 

3 

1 

2 

3 

5 

U 

U 

u 2 

3 

- 

1 

2 

U 

U 

3 

U 

2-3 

3 

2 

2 

9 

1 

1 

2 

2 

_ 

_ 



Character sutes of characters not included in the comparison: 10,1; 11,2; 19,2; 21,1; 22,1; 30,1; 33 2; 53 1- 

56,-; 76,3; 69,1; 71,2; 72,2; 73,2; 76,1; 77,2; 79,2. 

Species listed: 8 E. amphiheliae; 30 E. bipapillata; 31 E. borneensis; 57 E. eimeorum; 70 E.franklini; 81 E. 

guildingi; 83 E. harassii; 110 E. macrobranchia; 130 E. mutilata; 133 E. nesiotes; 141 E. ovalifera; 149 E. 

pauroneurata; 162 E. prognatha; 163 E. pulvinopalpata; 189 E. tibiana. 

2 

1 

2 

2 

2 

U 

2 

U 

1 

1 

163 

3 

3 

3 

2 

V 

U 

1 

1 

1 

3 

1 

1 

1 

3 

3 

2 

1 

1 

1 

3 

2 

I 

1 

1 

0 

25 

15 

2 

3 

2 

2 

5 

3 

1 

2 

2 

_ 

_ 

1 

3 

2 

1 

2 

2 

1 

2 

1 

1 

189 

2 

2 

2 

1 

1 

2 

5 

1 

1 

3 

1 

2 

2 

3 

•» 

3 

1 

1 

1 

1 

1 

1 

1 

1 

10-15 

5 

U 

2-3 

2-3 

2 

3 

4 

1/3 

2 

2 

2 

_ 

_ 

1 

1 

1 

2 

2 

U 

1 

u 

1 

5


TABLE 32.—Comparison of the Eunice species from group B-2 with disially drop-shaped or moniliform 

articulations of the ceratostyles. Stub (C) gives character number, column heads give species number. Characters 

and character states are listed in Appendix A. Species are in order by number, corresponding species names are 

listed at the end of the table. 

c 

28 

33 

69 71 79 101 107 127 

136 

165 

166 

170 

7 

8 

9 

12 

13 

14 

15 

16 

17 

18 

20 

23 

24 

25 

26 

27 

28 

29 

31 

32 

33 

34 

39 

40 

41 

42 

43 

44 

48 

51 

52 

53 

54 

55 

56 

60 

61 

64 

3 

1 

3 

1 

U 

U 

3 

2 

3 

3 

1 

1 

1 

1 

1 

2 

1 

1 

5 

7 

10 

1-2 

3 

2 

3 

3 

1 

1 

2 

_ 

2 

3 

1 

3 

1 

3 

1 

U 

U 

3 

1 

1 

3 

1 

1 

1 

3 

6 

1 

2 

1 

1 

1 

2 

2 

0 

60 

15 

12 

3 

2 

3 

- 

3 

2 

_ 

_ 

2 

2 

1 

2 

2 

2 

2 

1 

1 

3 

1 

2 

1 

3 

3 

2 

1 

3 

4 

2 

1 

1 

1 

1 

1 

1 

1 

0 

8 

U 

2-3 

3 

3 

7-9 

1 

1 

2 

2 

- 

4 

4 

2 


2 3 3 3 

3 1 2 1 

2 2 3 2 

2 2 1 2 

U U 1 1 

U U 3 2 

1 

3 1 

1 

1 3 

1 

1 3 

3 :J 3 2 

1 

1 1 

2 

2 U 

1 

2 2 

3 3 3 3 

3 5 5 3 

1 1 2 3 

1 2 1 1 

1 1 1 1 

2 1 1 1 

1 1 3 3 

2 2 2 2 

- 1 1 1 

1 1 1 

U U 0 

3 8 10 

15 20 15 

U 

u7 

15 

2 1 2 3 

2 2 3 3 

2 1 2 2 

4 5 9 4 

1 2 1 1 

1 1 1 1 

2 1 1 1 

2 2 2 2 

- - - - 

1 4 4 1 

3 1 3 4 

2 2 1 1 

3 

3 

2 

1 

1 

3 

U 

1 

1 

3 

1 

U 

1 

3 

5 

1 

2 

1 

1 

1 

2 

1 

8-10 

0 

12 

U 

2 

3 

2 

4 

3 

1 

2 

2 

- 

2 

3 

2 

3 

1 

3 

1 

1 

4 

1 

2 

1 

3 

1 

2 

1 

3 

3 

1 

3 

2 

1 

1 

2 

1 

3 

U 

4 

10 

3 

2 

2 

4 

1 

1 

1 

3 

- 

1 

4 

2 

3 

1 

3 

1 

U 

U 

1 

3 

2 

3 

1 

1 

1 

3 

5 

1 

2 

1 

1 

2 

2 

U 

U 

U 

4 

U 

2 

3 

3 

U 

3 

1 

1 

3 

- 

U 

4 

2 

3 

1 

3 

1 

1 

3 

1 

2 

1 

3 

1 

U 

1 

3 

5 

1 

1 

1 

1 

1 

2 

1 

0 

u 5 

15 

3 

2 

2 

5-6 

1 

1 

1 

2 

- 

1 

3 

3 

1 

3 

2 

2 

1 

3 

1 

1 

1 

3 

3 

4 

2 

1 

3 

1 

1 

1 

2 

1 

2 

- 

2 

U 

2 

7 

1 

2 

2 

7 

1 

1 

1 

2 

- 

4 

2 

3 

65 

66 

74 

75 

76 

77 

78 

82 

2 

2 

U 

1 

1 

2 

U 

1 

2 

1 

U 

1 

1 

2 

U 

2 

1 

2 

U 

12 

1 

2 

U 

2 


2 

2 

1 

1 

1 

2 

1 

2 

1 

1 

U 

1 

1 

2 

U 

2 

2 

1 

U 

1 

1 

2 

U 

1 

2 

1 

2 

1 

1 

2 

1 

1 

1 

1 

1 

1 

1 

2 

1 

1 



45,2; 57,2; 58,-; 59,-; 69,1; 70,2; 71,2; 72,2; 73,2; 79,2; 81,1. 

Species included: 28 E. biformicirrata; 33 E. bowerbanki; 69 E.floridana; 71 E.frauenfeldi; 79 E. grubei; 101 

E. laurillardi; 107 E. longicirris; 127 E. multipectinata; 136 E. northioidea; 165 E. pycnobranchiata; 166 E. 

quinquefida; 170 E. rubella. 

2 

1 

1 

1 

1 

2 

1 

1 

2 

2 

U 

1 

1 

2 

1 

1 

2 

1 

1 

1 

1 

2 

1 

3 

2 

1 

U 

1 

1 

2 

U 

1

358 


TABLE 33.—Comparison of Eunice species in groups B-3 and B-4. Column heads are the same as the character 



1 

4 

34 

35 

37 

39 

40 

43 

44 

45 

47 

49 

50 

51 

52 

56 

61 

62 

63 

66 

73 

74 

75 

80 

88 

93 

103 

104 

106 

112 

115 

124 

129 

138 

142 

147 

148 

154 

155 

157 

162 

9 

181 

182 

187 

203 

Species 

E. aciculala 

E. afra 

E. brevis 

E. bucciensis 

E. cariboea 

E. cincta 

E. cirrobranchiata 

E. collaris 

E. collini 

E. complanata 

E. conglomerans 

E. crassitentaculata 

E. curticirris 

E. denticulata 

E. dilatata 

E. ehiersi 

E. excariboea 

E. fauveli 

E. fdamentosa 

E. flavapunctata 

E. fuscafasciata 

E. gagzoi 

E. goodei 

E. guanica 

E. imogena 

E. jagori 

E. leucosticta 

E. levibranchia 

E.Vila 

E. magnified 

E. maremelleri 

E. modesla 

E. mutabilis 

E. notala 

E. pacifica 

E. parvibranchis 

E. paupera 

E. perimensis 

E. perrieri 

E. peter si 

E. prog nut ha 

E. pruvoti 

E. sonorae 

E. spongicola 

E. lenuis 

E. wasinensis 

1 

202 

U 

130 

U 

127 

U 

U 

U 

u 

100 

166 

U 

55 

U 

U 

U 

u 

216 

119 

U 

U 

160 

U 

224 

U 

U 

131 

105 

240 

U 

u 

80 

196 

95 

102 

U 

127 

U 

185 

302 

122 

115 

586 

152 

695 

84 

2 

129 

U 

18 

U 

24 

U 

U 

u 

u 

72 

163 

U 

12 

U 

u 

302 

25 

U 

u 

32.5 

u 

100 

u 

u 

130 

94 

78 

U 

u 

28 

95 

40 

33 

U 

U 

U 

135 

130 

80 

34 

345 

68 

460 

17 

3 

4.5 

6 

1.5 

3.5 

1.2 

1.1 

3 

6 

4 

U 

3.5 

3.5 

1 

3 

10 

4 

2 

S 

1.5 

2 

1.3 

1.5 

0.75 

4 

2 

6 

U 

8 

4 

U 

5 

1 

6 

1 

3 

4 

U 

5 

5 

5 

7.5 

3 

6 

1.75 

2 

2.5 

5 

11 

10 

2.5 

7.5 

2.5 

3 

6 

12 

7 

U 

6.5 

9.5 

2.25 

7 

16 

6 

3.7 

18 

3 

7 

3 

2.75 

4 

7 

3.5 

6 

U 

9 

4 

U 

11 

3 

8 

U 

6 

10 

U 

10 

12 

7 

16 

5 

14 

6 

4.5 

6 

35 

18 

16 

- 

18 

- 

- 

22 

17-18 

16 

19 

23-29 

28 

- 

27 

19 

14 

78-79 

18 

23-27 

13 

- 

85-120 

- 

19 

- 

17 

14-15 

69 

15 

22 

28 

17-18 

17-18 

14 

17-21 

19-20 

23 

17 

18 

13 

10/13 

15 

41 

24 

81 

- 

36 

190 

U 

- 

U 

- 

- 

U 

U 

85 

U 

166 

U 

- 

u uu 

216 

119 

U 

- 

160 

- 

224 

- 

U 

113 

85-95 

120 

U 

U 

70 

196 

70 

102 

U 

U 

U 

170 

200 

110 

115 

530 

150 

695 

- 

80 

22 

22 

27 

34 

24-31 

21 

17/24 

27/33 

26-27 

28 

22 

12 

16 

18-19 

28 

27 

78/87 

41 

17/22 

25 

23 

27-38 

23 

36 

50 

23 

U 

28 

23 

U 

38 

U 

22-23 

U 

23-28 

17-19 

23 

28 

25 

26 

35 

19 

54 

26 

124 

27-31 


numbers separated by a slash indicate paired alternative observations. Explanation of column heads: 









TABLE 34.—Comparison of Eunice species in groups B-3 and B-4 with articulated ceratostyles. Stub (C) gives 


Species are in order by number, corresponding species names are listed at the end of the table. 

c 

4 

40 

43 

47 56 61 62 80 93 

124 

142 

157 

162 

181 

7 

8 

9 

11 

12 

14 

15 

16 

17 

18 

20 

22 

23 

24 

25 

26 

27 

28 

29 

31 

32 

33 

34 

37 

38 

39 

40 

41 

42 

43 

44 

48 

51 

52 

53 

54 

55 

56 

57 

58 

59 

60 

61 

64 

3 

1 

3 

1 

2 

2 

1 

1 

3 

3 

3 

1 

1 

1 

3 

3 

1 

2 

2 

1 

1 

2 

1 

1 

1 

0 

U 

6 

30 

2 

2 

2 

5 

2 

1 

3 

2 

_ 

2 

_ 

_ 

1 

1 

3 

3 

3 

2 

2 

2 

2 

1 

1 

1 

3 

3 

1 

4 

1 

1 

9 

1 

1 

2 

2 

1 

2 

_ 

1 

1 

U 

u 

2 

U 

14 

2 

2 

4 

3 

1 

2 

1 

_ 

1 

2 

1 

_ 

2 

3 

3 

2 

3 

2 

2 

2 

U 

1 

1 

3 

3 

2 

2 

1 

2 

3 

1 

2 

2 

1 

1 

2 

1 

1 

1 

0 

U 

12 

50 

2 

2 

3 

3-4 

1 

1 

1 

2 

_ 

2 

_ 

_ 

1 

1 

3 

2 

2 

3 

1 

2 

2 

3 

1 

1 

3 

3 

1 

4 

1 

1 

4 

1 

1 

2 

2 

1 

2 

_ 

1 

1 

U 

u 

4 

130 

3 

2 

2 

4 

1 

1 

2 

2 

_ 

2 

_ 

_ 

2 

1 

3 


3 3 3 

2 2 1 

2 3 2 

1 2 2 

1 1 2 

2 5 U 

1 

2 

3 

1 

1 

1 

3 3 3 

3 

3 

1 

1 

1 2 4 

2 2 2 

3 3 3 

3 4 2 

1 1 1 

2 2 1 

2 2 2 

2 1 2 

2 1 1 

2 2 2 

_ 1 - 

1 1 1 

1 1 1 

U 1 

u 0 

3 14 

U 35 

10 

u2 

30 

24 2-3 2-3 

2 2 2 

1 3 3 

4 9 6-7 

1 1 3 

1 1 

2 1 

2 2 

_ - 

1 2 

2 - 

1 - 

_ 1 - 

3 1 3 

1 2 3 

2 

2 

3 

2 

1 

2 

1 

1 

1 

3 

3 

1 

1 

2 

4 

2-3 

1 

2 

2 

1 

1 

2 

2 

1 

1 

1 

U 

6 

U 

2 

2 

2 

4 

1 

2 

2 

1 

_ 

U 

u 

u1 

3 

3 

2 

3 

1 

1 

6 

1 

2 

1 

3 

1 

1 

2 

2 

3 

5 

2 

2 

;> 

U 

U 

8 

25 

2 

3 

3 

- 

1 

2 

- 

- 

1 

U 

U 

u 1 

1 

3 

3 

3 

2 

1 

3 

1 

1 

1 

1 

3 

1 

4 

1 

3 

3 

1 

5 

2 

3 

1 

- 

- 

2 

1 

- 

- 

1 

- 

U 

u 4 

U 

1 

1 

u 

-2 

- 

- 

1 

1 

3 

3 

3 

2 

2 

1 

2 

1 

3 

3 

1 

1 

1 

2 

2 

3 

4 

1 

3 

2 

1 

1 

2 

2 

1 

1 

U 

u 5 

40 

2 

2 

2 

9 

1 

1 

2 

3 

- 

2 

- 

- 

2 

2 

3 

3 

1 

3 

1 

2 

2 

3 

1 

1 

1 

3 

1 

4 

1 

4 

3 

1 

2 

1 

1 

1 

2 

1 

1 

1 

U 

130 

4 

U 

2 

2 

3 

4 

1 

1 

2 

1 

- 

2 

- 

- 

1 

1 

2 

3 

1 

2 

2 

1 

U 

1 

2 

1 

3 

3 

1 

1 

2 

3 

5 

U 

U 

U 

2 

3 

2 

- 

2 

1 

U 

U 

3 

U 

2-3 

3 

2 

9 

1 

1 

1 

2 

- 

2 

- 

- 

2 

1 

3 

2 

2 

3 

2 

2 

2 

1 

2 

1 

4 

2 

1 

1 

2 

1 

1 

2 

2 

2 

2 

0 

55 

6 

65 

1-2 

2 

3 

4 

1 

1 

2 

2 

- 

1 

2 

2 

- 

2 

3 

65 

66 

70 

75 

76 

79 

81 

1-2 

2 

2 

1 

1 

2 

1 

1 

2 

2 

1 

4 

4 

1 

1 

2 

2 

1 

1 

2 

1 

1 

2 

2 

1 

4-5 

2 

1 


1 

2 

2 

1 

1 

2 

U 

1 

2 

2 

1 

1 

2 

3 

1 

1 

1 

1 

1 

2 

1 

2 

2 

2 

1 

1 

2 

1 



73,2; 74,1; 77,2; 78.1. 82,1. 

Species listed: 4 E. afra; 40 E. cirrobranchiata; 43 E. collaris; 47 E. conglomerate; 56 E. ehlersi; 61 E. 

excariboea; 62 E.fauveli; 80 E. guanica; 93 E.jagori; 124 E. modesta; 142 E. pacifica; 157 E. petersi; 162 E. 

prognatha; 181 E. sonorae. 

1 

2 

2 

1 

1 

2 

1 

2 

2 

2 

1 

1 

2 

1 

2 

1 

2 

1 

1 

2 

1 

2 

1 

2 

1 

1 

2 

1 

2 

2 

2 

2 

1 

2 

1 

- 

- 

2 

2 

1 

1 

1

360 


TABLE 35.—Comparison of Eunice species in groups B-3 and B-4 with 

articulated ceratostyles; branchiae pectinate. Stub (C) gives character number, 

column heads give species number. Characters and character sutes are listed in 

Appendix A. Species are in order by number, corresponding species names are 


c 

7 

8 

9 

11 

12 

14 

15 

16 

17 

18 

20 

22 

23 

24 

25 

26 

27 

28 

29 

34 

37 

38 

39 

40 

41 

42 

43 

44 

48 

51 

52 

53 

54 

55 

56 

57 

58 

59 

60 

61 

64 

65 

66 

70 

75 

79 

4 

3 

1 

3 

1 

2 

2 

1 

1 

3 

3 

3 

1 

1 

1 

3 

3 

1 

2 

2 

1 

1 

1 

0 

u 

6 

30 

2 

2 

2 

5 

2 

1 

3 

2 

- 

2 

- 

- 

1 

1 

3 

1-2 

2 

2 

1 

2 

43 

3 

2 

3 

2 

2 

2 

U 

1 

1 

3 

3 

2 

2 

1 

2 

3 

1 

2 

2 

1 

1 

1 

0 

u 

12 

50 

2 

2 

3 

3-4 

1 

1 

1 

2 

- 

2 

- 

- 

1 

1 

3 

1 

2 

2 

1 

2 

56 62 93 142 


2 

2 

3 

2 

1 

2 

1 

1 

1 

3 

3 

1 

1 

2 

4 

2-3 

1 

2 

2 

2 

1 

1 

1 

U 

6 

U 

2 

2 

2 

4 

1 

2 

2 

1 

- 

U 

3 

2 

3 

2 

1 

5 

1 

1 

1 

3 

1 

1 

2 

2 

3 

4 

1 

2 

2 

1 

1 

1 

1 

0 

14 

35 

2-3 

2 

3 

9 

1 

1 

1 

2 

- 

2 

3 

2 

3 

1 

1 

6 

1 

2 

1 

3 

1 

1 

2 

2 

3 

5 

2 

2 

1 

1 

1 

1 

U 

U 

8 

25 

2 

3 

3 

_ 

u 

_ 

uu1 

_ 

1 

1 

3 2 

1 

2 

_ 

_ 

1 

U 

U 

u 

u 

1 

1 


1 

2 

2 

1 

2 

1 

1 

1 

1 

2 

1 

2 

2 

1 

2 

3 

3 

2 

2 

1 

2 

1 

3 

3 

1 

1 

1 

2 

2 

3 

4 

1 

3 

2 

2 

1 

1 

U 

U 

5 

40 

2 

2 

2 

9 

1 

1 

2 

3 

_ 

2 

_ 

_ 

2 

2 

3 

2 

1 

2 

1 

2 

157 

3 

1 

3 

1 

2 

2 

3 

1 

1 

1 

3 

1 

4 

1 

4 

3 

1 

2 

1 

1 

1 

1 

u 

130 

4 

U 

2 

2 

3 

4 

1 

1 

2 

1 

_ 

2 

_ 

_ 

1 

1 

2 

2 

1 

2 

1 

2 

181 

2 

2 

3 

2 

2 

2 

2 

1 

4 

2 

1 

1 

2 

2 

2 

2 

0 

55 

6 

65 

1-2 

2 

3 

4 

1 

1 

2 

2 

_ 



Character sutes of characters not included in the comparisons: 10,1; 13,1; 

19,2 21,1; 30,1; 31,1; 32,1; 33,2; 45,2; 69,1; 71,2; 72,2; 73,2; 74,1; 76,1; 78,1; 

81,1; 82,1. 

Species listed: 4 E. afra; 43 E. collaris; 56 E. ehlersi; 62 E.fauveli; 93 E. 

jagori; 142 E. pacifica; 181 E. sonorae. 

1 

2 

2 

_ 

2 

3 

_ 

_ 

2 

2 

1 

TABLE 36.—Comparison of Eunice species in groups B-3 and B-4 with 

articulated ceratostyles; branchiae palmate or single filaments. Stub (C) gives 

character number, column heads givespecies number. Characters and character 

states are listed in Appendix A. Species are in order by number; corresponding 

species names are listed at the end of the table. 

c 

7 

8 

9 

11 

12 

14 

15 

16 

18 

23 

24 

25 

26 

28 

31 

32 

33 

37 

39 

40 

41 

42 

43 

44 

48 

51 

52 

54 

55 

57 

58 

59 

60 

61 

64 

65 

74 

75 

76 

78 

79 

81 

9 

1 

2 

3 

2 

1 

3 

1 

1 

3 

4 

1 

3 

4 

1 

3 

U 

U 

1 

- 

- 

- 

- 

2 

2 

2 

9 

1 

1 

2 

2 

- 

- 

1 

1 

3 

2 

2 

1 

1 

2 

2 

1 

40 47 61 80 

SOFT BODY CHARACTKR STATES 

3 3 

2 1 

2 2 

1 2 

1 2 

2 i: 

1 2 

3 1 

3 

1 

2 

3 

3 

3 

2 

2 

2 

2 

1 

1 

3 

4 

1 

1 

y 

l 

2 

1 

2 

1 

U 

u 

2 

U 

14 

2 

2 

4 

3 

2 

1 

1 

2 

1 

_ 

2 

3 

2 

2 

3 

1 

2 

2 

3 

1 

3 

4 

1 

1 

4 

1 

2 

1 

2 

1 

U 

u 4 

130 

3 

2 

2 

4 

1 

2 

2 

2 

- 

_ 

2 

1 

3 

2 

2 

2 

2 

1 

u 

U 

3 

U 

24 

2 

1 

4 

1 

2 

2 

1 

2 

1 

_ 

3 

1 

4 

2 

3 

2 

1 

2 

1 

2 

1 

10 

u 2 

30 

2-3 

2 

3 

6-7 

3 

1 

1 

1 

1 

1 

_ 

3 

3 


1 

1 

1 

4 

1 

4 

1 

1 

U 

1 

4-5 

U 

2 

1 

1 

U 

1 

1 

u 

2 

3 

2 

U 

1 

1 

u 

2 

1 

124 

3 

3 

3 

2 

1 

1 

1 

I 

4 

1 

} 

5 

 

1 

- 

2 

- 

_ 

1 

- 

u 

U 

u4 

u 1 

u 2 

- 

_ 

1 

1 

3 

2 

U 

1 

1 

u 

2 

1 

162 

3 

1 

2 

2 

1 

U 

1 

2 

3 

1 

2 

3 

5 

U 

2 

3 

2 

2 

U 

U 

3 

U 

2-3 

3 

2 

9 

1 

1 

2 

2 

- 

_ 

2 

1 

3 



Character sutes of characters not included in the comparisons: 10,1; 13,1; 

19,2 21,1; 29,2; 30,1; 32,1; 33,2; 34,-; 45,2; 66,2; 69,1; 71,2; 72,2; 73.2; 74,1; 

76,1; 78,1; 81,1; 82,1. 

Species listed: 9 E. pruvoti; 40 E. cirrobranchiata; 47 E. conglomerate; 61 

E. excariboea; 80 E. guanica; 124 E. modesta; 162 E. prognatha. 

2 

U 

2 

1 

U 

2 

1


TABLE 37.—Comparison of Eunice species in groups B-3 and B-4 with branchiae present, without antennal 

articulations, and with the three median antennae of similar length. Stub (C) gives character number, column 

heads give species number. Characters and character sutes are listed in Appendix A. Species are in order by 

number, corresponding species names are listed at the end of the uble. 

c 

7 

8 

9 

11 

12 

13 

14 

16 

17 

18 

20 

24 

25 

26 

27 

28 

31 

32 

33 

34 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

57 

58 

59 

60 

61 

64 

65 

66 

70 

74 

76 

78 

79 

1 

2 

2 

3 

2 

2 

1 

2 

1 

1 

1 

3 

2 

3 

3 

1 

5 

1 

1 

2 

1 

1 

1 

10 

10 

5 

32 

2 

3 

2 

3 

15-20 

2 

1 

3 

2 

_ 

_ 

2 

4 

3 

2 

2 

2 

2 

1 

2 

2 

44 

3 

1 

3 

2 

2 

2 

- 

1 

1 

3 

1 

2 

3 

3 

1 

2 

2 

3 

2 

1 

2 

2 

U 

U 

2 

U 

1-2 

2 

2 

2 

4 

1 

1 

2 

2 

_ 

_ 

2 

1 

3 

2 

2 

2 

U 

1/2 

U 

2 

45 63 74 103 104 


U 2 3 U 3 

u 2 2 

1 

u 3 1 

2 

u 2 1 

2 

u 

1 

3 

1 

1 

2 

2 

1 

3 

1 

1 

3 

u 3 1 

1 

uuuu3 1 1 

1 

3 3 

3 

3 3 

3 

2 1 

2 

3 4 

3 

5 7 2 

2 

1 1 

1 

2 1 

1 

2 3 

2 

1 1 

3 

2 - 

2 

- - 

- 

3 

U 

1 

1 

2 

U 

U 

U 

2 

U 

6 

U 

u 

uu 

u 

u 

u 

u 

2 

U 

U 

U 

u 

u 

U 

U 

2 

U 

1 

U 

2 

1 

1 

U 

U 

2 

U 

2-3 

3 

2 

2 

4 

1 

2 

1 

2 

- 

- 

2 

1 

3 

1 

2 

_ 

_ 

- 

_ 

2 

2 

2 

3 

9 

1 

2 

2 

2 

_ 

- 

2 

1 

3 

u 

u 

u 

u 

u 

uuuuuuuuuuuuuu2 

u 

uu4 

u 

2 

3 

U 

U 

U 

u 

u 

u 

1 

1 

1 

u 

u 

2/3 

1-2 

2 

0 

0 

3 

U 

1 

2 

2 

2 

1-2 

1 

2 

1 

2 

_ 

- 

1 

3 

2 


1 2 u 

1 1 u 

1 2 2 

U U 1 

4 1 1 

U U 1 

2 2 2 

1 

2 

2 

U 

1 

U 

2 

112 

U 

U 

U 

u 

uuuuuuuuuuuu 

1 

1 

u 

1 

u 

uuu8 

u 

1 

2 

1 

3 

U 

2 

U 

U 

2 

_ 

- 

1 

3 

3 

2 

1 

2 

1 

1 

1 

2 

115 

138 

3 U 

2 u 

2 u 

2 u 

1 u 

1 u 

2 uuuu4 

3 

4 

3 

3 

2 u 

3 uu1 

4 

1 

5 u 

3 2-3 

1 U 

- 2 

- - 

1 2 

1 1 

- _ 

_ - 

1 1-3 

- U 

2 U 

2 U 

2 U 

3 U 

11 9 

1 U 

2 U 

2 u 

2 u 

_ 

u 

- 

u 

1 u 

2 3 

3 3 

2 1 

2 1 

2 2 

U 1 

1 1 

u 

1 



Character sutes of characters not included in the comparisons: 10,1; 15,1; 19,2; 21,2; 22,-; 23,1; 29,2; 30,1; 

53,1; 56,-; 69,1; 71,2; 72.2; 75,1; 77,2; 81,1; 82,1. 

Species listed: 1 E. aciculala, 44 E. collini; 45 E. complanata; 63 E. filamentosa; 74 E. gagzoi; 103 E. 

leucosticta; 104 E. levihranchia; 112 E. magnified; 115 E. marenzelleri; 138 E. noiata; 155 £. perrieri. 

1 

2 

155 

3 

3 

3 

2 

2 

1 

6 

1 

1 

1 

1 

1 

3 

3 

1 

2 

1 

1 

2 

1 

1 

1 

2-3 

4-5 

10 

30 

2 

2 

2 

3 

9 

2 

2 

2 

2 

- 

- 

2 

1 

3 

1 

2 

2 

1 

1 

1 

2


TABLE 38.—Comparison of Eunice species from group B-4 with branchiae, 

with the median antenna distinctly longer than the other antennae, without 

antennal articulations. Stub (Q gives character number, column heads give 

species number. Characters and character states are listed in Appendix A. 

Species are in order by number, corresponding species names are listed at the 

end of the table. 

c 

1 

44 

45 49 52 106 112 

129 

138 

187 

7 

8 

9 

12 

13 

14 

16 

17 

18 

20 

24 

25 

26 

27 

28 

31 

32 

33 

34 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

56 

57 

58 

59 

60 

61 

2 

2 

3 

2 

1 

2 

1 

1 

1 

3 

2 

3 

3 

1 

5 

1 

1 

2 

1 

1 

1 

10 

10 

5 

32 

2 

3 

2 

3 

15-20 

2 

1 

3 

- 

2 

- 

- 

2 

4 

3 

1 

3 

2 

2 

_ 

1 

1 

3 

1 

2 

3 

3 

1 

2 

2 

3 

2 

1 

2 

2 

U 

U 

2 

U 

1-2 

2 

2 

2 

4 

1 

1 

2 

- 

2 

- 

_ 

2 

1 


U 3 2 2 U 

uuu 1 2 3 U 

3 3 3 

u 

1 1 1 

1 1 1 1 u 

3 3 2 3 uuuuuui: 

U 1 3 2 

U 1 3 4 

U 1 3 3 

U 1 3 1 

u 2 2 2 

3 3 1 4 

5 1 3 5 u 

3 1 1 2 i; 

U 1 2 3 Li 

2 U 

2 

u 

U u 2 _ 1 

u 1 2 u 

u2 

1 

2 uuu8 

u 5 20 

U u2 U 50 

6 

6 2 

U U U 35 

u 

u 2-3 2 3 1 

u 2 2 2 2 

u 2 2 2 1 

u 3 3 2 3 

u 4 U 4 U 

uu2 13 2 1 2 

1 1 2 U 

2 2 1 u 

_ 2 _ _ 

u U U 2 2 

u U u 

_ _ 

u 

u 

_ _ 

u u u 3 1 

u 1 2 4 3 

1 

1 

2 

1 

2 

1 

1 

1 

3 

2 

3 

1 

1 

4 

1 

4 

3 

1 

1 

4 

12 

4 

1 

1 

2 

2 

1 

1 

2-3 

0 

12 

25 

1-2 

3 

2 

3 

9 

1 

2 

2 

_ 

1 

2 

2 

_ 

1 

U 

U 

U 

u 

u 

u 

uuu4 

u uu1 

u 

u 

2 

- 

2 

1 

- 

_ 

1-3 

U 

U 

u 

u u 

9 

u 

u u 

u uu 

u 

3 

u 

u3 

u uuuu3 

3 

u 4 

u 1 

5 

2 

1 

2 

- 

1 

1 

10 

U 

4 

400 

2 

2 

2 

2 

200 

3 

1 

2 

_ 

2 

_ 

_ 

3 

4 

65 

66 

74 

76 

78 

82 

2 

2 

2 

1 

2 

1 

2 

2 

U 

1/2 

U 

1 


U 2 1 2 

2 1 2 

1 u U 

1 1 1 

1 u U 

1 1 1 

u 

u1 

u 

u 





15,1; 19,2; 21,2; 22.-; 23,2; 29,2; 30,1; 53,1; 64,3; 69,1; 70,2; 71,2; 72,2; 73 2- 

75,1; 77,2; 79,2; 81,1. 

Species listed: 1 E. aciculata; 44 E. collini; 45 E. complanata; 49 E. 

crassitentaculata; 52 E. dilatata; 106 E. Vila; 112 E. magnified; 129 E. 

mutabilis; 138 E. noiata; 187 E. tenuis. 

2 

1 

1 

1 

1 

U 

1 

2 

1 

1 

1 

1 

U 

2 

1 

U 

1 

u 

3


TABLE 39.—Comparison of Eunice species in group B-4 with branchiae; with all antennae similar in length 

(short) and without antennal articulations. Stub (C) gives character number, column heads give species number. 

Characters and character states are listed in Appendix A. Species are in order by number, corresponding species 

names are listed at the end of the table. 

c 

35 

44 

45 51 66 112 138 

147 

148 

154 

182 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

23 

24 

25 

26 

27 

28 

29 

31 

32 

34 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

54 

55 

60 

61 

64 

3 

3 

3 

1 

2 

1 

U 

U 

1 

1 

3 

1 

2 

3 

4 

1 

4 

2 

1 

1 

2 

2 

1 

1 

1 

1 

3 

U 

3 

U 

1-2 

3 

2 

3 

20 

1 

1 

3 

2 

1 

3 

3 

1 

3 

1 

2 

2 

2 

- 

1 

1 

1 

3 

2 

1 

U 

2 

3 

3 

1 

2 

2 

2 

3 

1 

2 

2 

U 

u 2 

U 

1-2 

2 

2 

2 

4 

1 

1 

2 

2 

1 

3 


U 

U 

U 

u 

u 

u 

1 

3 

U 

u 

u 

u 

u 

u 

uu 

3 

5 

3 

U 

2 

1 

1 

U 

u 

u2 

U 

2 

3 

1 

2 

1 

1 

2 

3 

1 

1 

1 

2 

1 

4 

1 

3 

3 

1 

1 

2 

2 

1 

_ 

1 

1 

10-12 

U 

3 

U 

1 

3 

2 

3 

6 

U 

u 

u 

u 

uuu2 

u 

uu 

2 

9 

1 

2 

2 

2 

4 

3 

3 

1 

3 

2 

1 

2 

U 

U 

3 

2 

1 

3 

2 

3 

4 

2 

4 

2 

1 

2 

2 

1 

1 

1 

2 

1 

U 

u 

6 

50 

1 

3 

2 

2 

4 

1 

2 

2 

U 

4 

3 

U 

U 

u 

u 

u uuuuuuuuuuuuuuuu1 

1 

1 

u 

u 

u 

u 

8 

u 

1 

2 

1 

3 

u 

2 

U 

u 

1 

3 

3 

U 

u 

u 

u 

u 

uuuuuuuu4 

u uuu 

1 

u 

2 

2-3 

U 

- 

2 

1 

_ 

- 

1-3 

U 

u 

u 

u u 

9 

u 

u 

u 

u 

3 

3 

3 

1 

2 

1 

2 

1 

1 

2 

2 

1 

1 

3 

2 

3 

4 

2 

4 

4 

1 

1 

2 

2 

1 

_ 

1 

1 

U 

U 

3 

40 

1-2 

U 

2 

1 

4-5 

1 

1 

2 

U 

2 

3 

U 

u 

u 

u 

u 

uuuuuuuu 

3 

4 

u 

4 

1 

U 

u 

u 

2 

U 

U 

U 

u 

u 

u3 

U 

U 

U 

U 

U 

3 

U 

u 

5 

U 

4 

3 

1 

2 

3 

1 

2 

1 

1 

3 

1 

1 

1 

3 

2 

1 

4 

2 

3 

2 

1 

2 

2 

1 

1 

2 

1 

1 

1 

U 

8 

30 

1-2 

2 

2 

2 

9 

1 

1 

2 

U 

4 

3 

1 

2 

3 

1 

2 

2 

3 

2 

1 

4 

2 

2 

5 

1 

2 

2 

2 

1 

- 

1 

1 

5 

3 

3 

U 

1-3 

2 

2 

3 

3 

3 

1 

3 

2 

4 

3 

65 

66 

74 

76 

78 

2 

2 

1 

1 

1 

2 

2 

U 

12 

U 


U 

U 

U 

1 

1 

1 

U 

4 

2 

1 

U 

1 

2 

1 

1 

1 

1 

1 

1 

1 

U 

1 1 

u 

u 


Character states of characters not included in the comparisons: 21,2; 22,-; 30,1; 33,2; 53,1; 56,-; 57,2; 58,-; 

59,-; 69.1; 70.2; 71.2; 72,2; 73,2; 75,1; 77.2; 79.2; 81,1; 82,1. 

Species listed: 35 E. bucciensis; 44 E. collini; 45 E. complanata; 51 E. denticulata; 66 E.flavapunctata; 112 

E. magnified; 138 E. notata; 147 E. parvibranchis; 148 E. paupera; 154 E. perimensis; 182 E. spongicola. 

1 

2 

U 

1 

U 

U 

U 

U 

U 

u 

1 

2 

2 

1 

2 

2 

2 

U 

4 

U


TABLE 40.—Comparison of abranchiate Eunice species. Stub (C) gives 

character number; column heads give species number. Characters and character 

sutes are listed in Appendix A. Species are in order by number corresponding 


c 

34 

37 50 73 75 

88 

203 

7 

8 

9 

10 

11 

12 

14 

15 

16 

20 

21 

22 

24 

25 

26 

28 

43 

48 

51 

54 

55 

57 

58 

59 

60 

61 

64 

1 

2 

2 

1 

2 

2 

U 

3 

1 

3 

1 

1 

1 

1 

2 

2 

2-3 

2 

7 

2 

2 

2 

_ 

_ 

1 

2 

3 


2 

2 

2 

1 

2 

2 

2 

1 

3 

3 

2 

- 

2 

3 

3 

5 

2-3 

2 

4-5 

2 

3 

2 

- 

_ 

3 

3 

3 

3 

2 

1 

2 

1 

1 

3 

3 

3 

3 

1 

3 

2 

3 

3 

2 

14 

2 

11 

1 

2 

2 

_ 

1 

4 

3 

3 

1 

3 

1 

2 

2 

U 

3 

1 

2 

1 

1 

1 

1 

4 

2 

1 

2 

9 

1 

2 

2 

_ 

_ 

1 

3 

3 

3 

1 

2 

1 

2 

U 

2 

1 

3 

3 

2 

- 

U 

4 

2 

5 

13 

3 

9 

1 

1 

2 

_ 

_ 

2 

1 

3 

3 

2 

3 

1 

2 

2 

2 

1 

1 

2 

2 

- 

2 

4 

2 

2 

2 

2 

9 

1 

1 

1 

1 

2 

_ 

3 

3 

3 

1 

3 

2 

1 

2 

3 

1 

1 

3 

2 

- 

2 

3 

2 

1 

3 

3 

4 

2 

4 

2 

_ 

_ 

3 

3 

3 

65 

66 

2 

2 


2 

1 

2 

1 

2 

2 



Species listed: 34 E. brevis; 37 E. cariboea; 50 E. cwticirris; 73 E. 

fuscafasciata; 75 E. goodei; 88 E. imogena; 203 £. wasinensis. 

2 

1 

2 

1 

1 

2


TABLE 41.—Comparison of Eunice species in group C-l. Column heads are the same as character numbers used 

to prepare the key (see explanation below; for a complete listing of all characters and character slates see 

Appendix A). 

Species 

6 E. americana 

16 E. arenosa 

20 E. allantica 

21 E. aucklandica 

12 E. australis 

27 E. bicirrata 

38 E. cedroensis 

46 E. congesta 

60 E. eugeniae 

65 E.flaccida 

67 E. flavocuprea 

90 E. indica 

91 E. interrupta 

96 E. kerguelensis 

105 E. limosa 

117 E. marovoi 

119/:. medic ina 

125 E. mucronata 

126 E. multicylindri 

128 E. murrayi 

145 E. papeetensis 

206 E. parva 

152 E.pellucida 

161 E. pray ens is 

172 E. rubrivittata 

186 E. tentaculata 

197 E. unifrons 

201 E. vittata 

202 E. vitlatopsis 

205 E. woodwardi 

1 

123 

79 

U 

79 

109 

U 

U 

u uuuuu 

81-106 

65 

79 

116 

115 

101 

U 

59 

U 

U 

U 

U 

U 

73 

U 

U 

2 

86 

16 

U 

18 

70 

U 

U 

U 

u 

17 

U 

U 

u 

32-65 

14.5 

29 

58 

45 

37 

U 

17 

U 

U 

U 

U 

U 

23.5 

U 

U 

3 

2 

0.75 

2 

1 

4 

1 

2 

4 

1 

2.5 

1 

2 

2 

0.3 

2 

1 

1.3 

4 

1 

4 

2 

0.75 

2 

2 

1.2 

U 

1 

2 

2.5 

5 

5 

9 

2.5 

U 

3.2 

9 

U 

3 

6 

4 

8.5 

2 

4 

9 

1.2 

U 

2.5 

5 

8 

3.5 

7 

2.5 

4 

3.5 

4.5 

5.1 

U 

5 

4 

6 

7 

35 

3 

3 

3 

8 

7 

3 

3 

7 

4 

7 

4 

3 

5-7 

3 

3 

3 

3 

3 

3 

5 

6 

5 

5 

5 

3 

6 

3 

3 

3 

3 

36 

35 

25/26 

U 

21 

33 

48 

27 

47 

44 

U 

34 

21 

24-26 

U 

36 

17-22 

26 

34 

40 

28 

30/33 

20 

28 

33 

56 

28 

45-54 

23 

U 

39 

80 

25 

16 

U 

18 

31 

45 

17 

33 

18 

27 

17 

18 

24-27 

19 

U 

17-19 

17 

18-22 

27 

17 

21 

24 

21/23 

18 

31 

23 

25-31 

20-22 

39 

31 











c 

7 

8 

9 

10 

11 

12 

14 

15 

16 

18 

20 

23 

i< 

25 

26 27 

28 

29 

31 

32 

34 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

55 

60 

61 

64 

65 

66 

69 

70 

74 

78 

82 

TABLE 42.—Comparison of Eunice species of group C-1 with cylindrical articulations in the ceratostyles. Stub 

(Q gives character number, column heads give species number. Characters and character states are listed in 

Appendix A. Species are in order by number, corresponding species names are listed at the end of the table. 

6 

1 

1 

2 

1 

2 

2 

2 

3 

1 

3 

1 

1 

«j 

3 

2 

2 

3 

2 

1 

1 

1 

0 

2 

20 

10 

1 

2 

2 

2 

5 

3 

3 

2 

2 

1 

2 

2 

1 

1 

2 

2 

1 

16 

3 

2 

2 

2 

1 

2 

U 

1 

1 

3 

3 

2 

1 

2 

1 

1 

1 

1 

1 

5 

2-3 

5 

15 

1 

2 

2 

2 

4 

1 

2 

2 

3 

3 

U 

U 

1 

1 

2 

1 

1 

20 

3 

2 

2 

1 

U 

U 

u 

u 

u 

3 

1 

2 

2 

2 

1 

1 

1 

U 

u 

1 

u 

5 

U 

u 

uuuuuu1 

u u 

u u 

1 

1 

2 

2 

u 

21 

3 

2 

3 

1 

2 

2 

2 

1 

1 

3 

3 

2 

3 

1 

1 

2 

2 

1 

- 

U 

U 

2 

U 

2-3 

2 

2 

2 

5 

1 

2 

1 

1 

3 

2 

1 

1-2 

2 

2 

2 

3 

38 67 90 119 126 


1 

1 

2 

2 

1 

2 

2 

1 

1 

3 

1 

1 

2 

2 

2 

1 

2 

2 

U 

1 

1 

3 

3 

2 

3 

2 

2 

1 

1 

1 

2 

1 

2 

3 

1 

2 

2 

2 

2 

1 

2 

1 

2 

1 

1 

3 

3 

U 

3 

3 

3 

1 

2 

2 

4 

1 

3 

3 

3 

1 

2 

2 

1 

2 

1 

1 

1 

2 

3 

10 

10 

1 

3 

2 

2 

5 

1 

2 

U 

3 

1 

2 

1 

5 

2 

1 

1 

1 

5 

10 

4 

15 

U 

U 

u 

u 

4 

U 

1 

U 

2 

3 

2 

2 

1 

0 

8 

U 

2 

2 

2 

2 

2 

1 

2 

1 

2 

3 

2 

2 

2 

1 

1 

U 

1 

5 

0 

7 

U 

1-2 

3 

2 

1 

2 

3 

3 

2 

4 

2 

2 

1-2 

2 

10-15 

4 

U 

1-2 

2 

2 

2 

5-6 

1 

2 

1 

1 

3 


2 

1 

1 

1 

2 

2 

1 

U 

U 

1 

1 

2 

2 

2 

U 

U 

1 

1 

2 

2 

5 

2 

2 

1 

1 

2 

2 

1 

1 

1 

1 

2 

2 

2 

1 

161 

3 

3 

2 

1 

1 

2 

3 

1 

1 

1 

U 

U 

1 

i; 

u 1 

1 

1 

1 

0 

9 

12 

1 

2 

2 

2 

2 

1 

2 

1 

1 

3 

2 

1 

1 

1 

2 

2 

2 

172 

3 

2 

2 

2 

2 

1 

U 

1 

1 

3 

3 

1 

1 

U 

U 

u 1 

3 

2 

6 

11 

5 

25 

2/4 

2 

1 

1 

5-6 

3 

3 

1 

3 

2 



Character states of characters not included in the comparison: 13,1; 17,1; 19,2; 21,1; 22,1; 24,2; 30,1; 33,2; 

37,2; 38,2; 53,1; 54,1; 56.-; 57,2; 58.-; 59,-; 71,2; 72,2; 73,1; 75,1; 76.1; 77,1; 79,1; 81,1. 

Species listed: 6 E. americana; 16 E. arenosa; 20 E. atlantica; 21 E. aucklandica; 38 E. cedroensis; 67 E. 

flavocuprea; 90 E. indica; 119 E. medicina; 126 E. multicylindri; 161 E. prayensis; 172 E. rubrivittala; 186 E. 

tentaculala; 197 E. unifrons; 201 E. vittata; 205 E. woodwardi. 

U 

U 

1 

1 

2 

2 

1 

186 

3 

u 2 

U 

u 

u 

u 

u 

u3 

U 

2 

3 

2 

IJ 

9 

U 

U 

u 

u 

u 

u 

u 

u 

u 

uu 

2 

2 

1/2 

2 

2 

2 

5 

197 

3 

1 

1 

1 

2 

2 

U 

1 

2 

3 

3 

1 

2 

1 

2 

2 

1 

1 

1 

4 

10 

5 

15 

3 

2 

2 

2 

5 

1 

3 

1 

1 

1 

2 

1 

1 

2 

2 

2 

1 

201 

3 

1 

1 

1 

2 

2 

2 

1 

1 

3 

3 

2 

2 

2 

1 

1 

1 

2 

1 

0 

1 

12 

12 

3 

3 

2 

3 

4 

1 

2 

1 

1 

3 

2 

1 

1 

2 

2 

2 

5 

205 

3 

2 

3 

1 

1 

2 

U 

1 

2 

3 

3 

2 

2 

1 

2 

2 

1 

1 

1 

0 

4-5 

12 

15 

1 

2 

2 

2 

3 

1 

2 

1 

2 

3 

U 

U 

1 

1 

1 

2 

5


TABLE 43.—Comparison of Eunice species in group C-l with distally 

drop-shaped or moniliform articulations in the ceratostyles. Stub (C) gives 

character number, column heads give species number. Charaaers and character 



c 

7 

8 

9 

10 

11 

12 

18 

20 

23 

24 

25 

26 

27 

28 

29 

31 

32 

34 

39 

40 

41 

42 

43 

51 

52 

55 

60 

61 

64 

66 

70 

82 

22 

3 

2 

3 

1 

1 

2 

3 

1 

2 

2 

3 

3 

1 

2 

1 

1 

1 

1 

0 

2 

8 

10 

1-2 

6 

3 

2 

2 

4 

2 

2 

2 

1 

27 96 105 161 


U 3 3 

u 1 U 

u 2 2 

uuuu3 2 U 

1 U 

2 u 

3 3 

3 3 

2 1 

U 1 

U 4 

2 

u 2 1 

U 2 

2 

1 

2 

- 

U 

u 

2 

U 

u 

2 

U 

U 

u 

u 

u 

u 

uuu 

u 

u 

2 

3 

1 

2 

U 

1 

3 

2 

U 

4 

2 

1 

2 

2 

1 

1 

1 

U 

u 

12 

U 

U 

u 

u 

u 

4 

3 


U 

2 

U 

1 

2 

1 

U 

1 

U 




15,1; 16,1; 17,1; 19,2; 21,1; 22,2; 30.1; 33,2; 37,2; 38,2; 45,2; 48,2; 53.1; 54,1; 

56,-; 57,2; 58.-; 59,-; 65,2; 69.1; 71,2; 72,2; 73,1; 74,2; 75,1; 76,1; 77.1; 

78,2; 79.3; 81,1. 

Species listed: 22 E. australis; 27 E. bicirrata; 96 E. kerguelensis; 105 E. 

limosa; 161 E. prayensis; 202 E. vittatopsis. 

3 

3 

2 

1 

1 

2 

1 

U 

u 

2 

4 

1 

U 

u 

u 

1 

1 

1 

1 

0 

9 

12 

1 

2 

1 

2 

1 

1 

3 

1 

1 

2 

202 

2 

1 

2 

1 

2 

1 

3 

1 

1 

2 

4 

3 

3 

1 

1 

1 

1 

1 

2 

U 

10 

u 

2 

4 

1 

3 

2 

2 

1 

1 

2 

1 

TABLE 44.—Comparison of Eunice species of group C-l with moniliform 

articulations in the ceratostyles. Stub (C) gives character number, column heads 

give species number. Characters and character states are listed in Appendix A. 

Species are in order by number; corresponding species names are listed at the 

end of the table. 

c 

8 

9 

11 

12 

13 

14 

16 

18 

20 

23 

24 

25 

26 

27 

28 

31 

32 

33 

34 

37 

38 

39 

40 

41 

42 

43 

44 

51 

52 

53 

54 

55 

56 

60 

61 

64 

66 

70 

74 

76 

79 

82 

65 

1 

2 

2 

2 

U 

U 

1 

3 

1 

2 

1 

2 

5 

1 

2 

1 

3 

U 

1 

U 

u 

0 

u 

9 

12 

2 

3 

_ 

1 

2 

- 

_ 

2 

U 

2 

3 

1 

2 

2 

1 

4 

1 

91 128 145 152 


2 

3 

2 

2 

1 

1 

1 

1 

1 

1 

1 

2 

5 

1 

1 

2 

3 

1-2 

- 

1-2 

1 

U 

U 

3 

U 

2 

2 

6 

1 

1 

1 

2 

- 

2 

4 

1 

1 

2 

2 

2 

U 

U 

1 

3 

1 

U 

2 

3 

3 

1 

2 

1 

1 

2 

1 

1 

2 

2 

5 

9 

10 

1 

2 

4 

3 

1 

1 

3 

- 

2 

2 

2 

3 

2 

2 

1 

1 

2 

3 

3 

1 

2 

2 

3 

5 

1 

1 

2 

U 

2 

- 

2 

2 

2 

5 

3 

U 

2 

2 

3 

1 

1 

1 

2 

- 

U 

1 

2 

1 

2 

2 

2 

2 

- 

1 

3 

3 

2 

2 

3 

4 

1 

1 

1 

U 

2 

1 

2 

2 

U 

U 

8 

U 

2 

2 

4 

1 

1 

1 

3 

- 

2 

2 

3 


U 

2 

2 

1 

3 

1 

U 

2 

1 

1 

3 

U 

2 

2 

2 

3 

3 

U 

1 

2 

2 

4 

3 

1 

161 

3 

2 

1 

2 

1 

3 

1 

1 

U 

U 

2 

4 

1 

U 

u 

1 

1 

2 

1 

2 

2 

1 

0 

9 

12 

1 

2 

2 

1 

1 

1 

2 

- 

1 

1 

3 




15,1; 17,1; 19.2; 21,1; 22,3; 29,1; 30,1; 45.2; 48,2; 57,2; 58,-; 59,-; 65,2; 69,1; 

71,2; 72.2; 73,1; 75,1; 77,1; 78,2; 81.1. 

Species listed: 65 E. flaccida; 91 E. inlerrupta; 128 E. murrayi; 145 E. 

papeetensis; 152 E. pellucida; 161 E. prayensis; 206 E. parva. 

1 

1 

2 

1 

3 

2 

206 

1 

2 

2 

2 

1 

4 

1 

3 

1 

2 

2 

3 

2 

2 

2 

2 

2 

4 

10 

2 

3 

4 

1 

1 

1 

2 

- 

2 

3 

2 

U 

2 

2 

1 

3 

1


TABLE 45.—Comparison of Eunice species in group C-l without antennal 

articulations. Stub (C) gives character number, column heads give species 

number. Characters and character sutes are listed in Appendix A. Species are 

in order by number; corresponding species names are listed at the end of the 

table. 

c 

8 

9 

10 

11 

12 

13 

14 

16 

17 

18 

20 

26 

27 

28 

29 

32 

39 

40 

41 

42 

43 

44 

48 

51 

55 

57 

58 

59 

60 

61 

46 60 117 125 


2 

2 

1 

2 

2 

1 

3 

1 

2 

2 

1 

2 

2 

- 

1 

1 

1 

2 

2 

1 

3 

1 

2 

1 

2 

1 

1 

2 

U 1 1 2 

U 4 1 1 

3 3 3 3 

3 1 1 1 

2 1 2 2 

U 1 1 3 

U 

U 

1 

1 

0 

2 

2 

2 

4 

2-3 

2 

1 

1 

0 

1 

1 

2 

1 

2 

0 

18 5 6 15 

U U 

u U 

3 1-2 2 1-2 

3 2 3 3 

1 

7-8 

1 

2 

_ 

- 

1 

3 

2 

3 

2 

2 

_ 

- 

2 

1 

2 

2 

3 

1 

1 

3 

_ 

1 

2 

3 

2 

2 

_ 

- 

1 

3 

161 

3 

2 

1 

1 

2 

1 

3 

1 

U 

1 

u 1 

f 

u 

u1 

1 

0 

9 

12 

1 

2 

2 

2 

2 

2 

_ 

- 

1 

1 

66 

82 


2 

1 

2 

1 

U 

5 




19,2; 21,2; 22,-; 23,2; 24,2; 25,4; 30,1; 31,1; 33,2; 34,1; 37,2; 38,2; 45,2; 52,1; 

53,1; 54,1; 56,-; 64,3; 65,2; 69,1; 70,1; 71,2; 72,2; 73,1; 74,2; 75,1; 76,1; 77,1; 

78,2; 79,3; 81,1. 

Species listed: 46 E. congesta; 60 E. eugeniae; 117 E. marovoi; 125 E. 

mucronata; 161 E. prayensis. 

2 

1 

1 

2


TABLE 46.—Comparison of of Eunice species in group C-2. The column numbers are the same as the character 



2 

3 

12 

24 

27 

32 

59 

65 

77 

84 

87 

91 

96 

109 

113 

116 

118 

139 

140 

144 

171 

184 

191 

Species 

aedificatrix 

aequabilis 

antennata 

bassensis 

bicirrata 

bottae 

elseyi 

flaccida 

gracilis 

havaica 

hirschi 

interrupta 

ktrguelensis 

lucei 

makemoana 

marganlicacea 

martensi 

oliga 

ornata 

panamena 

rubra 

sligmatura 

torresiensis 

1 

270 

158 

98 

U 

U 

U 

120 

U 

81 

78 

112 

U 

U 

U 

92 

95 

168 

82 

110 

U 

U 

u 

72 

2 

128 

175 

54 

U 

U 

u 

64 

U 

66 

25 

55 

U 

U 

u 

20 

14 

220 

16.5 

45 

U 

U 

U 

47 

3 

4 

7 

3 

U 

1 

2 

7 

2.5 

2 

1 

3 

3 

0.3 

3 

1 

U 

6 

2 

2 

3.5 

3 

2 

3 

5 

7 

16 

7 

U 

u 

7 

13 

8.5 

6 

4 

7 

9 

1.2 

5.5 

3.5 

U 

14 

U 

6 

6 

7.5 

4 

6.5 

35 

4 

6 

7 

U 

3 

6 

6 

7 

4-5 

6 

4 

5-7 

3 

5 

7 

6 

6 

5 

5 

5 

4 

3 

5 

36 

250 

151 

92 

U 

48 

U 

104 

U 

81 

78 

102 

U 

u 

168 

86 

95 

163 

82 

110 

U 

U 

110-126 

70 

80 

35 

26/32 

25 

U 

45 

25 

30 

27 

28 

19-23 

22/25 

24-27 

19 

24-34 

28 

U 

28 

U 

22-25 

27 

27 

7-35 

24 


paired alternative observations. Explanation of column numbers: 









TABLE 47.—Comparison of Eunice species of group C-2 with all notopodial cirri articulated. Stub (C) gives 


Species are in order by number, corresponding species names arc listed at the end of the table. 

c 

7 

8 

9 

10 

12 

14 

15 

16 

18 

20 

22 

23 

24 

25 

26 

27 

28 

31 

32 

33 

34 

37 

39 

40 

41 

42 

43 

44 

51 

52 

54 

55 

57 

60 

61 

66 

69 

70 

76 

78 

79 

82 

12 24 

2 

2 

1 

j 

3 

1 

1 

; 

1 

; 

u uuuuuu 

> u u 

) U 

2 U 

u 

:) U 

; 

) U 

uu1 

1 

1 1 

1 

1 1 

() U 

5 U 

7 9 

u 

u 

1 1 

2 2 

• U 

1 U 

1 U 

3 U 

2 2 

2 2 

1 U 

2 U 

1 1 

2 2 

1 1 

2 2 

3 3 

1 U 

27 

U 

U 

U 

u uuu3 

2 

2 

U 

u 

2u2 

L 1 

U 

U 

u uuuuuuuuuuuu 

u 1 

2 

1 

u 3 

u 

32 59 91 109 113 


3 

2 

2 

1 

2 

U 

1 

U 

3 

3 

3 

U 

1 

4 

5 

1 

1 

1 

1 

U 

1 

1 

3 

1 

3 

1 

1 

U 

1 

1 

2 

1 

3 

2 

2 

3 

4 

1 

1 

1 

1 

2 

1 

1 

3 

2 

3 

1 

2 

1 

1 

1 

1 

1 

3 

1 

1 

2 

5 

1 

1 

2 

3 

1-2 

- 

1-2 

3 

1 

3 

1 

1 

2 

1 

2 

3 

1 

2 

1 

1 

4 

4 

2 

4 

1 

1 

2 

1 

1 

0 0 U 1 

u 0 U U 

8 18 3 8 

15 U U U 

1-2 

3 

U 

1 

1 

3 

2 

2 

1 

1 

2 

5 

1 

1 

3 

1 

2 

1 

2 

2 

6 

1 

1 

2 

2 

2 

4 

2 

3 

4 

1 

2 

1 

2 

1 

1 2 

U 

2 

2 

1 

2 

3 

1 

3 

1 

U 

1 

u 2 

U 

u 3 

U 

3 

2 

U 

u 

II 

u 3 

1 

- 

- 

1 

- 

- 

- 

- 

2 

U 

u 1uu2 

1 


2 

2 

2 

5 

2 

4 

1 

U 

1 

2 

1 

2 

3 

1 

1 

1 

1-2 

4-5 

2 

3 

1 

U - feature is unknown; dash (-) = character inappropriate. Numbers separated by a dash indicate a range; 


Character states of characters not included in the comparisons: 11,2; 13,1; 17,1; 19,2; 21,1; 29,1; 30 1; 38,1; 

45,2; 48,2; 53,1; 56,-; 58,-; 59,-; 64,1; 65,2; 81,1. 

Species listed: 12 E. antennata; 24 E. bassensis; 27 E. bicirrata; 32 E. bottae; 59 E. elseyi; 91 E interrupta; 

109 E. lucei; 113 E. makemoana; 116 E. margarilicacea; 118 E. mariensi; 139 E. oliga; 171 E. rubra; 191 E. 

torresiensis. 

1 

1 

2 

1/4 

2 

3 

U 

116 

u uuUu 

u 2uu2u1u1 

1 

1 

u 5 

10 

u 

uu1 

1 

u1_ 

u 

2 

2 

2 

1 

2 

3 

U 

118 

U 

U 

2 

1 

U 

3 

I) 

U 

3 

3 

1 

U 

2 

3 

3 

2 

2 

1 

1 

2 

1 

1 

0 

0 

16 

15 

1-2 

2 

U 

1 

1 

2 

2 

1 

1 

2 

2 

2 

1 

1 

3 

1 

139 

3 

3 

3 

U 

U 

u u 

I) 

3 

1 

3 

2 

i; 

3 

I 

2 

1 

2 

U 

1 

_ 

1 

1 

1 

3 

12 

1 

U 

U 

u uu2u3 

1 

1 

2 

1 

2 

3 

U 

171 

2 

2 

2 

2 

2 

3 

1 

1 

3 

3 

3 

U 

2 

3 

:> 

o 

u 

21 

15 

2 

2 

3 

1 

1 

2 

2 

4 

1 

1 

1 

2 

15 

2 

3 

2 

191 

3 

1 

1 

1 

2 

3 

3 

2 

1 

1 

3 

2 

2 

3 

2 

0 

7 

15 

1 

2 

4 

3 

1 

3 

2 

2 

4 

U 

1 

2 

1 

2 

3 

1


TABLE 48.—Comparison of Eunice species in group C-2 with articulations of 

the notopodia limited to the anterior end. Stub (C) gives character number, 

column heads give species number. Characters and character states are listed in 

Appendix A. Species are in order by number, corresponding species names are 


c 

7 

8 

9 

11 

12 

14 

16 

17 

20 

22 

23 

24 

25 

26 

27 

28 

31 

32 

33 

39 

40 

41 

42 

43 

44 

48 

51 

52 

55 

60 

61 

64 

65 

66 

69 

76 

79 

27 

U 

u 

u u 

u u 

u 

3 

2 

2 

U 

u 

u2 

U 

2 

U 

u 

u 

u 

u 

u 

uuuuuuuu 

u 

u 

1 

1 

3 

84 87 140 144 


2 

2 

2 

1 

1 

2 

1 

1 

1 

3 

2 

2 

4 

3 

1 

1 

1 

2 

1 

U 

1 

2 

3 

2 

2 

U 

1 

3 

1 

3 

2 

1 

3 

3 

2 

1 

1 

3 

2 

1 

3 

1 

3 

2 

1 

3 

1 

1 

1 

3 

2 

1 

3 

3 

1-2 

1 

1 

1 

1-2 

1 

u 

7 0 

5 7 20 

20 

1 

2 

2 

4 

3 

2 

1 

3 

2 

10 

2 

2 

2 

4 

1 

2 

1 

1 

1 

U 

2 

2 

3 

9 

1 

3 

2 

3 

2 


2 

1 

1/2 

1 

3 

2 

2 

1 

1 

3 

1 

1 

1 

1/5 

3 

3 

2 

3 

2 

2 

2 

3 

3 

2 

3 

1 

2 

3 

4 

1 

1 

1 

3 

2 

1 

U 

8 

15 

1 

2 

2 

3 

1 

1/3 

U 

1 

2 

2 

1 

1 

4 

3 

184 

3 

1 

2 

2 

1 

2 

1 

1 

3 

1 

1 

2 

4 

2 

2 

2 

1 

2 

2 

4 

11 

4 

15 

1 

3 

2 

6 

1 

2 

2 

3 

2 

2 

1 

1 

1 

2-3 

U = feature is unknown. Numbers separated by a dash indicate a range; 


Character states for characters not included in the comparisons: 10,1; 13,1; 

15,1; 18,3; 19,2; 21,1; 29,1; 30.1; 34,1; 37,1; 38,1; 45,2; 53,1; 54,1; 56,-; 57,2; 

58,-; 59,-; 70,2; 71,2; 72,2; 73,1; 74,2; 75,1; 77,1; 78,2; 81,1; 82,1. 

Species listed: 27 E. bicirrata; Tl E. gracilis; 84 E. havaica; 128 E. murrayi; 

140 E. ornata; 144 E. panamena; 184 E. stigmatura. 

TABLE 49.—Eunice species of group C-2 without articulated notopodial cirri. 

Stub (C) gives character number, column heads give species number. 

Characters and character states are listed in Appendix A. Species are in order by 

number, corresponding species names are listed at the end of the table. 

c 

7 

8 

9 

10 

11 

12 

13 

14 

16 

20 

22 

23 

24 

25 

26 

27 

28 

29 

31 

32 

34 

37 

38 

39 

40 

41 

42 

43 

44 

53 

54 

55 

56 

61 

64 

65 

69 

76 

79 

2 3 27 65 


2 U 

2 U 

2 u 

1 u 

2 u 

2 u 

2 u 

- 

u 

2 u 

1 3 

1 2 

2 2 

2 U 

3 U 

5 

u 

1 2 

1 U 

3 

1 

3 

1 

2 

1 

1 

2 

1 

1 

3 

2 

1 

4 

2 

2 

4 

1 

1 

1 

1 

1 

1 

10 

10 

11 

15 

1-2 

2 

1 

1 

2 

_ 

1 

3 

1 

1 

1/4 

3 

1 

0 

0 

17 

20 

1 

3 

1 

1 

2 

_ 

1 

3 

3 

1 

T 

1 

2 

2 

U 

U 

1 

1 

3 

2 

1 

2 

5 

1 

2 

u 1 

2 1 

U 3 

u 1 

uu U 

U 

u 0 

u 

uuuuuuuuuu 


2 

2 

3/5 

3 





18,3; 19,2; 21,1; 30,1; 33,2; 45,2; 48,2; 51,3; 52,1; 57,2; 58,-; 59,-; 60,1; 64,3; 

66,1; 70,2; 71,2; 72,2; 73,1; 74,2; 75,1; 77,1; 78,2; 81,1; 82,1. 

Species listed: 2 E. aedificatrix; 3 E. aequabilis; 27 E. bicirrata; 65 E. 

flaccida; 96 E. kerguelensis. 

u 

1 

1 

3 

u 

9 

12 

2 

3 

2 

- 

- 

2 

2 

3 

2 

1 

1 

4 

% 

3 

1 

2 

2 

1 

2 

1 

3 

1 

3 

2 

1 

1 

4 

2 

1 

2 

2 

1 

2 

- 

2 

2 

U 

U 

2 

U 

2 

2 

1 

1 

2 

_ 

1 

3 

U 

1 

1 

3


TABLE 50.—Comparison of Eunice species in group D. Column heads are the same as the character numbers used 

to prepare the key (see explanation below; for a complete listing of all characters and character states sec 

Appendix A). 

72 

no 

115 

178 

181 

196 

Species 

E. fucala 

E. macrobranchia 

E. marenzelleri 

E. sebasliani 

E. sonorae 

E. unidentata 

1 

250 

150 

U 

706 

586 

U 

2 

195 

200 

U 

1455 

345 

80-100 

3 

5 

U 

5 

23 

6 

2-3 

5 

10 

u11 

35 

14 

U 

35 

5 

7-8 

28 

7-14 

41 

23 

36 

170-180 

U 

U 

686 

530 

U 

80 

u 

U 

38 

51/56 

54 

20-25 






5 = Length through seliger 10 in mm. 




TABLE 51.—Comparison of Eunice species in group D. Stub (C) gives 

character number, column heads give species number. Characters and character 



c 

7 

8 

9 

11 

12 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

31 

32 

33 

34 

37 

38 

39 

40 

41 

42 

43 

44 

48 

51 

54 

72 

U 

U 

U 

U 

U 

U 

U 

U 

U 

U 

2 

- 

1 

U 

u 

3 

2 

1 

1 

1 

2 

1 

2 

1 

U 

30 

12 

U 

1 

3 

2 

U 

U 

110 115 178 181 


U 

U 

u 

u 

u 

u 

u 

u 

uu1 

1 

uuuu1 

u 

1 

2 

2 

1 

1 

1 

5 

U 

15-17 

U 

u 

u 

u 

u 

u 

3 

2 

2 

2 

1 

3 

4 

3 

2 

3 

2 

_ 

1 

2 

3 

4 

1 

5 

3 

1 

_ 

_ 

1 

1 

_ 

- 

1 

- 

2 

2 

3 

11 

2 

2 

1 

2 

2 

2 

3 

1 

2 

2 

1 

2 

_ 

4 

1 

3 

5 

1 

4 

1 

1 

2 

1 

1 

1 

0 

3 

40 

35 

15 

2 

1 

6-13 

2 

2 

2 

3 

2 

2 

1 

1 

1 

2 

1 

1 

1 

2 

1 

4 

2 

1 

1 

1 

1 

2 

2 

2 

2 

0 

55 

6 

65 

1-2 

2 

3 

4 

2 

196 

3 

U 

2 

1 

U 

1 

1 

1 

1 

1 

2 

_ 

4 

U 

u 

u 

1 

1 

1 

1 

2 

1 

2 

2 

U 

U 

5 

35 

1 

2 

U 

U 

1 

c 

55 

57 

58 

59 

60 

61 

65 

66 

67 

68 

74 

75 

79 

82 


72 110 115 178 181 

SOFT BODY CHARACTER STATES—Continued. 

U 

2 4 2 

2 u 2 1 1 

- 

- uuuu 

- 2 2 

- 4 2 

2 

1 _ _ 

1 

2 1 2 

U 

U 

1 

2 

1 

1 

1 

U 


U 

1 

2 

2 

1 

1 

2 2 

u 1 2 1 

U 2 

u 1 1 

1-2 

U 

1 

1 

1 

5 



Character states of characters not included in the comparison: 10,1; 13,1; 

14,2; 15,1; 29,2; 30,1; 45,2; 52,1 53,1; 56,-; 64,3; 69;1; 70,2; 71,2; 72,2; 73,2; 

76,1; 77,2; 78,1; 81,1. 

Species listed: 72 E. fucala; 110 E. macrobranchia; 115 E. marenzelleri; 178 

E. sebasliani; 181 E. sonorae; 196 E. unidentata. 

_ 

_ 

2 

2 

1 

2 

1 

1 

196 

2 

U 

uuu 

u 

2 

2 

2 

2 

1 

1 

1 

U


TABLE 52.—Comparison of Eunice species with various unusual kinds of setae present. Comparisons should be 

made only between members of the same section of the table. The column numbers are the same as the character 

numbers used to prepare the key (for a full listing see Appendix A). 

Species 

1 

2 3 5 35 

36 

80 

5 

151 

E. afuerensis 

E. pelamidis 

687 

113 

PSEUDOCOMPOUND HOOKS PRESENT 

720 

100 

12 

12 

19 

20 

3 

3 

686 

220 

100 

63 

89 

194 

195 

E. impexa 

E. lubicola 

E. tubifex 

U 

83 

U 

COMPOUND SPINIGERS PRESENT 

U 

26.5 

U 

5.5 

1.2 

2 

11 

4 

7 

22 

22/24 

17 

U 

69 

U 

60 

21 

29/32 

40 

59 

65 

168 

E. cirrobranchiata 

E. elseyi 

E.flaccida 

E. rosaurae 

SUBAC1CULAR HOOKS WITH TEETH IN TANDEM PRESENT 

U 

120 

U 

64 

3 

7 

6 

13 

22 

6 

U 

104 

U 2.5 8.5 7 U 

u 

142 152 4 18 3 48-55 

17/24 

30 

27 

32-38 



1 = Total number of seu'gers (complete specimens only). 



5 • Length through setiger 10 in mm. 




TABLE 53.—Comparison of Eunice species with various unusual setae. Comparisons should be made only 

between members of the same section of the table. Stub (C) gives character number, column heads give species 

number. Characters and character sutes are listed in Appendix A. Species are in order by number, corresponding 


c 

Pseudocompounds 

5 

151 

89 

Spinigers 

194 195 

40 

59 

Tandems 

65 

168 

7 

8 

9 

11 

12 

14 

15 

16 

18 

20 

21 

22 

23 

24 

25 

26 

28 

29 

31 

32 

33 

34 

35 

36 

3 

1 

3 

2 

1 

U 

3 

1 

3 

1 

1 

2 

4 

1 

3 

3 

3 

2 

1 

1 

2 

1 

3 

686 

2 

2 

2 

U 

U 

2 

U 

U 

3 

1 

1 

3 

1 

U 

3 

3 

1 

1 

1 

1 

2 

1 

3 

220 


3 

2 

2 

2 

1 

2 

3 

2 

1 

3 

3 

2 

2 

2 

1 

2 

2 

22 

U 

3 

2 

2 

2 

1 

U 

1 

1 

3 

3 

2 

- 

4 

2 

3 

4 

2 

2 

3 

2 

- 

- 

22/24 

69 

3 

3 

3 

1 

2 

2 

1 

3 

3 

3 

2 

- 

1 

1 

1 

4 

4 

2 

1 

1 

2 

2 

17 

U 

3 

3 

2 

2 

2 

2 

1 

1 

3 

3 

1 

1 

4 

1 

1 

9 

1 

2 

2 

1 

2 

- 

22 

U 

3 

1 

3 

2 

1 

U 

1 

1 

2 

1 

1 

3 

2 

2 

3 

4 

1 

1 

1 

1 

2 

1 

6 

104 

3 

1 

2 

2 

2 

U 

1 

1 

3 

1 

1 

3 

2 

1 

2 

5 

2 

1 

1 

3 

U 

1 

7 

U 

2 

2 

3 

2 

2 

3 

1 

1 

3 

2 

1 

2 

1 

2 

1 

3 

3 

2 

3 

3 

2 

- 

3 

48-55



c 

Pseudocompounds 

5 

151 

89 

Spinigcrs 

194 195 40 

59 

Tandems 

65 

168 

37 

38 

39 

40 

41 

42 

43 

44 

45 

48 

51 

52 

53 

54 

55 

56 

57 

58 

59 

60 

61 

64 

1 

1 

0 

0 

25 

U 

2 

2 

2 

2 

3 

1 

1 

2 

2 

- 

1 

2 

2 

- 

2 

2 

1 

1 

0 

0 

24 

20 

3 

2 

2 

2 

9 

1 

1 

1 

2 

_ 

2 

- 

- 

1 

1 

1 

SOFT BODY CHARACTER STATES—Continued. 

1 

1 

10 

u 

7 

U 

2 

3 

3 

2 

4 

1 

1 

2 

1 

- 

1 

2 

2 

_ 

1 

3 

2 

1 

- 

- 

- 

- 

3 

2 

2 

2 

5 

1 

1 

2 

1 

- 

1 

2 

1 

- 

2 

3 

1 

1 

U 

U 

4 

30 

1 

3 

2 

3 

5 

1 

1 

2 

1 

- 

1 

2 

1 

_ 

2 

3 

1 

1 

U 

u 

2 

U 

14 

2 

2 

2 

4 

3 

1 

2 

1 

- 

1 

2 

1 

- 

2 

3 

1 

1 

0 

0 

18 

U 

1 

2 

2 

2 

5 

1 

1 

1 

1 

- 

1 

- 

- 

2 

1 

1 

U 

u 0 

u 

9 

12 

2 

3 

2 

2 

- 

1 

2 

- 

- 

2 

U 

U 

u 

2 

2 

2 

2 

- 

- 

- 

- 

1-2 

1-2 

1 

1 

5 

3 

1 

1 

2 

- 

1 

2 

^ 

- 

l 

3 

65 

66 

69 

71 

72 

73 

74 

76 

77 

78 

79 

80 

81 

82 

1 

1 

1 

1 

2 

1 

2 

1 

1 

2 

2 

100 

1 

1 

1 

1 

1 

1 

2 

2 

U 

1 

2 

U 

3 

63 

2 

1 


1 

1 

1 

2 

1 

2 

1 

1 

2 

1 

2-3 

60 

1 

5 

2 

1 

1 

2 

1 

1 

U 

1 

2 

U 

2 

21 

1 

1 

1 

1 

1 

2 

1 

2 

U 

2 

2 

1 

2 

29/32 

1 

1 

1 

2 

1 

2 

2 

2 

1 

4 

2 

1 

4 

17/24 

1 

1 



Character states of characters not included in the comparisons: 10,1; 13,1; 17,1; 19 2- 271- 30 1- 56 -• 70 2- 

75.1. 

Species listed: 

Pseudocompound hooks present: 5 E. afuerensis; 151 E. pelamidis. 

Compound spinigers present: 89 E. impexa; 194 E. lubicola; 195 E. tubifex. 

Subacicular hooks with teeth in tandem present: 40 E. cirrobranchiata; 59 E. elseyi; 65 E.flaccida; 168 E. 

rosaurae. 

2 

2 

2 

2 

2 

1 

2 

5 

1 

2 

4 

30 

1 

1 

1 

1 

2 

2 

1 

2 

1 

1 

2 

4 

27 

1 

1 

2 

1 

1 

2 

2 

2 

1 

1 

2 

1 

2/4 

32-38 

1 

1

Appendix A 

Character List for Eunice 

1. number of sctigers 

2. total length (mm) 

3. maximum width (mm) 

4. first reached at at setiger 

5. length through setiger 10 (mm) 

6. width at setiger 10 (mm) 

7. prostomial lobes 

1. frontal ly truncate 

2. frontally obliquely truncate 

3. frontally rounded 

8. prostomial lobes 

1. dorsally inflated 

2. dorsally flattened 

3. dorsally excavate with thickened rim 

9. median sulcus 

1. median sulcus invisible dorsally 

2. median sulcus shallow 

3. median sulcus deep 


1. distinctly shorter than peristomium 

2. about as long as peristomium 


1. about as wide as peristomium 

2. distinctly narrower than peristomium 


1. less than 1 /i as deep as peristomium 

2. at least as deep as l li of peristomium 

13. eyes 

1. present 

2. absent 

14. eyes 

1. lateral to the bases of A-I 

2. behind bases of A-I 

3. between bases of A-I and A-I I 

4. behind bases of A-I I 

5. on ceratophores of A-I 

6. on ceratophores of A-I I 

15. antennae in a 

1. horseshoe 

2. semicircle 

3. transverse row 

16. antennae 

1. evenly spaced 

2. with A-I isolated by a gap 

3. with A-I 11 isolated by a gap 

17. antennae 

1. similar in thickness 

2. with A-I slimmer than other 3 

3. with A-I 11 slimmer than other 4 

4. with A-I thicker than other 3 

5. with A-I I thicker than other 3 

6. with A-III thicker than other 4 


1. long in all antennae 

2. long in A-I and ring-shaped in other 3 

3. ring-shaped in all antennae 





1. tapering 


3. digitiform 

4. club-shaped 




22. ceratostyle articulations 

1. long or short cylinders 

2. moniliform or drop-shaped distally 

3. moniliform 

23. length of 

1. A-II and III similar and A-I shorter 

2. antennae increasing from A-I to A-III 

3. A-II greater than A-I and A-III 

4. all antennae similar (short) 

24. peristomium 

1. with distinct, muscular lower lip 

2. cylindrical 

25. separation between peristomial rings 

1. visible dorsally only 

2. visible ventrally only 

3. visible both dorsally and ventrally 

4. present on all sides 

5. absent 

26. anterior ring makes up 



3. 3 A of total peristomial length 

4. 4 /s of total peristomial length 

5. s /6 of total peristomial length 




9. 9 /io of total peristomial length 

27. peristomial cirri reach 

1. middle or anterior end of peristomium 

2. middle or front of prostomium 

3. beyond prostomium 


1. tapering 

2. digitiform 



375


5. ovate 



2. without articulations 50. 

30. branchiae 

1. present 

2. absent 

31. branchiae 51. 

1. pectinate 52. 

2. palmate 

3. single filaments 

32. branchiae 

1. distinctly longer than notopodial cirri 53. 

2. about as long as notopodial cirri 

3. distinctly shorter than notopodial cirri 


1. reduced in mid-body region 

2. not reduced in mid-body region 

34. branchial stems 

1. erect 55. 

2. flexible 

35. branchiae from setiger 

to setiger 

36. to setiger 


1. present to near posterior end 

2. terminating well before posterior end 

38. branchiae present on 57. 

1. more than 65% of total number of setigers 

2. less than 55% of total number of setigers 

39. number of anterior setigers with single branchial filaments 58. 

40. number of posterior setigers with single branchial filaments 

41. maximum number of filaments 

42. maximum number of filaments first reached in setiger number 

43. median acicular lobes 

1. distally truncate 59. 

2. distally rounded 

3. triangular or conical 

4. withdrawn into body wall 

5. bilobed 

44. median acicular lobes with 60. 

1. acicula emerging ventral to midline 

2. acicula emerging at midline 

3. acicula emerging dorsal to midline 

45. anterior presetal lobes 

1. follow outline of acicular lobes closely 61. 

2. form low transverse folds 


46. median presetal lobes 



3. project as free lobes 62. 

47. posterior presetal lobes 




48. anterior postsetal lobes 

1. follow outline of acicular lobes closely 63. 



49. median postsetal lobes 




posterior postsetal lobes 




number of anterior setigers with ventral cirri without inflatioi 

anterior ventral cirri. 

1. tapering from narrow bases 

2. tapering from wide, triangular bases 

3. digitiform 

median ventral cirri 


2. without basal inflations 

bases of median inflated ventral cirri 

1. inflated bases ovate or spherical 

2. inflated bases thick, transverse wells 

3. inflated bases scoop-shaped 

median inflated ventral cirri 





median ventral cirri 

1. tapering 

2. digitiform 

posterior ventral cirri 


2. without basal inflation 

bases of posterior inflated ventral cirri 

1. inflated bases ovate or spherical 

2. inflated bases thick, transverse welts 

3. inflated bases triangular welts 

4. inflated bases scoop-shaped 

posterior inflated ventral cirri 





posterior ventral cirri 

1. tapering 

2. digitiform 

3. short, nearly tubercular 

4. broadly triangular, nearly scoop-shaped 

anterior notopodial cirri 



3. tapering 

4. digitiform 

5. clavate 

median notopodial cirri 



3. tapering 

4. digitiform 

5. clavate 

posterior notopodial cirri 



3. tapering


4. digitiform 

5. clavate 

64. notopodial cirri 

1. articulated throughout body 

2. articulated in anterior sctigers 



1. distally furled 

2. distally flat 


1. tapering 

2. flaring 


1. distally furled 

2. distally flat 


1. tapering 

2. flaring 

69. appendages of compound falcigers distally 

1. bidentate 

2. tridentate 

70. hoods of compound falcigers distally 

1. mucronate 

2. without mucros 

71. pseudocompound falcigers 

1. present 

2. absent 

72. compound spinigers 

1. present 

2. absent 

73. aciculae 

1. light yellow or translucent 


74. separation of acicular cores and sheaths 

1. distinct 

2. indistinct 

75. cross-section of aciculae 

1. round 

2. flattened or knife-edged 

76. aciculae distally 

1. pointed (sharp or blunt) 

2. flattened with rounded tabs 

3. expanded, knurled and knobbed 

4. hammer-headed 

5. bifid (bidentate) 

77. subacicular hooks 

1. light yellow or translucent 


78. separation of cores and sheaths 

1. distinct 

2. indistinct 

79. hooks 

1. tapering to simple falcate tips 

2. bidentate 

3. tridentate with teeth in a crest 

4. tridentate with teeth in tandem 

80. hooks first present from setiger in 

81. hooks 

1. present in all setigers posteriad 

2. missing in a some setigers 

3. missing in many setigers 

82. hooks 

1. always single (except for replacements) 

2. paired in some setigers 

3. paired in most setigers 

4. 3 or more in some setigers 

5. 3 or more in most setigers

Appendix B 

Characters and character states for each species of Eunice as set up for use with the DELTA 

program. (Numbers in column 1 (1.01,1.02, etc.) are line-numbers for each character—the 

digit(s) preceding the decimal is also the species number used in ccnterheads and 

comparison tables; paired numbers = character, state for all 82 characters using the 

character list in Appendix A (dash = character inappropriate, U = feature unknown); 

numbers separated by a dash indicate a range, by a slash, paired alternate observations.) 

1.01 aciculata 

1.02 1,202 2,129 3,4.5 4,15 5,11 6,4 7,2 8,2 9,3 10,1 11,2 

1.03 12,2 13,1 14,2 15,1 16,1 17,1 18,1 19,2 20,3 21,2 22- 

1.04 23.U 24,2 25,3 26,3 27,1 28,5 29,2 30,1 31,1 32,1 33.2 

1.05 34,1 35,18 36,190 37,1 38,1 39,10 40,10 41,5 42,32 43,2 

1.06 44,3 45,2 46,2 47,2 48,3 49,1 50,1 51,15-20 52,2 53,1 

1.07 54,1 55,3 56- 57,2 58- 59- 60,2 61,4 62.4 63,4 64.3 

1.08 65,2 66,2 67,2 68,2 69,1 70,2 71,2 72,2 73.2 74.2 75.1 

1.09 76,1 77,2 78,2 79,2 80,22 81,1 82,1 

2.01 aedificatrix 

2.02 1,270 2,128 3,4 4,10 5,7 6,4 7,3 8,1 9,3 10,1 11,2 12,1 

2.03 13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,3 23,2 

2.04 24,1 25,4 26,2 27,2 28,4 29,1 30,1 31,1 32,1 33,2 34,1 

2.05 35,4 36,250 37,1 38,1 39,10 40,10 41,11 42,15 43,1-2 44,2 

2.06 45,2 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 

2.07 56- 57,2 58- 59- 60,1 61,1 62,1 63,4 64,3 65,1 66,1 

2.08 67,1 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1/4 77,1 

2.09 78,2 79,3 80,35 81,1 82,1 

3.01 aequabilis 

3.02 1,158 2,175 3,7 4,15 5,16 6,6 7,2 8,2 9,2 10,1 11,2 12,2 

3.03 13,2 14- 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

3.04 24,2 25,3 26,5 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

3.05 35,6 36,151 37,1 38,1 39,0 40,0 41,17 42,20 43,1 44,3 45,2 

3.06 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 56- 

3.07 57,2 58- 59- 60,1 61,1 62,1 63,1 64,3 65,2 66,1 67,2 

3.08 68,1 69,2 70,2 71,2 72,2 73,1 74,2 75,1 76,3/5 77,1 78,2 

3.09 79,3 80,26/32 81,1 82,1 

4.01 afra 

4.02 1,U 2,U 3,6 4,U 5,10 6,U 7,3 8,1 9,3 10,1 11,1 12,2 13,1 

4.03 14,2 15,1 16,1 17,3 18,3 19,2 20,3 21,1 22,1 23,1 24,1 

4.04 25,3 26,3 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 35,16 

4.05 36,U 37,1 38,1 39,0 40,U 41,6 42,30 43,2 44,2 45,2 46,2 

4.06 47,2 48,2 49,2 50,2 51,5 52,2 53,1 54,3 55,2 56- 57,2 

4.07 58- 59- 60,1 61,1 62,2 63,4 64,3 65,1-2 66,2 67,2 68,2 

4.08 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78.U 79 2 

4.09 80,22 81,1 82,1 

5.01 afuerensis 

5.02 1,687 2,720 3,12 4,15 5,19 6,10 7,3 8,1 9,3 10,1 11,2 12,1 

378


5.03 13.U 14,U 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,4 

5.04 24,1 25,3 26,3 27,1 28,3 29,2 30,1 31,1 32,1 33,2 34,1 

5.05 35,3 36,686 37,1 38,1 39,0 40,0 41,25 42,U 43,2 44,2 45,2 

5.06 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,2 55,2 56- 

5.07 57,1 58,2 59,2 60- 61,2 62,2 63,2 64,2 65,1 66,1 67,1 

5.08 68,2 69,1 70,2 71,1 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

5.09 79,2 80,100 81,1 82,1 

6.01 americana 

6.02 1,123 2,86 3,2 4,10 5,9 6,2 7,1 8,1 9,2 10,1 11,2 12,2 

6.03 13,1 14,2 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,1 

6.04 24,2 25,3 26,2 27,2 28,3 29,2 30,1 31,1 32,1 33,2 34,1 

6.05 35,3 36,35 37,2 38,2 39,0 40,2 41,20 42,10 43,1 44,2 45,2 

6.06 46,2 47,2 48,2 49,2 50,2 51,5 52,3 53,1 54,1 55,3 56- 

6.07 57,2 58- 59- 60,2 61,2 62,2 63,4 64,1 65,2 66,2 67,2 

6.08 68,2 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

6.09 79,3 80,25 81,1 82,1 

7.01 amoureuxi 

7.02 l.U 2,13 3.U 4,U 5,U 6,U V 13,1 14.U 15.U 16,1 17,1 18,3 

7.03 19,2 20,3 21,2 22- 23,1 24,U 25.U 26,U 27,1 28.U 29,2 

7.04 30,1 31,3 32,1 33,2 34,U 35,9 36,24 37,2 38,U 39,U 40,U 

7.05 4l.U 42.U 43.U 44,U 45,U 46,U 47.U 48.U 49,U 50.U 51,U 

7.06 52.U 53.U 54.U 55,U 56.U 57.U 58,U 59.U 60,U 61,U 62.U 

7.07 63.U 64,U 65,U 66,U 67.U 68,U 69,1 70,1 71,2 72,2 73,2 

7.08 74.U 75,U 76,1 77,2 78.U 79,2 80,17 81,U 82.U 

8.01 amphiheliae 

8.02 1,140-150 2,180-200 3,7-10 4,U 5,U 6,U 7,1 8,2 9,3 10,1 

8.03 11,2 12,U 13,2 14- 15,3 16,U 17,1 18,3 19,2 20,1 21,1 

8.04 22,1 23,4 24,2 25,4 26,5 27,1 28,1 29,2 30,1 31,1 32,1 

8.05 33,2 34,1 35,4 36,140-150 37,1 38,1 39.U 40,U 41,7 42.U 

8.06 43,2-3 44,2 45,2 46,2 47,2 48,2 49,2 50,2 52,1 53.U 54.U 

8.07 55,U 56.U 57,U 58.U 59,U 60.U 61,3 62,3 63,3 64,3 65,U 

8.08 66,U 67.U 68.U 69,1 70,1 71,2 72,2 73,2 74,U 75,U 76,1 

8.09 77,2 78,U 79,2 80.U 81,U 82,U 

9.01 pruvoti 

9.02 1,115 2,34 3,3 4,U 5,5 6,U 7,1 8,2 9,3 10,1 11,2 12,1 13,1 

9.03 14,3 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,4 24,1 

9.04 25,3 26,4 27,1 28,1 29,2 30,1 31,3 32.U 33.U 34- 35,15 

9.05 36,115 37,1 38,1 39- 40- 41- 42- 43,2 44,2 45,2 46,2 

9.06 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 55,2 56,- 57,2 

9.07 58- 59- 60,1 61,1 62,4 63,4 64,3 65,2 66,2 67,2 68,2 

9.08 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,2 78,2 79,2 

9.09 80,19 81,1 82,1 

10.01 annulicornis 

10.02 1,145 2,135 3,4 4,U 5,12.5 6,U 7,3 8,1 9,3 10,1 11,2 12,2 

10.03 13,U 14.U 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,3 23,2 

10.04 24,2 25,1 26,3 27,2 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

10.05 35,3 36,134 37,1 38,1 39,0 40,2-3 41,9 42,15 43,1 44,3 

10.06 45,2 46,2 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,1 553 

10.07 56- 57,2 58- 59- 60,2 61,1 62,1 63,1 64,1 65,2 66,2 

10.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

10.09 78.U 79,2 80,35 81,1 82,1


11.01 antarctica 

11.02 1,122 2,102 3,3.5 4,U 5,11 6,U 

11.03 13,U 14.U 15,1 16,1 17,1 18,3 

11.04 24,2 25,3 26,4 27,2 28,2 29,2 

11.05 35,3 36,39-44 37,2 38,2 39,6-7 

11.06 45.1 46.1 47,1 48,1 49,1 50,1 

11.07 56- 57,2 58,- 59- 60.2 61,2 

11.08 67,2 68,1 69,1 70,2 71,2 72,2 

11.09 78,2 79,2 80,31^4 81,1 82,1 

7,3 

19.2 

30.1 

40.2 

51,3 

62.3 

73.1 

8.1 

20.3 

31,1 

41,5 

52.3 

63.4 

74,2 

9.2 110.1 1 

21.1 

32,3 

22.1 

33.2 

42,15 43. 

53,1 

64.1 

75.1 

54.1 

65.2 

76.1 

1.1 12,1 

23,2 

34,1 

I 44.2 

55,3 

66,1 

77.1 

12.01 antennata 

12.02 1,98 2.54 3,3 4.U 5,7 6.U 7,3 8.2 9,3 10.1 11.2 12,2 13.1 

12.03 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21.1 22.3 23.2 24.1 

12.04 25,3 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,1 34.1 35.7 

12.05 36,92 37,1 38,1 39,0 40,3 41,7 42.U 43,1 44,2 45.2 46.2 

12.06 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54.1 55.3 56.- 57.2 

12.07 58- 59- 60,2 61,1 62,1 63,1 64.1 65.2 66.2 67.2 68.2 

12.08 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76.1 77.1 78.2 79.3 

12.09 80,25 81,1 82,1 

13.01 antillensis 

13.02 1,U 2.U 3,3 4,U 5,7 6.U 7.1 8,2 9,2 10,1 11.2 12.2 13.1 

13.03 14,1 15,1 16,3 17,1 18,3 19.2 20,1 21,1 22,1 23,2 24,2 

13.04 25,3 26,4 27,3 28,2 29,1 30.1 31,1 32,3 33,2 34,1 35,4 

13.05 36,36 37,2 38,2 39,2 40,2 41,6 42.U 43,1 44,3 45,2 46,2 

13.06 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 56- 57,2 

13.07 58- 59- 60,1 61,4 62,4 63,4 64,1 65,2 66,1 67,U 68.U 

13.08 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,2 77,1 78,2 

13.09 79,2 80,33 81,1 82,2 

14.01 aphroditois 

14.02 1,U 2.U 3,22 4,25 5,55 6,U 7,3 8,1 9,2 10,1 11,2 12,1 

14.03 13.U 14.U 15,1 16,1 17,1 18,3 19,2 20,3 21,2 22- 23,4 

14.04 24,2 25,3 26,4 27,1 28,4 29,2 30,1 31,1 32,1 33,2 34,1 

14.05 35,6 36.U 37,1 38,1 39,0 40,U 41,30 42,30 43,1-2 44,2 

14.06 45,2 46,2 47,2 48,1 49,1 50,3 51,250 52,2 53,1 54,3 55,2 

14.07 56- 57,1 58,2 59,2 60- 61,1 62,1 63,1 64,3 65,1 66,1 

14.08 67,1 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

14.09 78,1 79,2 80,200 81,3 82,1 

15.01 arcturi 

15.02 1,110 2,80 3,U 4,U 5,U 6,U 7,U 8,U 9,U 10,U ll.U 12.U 

15.03 13.U 14.U 15,U 16,U 17.U 18.U 19.U 20.U 21,U 22.U 23.U 

15.04 24,U 25.U 26.U 27,U 28,U 29,U 30,1 31.U 32,U 33.U 34,U 

15.05 35,6 36,42 37,2 38,2 39.U 40.U 41,10 42.U 43.U 44.U 45.U 

15.06 46.U 47,U 48,U 49.U 50,U 51,U 52,U 53,U 54,U 55,U 56,U 

15.07 57,U 58,U 59,U 60,U 61,U 62.U 63.U 64,U 65.U 66.U 67,U 

15.08 68.U 69.U 70,U 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

15.09 79,2 80,36 81,U 82,U 

16.01 arenosa 

16.02 1,79 2,16 3,.75 4,10 5,2.5 6,.75 7,3 8,2 9,2 10,2 11,1 

16.03 12,2 13.U 14,U 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,1 

16.04 23,2 24,2 25,4 26,1 27,2 28,1 29,1 30,1 31,1 32,1 33,2 

16.05 34,1 35,3 36,25/26 37,2 38,2 39,5 40,2-3 41,5 42,15 43,1 

16.06 44,2 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1

NUMBER 523 

381 

16.07 

16.08 

16.09 

17.01 

17.02 

17.03 

17.04 

17.05 

17.06 

17.07 

17.08 

17.09 

18.01 

18.02 

18.03 

18.04 

18.05 

18.06 

18.07 

18.08 

18.09 

19.01 

19.02 

19.03 

19.04 

19.05 

19.06 

19.07 

19.08 

19.09 

20.01 

20.02 

20.03 

20.04 

20.05 

20.06 

20.07 

20.08 

20.09 

21.01 

21.02 

21.03 

21.04 

21.05 

21.06 

21.07 

21.08 

21.09 

22.01 

22.02 

22.03 

55,2 56- 57,2 58- 59- 60,2 61,3 62,3 63,4 64,3 65.U 

66.U 67.U 68.U 69,1 70,1 

77,1 78,1 79,3 80,16 81,1 

71,2 72,2 73,1 74,2 75,1 76,1 

82,1 

argentinensis 

l.U 2,U 3,5 4,10 5,12 6,5 7,3 8,2 9,3 10,1 11,2 12,2 13,2 

14- 15,3 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,2 24,2 

25.3 26,3 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 353 

36.U 37,1 38,1 39,0 40.U 41,10 42,10 43,1 44,3 45,2 46,2 

47,2 48,2 49,2 50,2 51,3 52,3 53,1 54,1 55,2 56- 57.U 

58.U 59.U 60.U 61,2 62,2 63.U 64,1 65,2 66,1 672 68,1 

69.1 70,2 71,2 72,2 73,2 74,U 75,2 76,1 77,2 78.U 792 

80,19-22 81,1 82,1 

armillata 

1,114 2,46 3,2 4,15 5,6 6,1.75 7,3 8,1 9,3 10,1 11,2 12,1 

13.U 14.U 15,3 16,1 17,1 18,1 19,2 20,1 21,1 22,3 23,2 

24^ 25,4 26,4 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 

35,6 36,109 37,1 38,1 39,1 40,U 41,3 42,U 43,1 44,3 452 

462 472 48,3 49,2 50,2 51,5 52,3 53,1 54,1 55,2 56- 

572 58- 59- 602 61,4 62,4 63,4 64,2 652 66,2 672 

68.2 69,1 70,2 712 72,2 73,1 74,2 75,1 76,1 77,1 782 

792 80,34 81,1 82,1 

articulata 

l.U 2,U 3,3 4.U 5,U 6,U 7,1 8,2 9,3 10,1 11,2 12,1 13,1 

14,1 15,3 16,1 17,6 18,3 192 20,1 21,1 222 23,2 242 

25,1 26,3 27,3 282 29,1 30,1 31,1 32,1 332 34,1 35,4 

36,55 37,2 382 39,1 402-3 41,6 42,15 43.U 44.U 45,U 

46,U 47.U 48.U 49,U 50.U 5l.U 52.U 53,1 54.U 55,U 56- 

572 58- 59- 602 61,4 623 633 64,2 652 66,1 672 

68,1 69,1 70,2 712 72,2 73,1 74,2 75,1 76,1 77,1 782 

792 80,40 81,1 82,1 

atlantica 

l.U 2,U 3,2 4,U 5,U 6,U 7,3 82 92 10,1 ll.U 12,U 13.U 

14,U 15.U 16.U 17,U 18,3 19,2 20,1 21.1 22,1 232 24,2 

25.4 26,2 27,2 28,1 29,1 30,1 31,1 32,U 332 34.U 35,3 

36,U 37,2 38,2 39,1 40.U 41,5 42.U 43.U 44,U 45.U 46,U 

47.U 48.U 49.U 50,U 51,U 52,U 53,1 54,U 55,U 56.U 57,2 

58,U 59,U 60,1 61,U 62.U 63,U 64,U 65,U 66.U 67,U 68,U 

69,1 70,1 71,2 722 73,1 742 75,1 76,1 77,1 78,2 79,3 

80.U 81.U 82.U 

aucklandica 

1,79 2,18 3,1 4,10 5,3.2 

13,1 

23,2 

14,2 15,1 16,1 17,1 

24,2 25,4 26,3 27,1 

34- 35,8 36,21 372 38,2 

45,2 46,2 47,2 48,2 49,2 

56- 57,2 58- 59,- 60,1 

672 68,1 69,1-2 70,2 712 

78,2 79,3 80,18 81,1 82,3 

australis 

1,109 2,70 

13,U 14,U 

6,1 7,3 8,2 9,3 10,1 112 12,2 

18,3 19,2 20,3 21,1 22,1 

28.1 29,2 30,1 31,2 32,1 33,2 

39,U 40.U 41,2 42,U 43,2-3 442 

50.2 51,5 52,1 53,1 54,1 55,2 

61,1 62,1 63,1 64,3 65,2 66,1 

72,2 73,1 74,2 75,1 76,1 77,1 

3,4 4,15 5,9 6,8.5 7,3 8,2 9,3 10,1 11,1 12,2 

15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,2

382 


22.04 

22.05 

22.06 

22.07 

22.08 

22.09 

23.01 

23.02 

23.03 

23.04 

23.05 

23.06 

23.07 

23.08 

23.09 

24.01 

24.02 

24.03 

24.04 

24.05 

24.06 

24.07 

24.08 

24.09 

25.01 

25.02 

25.03 

25.04 

25.05 

25.06 

25.07 

25.08 

25.09 

26.01 

26.02 

26.03 

26.04 

26.05 

26.06 

26.07 

26.08 

26.09 

27.01 

27.02 

27.03 

27.04 

27.05 

27.06 

27.07 

27.08 

27.09 

24,2 25,3 26,3 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

35,7 36,33 37,2 38,2 39,0 40,2 41,8 42,10 43,1-2 44,2 

45,2 46,2 47,2 48,2 49,2 50,2 51,6 52,3 53,1 54,1 55,2 

56- 57,2 58,- 59- 60,2 61,4 62,1 63,3 64,2 65,2 66.2 

67,2 68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

78,2 79,3 80,31 81,1 82,1 

barvicensis 

1,U 2,U 3,3.5 4,10 5,6 6,3.5 7,3 8,2 9,3 10,1 11,1 12,1 

13.1 14,3 15,1 16,1 17,1 18,1 19,2 20,1 21,2 22- 23,2 

24.2 25,3 26,3 27,1 28,1 29,2 30,1 31,1 32,2 33.2 34,1 

35,6 36,65 37,2 38,2 39,3 40,10 41,4 42,25 43,2-3 44,2 

45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,3 53,1 54.1 55.3 

56- 57,2 58- 59- 60.2 61,4 62,4 63,4 64,3 65.2 66,2 

67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76.1 77,2 

78.1 79,2 80,19 81,1 82,2 

bassensis 

1.U 2,U 3.U 4,U 5,U 6,U 7.U 8,U 9,U 10.U ll.U 12.U 13.U 

14.U 15.U 16.U 17.U 18.U 19.U 20.U 21.U 22.U 23.U 

24.U 25,U 26.U 27.U 28.U 29.U 30.1 31.1 32.1 33.1 34.1 

35.U 36.U 37,1 39.U 40,0 41.9 42.U 43.1 44,2 45.U 46.U 

47.2 48.U 49.U 50,2 51.U 52.U 53.U 54,U 55.U 56.U 57.2 

58- 59- 60,2 61,U 62.U 63,3 64,1 65.U 66.U 67.U 68.U 

69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 79.3 

80.U 81,U 82.U 

benedicti 

1,105 2,38 3,2.5 4,15 5,5 6,2 7,3 8,2 92 10.1 11,2 12,1 

13,1 14,2 15.1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,2 

24,2 25,4 26.2 27,2 28,2 29,2 30.1 31,1 32,1 33,2 34,2 

35,3 36,39 37,2 38,2 39,3 40,0 41,12 42.15 43,3^4 44.3 

45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,2 55.3 

56- 57,1 58,2 59,2 60- 61,2 62.2 63,4 64,3 65,2 66,1 

67,2 68,1 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76.1 77.1 

78,2 79,2 80,21 81,1 82,2 

biannulata 

1,139 2,105 3,3 4,15 5,12 6,2.75 7,1 8,2 9,3 10,1 11,1 

12,2 13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 

23,2 24,1 25,4 26,3 27,1 28,4 29,1 30,1 31,1 32,1 33,2 

34,1 35,3 36,50 37,2 38,2 39,1 40,2-3 41,8 42,10 43.1-2 

44,3 45,2 46,2 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,1 

55,3 56- 57,2 58- 59- 60,2 61,2 62,2 63,2 64,2 65,1 

66,1 67,1 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 

77,1 78,2 79,2 80,38 81,1 82,2 

bicirrata 

l.U 2,U 3,1 4,U 5,U 

14,U 15,U 16,U 17,U 

25.U 26,U 27,2 28.U 

36,48 37.U 38.U 39,U 

47,U 48,U 49.U 50.U 

58,U 59,U 60.U 61 ,U 

69,1 70,2 71,2 72,2 

80,45 81,U 82,U 

6,U 7,U 8.U 9,U 10,U ll.U 12,U 13.U 

18.U 19,U 20,3 21,1 22,2 23,2 24.U 

29,U 30,1 31,2 32,U 33,U 34.U 35,3 

40.U 41, U 42.U 43.U 44 ,U 45.U 46.U 

51.U 52,U 53,U 54 ,U 55.U 56,U 57.U 

62.U 63.U 64,U 65,U 66.U 67.U 68.U 

73,1 74,U 75,U 76,1 77,1 78,U 79,3


28.01 biformicirrata 

28.02 1,137 2,217 3,7 4,15 5,17 6,6.5 7,3 8,1 9,3 10,1 11,2 

28.03 12,1 13.U 14,U 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 

28.04 23,1 24,2 25,3 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,2 

28.05 34,1 35,3 36,132 37,1 38,1 39,1 40,5 41,7 42,10 43,1-2 

28.06 44,3 45,2 46,2 47,2 48,2 49,2 50,2 51,3 52,3 53,1 54,1 

28.07 55,2 56- 57,2 58- 59- 60,2 61,3 62,3 63,3 64,1 65,2 

28.08 66,2 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 

28.09 77,2 78.U 79,2 80,27 81,1 82,1 

29.01 bilobata 

29.02 1,U 2,U 3,6 4,U 5,7 6,U 7,2 8,2 9,3 10,1 11,2 12,2 13,2 

29.03 14- 15,U 16.U U 18,3 19,2 20,3 21,1 22,1 23.U 24,1 

29.04 25,3 26,9 27,2 28,2 29,2 30,1 31,1 32,3 33,2 34,2 35,7 

29.05 36,U 37,1 38,1 39,0 40.U 41,18 42,U 43,2 44,3 45,2 46,2 

29.06 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 56- 57.U 

29.07 58,U 59.U 60.U 61,1 62,3 63,3 64,1 65,2 66,1 67,2 68,1 

29.08 69,1 70,2 71,2 72,2 73,2 74 ,U 75,1 76,3 77,2 78.U 79,2 

29.09 80,25 81,1 82,1 

30.01 bipapillata 

30.02 1,U 2,U 3,U 4.U 5,U 6.U 7,U 8.U 9,U 10,U 11,U 12,U 13.U 

30.03 14.U 15.U 16,U 17,U 18.U 19.U 20,3 21,1 22,1 23,4 24,U 

30.04 25.U 26,U 27,U 28,U 29,U 30,1 31,1 32,3 33,2 34.U 35,4 

30.05 36.U 37,1 38,1 39.U 40.U 41,4 42,8 43,1 44,3 45,2 46,2 

30.06 47,2 48,2 49,2 50,2 51,U 52,U 53,1 54,1 55,3 56- 57,2 

30.07 58- 59- 60,2 61,1 62,1 63,1 64,3 65,2 66,2 67,2 68,2 

30.08 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78,U 79,2 

30.09 80.U 81,U 82,U 

31.01 borneensis 

31.02 1,120 2,42 3,6 4,50 5,7 6,4.5 7,1 8,2 9,3 10,1 11,2 12,1 

31.03 13.U 14.U 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,1 

31.04 24,2 25,3 26,6 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,2 

31.05 35,6 36,120 37,1 38,1 39,0 40,10 41,8 42,15 43,1 44,3 

31.06 45,2 46,2 47,2 48,3 49,3 50,3 51,4 52,1 53,1 54,2 55,1 

31.07 56- 57,1 58,2 59,1 60- 61,1 62,1 63,1 64,3 65,2 66,2 

31.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

31.09 78.U 79,2 80,30-35 81,1 82,4 

32.01 bottae 

32.02 1,U 2,U 3,2 4,40 5,7 6,1.75 7,3 8,2 9,2 10,1 11,2 12,2 

32.03 13,U 14,U 15,1 16,U 17,1 18,3 19,2 20,3 21,1 22,3 23,U 

32.04 24,1 25,4 26,5 27,1 28,1 29,1 30,1 31,1 32,1 33.U 34,1 

32.05 35,6 36,U 37,1 38,1 39,0 40,U 41,8 42,15 43,1-2 44,3 45,2 

32.06 46,2 47,U 48,2 49,2 50,U 51,U 52,1 53,1 54,1 55,3 56- 

32.07 57,2 58- 59- 60,2 61,1 62,3 63.U 64,1 65.U 66,U 67,U 

32.08 68,U 69,2 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

32.09 79,3 80,25 81,1 82,1 

33.01 bowerbanki 

33.02 1,142 2,95 3,7 4,20 5,11 6,6.5 7,3 8,1 9,3 10,1 11,2 12,1 

33.03 13.U 14,U 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,1 

33.04 24,1 25,3 26,6 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,2 

33.05 35,5 36,136 37,1 38,1 39,0 40,60 41,15 42,12 43,3 44,2 

33.06 45,2 46,2 47,2 48,3 49,2 50,2 51- 52,3 53,2 54- 55-

384 SMITHSONIAN CONTRIBUTIOXS TO ZOOLOGY 

33.07 56,2 57,2 58- 59- 60,2 61,1 62,3 63,4 64,2 65,2 66,1 

33.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

33.09 78.U 79,2 80,31 81,1 82,2 

34.01 brevis 

34.02 1,130 2,18 3,1.5 4,10 5,2.5 6,1.5 7,1 8,2 9,2 10,1 11,2 

34.03 12,2 13,U 14.U 15,3 16,1 17,1 18,3 19,2 20,3 21.1 22,1 

34.04 23,4 24,1 25,1 26,2 27,1 28,2 29,2 30,2 31- 32- 33- 

34.05 34,- 35- 36- 37- 38- 39- 40- 41- 42- 43,2-3 44,2 

34.06 45,2 46,2 47,2 48,2 49,2 50,2 51,7 52,1 53,1 54,2 55.2 

34.07 56- 57,2 58- 59,- 60,1 61,2 62,2 63,2 64,3 65.2 66,2 

34.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74 ,U 75.1 76,1 77,2 

34.09 78.U 79,2 80,27 81,1 82,1 

35.01 bucciensis 

35.02 l.U 2,U 3,3.5 4,15 5,7.5 6,3.25 7,3 8,3 9,3 10.1 11.2 

35.03 12,1 13,U 14,U 15,1 16.1 17,3 18,1 19,2 20,3 21.2 22,- 

35.04 23,4 24,1 25,4 26,2 27,1 28.1 29,2 30,1 31.2 32,1 33,2 

35.05 34,1 35,18 36.U 37,1 38,1 39,3 40.U 41.3 42.U 43.1-2 44,3 

35.06 45,2 46,2 47,2 48.3 49,3 50,3 51.20 52,1 53,1 54,1 55.3 

35.07 56,- 57,2 58- 59,- 60,2 61,1 62,1 63,1 64,3 65,2 66,2 

35.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

35.09 78,1 79,2 80,34 81,1 82,1 

36.01 caeca 

36.02 1,90 2,103 3,6 4,10 5,14 6,6 7,3 8,2 9,3 10,1 11,2 12,1 

36.03 13,1 14,1 15,1 16,2 17,1 18,3 19,2 20,3 21,1 22,1 23.U 

36.04 24,1 25,4 26,3 27,2 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

36.05 35,3 36,48 37,2 38,2 39,1 40,0 41,24 42,15 43,2 44,3 45,2 

36.06 46,2 47,2 48,3 49,2 50,2 51,3 52,1 53,1 54,1 55,2 56- 

36.07 57,2 58- 59,1 60,1 61,4 62,4 63,4 64,2 65,2 66,1 67,2 

36.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

36.09 79,2 80,42 81,1 82,1 

37.01 cariboea 

37.02 1,127 2,24 3,1.2 4,10 5,2.5 6,1.2 7,2 8,2 9,2 10,1 11,2 

37.03 12,2 13,1 14,2 15,1 16,3 17,1 18,3 19,2 20,3 21,2 22- 

37.04 23,4 24,2 25,3 26,3 27,1 28,5 29,2 30,2 31- 32- 33- 

37.05 34- 35- 36- 37- 38- 39- 40- 41- 42- 43,2-3 44,2 

37.06 45,2 46,2 47,2 48,2 49,2 50,2 51,4-5 52,1 53,1 54,2 55,3 

37.07 56- 57,2 58- 59- 60,3 61,3 62,3 63,3 64,3 65,2 66,1 

37.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

37.09 78.U 79,2 80,24-31 81,1 82,1 

38.01 cedroensis 

38.02 1,U 2,U 32 4,10 5,3 62 7,1 8,1 9,2 10,2 11,1 12,2 13,1 

38.03 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,1 24,2 

38.04 25,4 26,2 27,2 28,1 29,2 30,1 31,1 32,1 33,2 34,1 35,3 

38.05 36,27 37,2 38,2 39,2 40,3 41,10 42,10 43,1 44,3 45,2 46,2 

38.06 47.U 48,2 49,2 50,U 51,5 52,1 53,1 54,1 55,2 56- 57.U 

38.07 58,U 59,U 60.U 61,3 62,3 63,U 64,1 65,2 66,1 67,2 68,1 

38.08 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 79,3 

38.09 80,17 81,1 82,1 

39.01 cincta 

39.02 1,U 2,U 3,1.1 4,U 53 6,U 7,U 8,U 9,U 10.U ll.U 12,U 13.U


39.03 

39.04 

39.05 

39.06 

39.07 

39.08 

39.09 

40.01 

40.02 

40.03 

40.04 

40.05 

40.06 

40.07 

40.08 

40.09 

41.01 

41.02 

41.03 

41.04 

41.05 

41.06 

41.07 

41.08 

41.09 

42.01 

42.02 

42.03 

42.04 

42.05 

42.06 

42.07 

42.08 

42.09 

43.01 

43.02 

43.03 

43.04 

43.05 

43.06 

43.07 

43.08 

43.09 

44.01 

44.02 

44.03 

44.04 

44.05 

44.06 

44.07 

44.08 

44.09 

14.U 15.U 16.U 17.U 18,3 

25,4 26,1 27,1 28,5 29,2 

36- 37,- 38- 39- 40- 

47.U 48,3 49,1 50.U 51,U 

58.U 59.U 60.U 61,2 62,4 

69.1 70,2 71,2 72,2 73,2 

80,21 81,1 82,1 

cirrobranchiata 

1,U 2,U 3,3 4,U 5,6 6,U 

14.2 15,1 16,1 17,1 18,3 

25.1 26,9 27,1 

36.U 37,1 38,1 

47.2 48,2 49,2 

58,2 59,1 60- 

69,1 70,2 71,2 

80,17/24 81,1 82,1 

28,1 29,2 

39.U 40.U 

50,2 51,4 

61,2 62,2 

72,2 73,2 

cocanea 

l.U 2,U 3,5 4.U 5,10 6,U 7,3 8,1 

14.U 153 16,1 17,1 18,3 19,2 20,3 

25,3 26,5 27,1 28,2 29,1 30,1 31,1 

36.U 37,2 38,2 39,0 

47,2 48,2 49,2 50,2 

58,4 59,2 60- 61,3 

69,1 70,2 71,2 72,2 

79,2 80,38-39 81,1 

19,2 20,3 21,2 22,- 23,4 24.U 

30,2 31- 32- 33- 34- 35,- 

41,- 42- 43,2 44,1 45,2 46,2 

52,1 53,2 54- 55,- 56,1 57.U 

63.U 64,3 65,2 66,2 67,2 68,2 

74.U 75,1 76,1 77,2 78,U 79,2 

7,3 8,3 9,2 10,1 11,2 12,2 13,1 

19.2 20,3 21,1 22,1 23,4 24,1 

30,1 31,2 32,1 33,2 34- 35,22 

41,2 42.U 43,1/4 44,2 45,2 46,2 

52.3 53,1 54,2 55,1 56- 57,1 

63.U 64,3 65,1 66,2 67,1 68,1 

74,1 75,1 76,4 77,2 78,1 79,4 

9,3 10,1 11,2 12,2 13.U 

21,1 22,1 23,2 24,1 

32,1 33,2 34,1 35,6 

40.U 41,10 42,30 43,1 44,3 45,2 46,2 

51,4-5 52,1 53,1 54,1 55,4 56- 57,1 

62,3 63,3 64,2 65,2 66,2 67,2 68,2 

73,2 74.U 75,1 76,1 77,2 78.U 

82,1 

coccinoides 

1,U 2,U 3,5 4,20 5,8 6,U 7,3 8,1 9,3 10,1 11,1 12,1 13,1 

14,5 15,1 16,1 17,1 18,2 19,2 20,3 21,2 22- 23,2 24,2 

25,3 26,3 27,1 28,3 29,2 30,1 31,1 32,1 33,2 34,1 35,7-8 

36,78 37,2 38,2 39.U 40,U 41,9 42,U 43,2 44,3 45,2 46,2 

47,2 48,3 49,2 50,2 51,4 52,1 53,1 54,2 55,2 56- 57,2 

58- 59- 60,1 61,1 62,3 63,3 64,3 65,2 66,2 67,2 68,2 

69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78,U 79,2 

80,28/30 81,1 82,1 

collaris 

l.U 2,U 3,6 4,10 5,12 6,6 7,3 8,2 9,3 

10,1 

11,2 

12,2 13,1 

14,2 15.U 16,1 17,1 18,3 19,2 20,3 21,1 22,2 23,2 24,1 

25,2 26,3 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 

35,17-18 36.U 37,1 38,1 39,0 40.U 41,12 42,50 43,2 44,2 

45,2 46,2 47,U 48,3 49,2 50.U 51,3-4 52,1 53,1 54,1 55,2 

56- 57,2 58- 59- 60,1 61,1 62,1 63.U 64,3 65,1 66,2 

67,1 68,2 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

78,U 79,2 80,27/33 81,1 82,1 

collini 

l.U 2,U 3,4 4,15 5,7 6,U 7,3 8,1 9,3 10,1 11,2 12,2 13,2 

14- 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,U 24,2 

25,3 26,3 27,1 28,2 29,2 30,1 31,2 32,3 33,2 34,1 35,16 

36,85 37,2 38,2 

47,2 48,2 49,2 

58- 59- 60,2 

69,1 70,2 71,2 

79,2 80,26-27 81,1 82,1 

39,U 40,U 41,2 42,U 43,1-2 44,2 45,2 46,2 

50,2 51,4 52,1 53,1 54,1 55,2 56- 57,2 

61.1 62,1 63,1 64,3 65,2 66,2 67,2 68,2 

72.2 73,2 74,U 75,1 76,1/2 77,2 78,U


45.01 

45.02 

45.03 

45.04 

45.05 

45.06 

45.07 

45.08 

45.09 

46.01 

46.02 

46.03 

46.04 

46.05 

46.06 

46.07 

46.08 

46.09 

47.01 

47.02 

47.03 

47.04 

47.05 

47.06 

47.07 

47.08 

47.09 

48.01 

48.02 

48.03 

48.04 

48.05 

48.06 

48.07 

48.08 

48.09 

49.01 

49.02 

49.03 

49.04 

49.05 

49.06 

49.07 

49.08 

49.09 

50.01 

50.02 

50.03 

50.04 

50.05 

50.06 

complanata 

1,100 2,72 3,U 4,U 5.U 6,U 7,U 8,U 9.U 10.U ll.U 12.U 

13,1 14,3 15.U 16.U 17.U 18.U 19.U 20.U 21,2 22- 23.U 

24.U 25,3 26,5 27,3 28,U 29,2 30,1 31,1 32,1 33,2 34.U 

35,19 36.U 37.U 38.U 39,2 40,U 41,6 42.U 43.U 44.U 45.U 

46,U 47,U 48.U 49,U 50.U 51.U 52,U 53,1 54 ,U 55,2 56- 

57.U 58.U 59.U 60,U 61.U 62.U 63.U 64.U 65.U 66.U 67.U 

68.U 69,1 70,2 712 72,2 73,2 74.U 75,1 76,1 77,2 78.U 

792 8028 81.U 82.U 

congesta 

l.U 2,U 3,4 4,10 5,6 6,4 

14.3 15,1 16,U 17.U 18,3 

25.4 26,2 27.U 28.U 29.U 

36,47 37,2 38,2 

47,2 48,1 49,2 

39,1 

50,1 

58- 59- 60,1 61,3 

69,1 70,1 71,2 722 

80,33 81,1 82,1 

7,3 8,2 9,2 10,1 11,2 12,2 13,1 

19,2 20,3 21,2 22- 23.U 24,2 

30,1 31,1 32,1 33,2 34,1 35,7 

40,0 41,18 42.U 43,3 44,3 45,2 46,2 

51,7-8 52,1 53,1 54,1 55,1 56- 57,2 

62,3 63,3 64,3 65,2 66,2 67,2 6«,2 

73,1 742 75,1 76,1 77,1 78,2 793 

conglomerans 

1,166 2,163 3,3.5 4,10 5,6.5 6,3.5 7,2 8,2 9.3 10,1 11,1 

12,2 13,1 14,2 15,3 16,1 17,1 18,3 19,2 20,3 21,1 22,1 

23,4 24,1 25,1 26,4 27,1 

28,1 29,2 30,1 31,2 32,1 33,2 

34- 35,23-29 36,166 37,1 38,1 39.U 40,U 41,4 42,130 43,3 

44,2 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,2 

55,2 56- 57,2 58- 59- 60,2 61,1 62,4 63,4 64,3 65,1 

66,2 67,1 68,2 69,1 70,2 71,2 72,2 73,2 74 ,U 75,1 76,4-5 

77,2 78,U 79,2 80,22 81,1 82,1 

contingens 

1,U 2,U 3,U 4,U 5,U 6,U 7,U 8,U 9,U 10,U ll.U 12.U 13,U 

14,U 15.U 16.U 17,U 18,3 19,U 20.U 212 22- 23.U 

242 25,U 26,U 272 28,U 29,2 30,1 31,1 33,2 34,1 35,6 

36,U 37,1 38,1 39,1 40,U 41,18-19 42,15 43,1-2 44,2 45,2 

46,2 47,2 48,2 49,2 50,2 51.U 52,2 53,1 54,U 55,2 56- 

57.1 58- 59- 602 61,1 62,1 63,4 64,3 65,U 66,U 67,U 

68.U 69,1 70,2 712 72,2 732 74.U 75,1 76,1 77,2 78.U 

79.2 80,30 81.U 82.U 

crassitentaculata 

l.U 2,U 3,3.5 4,80 5,9.5 6,U 7,3 8,1 9,3 10,1 11,2 12,1 

13.1 14,3 15,1 16,1 17,1 18,1 19,2 20,1 21,2 22,- 23,2 

242 25,3 26,1 27,1 28,1 29,2 30,1 31,2 32,1 33,U 34,U 

35,28 36,U 37,1 38,1 39,U 40,U 41,2 42,U 43,2-3 44,2 45,2 

46.2 472 48,3 49,1 50,2 51,4 52,1/3 53,1 54,1 55,2 56,2 

57,U 58,U 59,U 60,U 61,1 62,1 63 ,U 64,3 65,2 66,2 67,2 

68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 

79,2 80,32 81,1 82,1 

curticirris 

1,55 2,12 3,1 4,10 5,2.25 6,1 7,3 8,2 9,1 10,2 11,1 12,1 

13.1 14,3 15,3 16,3 17,1 18,3 19,2 20,3 21,1 22,3 23,4 

242 25,3 26,3 27,1 28,2 29,2 30,2 31- 32- 33- 34- 

35- 36- 37- 38- 39- 40- 41- 42- 43,1/4 44,2 45,2 

46.2 47,2 48,2 49,2 50,2 51,11 52,1 53,1 54,1 55,2 56-


50.07 

50.08 

50.09 

51.01 

51.02 

51.03 

51.04 

51.05 

51.06 

51.07 

51.08 

51.09 

52.01 

52.02 

52.03 

52.04 

52.05 

52.06 

52.07 

52.08 

52.09 

53.01 

53.02 

53.03 

53.04 

53.05 

53.06 

53.07 

53.08 

53.09 

54.01 

54.02 

54.03 

54.04 

54.05 

54.06 

54.07 

54.08 

54.09 

55.01 

55.02 

55.03 

55.04 

55.05 

55.06 

55.07 

55.08 

55.09 

56.01 

56.02 

56.03 

57,2 

68,1 

79,2 

58- 

69,1 

80,16 

denticulata 

l.U 2,U • 5,3 < 4,10 5,7 6,3 7,3 8,2 9,3 10,1 11,2 12,1 13,1 

14,2 

25,3 

15,3 

26,3 

16,1 

27,1 

17,1 

28,1 

18,1 

29,2 

19,2 

30,1 

20,1 

31,2 

21,2 

32,1 

22- 

33,2 

23,4 

34- 

24,1 

35,27 

36.U 37,1 38,1 39,10-12 40,U * U,3 42.U 43,1 44,3 45,2 

46,2 

57,2 

68,1 

47,2 

58- 

69,1 

48,2 

59- 

70,2 

49,2 

60,2 

71,2 

50,2 

61,4 

72,2 

51,9 

62,4 

73,2 

52,1 

63,4 

74,U 

53,1 

64,3 

75,1 

54,2 

65,1 

76,4 

55,2 

66,1 

77,2 

56- 

67,1 

78.U 

79,2 80,18-19 81,1 82,1 

59- 

70,2 

81, 1 82,1 

60,1 61,4 

71,2 72,2 

62,4 

73,2 

63,4 

74,1 

64,3 

75,1 

65,2 

76,1 

66,1 

77,2 

67,2 

78,1 

dilatata 

l.U 2,U ; 1,10 4,85 5,16 6,5 % I 8,2 9,3 10,1 1U 12,1 

13,1 

24,2 

14,2 

25,1 

15,1 

26,3 

16,3 

27,1 

17,3 

28,2 

18,3 

29,2 

19,2 

30,1 

20,3 

31,1 

21,2 

32,1 

22- 

33,1 

23,2 

34,2 

35,19 

46,2 

36,U 

47,U 

37,1 

48,3 

38,1 

49,2 

39,5 

50,U 

40,U 

51, U 

41,6 

52,2 

42,U 

53,1 

43,2 

54,1 

44,2 

55,2 

45,2 

56- 

57.U 58,U 59.U 60.U 61,2 62,1 63 ,U 64,3 65, 1 66, 1 67,1 

68,1 

79,2 

69,1 

80,28 

70,2 

81, 1 

71,2 

82,1 

72,2 73,2 74.U 75,1 76,1 77,2 78,U 

djiboutiensis 

l.U 2,U ; 

uo 

4,10 5,20 6,10 7,3 8,2 9,3 10,1 11,2 12,2 

13,1 14,1 15,3 16,1 17,1 18,3 19,2 20,3 21,2 22- 23,4 

24,2 25,3 26,8 27,1 28,4 29,2 30,1 31,1 32,1 33,2 34,1 

35,5 36,U 37,1 38,1 39,0 40.U 4u: > 42.U 43, 1-2 44,3 45,: 

46,2 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,1 55,3 56- 

57,2 

68,2 

79,2 

58- 

69,1 

80,81 

59- 

70,2 

81,3 

60,4 

71,2 

82,1 

61,1 

72,2 

62,1 

73,2 

63,1 

74,1 

64,3 

75,1 

65,2 

76,1 

66,2 

77,2 

67,2 

78,1 

dubitata 

l.U 2,U : $.7 ' 4,10 5,16 6,7 7,3 8,2 9,2 10,1 11,2 

14,2 

25,3 

15,1 

26,5 

16,1 

27,2 

17,4 

28,2 

18,3 

29,1 

19,2 

30,1 

20,1 

31,2 

21,1 

32,3 

22,2 

33,2 

23,2 

34- 

36,38-48 37,2 38,2 39,5 40,10-15 41,3 42, ,10 43,1-2 

45,2 46,2 47,2 48,2 49,2 50,2 51,10 52,3 53,1 54,1 

56- 57,2 58- 59- 60,2 61,1 62,3 63,3 64,2 65,2 

67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

78,1 79,2 80,35-38 81,1 82,1 

edwardsi 

1,118 2,70 

12,1 13,1 

23,2 24,2 

34,1 35,3 

44,2 45,2 

55,2 56- 

66,U 67.U 

77,1 78,2 

3,3.5 4,10 5,8 

14,2 15,3 16,3 

25,4 26,2 27,2 

36,34 37,2 38,2 

46,2 47,2 48,2 

57,2 58- 59- 

68.U 69,1 70,2 

79,2 80,28 81,U 

6,3.5 

17,1 

28,2 

39,5 

49,2 

60,1 

71,2 

82,U 

ehlersi 

1.U 2,U ; 5,4 ' 4,10 5,6 6,4 7,2 

14,2 15,1 16,1 17,1 18,3 19,2 

7,3 

18,3 

29,2 

40,2 

50,2 

61,1 

72,2 

8,2 

20,3 

12,2 13, 

24,2 

35,3-5 

44,2 

55,2 

66,1 

77,2 

8,2 < ),2 10,1 11,2 

19,2 

30,1 

20,3 

31,1 

21,1 

32,2 

22,1 

33,2 

41,6 42,10 43,2-3 

51,5 

62,4 

73,1 

52,1 

63,4 

74,2 

53,2 

64,3 

75,1 

54,1 

65,U 

76,1 

9,3 10,1 11,2 12,1 13,1 

21,1 22,1 23,1 24,2

388 


56.04 25,4 26,2-3 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,2 35,14 

56.05 

56.06 

36.U 

47,2 

37,1 

48,2 

38,1 

49,2 

39,1 

50,2 

40.U 

51,4 

41,6 

52,1 

42,U 

53,2 

43,2 

54,2 

44,2 

55,1 

45,2 

56- 

46,2 

57.U 

56.07 58.U 59.U 60.U 61,1 62,4 63,4 64,3 65,1 66,2 67,1 68,2 

56.08 

56.09 

69,1 

80,27 

70,2 

81.U 

71,2 

82.U 

72,2 73,2 74,1 75,1 76,1 77,2 78,1 79,2 

57.01 eimeorum 

57.02 1,U 2,U 3,3.5 4,U 5,7.5 6,U 7,3 8,2 9,3 10,1 11,2 12,1 

57.03 13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,4 

57.04 24,2 25,3 26,5 27,1 28,3 29,2 30,1 31,1 32,1 33,2 34,1 

57.05 35,5 36,U 37,1 38,1 39,U 40.U 41,5 42,20 43,1-3 44,2 45,2 

57.06 46,2 47,2 48,2 49,2 50,2 51.U 52.1 53,1 54.1 55,3 56,- 

57.07 57,1 58,U 59.U 60- 61,2 62,2 63.2 64,3 65.2 66,1 67,2 

57.08 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 78.U 

57.09 79,2 80,38 81.U 82.U 

58.01 

58.02 

58.03 

58.04 

58.05 

58.06 

58.07 

58.08 

58.09 

59.01 

59.02 

59.03 

59.04 

59.05 

59.06 

59.07 

59.08 

59.09 

60.01 

60.02 

60.03 

60.04 

60.05 

60.06 

60.07 

60.08 

60.09 

61.01 

61.02 

61.03 

61.04 

61.05 

61.06 

61.07 

61.08 

61.09 

elegans 

1,133 2,75 

13,1 14,2 

24,2 

35,3 

46,2 

57,1 

68,1 

3,3 4,10 5,8 6,3 7,2 8,2 9,2 

15.1 16,1 17.U 18,1 19,2 20,3 

25,4 26,3 27,2 28,1 29,1 30,1 31,1 

36,33 37,2 38,2 39,1 40,1 

47,2 48,2 49,2 50,2 51,4 

58,4 59.2 60- 61,1 62,1 

69,1 70,2 71,2 72,2 73,2 

79,2 80,30 81,1 82,5 

10,1 11.2 12.2 

21.1 22.1 23.U 

32,1 33.2 34,1 

41,10 42,10 43,2 44,3 45.2 

52.1 53,1 54,1 55,2 56,- 

63,3 64,2 65,2 66,1 67,2 

74.2 75,1 76,1/3 77,2 78,2 

elseyi 

1,120 2,64 3,7 4.U 5,13 6.U 7,3 8,1 9,3 10,1 11,2 12,1 

13.U 14,U 15,1 16,1 17,1 18,2 19,2 20,1 21,1 22,3 23,2 

24,2 25,3 26,4 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

35,6 36,104 37,1 38,1 39,0 40,0 41,18 42.U 43,1 44,2 45,2 

46,2 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,1 55,3 56- 

57.1 58- 59,- 60,2 61,1 62,1 63,1 64,1 65,2 66,2 67,2 

68.2 69,2 70,2 71,2 72,2 73,1 74,2 75,1 76,5 77,1 78,2 

79,4 80,30 81,1 82,1 

eugeniae 

1,U 2,U 3,1 4,10 5,4 6,1 7,3 

14- 15,1 16,1 17,4 18,3 19,2 

25,4 26,1 27,1 28,2 29,2 30,1 

8,1 

20,1 

31,1 

36,44 37,2 38,2 39,4 40,2-3 41,5 42.U 43,1-2 

46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 

57,2 58- 59,- 60,2 61,1 62,4 63,U 64,3 65,2 

68,2 

79,3 

69,1 

80,18 

70,1 

81,1 

71,2 

82,1 

72,2 73,1 74,2 75,1 76,1 

excariboea 

1,U 2,U 3,2 4,10 5,3.7 6,2 7,3 8,2 

13.1 14,2 15,1 16,3 17,1 18,3 19,2 

24.2 25,3 26,3 27,1 28,2 29,2 30,1 

35,78-79 36,U 37,1 38,1 

45,2 46,1 47,1 48,1 49,1 

56- 57,1 58,2 59,1 60- 

67,1 68,2 69,1 70,2 71,2 

78.U 79,2 80,78/87 813 

39,U 

50,1 

61,3 

72,2 

82,1 

40,U 

51,4 

62,3 

73,2 

9,2 10,2 11,1 12,2 13,2 

21,2 22- 23,2 24,2 

32,2 33,2 34,1 35,4 

9,2 

20,3 

31,2 

41,3 

52,1 

63,4 

74,U 

10,1 

21,1 

32,2 

42,U 

53,1 

64,1 

75,1 

44,2 45,2 

55,2 56- 

66,2 67,2 

77,1 78,2 

11,1 12,1 

22.1 23,1 

33.2 34- 

43,2/4 44,2 

54,2 55,2 

65,1 66,2 

76,1 77,2


62.01 

62.02 

62.03 

62.04 

62.05 

62.06 

62.07 

62.08 

62.09 

63.01 

63.02 

63.03 

63.04 

63.05 

63.06 

63.07 

63.08 

63.09 

64.01 

64.02 

64.03 

64.04 

64.05 

64.06 

64.07 

64.08 

64.09 

65.01 

65.02 

65.03 

65.04 

65.05 

65.06 

65.07 

65.08 

65.09 

66.01 

66.02 

66.03 

66.04 

66.05 

66.06 

66.07 

66.08 

66.09 

67.01 

67.02 

67.03 

67.04 

67.05 

67.06 

fauveli 

1,216 2,302 3,8 

12,1 13,1 14,5 

23,2 

34,1 

44,2 

55,2 

66,1 

77,2 

4,20 5,18 6,7.5 73 8,2 9,3 10,1 11,2 

15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 

24,2 25,3 26,4 27,1 28,2 29,2 30,1 31,1 32,1 33,2 

35,18 36,216 37,1 38,1 39,1 40,0 41,14 42,35 43,2-3 

45,2 46,2 47,2 48,3 49,2 50,2 51,9 52,1 53,1 54,1 

56- 

67,1 

78,U 

filamentosa 

1,119 2,25 

12,1 13,1 

23,1 24,2 

34- 

44,3 45,2 

55,1 56- 

66,1 67,1 

57,2 58- 59- 60,1 61,1 62,1 63,1 64,2 65,1 

68,1 69,1 70,1 71,2 72,2 73,2 74.U 75,1 76,1 

79,2 80,41 81,1 82,1 

3,1.5 4,10 5,3 6,1.5 7,2 

14.2 15,1 16,3 17,1 18,3 

25.3 26,7 27,1 28,2 29,2 

35,23-27 37,1 

46,2 

57,2 

68,1 

36,119 

47,2 

58- 

69,1 

48,2 

59- 

70,1 

38,1 

49,2 

60,2 

71,2 

77,2 78,U 79,2 80,17/22 81,1 82,1 

39,U 

50,2 

61,1 

72,2 

8,2 9,3 10,1 11,2 

19.2 20,3 21,2 22- 

30.1 31,2 32,1 33,2 

40,U 41,2 42,U 43,2-3 

51,4 52,1 53,1 54,2 

62.3 63,3 64,3 65,1 

73.2 74.U 75,1 76,4 

fimbriata 

1,362 2,230 3,7 4,30 5,11 6,6 7,2 8,3 9,2 10,1 11,2 12,2 

13.U 14.U 15,3 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,4 

24.1 25,4 26,4 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,2 

35,9 36,122 37,2 38,2 39,0 40,10-12 41,5 42,15 43,2 44,3 

45.2 46,2 47,2 48,3 49,3 50,3 51,4-5 52,1 53,1 54,2 55,1 

56- 57,2 58- 59- 60,3 61,4 62,4 63,4 64,2 65,2 66,2 

67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

78,2 79,2 80,26 81,1 82,1 

flaccida 

1,U 2,U 3,2.5 4,10 5,8.5 

13,U 14,U 15,1 16,1 17,1 

24.1 25,2 26,5 27,1 28,2 

35,7 36,U 37.U 

46.2 47.U 48,2 

57,U 58,U 59,U 

68.1 69,1 70,2 

78.2 79,4 80,27 

6,2.5 7,3 8,1 9,2 10,1 11,2 12,2 

18,3 19,2 20,1 21,1 22,3 23,2 

29,1 30,1 31,1 32,3 33,U 34,1 

38,U 39,0 40,U 41,9 42,12 43,2 44,3 45,2 

49,2 50.U 51- 52,1 53,2 54- 55- 56,2 

60.U 61,2 62,2 63,U 64,3 65,2 66,1 67,2 

71,2 72,2 73,1 74,2 75,1 76,1 77,1 

81,1 82,1 

flavapunctata 

1,U 2,U 3,2 4,10 5,7 6,2 7,3 8,1 9,3 10,2 11,1 12,2 13,U 

14,U 15,3 16,2 17,1 18,3 19,2 20,3 21,2 22- 23,4 24,2 

25,4 26,2 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 35,13 

36,U 37,2 38,1 39.U 40,U 41,6 42,50 43,1 44,3 45,2 46,2 

47.U 48,2 49,2 50,U 51,4 52,1 53,1 54,2 55,2 56- 57,U 

58.U 59.U 60.U 61,4 62,4 63,U 64,3 65,2 66,1 67,2 68,1 

69,1 70,2 71,2 72,2 73,2 74 ,U 75,1 76,1 77,2 78.U 79,2 

80,25 81,1 82,1 

flavocuprea 

l.U 2,17 3,1 4,U 5,2 6,U 7,2 8,2 9,2 10,1 11,2 12,2 13,U 

14,U 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,2 24,2 

25,4 26,2 27,1 28,5 29,2 30,1 31,1 32,1 33,2 34,1 35,4 

36,34 37,2 38,2 39,5 40,10 41,4 42,15 43,U 44,U 45,U 46,U 

47,U 48,U 49,U 50,U 51,4 52.U 53,1 54,1 55,1 56- 57,U

390 


67 07 58 U 59.U 60.U 61,2 62,4 63,4 64,3 65.U 66.U 67.U 68.U 

67.08 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 79,3 

67.09 80,17 81,1 82,2 

68.01 

68.02 

68.03 

68.04 

68.05 

68.06 

68.07 

68.08 

68.09 

69.01 

69.02 

69.03 

69.04 

69.05 

69.06 

69.07 

69.08 

70.01 

70.02 

70.03 

70.04 

70.05 

70.06 

70.07 

70.08 

70.09 

71.01 

71.02 

71.03 

71.04 

71.05 

71.06 

71.07 

71.08 

71.09 

72.01 

72.02 

72.03 

72.04 

72.05 

72.06 

72.07 

72.08 

72.09 

flavopicta 

1,166 2,215 3,6 

13,1 14,1 15.U 

24.U 

35,5 

45.U 

56.U 

66,1 

77,2 

25.U 26.U 

36,166 37,1 

46.U 47.U 

57.U 58.U 

4,10 

16.U 

27,1 

38,1 

48,U 

59.U 

67,2 68,1 69,1 70,2 71,2 72,2 

78,U 79,2 80.U 81.U 82.U 

floridana 

1,111 2,80 3,6 4,10 

13,1 14,3 15,1 16,2 

24.1 25,3 26,4 27,2 

35,8-10 36,111 37,1 

45.2 46,2 47,2 48,3 

56- 57,2 58- 59- 

67,1 68,2 69,1 70,2 

5,U 6,6 7,U 8,U 9.U 10.U ll.U 12.U 

17.U 18,3 19,2 20.U 21,1 22.U 23,2 

28,U 29,2 30,1 31,1 32,1 33.2 34,1 

39,0 40,0 41,18-19 42,14 43.U 44.U 

49.U 50.U 51.U 52.U 53,1 54.U 55.U 

60.U 61,U 62,1 63.U 64,1/2 65,2 

73,2 74 ,U 75,1 76.U 

5,10.5 6,6 

17,1 18,3 

28,1 29,1 

38,1 

49,3 

60,4 

71,2 

39,1 

50,2 

61,4 

72,2 

69.09 78.U 79,2 80,29-40 81,1 82,2 

7,2 8,2 9,2 10,1 11.2 12.1 

19.2 20.3 21.1 22,2 23.2 

30,1 31,1 32,1 33,1 34.1 

40,0 41,8 42.U 43,2-3 44,3 

51,7-9 52.1 53.1 54.2 55.2 

62.1 63.1 64,2 65,1 66.2 

73.2 74.U 75,1/2 76,1 77.2 

franklini 

1,U 2,U 3,7 4,10 5,15 6,7 7,3 8,1 9,3 10,1 11,2 12,1 13.U 

14,U 15,3 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,2 24,2 

25,3 26,5 27,2 28,1 29,1 30,1 31,1 32,3 33,2 34,1 35,6 

36.U 37,1 38,1 39,0 40.U 41,25 42.U 43,3 44,2 45,3 46,2 

47,U 48,3 49,2 50.U 51.U 52,1 53,1 54,1 55,2 56- 57.U 

58,U 59.U 60,U 61,4 62,4 63.U 64,3 65,2 66,2 67,2 68,2 

69,1 70,1 71,2 72,2 73,2 74.U 75,1 76,1 77,2 78.U 79,2 

80,34 81,1 82,1 

frauenfeldi 

1,U 2,U 3,5 4,10 5,7 6,5 7,2 

14,U 15,1 16,1 17,1 18,3 19,2 

25,3 26,3 27,1 28,1 29,1 30,1 

36,U 37,1 38,1 39,1 40.U 41,3 

47.U 48,2 49,2 50,U 51,4 52,1 

58,- 59- 60,1 61,3 62,3 63 ,U 

69,1 70,2 71,2 72,2 73,2 74,1 

80,27 81,1 82,2 

fucata 

1,250 

13,1 

24,U 

35,5 

45,2 

56.U 

67,1 

78,1 

8,3 9,2 10,1 11,2 12,2 13.U 

20.1 21,1 22,2 23,2 24,1 

31.2 32,1 33,2 34,- 35,6 

42,15 43,2 44,2 45,2 46,2 

53.1 54,2 55,2 56- 57,2 

64.2 65,2 66,2 67,2 68,2 

75,1 76,1 77,2 78,1 79,2 

2,195 3,5 4,U 5,10 6,U 7,U 8,U 9,U 10,U ll.U 12.U 

14,U 15,U 16,U 17,U 18.U 19,U 20,U 21,2 22- 23,1 

25,U 26,3 27,2 28,1 29,2 30,1 31,1 32,1 33,2 34,1 

36,170-180 37,2 38,1 39,U 40,30 41,12 42,U 43,1 44,3 

46,2 47,2 48,2 49,2 50,2 51,U 52.U 53,1 54.U 55,U 

57,2 58- 59- 60,2 61,1 62,1 63,4 64,3 65,U 66.U 

68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

79,1 80,U 81.U 82,U 

73.01 fuscafasciata 

73.02 1,U 2,U 3,1.3 4,10 5,3 6,1.3 7,3 8,1 9,3 

73.03 13,U 14,U 15,3 16,1 17,1 18,3 19,2 20,2 

10,1 

21,1 

11,2 

22,1 

12,2 

23,4


73.04 

73.05 

73.06 

73.07 

73.08 

73.09 

74.01 

74.02 

74.03 

74.04 

74.05 

74.06 

74.07 

74.08 

74.09 

75.01 

75.02 

75.03 

75.04 

75.05 

75.06 

75.07 

75.08 

75.09 

76.01 

76.02 

76.03 

76.04 

76.05 

76.06 

76.07 

76.08 

76.09 

77.01 

77.02 

77.03 

77.04 

77.05 

77.06 

77.07 

77.08 

77.09 

78.01 

78.02 

78.03 

78.04 

78.05 

78.06 

78.07 

78.08 

78.09 

24.1 25,1 26,4 27,1 28,2 29,2 30,2 31- 32- 33- 34- 

35- 36- 37- 38- 39- 40,- 41- 42- 43,1 44,2 45,2 

46.2 47.U 48,2 49,2 50,U 51,9 52,1 53,1 54,1 55,2 56- 

57,2 58- 59- 60,1 61,3 62,2 63.U 64,3 65,2 66,2 67,2 

68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 

79,2 80,23 81,1 82,1 

gagzoi 

1,160 2,32.5 3,1.5 4,10 5,2.75 6,1.5 

11.1 12,2 13,1 14,3 15,1 16,1 17,1 

22- 23,1 24,1 25,4 26,2 27,1 28,1 

33- 34- 35,85-120 

43.2 44,2 45,2 46,1 

54,2 55,2 56- 57,2 

65,2 66,1 67,2 68,1 

76,1 77,2 78.U 79,2 

7,3 8,2 9,1 10,1 

18,3 19,2 20,3 21,2 

29,2 30,1 31,3 32,1 

36,160 37,1 38,2 39- 40- 41- 42- 

47,1 48,3 49,1 50,1 51,9 52,1 53,1 

58- 59- 60,2 61,1 62,3 63,4 64,3 

69,1 70,2 71,2 72,2 73,2 74,U 75,1 

80,27-38 81,1 82,1 

goodei 

2.U 3,.75 4,U 5,4 6,U 7,3 8,1 9,2 10,1 11,2 12,U 13,1 

14,2 15.1 16,3 17,1 18,3 19,2 20,3 21,2 22- 23,4 24 ,U 

25,4 26.2 27,1 28,5 29,2 30,2 31- 32- 33- 34- 35- 

36- 37- 38- 39- 40- 41- 42- 43,1/3 44,2 45,2 46,2 

47.U 48,3 49,2 50,U 51,9 52,1 53,1 54.1 55,1 56- 57,2 

58- 59- 60,2 61,1 62,3 63.U 64,3 65.2 66,1 67,2 68,1 

69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 79,2 

80,23 81,1 82,1 

gracilicirrata 

1,323 2,214 3,2.5 4,10 5,9 6,2.5 7,3 8,1 9,3 10,1 11,2 

12,2 13,U 14,U 15,1 16,3 17,3 18,3 19,2 20,1 21,1 22,1 

23,2 24,2 25,4 26,4 27,2 28,2 29,1 30,1 31,1 32,1 33,2 

34.1 35,3 36,181 37,2 38,2 39,2 40,36 41,7 42,20 43,1 

44.2 45,2 46,2 47,2 48,3 49,2 50,2 51,10 52,3 53,1 54,1 

55.3 56- 57,2 58- 59- 60,2 61,3 62,3 63,3 64,2 65,2 

66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 

77,1 78,2 79,2 80,63 81,1 82,1 

gracilis 

1,81 2,66 3,2 4,10 5,6 6,2 7,2 8,2 9,2 10,1 11,2 12,2 

13.1 14,3 15,1 16,1 17,4 18,3 19,2 20,3 21,1 22,3 23,U 

24.2 25,1 26,4 27,1 28,2 29,1 30,1 31,1 32,1 33,1 34,1 

35,4-5 36,81 37,1 38,1 39,0 40,0 41,6 42.U 43,2-3 44,2 

45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 55,2 

56- 57,2 58- 59- 60,1 61,3-4 62,4 63,4 64,2 65.U 66,U 

67,U 68.U 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

78.2 79,3 80,28 81,1 82,1 

gravieri 

l.U 2,28 3,2 4,U 5,U 6,U 73 8,U 9,U 10,U ll.U 12,U 13,1 

14.3 15,U 16.U 17.U 18,3 19,2 20,3 21,1 22,1 23,1 24.U 

25,5 26- 27,3 28,2 29,1 30,1 31,3 32,3 33- 34- 35,5 

36,20/26 37,2 38,2 39- 40- 41,1 42- 43,U 44,U 45,U 

46,U 47,U 48,U 49,U 50,U 51,U 52,1 53,1 54,U 55,U 56- 

57,2 58- 59- 60,1 61,4 62,4 63,4 64,2 65,U 66,1 67,U 

68.1 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78,U 

79.2 80,25 81,U 82.U


79.01 grubei 

79 02 1U 2.U 3,6 4,10 5,12 6,6 7,3 8,1 9,2 10,1 11,2 12,2 

79.03 13.U 14.U 15.1 16,1 17,1 18,3 19,2 20,1 21.1 22,2 23,1 

79.04 24,1 25,3 26,5 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

79.05 35,3 36.U 37,1 38,1 39,1 40,U 41,8 42,20 43,1 44,2 45,2 

79.06 46,2 47.U 48,1 49,1 50,U 51,5 52,2 53,1 54,1 55,2 56- 

79 07 57,2 58- 59- 60,4 61,1 62,1 63,U 64,2 65,1 66,1 67,1 

79.08 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1. 77,2 78.U 

79.09 79,2 80,24 81,1 82,2 

80.01 guanica 

80 02 1,224 2,100 3,4 4,10 5,7 6,4 7,3 8,1 9,2 10,1 11,2 12,2 

80.03 13.U 14.U 15,2 16.1 17,1 18,3 19,2 20,3 21.1 22,1 23.4 

80.04 24,2 25,3 26,2 27,1 28,1 29,2 30.1 31.2 32.1 33.2 34- 

80.05 35,19 36,224 37,1 38,1 39,10 40.U 41,2 42,30 43.2-3 44,2 

80 06 45,2 46,2 47,2 48,3 49,2 50,2 51.6-7 52,3 53,1 54.1 55.1 

80.07 56- 57,1 58,1 59,1 60- 61,3 62,3 63,3 64,3 65.2 66.2 

80.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75.1 76,1 77.2 

80.09 78.U 79,2 80,36 81,1 82,1 

81.01 guildingi 

81.02 1,258 2,138 3,6 4,10 5,7 6,6 7,3 8,1 9,3 10.1 

81.03 13.U 14.U 15,1 16,1 17,1 18,3 19,2 20,3 21,1 

81.04 24,1 25,4 26,3 27,2 28,2 29,2 30,1 31,1 32,1 

81.05 35,5 36,236 37,2 38,1 39,0 40,20 41,11 42,15 

81.06 45,2 46,2 47,2 48,1 49,2 50,2 51,4 52,1 53,1 

81.07 56- 57,1 58,1 59,2 60- 61,3 62,2 63,4 64.3 

81.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 

81.09 78,U 79,2 80,24 81,1 82,1 

11.2 

22,1 

33.2 

43,2 

54,1 

65,2 

76,1 

12,1 

23.2 

34.1 

44.3 

55.2 

66,2 

77,2 

82.01 guttata 

82.02 1,115 2,52 3,3 4,10 5,8 6,3 7,2 8,1 9,3 10,1 11,2 12,2 

82.03 13,U 14.U 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,2 

82.04 24,2 25,1 26,4 27,1 28,1 29,2 30,1 31,1 32,1 33,2 34,1 

82.05 35,6 36,105 37,1 38,1 39,U 40.U 41,10 42,U 43,3 44,2 45,2 

82.06 46,2 47,2 48,3 49,2 50,2 51,19 52,1 53,1 54,1 55,1 56- 

82.07 57,1 58,1 59,1 60- 61,1 62,1 63,1 64,3 65,2 66,1 67,2 

82.08 68,1 69,1 70,1 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78.U 

82.09 79,2 80,32 81,1 82,2 

83.01 harassii 

83.02 1,U 2,U 3,2.5 4,10 5,5 6,2.5 7,2 8,2 9,3 10,1 11,2 12,1 

83.03 13,1 14,1 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

83.04 24,2 25,4 26,4 27,1 28,1 29,2 30,1 31,1 32,1 33,2 

83.05 34,1 35,4 36,U 37,1 38,1 39,3 40.U 41,10 42,15 43,1/3 

83.06 44,2 45,2 46,2 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,1 

83.07 55,2 56- 57,2 58- 59- 60,4 61,1 62,3 63,3 64,3 65,2 

83.08 66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 

83.09 77,2 78,2 79,2 80,28 81,1 82,1 

84.01 havaica 

84.02 1,78 2,25 3,1 4,10 5,4 6,1 7,2 8,2 9,2 10,1 11,1 12,1 

84.03 13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,3 23,2 

84.04 24,2 25,4 26,3 27,1 28,1 29,1 30,1 31,1 32,2 33,1 34,1 

84.05 35,6 36,78 37,1 38,1 39,U 40,U 41,5 42,20 43,1 44,2 45,2 

84.06 46,2 47,2 48,2 49,2 50,2 51,4 52,3 53,1 54,1 55,2 56-

NUMBER 523 

393 

84.07 

84.08 

84.09 

85.01 

85.02 

85.03 

85.04 

85.05 

85.06 

85.07 

85.08 

85.09 

86.01 

86.02 

86.03 

86.04 

86.05 

86.06 

86.07 

86.08 

86.09 

87.01 

87.02 

87.03 

87.04 

87.05 

87.06 

87.07 

87.08 

87.09 

88.01 

88.02 

88.03 

88.04 

88.05 

88.06 

88.07 

88.08 

88.09 

89.01 

89.02 

89.03 

89.04 

89.05 

89.06 

89.07 

89.08 

89.09 

90.01 

90.02 

90.03 

57.2 58- 59- 60,1 61,3 62,3 63,3 64,2 65,2 66,1 67,2 

68,1 69,1/2 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

79.3 80,19-23 81,1 82,1 

hawaiensis 

l.U 2,U 3,7 4,10 5,13 6,7 7,3 8,1 

14,3 15,1 16,1 17,1 18,3 19,2 20,1 

25.U 

36,54 

47.U 

58- 

69,1 

26,2 27,1 28,1 29,2 30,1 

37,2 39,0 40,4 

48,2 50.U 513 

59- 

70,1 

61.1 62,1 

72.2 73,1 

80,30 81,1 

heterochaeta 

1,105 2,27 

12.1 13.2 

23,2 24,1 

34,1 35,3 

44,2 45,2 

55,2 56,- 

66,1 67,2 

77,1 78,2 

38,2 

49,2 

60,1 

71,2 

82,1 

3,3.5 4,10 5,4 6,3.5 7,1 

14,- 15,1 16,1 17,2 18,3 

25,3 26,2 27,1 28,1 29,2 

38,1 

47,2 48,2 

58,- 59- 

69,1 70,2 

80,22 81,1 

36,85 37,2 

46,2 

57,2 

68,1 

79,2 

9,3 10,1 11,2 12.2 13,1 

21,2 22- 23,2 24.U 

31,1 32,1 33,2 34,1 35,3 

41,30 42.U 43,2 44,2 45,2 46,2 

52,1 53,1 54,1 55,2 56- 57,2 

63.U 64,3 65,2 66,1 67,2 68,1 

74,2 75,1 76,1 77,1 78,2 79,2 

39,3 

49,2 

60,2 

71,2 

82,1 

8,2 9,2 10,1 11,2 

19,2 20.1 21,1 22,1 

30,1 31,1 32,1 33,2 

40,15 41,7 42,25 43,1-2 

50,2 51,4 52,3 53,1 54,2 

61.1 62,1 63,1 64,2 65,2 

72.2 73,1 74,2 75,1 76,1 

hirschi 

1,112 2,55 3,3 4,10 5,7 6,3 7,1 8,2 9,3 10,1 11,2 12,2 

13.U 14.U 15.1 16,1 17,3 18,3 19,2 20,1 21,1 22,3 23,2 

24.1 25,3 26,3 27,2 28,1 29,1 30,1 31,1 32,3 33,2 34,1 

35,4 36,102 37,1 38,1 39,1 40,7 41,7 42,10 43,2 44,2 45,2 

46.2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1 55,2 56- 

57,2 58.- 59,- 60,1 61,1 62,1 63,1 64,1 65,2 66,2 67,2 

68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

793 80,22/25 81,1 82,1 

imogena 

l.U 2,U 

13,1 14,2 

24,2 25,4 

35- 36- 

46,2 47,2 

57,1 58,1 

68,1 69,1 

79,2 

3,2 4,10 5,3.5 6,2 7,3 8,2 

15,1 16,1 17,1 18,3 19,2 

26,2 27,1 28,2 29,2 30,2 

37- 38- 39- 40- 41- 

48,2 49,2 50,2 51,9 52,1 

59,2 60- 61,3 62,4 63,4 

70,2 71,2 72,2 73,2 74,1 

80,50 81,1 82,1 

9,3 

20,2 

31- 

42- 

53,1 

64,3 

75,1 

10,1 

21,2 

32- 

43,2 

54,1 

65,2 

76,1 

22- 

33- 

44,3 

55,1 

66,1 

12,2 

23,4 

34- 

45,2 

56- 

67,2 

77,2 78,1 

impexa 

1,U 2,U 3,5.5 4,10 5,11 6,5.5 7,3 8,2 9,2 10,1 11,2 12,1 

13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

24,1 25,3 26,3 27,1 28,2 29,2 30,1 31,2 32,1 33,2 34,2 

35,22 36.U 37,1 38,1 39,10 40,U 41,7 42,U 43,2 44,3 45,3 

46,3 47.U 48,2 49,2 50,U 51,4 52,1 53,1 54,2 55,1 56- 

57,1 58,2 59,2 60- 61,1 62,1 63,U 64,3 65,1 66,1 67,1 

68.1 69,1 70,2 71,2 72,1 73,2 74,1 75,1 76,1 77,2 78,1 

79,2-3 80,60 81,1 82,5 

indica 

1,U 2,U 3,2 4,U 5,4 6,U 7,3 8,2 9,2 10,1 11,1 12,1 13,1 

14.2 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,1 23,2 24,2


90.04 25,4 26,2 27,2 28,1 29,1 30,1 31,1 32,1 33,2 34,1 35,3 

90.05 36,21 37,2 38,2 39,1 40,0 41,8 42,U 43,2 44,2 45,2 46,2 

90.06 47.U 48,2 49,2 50.U 51,2 52,1 53,1 54,1 55,2 56- 57,2 

90.07 58- 59- 60,1 61,2 62,4 63.U 64,3 65,U 66.U 67.U 68.U 

90.08 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 79,3 

90.09 80,18 81,1 82,5 

91.01 interrupta 

91.02 1,U 2,U 3,2 4,U 5,9 6,U 7,3 8,2 9,3 10,1 11,2 12,2 13,1 

91.03 14,1 15,1 16,1 17,1 18,1 19,2 20,1 21,1 22,3 23,1 24,1 

91.04 25,2 26,5 27,1 28,1 29,1 30,1 31,2 32,3 33,1-2 34- 

91.05 35,5-7 36,24-26 37,1-2 38,1 39.U 40.U 41,3 42.U 43,2 44,2 

91.06 45,2 46,2 47.U 48,2 49,2 50.U 51,6 52,1 53,1 54,1 55,2 

91.07 56- 57,2 58,- 59- 60,2 61,4 62,4 63.U 64,1 65 ,U 66.U 

91.08 67.U 68.U 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

91.09 78,2 79,3 80,24-27 81,1 82,1 

92.01 investigatoris 

92.02 1,U 2,110 3,7 4,U 5,U 6,U 7,U 8.U 9.U 10.U ll.U 12.U 13.U 

92.03 14,U 15.U 16,U 17,1 18,3 19,2 20,1 21,2 22,- 23,1 24.U 

92.04 25,U 26,U 27.U 28.U 29.U 30,1 31,1 32,1 33,1 34,1 35,6 

92.05 36.U 37,1 38,1 39.U 40.U 41,18-20 42,14 43.U 44.U 45.U 

92.06 46.U 47.U 48.U 49.U 50,U 51.U 52,1 53,1 54.U 55,2 56- 

92.07 57,2 58- 59- 60,2 61,4 62,4 63,4 64,3 65,11 66.U 67.U 

92.08 68.U 69,1 70,2 71,2 72,2 73,2 74 ,U 75,1 76,1 77,2 78.U 

92.09 79,2 80,44-45 81,1 82,1 

93.01 jagori 

93.02 1,U 2,U 3,6 4,15 5,6 6,U 7,3 8,2 9,3 10,1 11,1 12,1 13,1 

93.03 14,6 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,1 23,2 24,2 

93.04 25,3 26,5 27,2 28,2 29,1 30,1 31,1 32,1 33,2 34,1 35,17 

93.05 36.U 37,1 38,1 39,U 40,U 41,8 42,25 43,2 44,3 45,2 46,2 

93.06 47.U 48,3 49,1 50.U 51- 52,1 53,2 54,- 55- 56,1 57.U 

93.07 58,U 59.U 60,U 61,1 62,1 63.U 64,1 65,1 66,2 67,1 68,2 

93.08 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 79,2 

93.09 80,23 81,1 82,1 

94.01 japonica 

94.02 1,110 2,50 3,2.5 4,U 5,7 6,U 7,3 8,2 9,2 10,1 11,2 12,1 

94.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,2 

94.04 24,2 25,4 26,3 27,2 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

94.05 35,4 36,46 37,2 38,2 39,2 40,2-3 41,11 42,U 43,3 44,2 

94.06 45,2 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 

94.07 56- 57,2 58- 59- 60,2 61,1 62,3 63,4 64,2 65,U 66.U 

94.08 67,U 68,U 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

94.09 78,2 79,2 80,34 81,1 82,2 

95.01 johnsoni 

95.02 1,160 2,75 3,3 4,U 5,7 6,U 7,3 8,1 9,3 

95.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,4 

95.04 24,2 25,4 26,3 27,1 28,2 29,2 30,1 31,1 

95.05 35,24 36,130 37,2 38,1 39,3 40,20 41,10 

95.06 45,2 46,2 47,2 48,2 49,2 50,2 51- 52,1 

95.07 56,1 57,2 58- 59- 60,1 61,1 62,1 63,4 

95.08 67,1 68,1 69,1 70,2 71,2 72,2 73,1 74,2 

95.09 78,2 79,2 80,31 81,1 82,1 

10,1 

21,1 

32,1 

42,U 

53,2 

64,3 

75,1 

11,2 

22,1 

33,2 

43,2 

54- 

65,1 

76,1 

12,1 

23,2 

34,1 

44,2 

55- 

66,1 

77,1

NUMBER 523 

395 

96.01 kerguelensis 

96.01 1,U 2,U 3,0.3 

96.03 13,1 14,3 

96.04 24,1 

96.05 35,3 

96.06 46,2 

96.07 57.U 

96.08 68,1 

96.09 79,3 

25,4 

36.U 

47.U 

58.U 

69,1 

80,19 

37,2 

48,2 

59,U 

70,2 

81,1 

4,U 5,1.2 6,U 7,3 8,1 9,2 10,2 11,1 12,2 

15.1 16,1 17,1 18,3 19,2 20,3 21,1 22,2 23,1 

26.2 27,1 28,2 29,2 30,1 31,1 32,2 33,2 34,- 

97.01 kinbergi 

97.02 1,380 2,398 3,15 4,U 5,16 6,U 7,3 

97.03 13,1 14,3 15,3 16,1 17,1 18,3 19,2 

97.04 24,1 25,3 26,5 27,1 28,1 29,1 30,1 

38,2 39.U 40.U 41,2 42,U 43,2 44,2 45,2 

49,2 50,U 51,3 52,1 53,1 54,1 55,2 56- 

60,U 61,1 62,1 63 ,U 64,3 65.U 66,1 67.U 

71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

82,1 

97.05 35,8-9 36,365 37,1 38,1 39,2 40,0 41,22 

97.06 45,2 46,2 47,2 48,1 49,1 50,2 51,4 52,1 

97.07 56- 57,1 58,4 59,2 60- 61,1 62,1 63,1 

97.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 

97.09 78,1 79,2 80,123 81,3 82,2 

8,1 9,3 10,1 11,2 12,2 

20,1 21,1 22,1 23,1-2 

31,1 32,2 33,2 34,1 

42,25 43,3 44,3 

53,1 54,2 55,2 

64,1 65,1 66,1 

75,1 76,1 77,2 

98.01 kobiensis 

98.02 1,U 2,U 3,3.5 4,U 5,6.5 6,U 7,3 8,2 9,3 10,1 11,2 12,1 

98.03 13.U 14,U 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

98.04 24,2 25,4 26,2 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

98.05 35,3 36,41 37,2 38,2 39,3 40,2 41,8 42,U 43,3 44,2 45,2 

98.06 46,2 47.U 48,2 49,2 50,U 51,3 52,1 53,1 54,1 55,2 56- 

98.07 57,2 58- 59- 60,1 61,2 62,3 63.U 64,2 65,2 66,1 67,2 

98.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

98.09 79,2 80,30 81,1 82,1 

99.01 langi 

99.02 1,71 2,42 3,2 4,U 5,5.5 6.U 7,2 8,2 9,2 10.U ll.U 12,U 

99.03 13,U 14,U 15.U 16,U 17.U 18,3 19.U 20,U 21,2 22.U 23,U 

99.04 24,U 25,3 26,3 27,3 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

99.05 35,3 36,39 37,2 38,2 39,U 40,U 41,8 42,15 43,1 44,2 45,2 

99.06 46,2 47,2 48,2 49,2 50,2 51,4 52,3 53,1 54,1 55,2 56- 

99.07 57,2 58- 59- 60,3 61,4 62,3 63,3 64,3 65,2 66,2 67,2 

99.08 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78.U 

99.09 79,2 80,30 81,1 82,1 

100.01 laticeps 

100.02 1,U 2,U 3,10 4,U 5,21 6,U 7,3 8,1 9,3 10,1 11,2 12,1 

100.03 13.U 14,U 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,1 

100.04 23,1 24,2 25,3 26,3 27,1 28,2 29,1 30,1 31,1 32,1 33,2 

100.05 34,1 35,4 36.U 37,1 38,1 39,0 40,U 41,18 42,20 43,1 44,3 

100.06 45,2 46,2 47.U 48,2 49,2 50,U 51,3 52,1 53,1 54,1 55,2 

100.07 56- 57,2 58- 59- 60,1 61,4 62,4 63.U 64,2 65,2 66,1 

100.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

100.09 78,U 79,2 80,38 81,1 82,2 

101.01 laurillardi 

101.02 1,189 2,150 3,5 4.U 5,9 6,U 7,3 8,2 9,3 10,1 11,2 12,1 

101.03 13,1 14,3 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,2 

101.04 24,2 25,3 26,5 27,2 28,1 29,1 30,1 31,1 32,3 33,2 34,1 

101.05 35,3 36,184 37,1 38,1 39,1 40,0 41,10 42,15 43,2 44,3 

101.06 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 55,2

3% SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

101.07 56- 57,2 58- 59- 60,4 61,3 62,3 63,3 64,1 65,2 66.1 

101.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

101.09 78.U 79,2 80,32-33 81,1 82,1 

102.01 leptocirris 

102.02 1,U 2,U 3,3 4,U 5,9 6,U 7,2 8,2 9,3 10,1 11,2 12,2 13.U 

102.03 14.U 15,1 16,3 17,1 18,1 19,2 20,3 21,2 22- 23,2 24,2 

102.04 25,4 26,2 27,3 28,1 29,2 30,1 31,1 32,3 33,2 34,1 35,3 

102.05 36.U 37,1 38,1 39.U 40.U 41,6 42,15 43,2-3 44,3 45,2 

102.06 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1 55,2 56- 

102.07 57,2 58- 59- 60,2 61,1 62,3 63,3 64,2 65,2 66,1 67,2 

102.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78.2 

102.09 79,2 80,51 81,1 82,1 

103.01 leucosticta 

103.02 1,131 2,130 3,U 4,U 5,U 6.U 7,U 8,U 9.U 10.U ll.U 12.U 

103.03 13,U 14.U 15,U 16.U 17.U 18,U 19.U 20.U 21.U 22.U 23.1 

103.04 24.U 25.U 26.U 27.U 28.U 29.U 30,1 31.U 32.U 33.U 34.U 

103.05 35,14-15 36,113 37,2 38.U 39.U 40.U 41.4 42.U 43.2 44,3 

103.06 45.U 46,U 47,2 48.U 49.U 50,2 51.U 52.U 53.U 54.U 55.U 

103.07 56.U 57,1 58,1 59,1 60.U 61,U 62.U 63.1 64,2/3 65.U 66.U 

103.08 67,U 68.U 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76.1 77,2 

103.09 78,1 79,2 80,U 81.U 82.U 

104.01 levibranchia 

104.02 1,105 2,94 3,8 4,U 5,9 6,U 7,3 8,1 9,2 10,1 11,2 12,1 

104.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,3 21,2 22- 23,1 

104.04 24,2 25,3 26,2 27,1 28,1 29,2 30,1 31,2 32,3 33,2 34- 

104.05 35,69 36,85-95 37,1-2 38,2 39,0 40,0 41,3 42.U 43,1 44,2 

104.06 45,2 46,2 47,2 48,2 49,2 50,2 51,1-2 52,1 53,1 54,2 55,1 

104.07 56- 57,2 58- 59- 60,1 61,3 62,3 63,3 64,2 65,1 66,2 

104.08 67,1 68,2 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

104.09 78.U 79,2 80,28 81,1 82,1 

105.01 limosa 

105.02 1,81-106 2,32-65 3,2 4,U 5,U 6,U 7,3 8,U 9,2 10,U ll.U 

105.03 12,U 13.U 14.U 15,U 16,U 17.U 18,3 19,2 20,3 21,1 22,2 

105.04 23,2 24.U 25,4 26,2 27,1 28,2 29,2 30,1 31,1 32,1 33,2 

105.05 34,1 35,3 36,36 37,2 38,2 39.U 40.U 41,12 42.U 43,U 44,U 

105.06 45,U 46.U 47,U 48.U 49.U 50.U 51.U 52.U 53,1 54.U 55.U 

105.07 56,U 57,2 58- 59- 60.U 61,4 62,4 63,4 64,3 65,U 66,U 

105.08 67.U 68.U 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

105.09 78,2 79,3 80,U 81.U 82.U 

106.01 lita 

106.02 1,240 2,78 3,4 4,U 5,4 6,U 7,2 8,3 9,3 10,1 11,2 12,1 

106.03 13,1 14,3 15,1 16,2 17,4 18,3 19,2 20,1 21,2 22- 23,2 

106.04 24,2 25,4 26,5 27,2 28,3 29,2 30,1 31,2 32,1 33,2 34- 

106.05 35,15 36,120 37,2 38,2 39,20 40,50 41,2 42,35 43,3 44,2 

106.06 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,2 55,1 

106.07 56- 57,2 58- 59- 60,3 61,4 62,4 63,4 64,3 65,2 66,2 

106.08 672 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

106.09 78,U 79,2 80,23 81,1 82,1 

107.01 longicirris 

107.02 l.U 2,U 3,5 4,U 5,16 6,U 7,3 8,1 9,2 10,1 11,2 12,2 13,1 

107.03 14,2 15,1 16,3 17,3 18,2 19,2 20,1 21,1 22,2 23,U 24,2


107.04 25,3 26,3 27,3 28,1 29,1 30,1 31,1 32,3 33,2 34,1 35,3 

107.05 36,U 37,1 38,1 39.U 40.U 41,7 42,15 43,3 44,3 45,2 46,2 

107.06 47.U 48,2 49,2 50.U 51,4 52,1 53,1 54,1 55,2 56- 57,2 

107.07 58- 59- 60,1 61,4 62,4 63.U 64,1 65,2 66,1 67,2 68,1 

107.08 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,2 78,1 79,2 

107.09 80,30 81,1 82,1 

108.01 longisetis 

108.02 1,108 2,40 3,3.5 4,U 5,6.4 6,U 7,3 8,2 9,2 10,1 11,2 

108.03 12,2 13.1 14,3 15,1 16,1 17,4 18,3 19,2 20,3 21,1 22,1 

108.04 23.U 24.2 25,3 26,5 27,1 28,2 29,1 30,1 31,1 32,1 33,2 

108.05 34,1 35,6 36,104 37,1 38,1 39,1 40,15 41,8 42,U 43,3 

108.06 44,3 45,2 46,2 47,2 48,3 49,3 50,3 51,9 52,1 53,1 54,2 

108.07 55,3 56,- 57,2 58- 59- 60,2 61,4 62,4 63,4 64,2 65,2 

108.08 66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

108.09 77,2 78,1 79,2 80,18 81,1 82,1 

109.01 

109.02 

109.03 

109.04 

109.05 35,5 

109.06 46,2 

109.07 57,2 

109.08 68,1 

109.09 78,2 

110.01 

110.02 

110.03 

110.04 

110.05 

110.06 

110.07 

110.08 

110.09 

lucei 

1,U 2,U 3,3 4,40 5,5.5 6,U 7,3 8,1 9,3 10,1 11,2 12,1 

13,1 14,2 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,2 23,1 

24,1 25,4 26,4 27,2 28,4 29,1 30,1 31,1 32,1 33,2 34,1 

36,168 37,1 38,1 39,1 40.U 41,8 42.U 43,2 44,3 45,2 

47.2 48,2 49,2 50,2 51,4 52,1 53,1 54,2 55,1 56- 

58- 59- 60,1 61,1 62,1 63,1 64,1 65,2 66,1 67,2 

69,1 70,1-2 71,2 72,2 73,1 74,2 75,1 76,4-5 77,1 

79.3 80,24-34 81,1 82,1 

macrobranchia 

1,150 2,200 3,U 

13.U 14.U 15,U 

24.U 25.U 26.U 

35,7-8 36.U 37,1 

45.U 46.U 47.U 

56.U 57,U 58.U 

4.U 5,U 6,U 7,U 8,U 9,U 10,U 11,U 12,U 

16.U 17.U 18,U 19.U 20.U 21,1 22,1 23.U 

27,1 28.U 29.U 30,1 31,1 32,2 33,2 34,1 

38,1 

48,U 

59.U 

67,U 68,1 69,1 70,2 71,2 

78,U 79,1-2 80.U 81.U 

39,5 40.U 41,15-17 42.U 43,U 44,U 

49.U 50.U 51.U 52,U 53,U 54.U 55,U 

60.U 61,U 62.U 63,U 64,U 65,U 66,1 

72,2 73,2 74.U 75.U 76,U 77,2 

111.01 magellanica 

111.02 1,130 2,140 3,7 4,U 5,17 6,U 

111.03 13,1 14,2 15.U 16,1 17,1 18,3 

111.04 24,1 25,3 26,5 27,1 28,5 29,2 

111.05 35,7-8 36,120 37,1 38,1 39,0 40,0 41,20 

111.06 44,2 45,2 46,2 47,2 48,2 49,2 50,2 51,5 

111.07 55,2 56- 57,2 58- 59- 60,2 61,4 62,3 

111.08 66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,2 

111.09 77,2 78,1 79,2 80,26/30 81,1 82,1 

7,3 8,1 9,3 10,1 11,2 12,1 

19,2 20,3 21,1 22,1 23,2 

30,1 31,1 32,1 33,2 34,1 

42,85 

52,3 

63,3 

74,1 

43,1-2 

53,1 54,1 

64,1 65,2 

75,1 76,1 

112.01 magnifica 

112.02 1,U 2,U 3,U 4,U 5,U 6,U 7,U 8,U 9,U 10.U ll.U 12,U 13.U 

112.03 14,U 15,U 16.U 17.U 18,U 19.U 20,U 21.U 22.U 23,U 24,U 

112.04 25,U 26.U 27,U 28,U 29,U 30,1 31,1 32,1 33.U 34,1 35,22 

112.05 36,U 37,U 38,U 39.U 40.U 41,8 42,U 43,1 44,2 45,1 46,U 

112.06 47,U 48,3 49,U 50,U 51,U 52,2 53.U 54,U 55,U 56,U 57,2 

112.07 58,- 59- 60,1 61,3 62,U 63,3 64,3 65,2 66,1 67,2 68,1 

112.08 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 79,U 

112.09 80,U 81,U 82,U

398 

SM m 

ISONIAN CON TRIB UTIONS TO ZOOLOGY 

113.01 makemoana 

113.02 1,92 2,20 3,1 4,U 5,3.5 6,U 7,3 8,1 9.U 10,1 ll.U 12.U 

113.03 13,1 14,2 15.U 16,U 17.U 18,3 19,2 20.U 21,1 22,3 23,2 

113.04 24,U 25.U 26.U 27,U 28.U 29.U 30,1 31,3 32,1 33- 34- 

113.05 35,7 36,86 37,1 38,1 39- 40- 41,1 42- 43,2 44.U 45.2 

113.06 46,2 47,2 48,2 49,2 50,2 51.U 52,1 53,1 54.U 55.U 56.U 

113.07 57,2 58- 59- 60,1 61,2 62,2 63,2 64,1 65.U 66,1 67.U 

113.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1/4 77,1 78,2 

113.09 79,3 80,28 81.U 82.U 

114.01 manihine 

114.02 1,127 2,75 3,4 4,U 5,9 6,U 7,2 8,2 9,3 10,1 11,2 12,1 

114.03 13,2 14- 15,1 16,1 17,U 18,3 19,2 20,1 21,2 22- 23.U 

114.04 24,1 25,3 26,2 27,2 28,2 29,2 30,1 31,2 32,1 33,2 34- 

114.05 35,8 36,43 37,2 38,2 39- 40- 41,3 42,U 43,1 44,2 45,2 

114.06 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,2 55.1 56,- 

114.07 57,2 58- 59- 60,1 61,1 62,3 63,4 64,3 65.2 66,2 67,2 

114.08 68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76.1 77.1 78.2 

114.09 79,2 80,29 81,1 82,5 

115.01 marenzelleri 

115.02 1,U 2.U 3,5 4,U 5,11 6,U 7,3 8,2 9,2 10,1 11,2 12,1 13.1 

115.03 14,2 15,1 16,3 17,4 18,3 19,2 20,3 21,2 22- 23,1 24,2 

115.04 25,3 26,4 27,1 28,5 29,2 30,1 31,3 32,1 33- 34- 35,28 

115.05 36.U 37,1 38,1 39- 40- 41,1 42- 43,2 44,2 45,2 46,2 

115.06 47,2 48,3 49,1 50,2 51,11 52,1 53,1 54,2 55,2 56,- 57,2 

115.07 58,- 59- 60,1 61,2 62,3 63,3 64,3 65,2 66,2 67,2 68,2 

115.08 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78.U 79,1 

115.09 80,38 81,1 82,1 

116.01 margariticacea 

116.02 1,95 2,14 3,U 4.U 5,U 6,U 7,U 8,U 9,U 10,U ll.U 12.U 

116.03 13,U 14.U 15,U 16.U 17.U 18.U 19,U 20.U 21,1 22,3 23.U 

116.04 24,2 25,U 26,U 27,2 28.U 29,1 30,1 31,1 32,U 33,1 34,1 

116.05 35,6 36,95 37,1 38,1 39,U 40,U 41,5 42,10 43,U 44,U 45.U 

116.06 46,U 47,U 48,U 49,U 50.U 51.U 52,1 53,1 54,1 55.U 56.U 

116.07 57,1 58.U 59,U 60,U 61.U 62,U 63,U 64,1 65,2 66,2 67,2 

116.08 68,2 69,2 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

116.09 79,3 80,U 81,U 82,U 

117.01 marovoi 

117.02 1,65 2,14.5 3,1 4,U 5,2.5 6,U 7,3 8,1 9,1 10,1 11,2 12,2 

117.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,2 

117.04 24,2 25,4 26,2 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

117.05 35,3 36,17-22 37,2 38,2 39,0 40,1 41,6 42,U 43,2 44,3 

117.06 45,2 46,2 47,2 48,2 49,2 50,2 51,2 52,1 53,1 54,1 55,3 

117.07 56- 57,1 58,1 59,3 60- 61,1 62,1 63,4 64,3 65,U 66.U 

117.08 67,U 68,U 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

117.09 78,2 79,3 80,17-19 81,1 82,5 

118.01 martensi 

118.02 1,168 2,220 3,6 4,U 5,14 6,U 7,U 8,U 9,2 10,1 ll.U 12,U 

118.03 13,1 14,3 15,U 16.U 17.U 18,3 19,2 20,3 21,1 22,1 23,U 

118.04 24,2 25,3 26,3 27,2 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

118.05 35,6 36,163 37,1 38,1 39,0 40,0 41,16 42,15 43,1-2 44,2 

118.06 45,2 46,2 47,2 48,2 49,2 50,2 51,U 52,1 53,1 54,1 55,2


118.07 

118.08 

118.09 

56- 

67,2 

78,1 

57,2 

68,2 

79,3 

58,- 

69,2 

80,28 

59- 60,1 61,1 

70,2 71,2 72,2 

81,1 82,1 

62,1 

73, 1 

63,1 

74,2 

64,1 

75, 1 

65,2 

76,1 

66,2 

77,1 

119.01 medicina 

119.02 1,79 2,29 3,1.3 4,U 5,5 6,U 7,2 8,2 9,2 10,1 11,2 12,1 

119.03 13,1 14,2 15,1 16,1 17,U 18,3 19,2 20,3 21,1 22,1 23,U 

119.04 24,2 25,4 26,2 27,2 28,2 29,1 30,1 31,1 32,U 33,2 34,1 

119.05 35,3 36,26 37,2 38,2 39,5 40,0 41,7 42,U 43,1-2 44,3 

119.06 45,2 46,2 47,2 48,1 49,2 50,2 51,2 52,3 53,1 54,1 55,3 

119.07 56- 57,1 58- 59- 60,2 61,4 62,4 63,4 64,2 65,2 66,2 

119.08 67,2 68,2 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

119.09 78,2 79,3 80,17 81,1 82,1 

120.01 megabranchia 

120.02 1,U 2,U 3,7 4,U 5,12 6,U 7,3 8,1 9,2 10,1 11,2 12,2 13,1 

120.03 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,2 24,2 

120.04 25,4 26,3 27,3 28,1 29,2 30,1 31,1 32,1 33,2 34,1 35,3 

120.05 36,54 37,2 38,2 39,0 40,7 41,47 42,15 43,1 44,3 45,2 

120.06 46,2 47.U 48,2 49,2 50.U 51,3 52,1 53,1 54,1 55,2 56- 

120.07 57,2 58- 59- 60,1 61,3 62,2 63,U 64,2 65,1 66,2 67,2 

120.08 68,2 69,1 70,1 71,2 72,2 73,1 74.U 75,1 76,1 77,1 78,U 

120.09 79,2 80,35 81,1 82,1 

121.01 mexicana 

121.02 1,106 2,87 3,3.5 4,15 5,10.5 6,3.2 7,2 8,1 9,2 10,1 11,2 

121.03 12,2 13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 

121.04 23,2 24,1 25,3 26,3 27,2 28,1 29,1 30,1 31,1 32,1 33,2 

121.05 34,1 35,3 36,40 37,2 38,2 39,0 40,0-3 41,18 42,15 43,2 

121.06 44,3 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 

121.07 55,3 56- 57,2 58- 59- 60,2 613 62,3 63,4 64,2 65,1 

121.08 66,1 67,1 68,1 69,1 70,2 71,2 72,2 73,1 74,1 75,1 76,1 

121.09 77,1 78,1 79,2 80,32 81,1 82,5 

121.01 microprion 

122.02 1,127 2,82 3,5 4,U 5,10 6,U 7,2 8,3 9,3 10,1 11,2 12,1 

122.03 13,1 14,4 15,1 16,3 17,1 18,3 19,2 20,1 21,1 22,1 23,1 

122.04 24,2 25,3 26,5 27,2 28,1 29,1 30,1 31,1 32,3 33,1 34,2 

122.05 35,7-8 36,127 37,1 38,1 39,0 40,0 41,8 42,15 43,2-3 44,2 

122.06 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,2 55,2 

122.07 56,- 57,2 58- 59- 60,4 61,4 62,4 63,4 64,1 65,2 66,1 

122.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

122.09 78,1 79,2 80,38/41 81,1 82,4 

123.01 mindanavensis 

123.02 1,63 2,40 3,4 4,U 5,7 6,U 7,3 8,2 9,3 10,1 11,2 12,1 

123.03 13.U 14.U 15,U 16,U 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

123.04 24,2 25,3 26,6 27,2 28,1 29,1 30,1 31,1 32,3 33,2 34,1 

123.05 35,6 36,63 37,1 38,1 39,0 40,5 41,3 42,15 43,1-2 44,3 

123.06 45,2 46,2 47,2 48,1 49,2 50,2 51,5 52,1 53,1 54,3 55,2 

123.07 56- 57,1 58,4 59,2 60- 61,2 62,2 63,2 64,2 65,2 66,2 

123.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

123.09 78,1 79,2 80,21 81,1 82,1 

124.01 modesta 

124.02 1,80 2,28 3,1 4,U 5,3 6,U 7,3 8,3 9,3 10,1 11,2 12,1 

124.03 13,1 14,3 15,1 16,1 17,1 18,1 19,2 20,3 21,1 22,1 23,4


124.04 24,1 25,3 26,3 27,1 28,5 29,2 30,1 31,3 32,1 33- 34- 

124.05 35,17-18 36,70 37,2 38,1 39- 40- 41,1 42- 43.U 44,U 

124.06 45,U 46.U 47,U 48,U 49,U 50.U 51,4 52.U 53,1 54,1 55.U 

124.07 56- 57,2 58- 59- 60,1 61,1 62,1 63,1 64,3 65,2 66,2 

124.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

124.09 78,U 79,2 80,U 81,1 82,1 

125.01 mucronata 

125.02 1,116 2,58 3,4 

125.03 13,1 14,2 15,1 

125.04 24,2 25,4 26,2 

125.05 35,3 36,34 37,2 

4,25 5,8 6,U 7,3 8,1 9,2 10,1 11,2 12,1 

16.2 17,1 18,3 19,2 20,1 21,2 22,- 23,2 

27.3 28,1 29,2 30,1 31,1 32,1 33,2 34,1 

38,2 39,2 40,0 41,15 42.U 43,1-2 44,3 

125.06 45,2 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53.1 54,1 55,2 

125.07 56- 57,2 58- 59- 60,1 61,3 62,3 63,3 64.3 65,2 66,2 

125.08 67,2 68,2 69,1 70,1 71,2 72,2 73.1 74,2 75,1 76,1 77,1 

125.09 78,2 79,3 80,18-22 81,1 82,1 

126.01 multicylindri 

126.02 1,115 

126.03 13,1 

126.04 24,2 

126.05 35,3 

126.06 45,2 

2,45 3,1 

14,4 15,1 

25,4 26,2 

36,40 37,2 

46,2 47,2 

4,U 5,3.5 6,U 7,3 8,3 9.3 10.1 11.2 12,2 

16,3 17,1 18,3 19.2 20.3 21.1 22,1 23.1 

27,1-2 28,1 29,1 30.1 31.1 32.1 33.2 34,1 

38,2 39,2 40.10-15 41,4 42.U 43.1-2 44,2 

48,2 49,2 50,2 51,5-6 52,1 53.1 54.1 55.2 

126.07 56- 57,2 58- 59- 60,1 61,1 62,1 63,1 64,3 65,1 66,1 

126.08 67,1 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

126.09 78,2 79,3 80,27 81,1 82,1 

127.01 multipectinata 

127.02 1,196 2,172 3,7 4,U 5,16 6.U 7,3 8,3 9,2 10,1 11,2 12,1 

127.03 13,1 14,3 15,U 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,U 

127.04 24,1 25,3 26,5 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

127.05 35,7-8 36,196 37,1 38,1 39,8-10 40,0 41,12 42.U 43,2 

127.06 44,3 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,3 53,1 54,2 

127.07 55,2 56- 57,2 58- 59- 60,2 61,3 62,3 63,3 64,2 65,1 

127.08 66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

127.09 77,2 78,1 79,2 80,37-45 81,1 82,1 

128.01 murrayi 

128.02 1,101 2,37 

128.03 13,U 

128.04 24,2 

128.05 35,5 

128.06 46,2 

128.07 

128.08 

128.09 

57,2 

68,U 

79,3 

3,4 

14,U 15,1 

25,3 26,3 

36,28 37,1 

47,2 48,2 

58,- 59,- 

69,1 70,2 

4,U 5,7 6,U 7,3 8,1 9,2 

16,1 17,1 18,3 19,2 20,1 

27,1 28,2 29,1 30,1 31,1 

38,2 39,2 

50,2 

61,2 

72,2 

49,2 

60,2 

71,2 

80,17 81.U 82,U 

129.01 mutabilis 

129.02 1,196 2,95 3,6 

129.03 13,1 14,4 15,1 

129.04 24,1 25,3 26,4 

129.05 35,17-18 36,196 

129.06 44,3 45,2 46,2 

129.07 55,2 56- 57,1 

129.08 66,2 67,1 68,2 

129.09 77,2 78,1 79,2 

40,5 

51,4 

62,2 

73,1 

41,9 

52,3 

63,2 

74,1 

10,1 11,2 12,2 

21,1 22,3 23 ,U 

32,1 33,2 34,1 

42,10 43,1 44,2 45,2 

53,1 

64,2 

75,1 

54,1 

65.U 

76,1 

55,3 56- 

66,U 67,U 

77,1 78,2 

4,U 5,8 6,U 7,3 8,2 9,3 10,1 11,2 12,1 

16.1 17,4 18,3 19,2 20,1 21,2 22- 23,2 

27,1-2 28,4 29,2 30,1 31,1 32,1 33,2 34,2 

37,1 38,1 39,2-3 40,0 41,12 42,25 43,1-2 

47.2 48,3 49,2 50,2 51,9 52,1 53,1 54,2 

58,2 59,2 60- 61,1 62,1 63,1 64,3 65,1 

69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

80,22-23 81,1 82,1


130.01 mutilata 

130.02 1,U 2,U 33.5 4,U 5,10 6,U 7,3 8,1 9,3 10,1 11,2 12,2 

130.03 13,1 14,3 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

130.04 24,2 25,4 26,3 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 

130.05 35,7 36.U 37,1 38,1 39,3 40.U 41,4 42,25 43,3 44,2 45,2 

130.06 46,2 47,2 48,3 49,3 50,2 51,9 52,1 53,1 54,2 55,3 56- 

130.07 57,2 58,- 59,- 60,2 61,4 62,4 63,4 64,3 65,1 66,1 67,1 

130.08 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 78.U 

130.09 79,2 80,27 81,1 82,1 

131.01 mutilatoides 

131.02 1,110 2,32 3,1 4,U 5,5 6.U 7,U 8,U 9,2 10,1 11,2 12,2 

131.03 13,1 14,2 15,3 16,1 17,1 18,3 19,2 20,3 21,2 22,- 23,4 

131.04 24,2 253 26,3 27,1 28,5 29,2 30,1 31,2 32,3 33,2 34- 

131.05 35,8 36,48 37,2 38,2 39.U 40.U 41,2 42,U 43,1-2 44.U 

131.06 45.U 46.U 47.U 48.U 49.U 50,U 51,7 52,1 53,1 54,2 55,2 

131.07 56,- 57,2 58- 59,- 60,1 61,2 62,2 63,2 64,3 65,1 66,1 

131.08 67,1 68,1 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

131.09 78.U 79,2 80,19 81,1 82,1 

132.01 narconi 

132.02 l.U 2,U 3,2 4.U 5,6 6,U 7,3 8,1 9,2 10,1 11,2 12.U 13,U 

132.03 14.U 15,U 16.U 17.U 18,U 19.U 20.U 21,U 22.U 23.U 24,2 

132.04 25.U 26,3 27,1 28,5 29,2 30,1 31,2 32.U 33,U 34.U 35,8 

132.05 3633-35 37,2 38,2 39.U 40,U 41,3 42,15 43.U 44.U 45.U 

132.06 46.U 47.U 48.U 49.U 50.U 51.U 52.U 53,1 54.U 55.U 56- 

132.07 57.U 58.U 59.U 60,U 61,U 62,U 63.U 64.U 65,2 66,1 67,2 

132.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

132.09 79,2 80,35-38 81.U 82.U 

133.01 nesiotes 

133.02 l.U 2,U 3,6 4,U 5,10 6,U 7,2 8,2 9,2 10,1 11,2 12,1 

133.03 13.U 14.U 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

133.04 24,2 25,3 26,4 27,2 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

133.05 35,6 36.U 37,1 38,1 39,2 40.U 41,14 42.U 43,1-2 44,3 

133.06 45,2 46,2 47,U 48,2 49,2 50.U 51,5 52,1 53,1 54,2 55,3 

133.07 56- 57.U 58,U 59.U 60.U 61,1 62,1 63,U 64,3 65,2 66,1 

133.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

133.09 78.U 79,2 80,31 81,1 82,1 

134.01 nicidioformis 

134.02 1,97 2,45 3,2 4,U 5,6 6,U 7,3 8,3 9,3 10,1 11,2 12,1 

134.03 13,1 14,4 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,2 23,1 

134.04 24,1 25,3 26,5 27,2 28,1 29,1 30,1 31,3 32,3 33- 34- 

134.05 35,6-8 36,40-50 37,2 38,2 39- 40- 41,1 42- 43,3 44,2 

134.06 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 55,3 

134.07 56- 57,2 58- 59- 60,1 61,4 62,4 63,4 64,2 65,2 66,1 

134.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

134.09 78,U 79,2 80,19-26 81,1 82,1 

135.01 nigricans 

135.02 1,180 2,100 3,7 4,25-30 5,10 6,6 7,2 8,3 9,3 10,1 11,2 

135.03 12,2 13,1 14,3 15,1 16,1 17,6 18,3 19,2 20,1 21,1 22,1 

135.04 23,2 24,2 25,1 26,4 27,2 28,4 29,1 30,1 31,1 32,1 33,2 

135.05 34,1 35,6-7 36,180 37,1 38,1 39,0 40,0 41,18 42.U 43,1 

135.06 44,3 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1

402 SMmISONI AN CONTRIBUTIONS TO ZOOLOGY 

135.07 55,2 56- 57,2 58- 59- 60,4 61,1 62,1 63,1 64.1 65,2 

135.08 66,1 67,2 68,1 69,1 70,1 71,2 72,2 73,2 74,1 75,1 76,1 

135.09 77,2 78,1 79,2 80,30 81,1 82,1 

136.01 northioidea 

136.02 1,U 2,U 3,2.5 4,U 5,7 6,U 7,3 8,1 9,3 10,1 11,2 12,1 

136.03 13,1 14,4 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,2 23,2 

136.04 24,1 25,3 26,3 27,1 28,3 29,2 30,1 31,1 32,1 33,2 34,1 

136.05 35,3 36,U 37,1 38,1 39,3 40.U 41,4 42,10 43.3 44,2 45,2 

136.06 46,2 47.U 48,2 49,2 50.U 51,4 52.1 53,1 54.1 55.3 56- 

136.07 57,2 58,- 59- 60,1 61,4 62,4 63.U 64.2 65,2 66.1 67.2 

136.08 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78.1 

136.09 79,2 80,25 81,1 82,1 

137.01 norvegica 

137.02 1,157 2,200 3,9 4,U 5,13 6.U 7.1 8.3 9.3 10.1 11.2 12,2 

137.03 13,1 14,4 15,1 16,1 17,1 18,3 19,2 20.3 21.1 22.1 23.2 

137.04 24,2 25,3 26,4 27,2 28,1 29,1 30.1 31.1 32,3 33.2 34.1 

137.05 35,7 36,155 37,1 38,1 39.1 40.3 41,7 42.U 43.1-2 44.2 

137.06 45,2 46,2 47,2 48,2 49,2 50,2 51,6 52.1 53,1 54,1 55,3 

137.07 56- 57,2 58- 59- 60.2 61,3 62,3 63,3 64,2 65.2 66.1 

137.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75.1 76.1 77.2 

137.09 78.U 79,2 80,42 81,1 82,2 

138.01 notata 

138.02 1,95 2,40 3,1 4,U 5,U 6,U 7,U 8,U 9,U 10.U ll.U 12.U 

138.03 13.U 14.U 15.U 16.U 17,U 18.U 19.U 20,4 21,2 22- 23.U 

138.04 24.U 25.U 26.U 27,1 28.U 29,2 30,1 3 32.U 33,2 34- 

138.05 35,14 36,70 37,2 38,1 39- 40- 41,1-3 42,U 43.U 44.U 

138.06 45.U 46,U 47,U 48,U 49.U 50.U 51,9 52,U 53,1 54,U 55.U 

138.07 56.U 57,U 58,U 59.U 60.U 61,3 62,3 63,3 64,3 65,1 66,1 

138.08 67,1 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

138.09 78,1 79,2 80,U 81.U 82.U 

139.01 oliga 

139.02 1,82 2,16.5 3,2 4,U 5,U 6,U 7,3 8,3 9,3 10.U ll.U 12.U 

139.03 13.U 14.U 15,U 16.U 17,U 18,3 19,2 20,1 21,1 22.3 23,2 

139.04 24,U 25,3 26,U 27,2 28,1 29,1 30,1 31,2 32,U 33,1 34,- 

139.05 35,5 36,82 37,1 38,1 39,1 40,1 41,3 42,12 43,1 44,U 45,U 

139.06 46,U 47,U 48,U 49,U 50,U 51,U 52,U 53,1 54,U 55.U 56.U 

139.07 57,2 58- 59- 60,U 61,3 62,3 63,3 64,1 65,2 66,1 67,2 

139.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

139.09 79,3 80.U 81,U 82,U 

140.01 ornata 

140.02 1,110 2,45 3,2 4,U 5,6 6,U 7,3 8,1 9,3 10,1 11,2 12,1 

140.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,3 23,2 

140.04 24,1 25,3 26,3 27,1-2 28,1 29,1 30,1 31,1 32,1 33,1-2 

140.05 34,1 35,5 36,110 37,1 38,1 39,1 40,0 41,20 42,U 43,2 

140.06 44,2 45,2 46,2 47,2 48,3 49,2 50,2 51,9 52,1 53,1 54,1 

140.07 55,3 56- 57,2 58- 59- 60,2 61,3 62,3 63,3 64,2 65,1 

140.08 66,1 67,2 68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1/5 

140.09 77,1 78,2 79,3 80,22-25 81,1 82,1 

141.01 ovalifera 

141.02 1,U 2,U 3,5 4,U 6,U 7,U 8,U 9,U 10,U ll.U 12,U 13,U 14.U

NUMBER 523 

403 

141.03 15.U 16,U 17.U 18,U 19.U 20,4 21,1 22,1 23,U 24.U 25.U 

141.04 26,U 27.U 28,U 29.U 30,1 31,1 32,U 33,U 34,U 35,5 36,U 

141.05 37,U 38.U 39.U 40,U 41,5-6 42.U 43,U 44.U 45,U 46,U 47,U 

141.06 48.U 49,U 50.U 51,U 52.U 53,1 54,U 55,U 56.U 57,1 58,U 

141.07 59.U 60- 61,5 62,5 63,5 64,3 65,U 66,1 67.U 68,1 69,1 

141.08 70,2 71,2 72,2 73,2 74,U 75,U 76.U 77,2 78,2 79,2 80,35 

141.09 81,U 82,U 

142.01 pacifica 

142.02 1,102 2,33 3,3 4.U 5,6 6,U 7,3 8,3 9,2 10,1 11,2 12,1 

142.03 13,1 14,2 15,1 16,3 17,3 18,1 19,2 20,1 21,1 22,1 23,2 

142.04 24,2 25,3 26,4 27,1 28,3 29,2 30,1 31,1 32,1 33,2 34,2 

142.05 35,17-21 36,102 37,1 38,1 39,U 40,U 41,5 42,40 43,2 44,2 

142.06 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,2 55,3 

142.07 56- 57,2 58- 59- 60,2 61,2 62,4 63,4 64,3 65,2 66,1 

142.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

142.09 78.U 79,2 80,23-28 81,1 82,1 

143.01 palauensis 

143.02 1,U 2,U 3,U 4,U 5,U 6,U 7,3 8,1 9,2 10,1 11,1 12,U 13,U 

143.03 14.U 15,1 16,1 17,1 18,3 19,2 20,1 21,1-2 22- 23,2 24,U 

143.04 25,4 26,3 27,1 28,1 29,2 30,1 31,2 32,1 33,U 34- 35,6-8 

143.05 36.U 37,2 38.U 39,5-9 40.U 41,3 42,U 43,2 44,2 45,1-2 

143.06 46,1-2 47,1-2 48,1-2 49,1-2 50,1-2 51.U 52,U 53,1 54,1 

143.07 55,U 56- 57,2 58- 59.U 60.U 61,1 62,1 63,U 64,3 65,2 

143.08 66,2 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 

143.09 77,2 78.U 79,2 80,19-22 81.U 82,U 

144.01 panamena 

144.02 l.U 2,U 3,3.5 4,U 5,6 6,U 7,3 8,2 9,3 10,1 11,2 12,2 

144.03 13,1 14,2 15,1 16,3 17,3 18,3 19,2 20,2 21,1 22,3 23,1 

144.04 24,2 25,3 26,4 27,1 28,1 29,1 30,1 31,1 32,3 33,2 34,1 

144.05 35,5 36,U 37,1 38,1 39,1 40.U 41,8 42,15 43,1 44,2 45,2 

144.06 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,1/3 56- 

144.07 57,U 58.U 59.U 60,U 61,1 62,1 63.U 64,2 65,2 66,1 67,2 

144.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,4 77,1 78,2 

144.09 79,3 80,27 81,1 82,1 

145.01 papeetensis 

145.02 1,U 2,U 3,2 4,U 5,2.5 6,U 7,3 8,3 9,2 10,1 11,2 12,1 

145.03 13,1 14,2 15,1 16,3 17,1 18,3 19,2 20,1 21,1 22,3 23,2 

145.04 24,2 25,3 26,5 27,1 28,1 29,1 30,1 31,2 32.U 33,2 34- 

145.05 35,6 36,30/33 37,2 38,2 39,2 40,5 41,3 42,U 43,2 44,2 

145.06 45,2 46,2 47,U 48,2 49,2 50.U 51,3 52,1 53,1 54,1 55,2 

145.07 56- 57.U 58,U 59.U 60.U 61,1 62,1 63,U 64,2 65,2 66,2 

145.08 67,2 68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,3 77,1 

145.09 78,2 79,3 80,21 81.U 82,U 

146.01 parasegregata 

146.02 1,U 2,U 3,3 4,U 5,8 6,U 7,3 8,1 9,2 10,1 11,2 12,2 13,1 

146.03 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,2 24,2 

146.04 25,4 26,2 27,2 28,2 29,2 30,1 31,1 32,1 33,2 34,1 35,3 

146.05 36,U 37,2 38,2 39,0 40.U 41,18 42,10 43,1 44,3 45,2 

146.06 46,1 47,1 48,2 49,1 50,1 51,5 52,1 53,1 54,1 55,3 56- 

146.07 57,2 58- 59- 60,2 61,1 62,1 63,1 64,2 65,2 66,1 67,2


146.08 68,1 69.1 70.2 71,2 72,2 73,1 74,2 75,1 76.1 77,1 78,2 

146.09 79,2 80.U 81,1 82,1 

147.01 parvibranchis 

147.02 1,U 2,U 3,4 4.U 5,10 6,U 7,3 8,1 

147.03 14,2 15.2 16,1 17.1 18,3 19,2 20,3 

147.04 25,4 26,4 27,1 28,1 29,1 30,1 31,2 

147.05 35,19-20 36.U 37,1 38,1 39.U 40.U 

9.2 

21.2 

32,1 

41.3 

10.1 

22.- 

33.2 

42.40 

11.2 

23.4 

34,- 

12.1 13 

24.2 

43. 1-2 44 ,U 

147.06 45,2 46,2 47,U 48,1 49,2 50,U 51,4-5 52,1 53,1 54,1 55,2 

147.07 56- 57.U 58.U 59.U 60.U 61,2 62,2 63.U 64,3 65,1 66,2 

147.08 67,1 68,2 69,1 70,2 71.2 72,2 73,2 74.U 75.1 76.1 77,2 

147.09 78,U 79,2 80,17-19 81,1 82,1 

148.01 paupera 

148.02 1,127 2,U 3,U 4,U 5.U 6.U 7.U 8.U 9.U 10.U ll.U 12.U 

148.03 13.U 14.U 15.U 16.U 17.U 18.U 19.U 20,3 21,2 22,- 23,4 

148.04 25,U 26,4 27,1 28.U 29.U 30.1 31,2 32.U 33.U 34,U 35,23 

148.05 36,U 37.U 38,U 39.U 40.U 41,3 42.U 43.U 44.U 45.U 46.U 

148.06 47,U 48.U 49.U 50.U 51,3 52.U 53.1 54.U 55.U 56.U 57.2 

148.07 58.U 59.U 60.U 61,4 62,4 63.4 64.3 65.U 66.U 67.U 68.U 

148.08 69,1 70.U 71,2 72,2 73,2 74,U 75.U 76.U 77,2 78.U 79,2 

148.09 80,23 81,U 82.U 

149.01 pauroneurata 

149.02 1,U 2,U 3,5.5 4.U 5,6 6,U 7,3 8.1 9.3 10.1 11.2 12.1 

149.03 13.U 14,U 15.1 16,1 17,6 18,3 19,2 20,1 21,1 22,1 23.U 

149.04 24,2 25,3 26,3 27,1 28,2 29,2 30,1 31,1 32,3 33,2 34.2 

149.05 35,8 36.U 37,1 38,1 39,1 40.U 41,5 42.15 43.1-2 44.2 

149.06 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1 55,3 

149.07 56- 57,2 58- 59- 60,2 61,1 62,1 63,1 64,3 65,2 66,1 

149.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

149.09 78.U 79,2 80,27 81,1 82,1 

150.01 pectinata 

150.02 l.U 2,U 3,2 4,U 5,6 6,U 7,3 8,1 93 10,1 11,2 12,1 13,1 

150.03 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,2 23,2 24,1 

150.04 25,4 26,3 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,2 35,5 

150.05 36,U 37,2 38,2 39,2 40,U 41,10 42,U 43,1-2 44,2 45,1 

150.06 46,1 47,U 48,1 49,1 50,U 51,3 52,1 53,1 54,1 55,3 56- 

150.07 57.U 58.U 59.U 60,U 61,2 62,2 63,U 64,1 65,2 66,2 67,2 

150.08 68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

150.09 79,2 80,41 81,1 82,1 

151.01 pelamidis 

151.02 1,113 2,100 3,12 4,U 5,20 6,U 7,2 8,2 9,2 10,U ll.U 12.U 

151.03 13,1 14,2 15,U 16,U 17.U 18,3 19,2 20,1 21,1 22,3 23,1 

151.04 24,U 25,3 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

151.05 35,3 36,220 37,1 38,1 39,0 40,0 41,24 42,20 43,3 44,2 

151.06 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 55,2 

151.07 56- 57,2 58- 59- 60,1 61,1 62,1 63,1 64,1 65,1 66,1 

151.08 67,2 68,2 69,1 70,U 71,1 72,2 73,2 74,U 75,1 76,1 77,2 

151.09 78.U 79,3 80,63 81,2 82,1 

152.01 pellucida 

152.02 1,U 2,U 3,2 4.U 5,3.5 6,U 7,3 8,1 9,2 10,1 11,2 12,2 

152.03 13,2 14- 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,3 23,2 

152.04 24,2 25,3 26,4 27,1 28,1 29,1 30,1 31,1 32,U 33,2 34,1

NUMBER 523 

405 

152.05 35,5 36,28 37,2 38,2 39.U 40.U 41,8 42.U 43,2 44,2 45,2 

152.06 46,2 47.U 48,2 49,2 50.U 51,4 52,1 53,1 54,1 55,3 56- 

152.07 57,2 58- 59- 60,2 61,2 62,2 63.U 64,3 65,2 66,1 67,2 

152.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,4 77,1 78,2 

152.09 79,3 80,21/23 81,1 82,1 

153.01 pennata 

153.02 1,114 2,73 3,3 4.U 5,7.5 6,U 7,2 8,1 9,2 10,1 11,2 12,1 

153.03 13,1 14,1 15,1 16,3 17,4 18,3 19,2 20,1 21,1 22,2 23.U 

153.04 24,2 25,4 26,2 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

153.05 35,3 36,39-41 37,2 38,2 39,5 40,2 41,12 42,15 43,1 44,3 

153.06 45,2 46,2 47,2 48,3 49,1 50,1 51,5-6 52,1 53,1 54,1 55,2 

153.07 56,- 57,2 58,- 59- 60,1 61,4 62,3 63,3 64,2 65,2 66,1 

153.08 67,2 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

153.09 78,2 79,2 80,35^3 81,1 82,2 

154.01 perimensis 

154.02 l,U 2,U 3,5 4,70 5,10 6,U 7,1 8,2 9,3 10,1 11,2 12,1 

154.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,4 

154.04 24,2 25,3 26,2 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,2 

154.05 35,17 36.U 37,1 38,1 39,1 40,U 41,8 42,30 43,1-2 44,2 

154.06 45,2 46,2 47.U 48,2 49,2 50.U 51,9 52,1 53,1 54,1 55,2 

154.07 56,- 57.U 58,U 59.U 60.U 61,4 62,4 63,U 64,3 65,1 66,2 

154.08 67,1 68,2 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,2 

154.09 78,2 79,2 80,28 81,1 82,1 

155.01 perrieri 

155.02 1,185 2,135 3,5 4,U 5,12 6,U 7,3 8,3 9,3 10,1 11,2 12,2 

155.03 13,1 14,6 15,1 16,1 17,1 18,1 19,2 20,1 21,2 22- 23,1 

155.04 24,1 25,3 26,3 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 

155.05 35,18 36,170 37,1 38,1 39,2-3 40,4-5 41,10 42,30 43,2 

155.06 44,2 45,2 46,2 47,2 48,3 49,1 50,2 51,9 52,2 53,1 54,2 

155.07 55,2 56- 57,2 58- 59- 60,2 61,1 62,1 63,4 64,3 65,1 

155.08 66,2 67,1 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

155.09 77,2 78,1 79,2 80,25 81,1 82,1 

157.01 peter si 

157.02 1,302 2,130 3,5 4,U 5,7 6,U 7,3 8,1 9,3 10,1 11,1 12,2 

157.03 13,1 14,2 15,3 16,1 17,1 18,1 19,2 20,3 21,1 22,1 23,4 

157.04 24,1 25,4 26,3 27,1 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

157.05 35,13 36,200 37,1 38,1 39.U 40,130 41,4 42,U 43,2 44,2 

157.06 45,2 46,2 47,2 483 49,3 50,1 51,4 52,1 53,1 54,2 55,1 

157.07 56- 57,2 58- 59- 60,1 61,1 62,1 63,1 64,2 65,2 66,1 

157.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

157.09 78,1 79,2 80,26 81,1 82,1 

158.01 philocorallia 

158.02 1,136 2,115 3,6 4,U 5,6 6,U 7,3 8,1 9,2 10,1 11,2 12,2 

158.03 13,1 14,3 15,3 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,2 

158.04 24,2 25,3 26,3 27,2 28,1 29,1 30,1 31,1 32,3 33,2 34,2 

158.05 35,6 36,135 37,1 38,1 39,1 40,0 41,4 42,U 43,3 44,3 45,2 

158.06 46,2 47,2 48,1 49,1 50,1 51,3 52,2 53,1 54,1 55,2 56- 

158.07 57,2 58- 59- 60,1 61,2 62,3 63,3 64,1 65,2 66,1 67,2 

158.08 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 

158.09 79,2 80,34 81,1 82,4

406 


159.01 plicata 

159.02 1,126 2,80 3,4 4,U 5,8 6,U 7,3 8,3 9,3 10,1 11,2 12,1 

159.03 13.U 14.U 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,1 

159.04 24,1 25,3 26,9 27,1 28,4 29,2 30,1 31,2 32,1 33,2 34,2 

159.05 35,7 36,115 37,1 38,1 39,1 40,30 41,5 42,U 43,3 44,2 

159.06 45,2 46,2 47,2 48,3 49,1 50,2 51.U 52,1 53,1 54,2 55,1 

159.07 56- 57,1 58,2 59,4 60- 61,1 62,1 63,1 64,1 65,2 66,2 

159.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

159.09 78.U 79,2 80,25 81,1 82,1 

160.01 polybranchia 

160.02 l.U 2.U 3,3 4,U 5,8 6,U 7,3 8,3 9,2 10,1 11,1 12,2 13,1 

160.03 14,3 15,1 16,2 17,1 18,3 19,2 20,1 21,2 22- 23,2 24,1 

160.04 25,3 26,4 27,2 28,2 29,2 30,1 31,1 32,1 33,2 34,1 35,7-8 

160.05 36,U 37,1 38,1 39,1 40.U 41,5 42,15 43,3 44,2 45,2 46,2 

160.06 47.U 48,2 49,2 50,U 51,4 52,1 53,1 54,1 55,1 56- 57,2 

160.07 58- 59- 60,1 61,4 62,4 63,U 64,3 65,1 66,2 67,1 68,2 

160.08 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 78,1 79,2 

160.09 80,24-36 81,1 82,1 

161.01 prayensis 

161.02 1,U 2,U 3,2 4,U 5,4.5 6,U 7,3 8,3 9,2 10,1 11,1 12,2 

161.03 13,1 14,3 15,1 16,1 17.U 18,1 19,2 20.U 21.U 22,U 23.U 

161.04 24,2 25,4 26,1 27.U 28,U 29,U 30,1 31,1 32,1 33,2 34,1 

161.05 35,5 36,33 37,2 38,2 39,1 40,0 41,9 42,12 43,1 44,2 45,2 

161.06 46,2 47,U 48,2 49,2 50.U 51,2 52,1 53,1 54,1 55,2 56,- 

161.07 57,2 58- 59- 60,1 61,1 62,1 63,U 64,3 65,2 66,1 67,2 

161.08 68,1 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

161.09 79,3 80,18 81,1 82,2 

162.01 prognatha 

162.02 1,122 2,80 3,7.5 4,U 5,16 6,U 7,3 8,1 9,2 10,1 11,2 12,1 

162.03 13,U 14.U 15,1 16,2 17,1 18,3 19,2 20,3 21,1 22,1 23,1 

162.04 24,2 25,3 26,5 27.U 28,U 29,U 30,1 31,2 32,3 33,2 34- 

162.05 35,10/13 36,110 37,1 38,1 39,U 40,U 41,3 42,U 43,2-3 

162.06 44,3 45,2 46,2 47,2 48,2 49,2 50,2 51,9 52,1 53,1 54,1 

162.07 55,2 56- 57,2 58- 59- 60,2 61,1 62,4 63,4 64,3 65,2 

162.08 66,2 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,2 76,1 

162.09 77,2 78.U 79,2 80,35 81,1 82,1 

163.01 pulvinopalpata 

163.02 1,U 2,U 3,11 4,U 5,23 6,U 7,3 8,3 9,3 10,1 11,2 12,2 

163.03 13.U 14.U 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,1 

163.04 24,1 25,3 26,3 27,2 28,1 29,1 30,1 31,1 32,3 33,2 34,2 

163.05 35,3 36,U 37,1 38,1 39,1 40,0 41,25 42,15 43,2 44,3 45,2 

163.06 46,2 47,2 48,2 49,2 50,2 51,5-6 52,3 53,1 54,1 55,2 56- 

163.07 57,2 58- 59- 60,1 61,3 62,3 63,3 64,3 65,2 66,1 67,2 

163.08 68,1 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,2 78,2 

163.09 79,2 80,62-72 81,1 82,1 

164.01 purpurea 

164.02 1,106 2,53 3,4 4,U 5,9.5 6,U 7,3 8,1 9,3 10,1 11,2 12,2 

164.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,1 

164.04 24,1 25,3 26,4 27,1 28,2 29,1 30,1 31,1 32,1 33^2 34^1 

164.05 35,5 36,100 37,1 38,1 39,2 40,0 41,14 42,20 43,2 44,2* 

164.06 45,1 46,1 47,1 48,1 49,1 50,1 51,4 52,1 53,1 54,2 55,1


164.07 

164.08 

164.09 

165.01 

165.02 

165.03 

165.04 

165.05 

165.06 

165.07 

165.08 

165.09 

166.01 

166.02 

166.03 

166.04 

166.05 

166.06 

166.07 

166.08 

166.09 

167.01 

167.02 

167.03 

167.04 

167.05 

167.06 

167.07 

167.08 

167.09 

168.01 

168.02 

168.03 

168.04 

168.05 

168.06 

168.07 

168.08 

168.09 

169.01 

169.02 

169.03 

169.04 

169.05 

169.06 

169.07 

169.08 

169.09 

170.01 

170.02 

170.03 

56- 

67,2 

78,1 

57,2 58- 

68,2 i69,1 

79,2 80,19 

pycnobranchiata 

l.U 2,U 3,8 4,U 5,12 6,U 7,3 

14.U 

25,3 

36.U 

47.U 

58- 

69,1 

15,1 

26,5 

37,1 

48,3 

59- 

70,2 

16,3 

27,1 

38,1 

49,1 

60.U 

71,2 

17,2 

28,2 

39.U 

50.U 

61,4 

72,2 

18,3 

29,1 

40.U 

51,U 

62,4 

73,2 

19,2 

30,1 

41,4 

52,3 

63,U 

74.U 

80,28/29 81 ,1 8: 

quinquefida 

l.U 2,U 3 

13,1 

24,1 

35,7 

46,2 

57,2 

68,1 

14,3 

25,3 

36.U 

47.U 

58- 

69,1 

79,2 80,30 

59- 

70,2 

81,1 

U 

60,1 61,1 

71,2 72,2 

82,1 

,2.5 . 411 * i,8 6,U 7.3 

15,1 

26,5 

37,1 

48,2 

59- 

70,2 

81,1 

16,2 

27,1 

38,1 

49,2 

60,1 

71,2 

82,3 

17,1 

28,1 

39,0 

50,U 

61,3 

72,2 

18,3 

29,1 

40.U 

62,3 

73,2 

62,1 

73,2 

8,1 

20,1 

31,1 

63,4 64,2 65,2 66,2 

74,1 75,1 76,1 77,2 

9,3 10,1 11,2 12,1 13.U 

21,1 22,2 23,1 24,1 

32,2 33,2 34.U 35,5-6 

42.U 43,2 44,3 45,2 46,2 

53,1I 54,1 55,3 56- 57,2 

64,: I 65,2 66,2 67,2 68,2 

75,1 76,1 77,2 78,1 79,2 

8,1 

19,2 

30,1 

9,3 

20,1 

31,1 

10,1 

21,1 

32,1 

11,2 

22,2 

33,2 

12,1 

23.U 

34,1 

41,5 42,15 43,3 44,2 45,2 

51,5-6 52,1 53,1 54,1 55,2 56- 

63,U 

74,1 

64,3 65,2 66,1 67,1 

75,1 76,1 77,2 78,1 

reducta 

1,298 2,3 IS; 3,6 4,U 5,15 6,U 12 8,3 9,2 10,1 11,2 12,2 

13,1 

24,1 

14,2 

25,3 

15,1 

26,4 

16,1 

27,1 

17,1 

28,1 

18,3 

29,2 

19,2 

30,1 

20,3 

31,1 

21,1 

32,1 

22,1 

33,2 

23,1 

34,1 

35,4 36,131 37,2. 38,2: 39,0i 40,6. 41,21 42.U 43,1 44,2 

45,2 

56- 

67,2 

78,U 

46,2 

57,2 

68,2 

79,2 

47,2 

58- 

69,1 

80,45 

48,2 

59- 

70,2 

81,1 

49,2 

60,2 

71,2 

82,1 

50,2 

61,2 

72,2 

51,4 

62,3 

73,2 

52,1 

63,4 

74.U 

53,1 

64,2 

75,1 

54,1 

65,2 

76,1 

55,4 

66,2 

77,2 

rosaurae 

1,142 2,152• 3,4 4,U 5,18 6,U 12 8,2 9,3 10,1 11,2 12,2 

13,1 

24,2 

14,3 

25,1 

15,1 

26,3 

16,1 

27,1 

17,1 

28,3 

18,3 

29,2 

19,2 

30,1 

20,2 

31,3 

21,1 

32,3 

22,2 

33,2 

23,1 

34- 

35,3 36,48-55 37,2 38,2 39- 40- 41- 42- 43,1-2 

44,1-2 45,1I 46,2 47,: \ 48,1 49,2\ 50,2 51,5 52,3 53,1 54,1 

55,2 56- 57,1 58,2 59,3 60- 61,1 62,1 63,4 64,3 65,2 

66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

112 78,1 79,2/4 80,32-38 81,1 82,1 

roussaei 

l.U 2,U 3,U 4,U 5,U 6.U 

14,U 

25 ,U 

36,U 

47.U 

58.U 

69,1 

80,U 

15,U 

26,U 

37,1 

48,3 

59.U 

70,2 

81.U 

16.U 

27,1 

38,1 

49,1 

60,U 

71,2 

82,3 

17.U 

28.U 

39,U 

50.U 

61,1 

72,2 

18,U 

29,U 

40,U 

51,U 

62,1 

73,2 

rubella 

l.U 

14,3 

2,U 3,2 4,U 5,8 6,U 

15,1 16,1 17,1 18,3 

7,U 8,U 9,U 10,U 11,U 12.U 13,U 

19,1J 20.U 21,1-2 22,1 23,1 24,U 

30,1 31,,1 32,1 33,U 34,1 35,6-8 

41,47 42,U 43,5 44,2 45,2 46,2 

52,U 53, ,1 54,2 55,1 56- 57,U 

63,1J 64,3 65,2 66,1 67,2 68,2 

74,1 75,1 76,1 77,2 78,1 79,2 

7,1 

19,2 

8,3 

20,3 

9,2 10,1 11,2 12,2 13,1 

21,1 22,2 23,4 24,2


170.04 25,1 26,3 27,1 28,1 29,1 30,1 31,2 32,1 33,2 34- 35,5 

170.05 36,U 37,1 38,1 39,2 40,U 41,2 42,7 43,1 44,2 45,2 46,2 

170.06 47,2 48,2 49,2 50,2 51,7 52,1 53,1 54,1 55,2 56- 57,2 

170.07 58- 59- 60,4 61,2 62,2 63,4 64,3 65,2 66,1 67,2 68,1 

170.08 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78.U 79,2 

170.09 80,28 81,1 82,1 

171.01 rubra 

171.02 1,U 2,U 3,3 4,U 5,7.5 6,U 7,2 8,2 9,2 10,2 11,2 12,2 

171.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,3 23.U 

171.04 24,2 25,3 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

171.05 35,4 36.U 37,1 38,1 39,0 40.U 41,21 42,15 43,2 44,2 45,2 

171.06 46,1 47,1 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 56- 

171.07 57,2 58- 59- 60,4 61,1 62,1 63,1 64,1 65,2 66,1 67,2 

171.08 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1/5 77,1 78,2 

171.09 79,3 80,27 81,1 82,2 

172.01 rubrivittata 

172.02 1,U 2,U 3,1.2 4,U 5,5.1 6,U 7,3 8,2 9,2 10,2 11,2 12,1 

172.03 13,U 14,U 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,1 

172.04 24,2 25,4 26,1 27,U 28.U 29,U 30,1 31,1 32,3 33,2 34,2 

172.05 35,3 36,56 37,2 38,2 39,6 40,11 41,5 42,25 43,2/4 44,2 

172.06 45,1 46,1 47,1 48,1 49,1 50,1 51,5-6 52,3 53,1 54,1 55,3 

172.07 56- 57,2 58- 59- 60,1 61,3 62,3 63,4 64,2 65.U 66.U 

172.08 67.U 68.U 69,1 70,1 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

172.09 78,2 79,3 80,31 81,1 82,1 

173.01 rubrocincta 

173.02 1,100 2,68 3,3 4,U 5,U 6,U 7,U 8,U 9,U 10.U ll.U 12.U 

173.03 13,U 14.U 15,U 16,U 17,U 18,U 19.U 20,3 21,2 22- 23,4 

173.04 24.U 25,4 26,U 27,2 28,2 29,2 30,1 31,1 32,1 33,U 34,1 

173.05 35,3 36,80 37,2 38,1 39.U 40,11 41,8 42.U 43.U 44,U 45.U 

173.06 46.U 47.U 48,U 49.U 50.U 51,U 52,1 53,1 54,U 55,U 56,U 

173.07 57,2 58- 59- 60.U 61,2 62,2 63,2 64,3 65,1 66,2 67,1 

173.08 68,2 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

173.09 79,2 80.U 81,U 82.U 

174.01 rullieri 

174.02 1,72 2,24 3,3 4,U 5,5 6,U 7,3 8,1 9,3 10,1 11,2 12,2 

174.03 13,1 14,4 15,1 16,1 17,1 18,3 19,2 20,3 21,2 22- 23,1 

174.04 24,1 25,1 26,5 27,1 28,5 29,2 30,1 31,1 32,1 33,2 34,2 

174.05 35,5 36,70 37,1 38,1 39,0 40,4-5 41,5 42,10 43,2 44,3 

174.06 45,2 46,2 47,2 48,3 49,2 50,2 51,4 52,1 53,1 54,1 55,2 

174.07 56- 57,2 58- 59- 60,4 61,2 62,2 63,2 64,3 65,2 66,1 

174.08 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

174.09 78,1 79,2 80,19 81,1 82,3 

175.01 samoae 

175.02 l.U 2,U 3,4 4,15 5,5 6,U 7,2 8,2 9,3 10,1 11,2 12,2 13.U 

175.03 14.U 15,1 16,1 17,1 18,3 19,2 20,2 21,1 22,1 23,4 24,2 

175.04 25,4 26,3 27,1 28,3 29,2 30,1 31,1 32,1 33,2 34,1 35,6 

175.05 36,51 37,2 38,2 39,2 40,10 41,6 42,15 43,2 44,3 45,2 

175.06 46,2 47,2 48,3 49,1 50,2 51,5 52,1 53,1 54,2 55,2 56- 

175.07 57,2 58- 59- 60,1 61,1 62,1 63,1 64,3 65,1 66,2 67,1 

175.08 68,2 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,2 78,1 

175.09 79,2 80,26 81,1 82,1


176.01 savignyi 

176.02 1,U 2,U 3,U 4,U 5,U 6,U 7,U 8,U 9,U 10.U 11,U 12,U 13,U 

176.03 14.U 15,U 16.U 17,U 18.U 19,U 20,U 21,1 22,1 23.U 24,U 

176.04 25.U 26.U 27,2 28.U 29,1 30,1 31,1 32.U 33.U 34.U 35,4 

176.05 36,45 37,2 38,2 39.U 40.U 41,12 42.U 43,U 44.U 45.U 46,U 

176.06 47.U 48,U 49.U 50.U 51.U 52,U 53.U 54,U 55,U 56.U 57,U 

176.07 58.U 59,U 60.U 61,U 62.U 63,U 64.U 65.U 66.U 67.U 68,U 

176.08 69,1 70,U 71,2 72,2 73,1 74,U 75,U 76.U 77,1 78,U 79,2 

176.09 80.U 81.U 82.U 

177.01 schizobranchia 

177.02 1,731 2,655 3,5 4,10 5,10 6,5 7,2 8,2 9,2 10,1 11,2 12,1 

177.03 13.U 14,U 15,1 16,3 17,1 18,3 19,2 20,1 21,2 22- 23,4 

177.04 24,1 25,3 26,5 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 

177.05 35,67 36,730 37,1 38,1 39,100 40,0 41,7 42.U 43,3 44,2 

177.06 45,2 46,2 47,2 48,3 49,1 50,1 51,9 52,1 53,1 54,1 55,2 

177.07 56- 57,1 58,2 59,1 60- 61,1 62,1 63,1 64,3 65,1 66,1 

177.08 67.1 68,1 69,1 70,2 71,2 72,2 73,2 74,U 75,1 

177.09 76,1 77,1 78.U 79,2 80,60 81,2 82,1 

178.01 sebastiani 

178.02 1,706 2,1455 3,23 4,25 5,35 6,21 7,2 8,1 9,2 10,1 11,2 

178.03 12,2 13.U 14.U 15,1 16,3 17,1 18,2 19,2 20,1 21,2 22- 

178.04 23,4 24,1 25,3 26,5 27,1 28,4 29,2 30,1 31,1 32,1 33,2 

178.05 34,1 35,7-14 36,686 37,1 38,1 39,0 403 41,40 42,35 

178.06 43,1/5 44,2 45,2 46,2 47,2 48,1 49,1 50,1 51,6-13 52,1 

178.07 53,1 54,2 55,4 56- 57,1 58,2 59,4 60,- 61,1 62,1 63,3 

178.08 64,3 65,1 66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,2 74,2 

178.09 75,1 76,1 77,2 78,1 79,1 80,51/56 81,1 82,5 

179.01 segregata 

179.02 l.U 2,U 3,4 4,15 5,5 6,3.5 7,1 8,2 9,2 10,1 11,2 12,1 

179.03 13,1 14,3 15,3 16,2 17,1 18,3 19,2 20,3 21,1 22,1 23,2 

179.04 24,1 25,4 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

179.05 35,3 36,39 37,2 38,2 39,2 40,0 41,15 42,25 43,3 44,2 

179.06 45,2 46,2 47,U 48,3 49,1 50.U 51,7 52,1 53,1 54,1 55,3 

179.07 56- 57,2 58- 59- 60,2 61,3 62,3 63,U 64,2 65,1 66,1 

179.08 67,2 68,2 69,1 70,2 71,2 72,2 73,1 74,1/2 75,1 76,1 77,1 

179.09 78,2 79,2 80,36 81,1 82,1 

180.01 semisegregata 

180.02 1,U 2,U 3,6 4.U 5,9 6,U 7,3 8,1 9,3 10,1 11,2 12,1 13,1 

180.03 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,2 24,2 

180.04 25,4 26,2 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 35,3 

180.05 36,65/69 37,2 38,2 39,0 40,0 41,38 42,15 43,2 44,2 45,2 

180.06 46,2 47.U 48,2 49,2 50,U 51,4 52,1 53,1 54,1 55,2 56- 

180.07 57,2 58- 59- 60,1 61,3 62,3 63,U 64,3 65,U 66.U 67,U 

180.08 68,U 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

180.09 79,2 80,51/55 81,1 82,1 

181.01 sonorae 

181.02 1,586 2,345 3,6 4,U 5,14 6,U 7,2 8,2 9,3 10,1 11,2 12,2 

181.03 13,1 14,2 15,1 16,1 17,1 18,1 19,2 20,1 21,1 22,1 23,2 

181.04 24,1 25,4 26,2 27,1 28,1 29,2 30,1 31,1 32,1 33,2 34,2 

181.05 35,41 36,530 37,2 38,1 39,0 40,55 41,6 42,65 43,1-2 

181.06 44,2 45^ 46,2 47,2 48,3 49,3 50,3 51,4 52,1 53,1 54,2

410 


181.07 55,2 56- 57,1 58,2 59,2 60- 61,2 62,3 63,3 64,3 65- 

181.08 66- 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 

181.09 77,2 78,1 79,1 80,54 81,1 82,1 

182.01 spongicola 

182.02 1,152 2,68 3,1.75 4,10 5,6 6,1.75 7,1 8,2 9,3 10,1 11,2 

182.03 12,1 13,1 14,2 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 

182.04 23,4 24,2 25,2 26,5 27,1 28,2 29,2 30,1 31,2 32,1 33,2 

182.05 34- 35,24 36,150 37,1 38,1 39,5 40,3 41,3 42.U 43,1-3 

182.06 44,2 45,2 46,2 47,2 48,3 49,1 50,2 51,3 52,3 53,1 54,1 

182.07 55,3 56- 57,2 58- 59- 60,2 61,4 62,4 63,4 64,3 65,2 

182.08 66,2 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,4 

182.09 77,2 78.U 79,2 80,26 81,1 82,1 

183.01 stanleyi 

183.02 1,130 2,58 3,4 4,U 5,7 6,U 7,3 8,1 9,3 10,1 11,2 12,2 

183.03 13,1 14,2 15,1 16,2 17,1 18,3 19,2 20,3 21,1 22,2 23,2 

183.04 24,2 25,3 26,3 27,1 28,1 29,1 30,1 31,2 32,1 33,2 34- 

183.05 35,4 36,33 37,2 38,2 39,8 40,8 41,2 42,12 43,1-2 44,2 

183.06 45,2 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 

183.07 56- 57,2 58- 59- 60,1 61,1 62,1 63,1 64,1 65,2 66,2 

183.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

183.09 78,U 79,2 80,29 81,1 82,1 

184.01 stigmatura 

184.02 1,U 2,U 3,2 4,U 5,4 6,U 7,3 8,1 9,2 10,1 11,2 12,1 13,1 

184.03 14,2 15,1 16,1 17,1 18,3 19,2 20,3 21,1 22,1 23,1 24,2 

184.04 25,4 26,2 27,2 28,2 29,1 30,1 31,1 32,2 33,2 34,1 35,3 

184.05 36,110-126 37,1 38,1 39,4 40,40 41,5 42,15 43,1 44,3 

184.06 45,2 46,2 47,2 48,2 49,2 50,2 51,6 52,1 53,1 54,1 55,2 

184.07 56- 57,2 58- 59- 60,2 61,3 62,3 63,3 64,2 65,2 66,1 

184.08 67,2 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

184.09 78,2 79,2-3 80,27-35 81,1 82,1 

185.01 suviensis 

185.02 1,U 2,370 3,9 4,U 5,U 6,U 7,U 8,U 9,U 10.U ll.U 12,U 

185.03 13,1 14,6 15,1 16,U 17,U 18,3 19,2 20.U 21,2 22- 23,4 

185.04 24,U 25,U 26,U 27,1 28,U 29.U 30,1 31,1 32,1 33,2 34,1 

185.05 35,9 36,160 37,1 38,1 39,U 40,U 41,9 42,U 43.U 44,U 45,U 

185.06 46,U 47,U 48,U 49,U 50,U 51,U 52,U 53,1 54,U 55.U 56.U 

185.07 57,2 58- 59- 60,U 61,3 62,3 63,3 64,3 65.U 66,2 67,U 

185.08 68,2 69,1 70,2 71,2 72,2 73,2 74.U 75,U 76,1 77,2 78.U 

185.09 79,2 80,U 81,U 82,U 

186.01 tentaculata 

186.02 1,U 2,U 3,U 4,U 5,U 6,U 73 8,U 9,2 10.U ll.U 12,U 13,1 

186.03 14,U 15.U 16,U 17,U 18,3 19,2 20,U 21,1 22,1 23,2 24,2 

186.04 25,4 26,3 27,2 28,1 29,1 30,1 31,1 32,U 33,2 34,1 35,6 

186.05 36,28 37,2 38,2 39,1 40,1 41,9 42,U 43.U 44,U 45.U 46,U 

186.06 47,U 48.U 49,U 50,U 51,U 52,U 53,1 54,U 55,U 56,U 57,2 

186.07 58- 59- 60,U 61,U 62,2 63,U 64,U 65,2 66,2 67,2 68,2 

186.08 69,1/2 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78 2 79 3 

186.09 80,23 81,1 82,5 

187.01 tennis 

187.02 1,695 2,460 3,2 4,U 5,4.5 6,U 7,U 8,U 9,3 10,U 11 U 12 U 

187.03 13,U 14,U 15,U 16,U 17,U 18,3 19,2 20,3 21,2 22- 23,2


187.04 24,U 25,4 26.U 27,1 28,5 29,2 30,1 31,2 32,1 33,2 34- 

187.05 35,81 36,695 37,1 38,1 39,10 40.U 41,4 42,400 43,2 44,2 

187.06 45,2 46,2 47,2 48,2 49,2 50,2 51,200 52,3 53,1 54,1 55,2 

187.07 56- 57,2 58- 59- 60,3 61,4 62,4 63,4 64,3 65,2 66,1 

187.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 

187.09 78.U 79,2 80,124 81,1 82,3 

188.01 thomasiana 

188.02 1,99 2,45 3,3 4,U 5,6 6,U 7,2 8,2 9,2 10,1 11,2 12,1 

188.03 13,1 14,3 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,2 23,2 

188.04 24,2 25,1 26,4 27,1 28,2 29,1 30,1 31,2 32,3 33,2 34- 

188.05 35,4 36,79 37,2 38,2 39,U 40,U 41,3 42,15 43,2 44,2 45,2 

188.06 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,2 55,2 56- 

188.07 57,2 58- 59- 60,1 61,4 62,5 63,4 64,1 65,1 66,2 67,1 

188.08 68,1 69,1 70,2 71,2 72,2 73,2 74.U 75,2 76,1 77,2 78,U 

188.09 79,2 80,22/24 81,1 82,2 

189.01 libiana 

189.02 1,165 2,73 3,3 4,U 5,5.5 6,U 7,2 8,2 9,2 10,1 11,2 12,1 

189.03 13,1 14,2 15,3 16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,2 

189.04 24,2 25,3 26,4 27,3 28,1 29,1 30,1 31,1 32,3 33,2 34,1 

189.05 35,6 36,135 37,2 38,1 39,1 40,10-15 41,5 42,U 43,2-3 

189.06 44,2-3 45,2 46,2 47,2 48,3 49,1 50,2 51,4 52,1/3 53,1 

189.07 54,2 55,2 56- 57,2 58- 59- 60,1 61,1 62,1 63,1 

189.08 64,3 65,1 66,2 67,1 68,1 69,1 70,2 71,2 72,2 73,2 74.U 

189.09 75,1 76,1 77,2 78.U 79,2 80,26 81,1 82,5 

190.01 torquata 

190.02 1,U 2,U 3,4 4,U 5,10.5 6,U 7,3 8,1 9,3 10,1 11,2 12,2 

190.03 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,3 23,2 

190.04 24,2 25,3 26,5 27,1 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

190.05 35,3 36.U 37,1 38,1 39,0 40,U 41,7 42,15 43,2 44,2 45,2 

190.06 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1 55,2 56- 

190.07 57,2 58- 59- 60,1 61,1 62,1 63,1 64,1 65,2 66,1 67,2 

190.08 68,1 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,2 78,1 

190.09 79,2 80,32 81,1 82,2 

191.01 torresiensis 

191.02 1,72 2,47 3,3 4,10 5,6.5 6,3 7,3 8,1 9,3 10,1 11,2 12,2 

191.03 13,1 14,3 15,3 16,2 17,1 18,1 19,2 20,1 21,1 22,3 23,2 

191.04 24,2 25,3 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,1 34,2 

191.05 35,5 36,70 37,1 38,1 39,1 40,0 41,7 42,15 43,1 44,2 45,2 

191.06 46,2 47,2 48,2 49,2 50,2 51,4 52,3 53,1 54,1 55,3 56- 

191.07 57,2 58- 59- 60,2 61,4 62,4 63,4 64,1 65.U 66,U 67,U 

191.08 68,U 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 78,2 

191.09 79,3 80,24 81,1 82,1 

192.01 tribranchiata 

192.02 l.U 2,U 3,7 4,U 5,13 6,U 7,3 8,1 9,3 10,1 11,2 12,1 13,U 

192.03 14,U 15,1 16,2 17,1 18,3 19,2 20,1 21,1 22,1 23,1 24,1 

192.04 25,3 26,4 27,1 28,4 29,1 30,1 31,1 32,1 33,2 34,2 35,4-5 

192.05 36.U 37,1 38,1 39,0 40,U 41,8 42,U 43,3 44,2 45,2 46,2 

192.06 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,3 55,2 56- 57,U 

192.07 58,U 59,U 60,U 61,1 62,1 63,U 64,1 65,2 66,2 67,2 68,2 

192.08 69,1 70,2 71,2 72,2 73,2 74,U 75,1 76,1 77,2 78,U 79,2 

192.09 80,30 81,1 82,1


193.01 tridentata 

193.02 1,185 2,170 3,7 4,10 5,12 6,7 7,3 8,1 9,3 10,1 11,2 12,2 

193.03 13.U 14.U 15,3 16,1 17,1 18,3 19,2 20,3 21,1 22,2 23,2 

193.04 24,1 25,3 26,4 27,2 28,1 29,2 30,1 31,1 32,1 33,2 34,1 

193.05 35,3 36,80 37,2 38,2 39,1 40,15 41,14 42,30 43,2-3 

193.06 44,2-3 45,2 46,1 47,1 48,2 49,1 50,1 51,4 52,1 53,1 54,1 

193.07 55,2 56- 57,2 58- 59- 60,2 61,2 62,1 63,2 64,3 65,2 

193.08 66,2 67,2 68,2 69,1 70,2 71,2 72,2 73,1 74,1 75,1 76,1 

193.09 77,1 78,2 79,2 80,51 81,2 82,1 

194.01 tubicola 

194.02 1,83 2,26.5 3,1.2 4,10 5,4 6,1.2 7,3 8,2 9,2 10,1 11,2 

194.03 12,1 13.U 14.U 15,1 16,1 17,1 18,3 19,2 20,3 21,2 22- 

194.04 23,4 24,2 25,3 26,4 27,1 28,2 29,2 30,1 31,3 32,2 33- 

194.05 34- 35,22/24 36,69 37,2 38,1 39- 40- 41,- 42- 43,3 

194.06 44,2 45,2 46,1 47,1 48,2 49,1 50,1 51,5 52,1 53,1 54,2 

194.07 55,1 56- 57,1 58,2 59,1 60- 61,2 62,4 63,4 64,3 65,2 

194.08 66,1 67,2 68,1 69,1 70,2 71,2 72,1 73,1 74,U 75,1 76,1 

194.09 77,2 78,U 79,2 80,21 81,1 82,1 

195.01 tubifex 

195.02 1,U 2,U 3,2 4,U 5,7 6,U 7,3 8,3 9,3 10,1 11,1 12,2 13.1 

195.03 14,2 15,1 16,3 17,1 18,3 19,2 20,3 21,2 22- 23,1 24,1 

195.04 25,1 26,4 27,1 28,4 29,2 30,1 31,1 32,1 33,2 34,2 35,17 

195.05 36,U 37,1 38,1 39.U 40,U 41,4 42,30 43,1 44,3 45,2 46,2 

195.06 47.U 48,3 49,2 50.U 51,5 52,1 53,1 54,2 55,1 56,- 57,1 

195.07 58,2 59,1 60- 61,2 62,1 63.U 64,3 65,1 66,1 67,1 68,1 

195.08 69,1 70,2 71,2 72,1 73,2 74.U 75,1 76,2 77,2 78,1 79,2 

195.09 80,29/32 81,1 82,1 

196.01 unidentata 

196.02 1,U 2,80-100 3,2-3 4,U 5,U 6,U 7,3 8,U 9,2 10,1 11,1 

196.03 12.U 13,1 14.U 15,1 16,1 17,1 18,1 19,1 20,1 21,2 22,- 

196.04 23,4 24,U 25.U 26.U 27,1 28,1 29,2 30,1 31,1 32,1 33,2 

196.05 34,1 35,23 36,U 37,2 38,2 39.U 40,U 41,5 42,35 43,1 44,2 

196.06 45.U 46,2 47.U 48.U 49,2 50,U 51.U 52,U 53,1 54,1 55,2 

196.07 56- 57.U 58.U 59.U 60,U 61.U 62,3 63.U 64,U 65,2 66,2 

196.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

196.09 78,1 79,1 80,20-25 81,U 82,U 

197.01 unifrons 

197.02 l.U 2,U 3,1 4,10 5,5 6,1 7,3 8,1 9,1 10,1 11,2 12,2 

197.03 13.U 14,U 15,1 16,2 17,1 18,3 19,2 20,3 21,1 22,1 23,1 

197.04 24,2 25,4 26,2 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 

197.05 35,3 36,45-54 37,2 38,2 39,4 40,10 41,5 42,15 43,3 44,2 

197.06 45,2 46,2 47,2 48,2 49,2 50,2 51,5 52,1 53,1 54,1 55,3 

197.07 56- 57,2 58- 59- 60,1 61,1 62,4 63,U 64,1 65,2 66,1 

197.08 67,2 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 77,1 

197.09 78,2 79,3 80,25-31 81,1 82,1 

198.01 valens 

198.02 1,179 2,140 3,9 4,15 5,12.5 6,8.5 7,2 8,2 9,2 10,1 11,2 

198.03 12,1 13,1 14,1 15,1 16,3 17,1 18,3 19,2 20,1 21,1 22,1 

198.04 23,2 24,2 25,3 26,3 27,1 28,1 29,1 30,1 31,1 32,1 33,2 

198.05 34,1 35,3 36,76 37,2 38,2 39,2 40,10 41,11 42,U 43,2-3 

198.06 44,3 45,2 46,2 47,2 48,3 49,1 50,1 51,9 52,1 53,1 54,2


198.07 55,2 56- 58- 59- 60,1 61,1 62,1 63,3 64,2 65,2 

198.08 66,1 67,2 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1 

198.09 77,1 78,2 79,2 80,43 81,1 82,2 

199.01 

199.02 

199.03 

199.04 

199.05 

199.06 

199.07 

199.08 

199.09 

200.01 

200.02 

200.03 

200.04 

200.05 

200.06 

200.07 

200.08 

validobranchiata 

l.U 2,U 3,4 4,U 5,4 6,U 7,3 

14.U 15,1 16,2 17,1 18,3 19,2 

25,3 26,3 27,3 28,1 29,2 30,1 

36,37^40 37,2 38,2 39,0 40,2-3 

45,2 46,2 47,2 48,2 49,2 50,2 

56- 57,2 58- 59- 60,2 61,3 

67,2 68,2 69,1 70,2 71,2 72,2 

78,2 79,2 80,28-37 81,1 82,1 

violaceomaculata 

1,215 2,170 3,7 

13,1 14,3 15,1 

24.1 25,3 26,5 

35,6-7 36,215 

45.2 46,2 47,2 

56- 57,2 58- 

67,2 68,1 69,1 

200.09 78.U 79,2 80,25-39 81,1 82,1 

201.01 vittata 

201.02 1,73 2,23.5 3,2 4,10 5,4 

201.03 13,1 14,2 15,1 16,1 17,1 

201.04 24,2 25,4 26,2 27,2 28,1 

201.05 35,3 36,23 37,2 38,2 39,0 

201.06 45,2 46,2 47,2 48,3 

201.07 56- 57,2 58- 59- 

201.08 67,2 68,1 69,1 70,2 

8,1 9,3 10,1 11,2 12,1 13.U 

20,1 21,2 22- 23,2 24,2 

31,1 32,1 33,2 34,1 35,3 

41,33 42,15 43,1 44,2 

51,3 52,1 53,1 54,1 55,3 

62,3 63.U 64,3 65,2 66,2 

73,1 74,2 75,1 76,1 77,1 

4,10 5,12 6,7 7,2 8,2 9,3 10,1 11,2 12,2 

16,1 17,1 18,3 19,2 20,1 21,1 22,1 23,1 

27,1-2 28,1 29,1 30,1 31,1 32,1 33,2 34,1 

37,1 38,1 39.U 40.U 41,20 42,15 43,1 44,3 

48.1 49,1 50,3 51,4 52,1 53,1 54,2 55,2 

59- 60,4 61,3 62,3 63,3 64,2 65,2 66,1 

70.2 71,2 72,2 73,2 74.U 75,1 76,1 77,2 

49,1 

60,1 

71,2 

201.09 78,2 79,3 80,20-22 81,1 82,5 

202.01 

202.02 

202.03 

202.04 

202.05 

202.06 

202.07 

202.08 

vittatopsis 

1,U 2,U 

13,1 14,3 

24,2 25,4 

35,3 36,U 

46,2 47,2 

57,2 58- 

68,1 69,1 

3,2.5 

15,1 

26,3 

37,2 

48,2 

59- 

70,2 

4,10 

16,1 

27,3 

38,2 

49,2 

60,2 

71,2 

202.09 79,3 80,39 81,1 82,1 

5,6 6,2.5 

17,1 18,3 

28,1 29,1 

39,2 

50,2 

61,2 

72,2 

6,2 7,3 8,1 9,1 10,1 11,2 12,2 

18,3 19,2 20,3 21,1 22,1 23,2 

29,1 30,1 31,1 32,2 33,2 34,1 

40,1 41,12 42,12 43,3 44,3 

50,1 51,4 52,1 53,1 54,1 55,2 

61.1 62,3 63,3 64,3 65,2 66,1 

72.2 73,1 74,2 75,1 76,1 77,1 

40.U 

51,4 

62,3 

73,1 

7,2 8,1 9,2 10,1 11,2 12,1 

19,2 20,1 21,1 22,2 23,1 

30,1 31,1 32,1 33,2 34,1 

41,10 42,U 43,2 44,2 45,2 

52.1 53,1 54,1 55,3 56- 

63 ,U 64,1 65,2 66,1 67,2 

74.2 75,1 76,1 77,1 78,2 

203.01 wasinensis 

203.02 1,84 2,17 3,2.5 4,10 5,6 6,2.5 7,3 8,1 9,3 10,2 11,1 

203.03 12,2 13,1 14,3 15,1 16,1 17,1 18,3 19,2 20,3 21,2 22- 

203.04 23,4 

203.05 34- 

203.06 45,2 

203.07 56- 

203.08 67,1 

24,2 

35- 

46,2 

57,2 

68,2 

25,3 

36- 

47,2 

58- 

69,1 

26,2 

37- 

48,3 

59- 

70,2 

27,1 

38- 

49,1 

60,3 

71,2 

28,1 

39- 

50,1 

61,3 

72,2 

203.09 78,U 79,2 80,27-31 81,1 82,1 

204.01 websteri 

204.02 1,150 2,120 3,4 4,10 5,8 6,4 

204.03 13,1 14,2 15,1 16,1 17,1 18,3 

29,2 

40- 

51,4 

62,4 

73,2 

30,2 

41- 

52,1 

63,4 

74,U 

31- 32- 

42- 43,3 

53,1 54,2 

64,3 

75,1 

65,1 

76,1 

33- 

44,2 

55,4 

66,2 

77,2 

7,1 8,2 9,2 10,1 11,2 12,2 

19,2 20,3 21,1 22,2 23,2


204.04 24,2 25,3 26,3 27,2 28,2 29,1 30,1 31,1 32,1 33,2 34,1 

204.05 35,3 36,53 37,2 38,2 39,1 40,10 41,15 42.U 43,2 44,2 

204.06 45,2 46,2 47,2 48,3 49,2 50,2 51,6 52,1 53,1 54,1 55,3 

204.07 56- 57,2 58- 59- 60,2 61,3 62,4 63,4 64,1 65,2 66,1 

204.08 67,2 68,1 69,1 70,2 71,2 72,2 73,1 74,2 75,1 76,1/3 77,1 

204.09 78,2 79,2 80,31 81,1 82,3 

205.01 woodwardi 

205.02 1,U 2,U 3,5 4,10 5,7 6,5 7,3 8,2 9,3 10,1 11,1 12,2 13.U 

205.03 14,U 15,1 16,2 17,1 18,3 19,2 20,3 21,1 22,1 23,2 24,2 

205.04 25,3 26,2 27,1 28,2 29,2 30,1 31,1 32,1 33,2 34,1 35,3 

205.05 36,39 37,2 38,2 39,0 40,4-5 41,12 42,15 43,1 44,2 45,2 

205.06 46,2 47,2 48,2 49,2 50,2 51,3 52,1 53,1 54,1 55,2 56- 

205.07 57,2 58- 59- 60,1 61,2 62,4 63.U 64,3 65.U 66.U 67.U 

205.08 68,U 69,1 70,1 71,2 72,2 73,1 74,1 75,1 76,1 77,1 78,2 

205.09 79,3 80,31 81,1 82,5 

206.01 parva 

206.02 1,59 2,17 3,.75 4,10 5,4 6,.75 7,3 8,1 9,2 10,1 11,2 

206.03 12,2 13,1 14,4 15,1 16,1 17,1 18,3 19,2 20,1 21,1 22,3 

206.04 23,2 24,2 25,3 26,2 27,2 28,1 29,1 30,1 31,1 32,1 33,1 

206.05 34,1 35,5 36,20 37,2 38,2 39,2 40,2 41,4 42,10 43,2 44,3 

206.06 45,2 46,2 47,2 48,2 49,2 50,2 51,4 52,1 53,1 54,1 55,2 

206.07 56- 57,2 58- 59- 60,2 61,3 62,4 63,4 64,2 65.U 66.U 

206.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,2 75,1 76,1 77,1 

206.09 78,2 79,3 80,24 81,1 82,1 

207.01 goodsiri 

207.02 1.N 2,N 3,2.5 4.N 5.N 6,N 7.N 8.N 9,3 10JM 11.N 12.N 

207.03 13,1 14,N 15,N 16JST 17,1 18.N 19JST 20,1 21,1 22,2 23,1 

207.04 24.N, 25,N 26.N 27,1 28,N 29,N 30,1 31,2 32,1 33,2 34- 

207.05 35,25 36,N 37.N 38Jsf 39,11 40,N 41,2 42,2 43,3 44,2 45.N 

207.06 46JSf 47,N 48,N 49,N 50JSf 51JST 52,N 53,1 54,1 55,2 56- 

207.06 5ZN 58.N 59.N 60.N 61,3 62,3 63 ,N 64,3 65,N 66,N 67.N 

207.07 68JST 69,1 70,2 112 72,2 73,2 74,1 75,1 76,1 77,2 78,1 

207.08 79,2 80.N 81,1 82,1 

208.01 scombrinis 

208.02 1,130 2,47 3,4 4,10 5,7 6,10 7,3 8,1 9,3 10,1 11,1 12,1 

208.03 13,U 14,U 15,1 16,1 17,1 18,3 19,2 20,1 21,2 22- 23,2 

208.04 24,1 25,3 26,9 27,1 28,1 29,2 30,1 31,1 32,1 33,2 34,2 

208.05 35,7 36,125 37,1 38,1 39,5 40,5 41,17 42,U 43,1 44,3 

208.06 45,2 46,2 47,2 48,3 49,1 50,1 51,5 52,1 53,1 54,2 55,1 

208.07 56- 57,1 58,2 59,1 60- 61,1 62,1 63,1 64,3 65- 66- 

208.08 67,2 68,2 69,1 70,2 71,2 72,2 73,2 74,1 75,1 76,1 77,2 

208.09 78,1 79,2 80,29 81,1 82,5

Appendix C 

Important publications on Eunice issued since 1985 and new taxa described. 

Dounas, C, and A. Koukouras 

1989. Some Observations on the Possible Synonymy of 

Eunice vittata (Delle Chiaje, 1825) and E. indica 

Kinberg, 1865 (Annelida, Polychaeta). Cahiers de 

Biologie Marine, 30(2):227-234. 

Hanlcy, J. Russell 

1986. Co-operative Effort in a New Species of Tube 

Dwelling Worm, Eunice metatropos (Polychaeta: 

Eunicidac). The Beagle, Occasional Papers of the 

Northern Territory Museum of Arts and Sciences, 

3(1):215-221. 

New species: Eunice metatropos. 

Lc6n-Gonzilcz, Jesus Angel de 

1988. A New Eunicid Polychaete from the Caribbean Coast 

of Mexico. Revista de Biologia Tropical, 36(1):75- 

79. 

New species: Eunice riojai. 

Miura, Tomoyuki 

1986. Japanese Polychaetes of the Genera Eunice and 

Euniphysa: Taxonomy and Branchial Distribution 

Patterns. Publications of the Seto Marine Biological 

Laboratory, 31(3/6):269-325. 

New species: Eunice masudai, E. yamamotoi, E. 

annulicirrata, E.fauchaldi, and E. tanseiae. 

New subspecies: E. northioidea brevibranchiata. 

1987. New or Little Known Species of the Family Eunicidae 

(Annelida Polychaeta) from Japan. Proceedings 

of the Japanese Society of Systematic 

Zoology, 36:1-9. 

New species: Eunice profunda, E. misakiensis. 

Nonato, E. 

1965. Eunice sebastiani sp. nov. (Annelida, Polychaeta). 

Boletin do Instituto Oceanogrdfico, Universidade de 

Sao Paulo, 14:133-139. 

Winsnes, Inger M. 

1989. Eunicid Polychaetes (Annelida) from Scandinavian 

and Adjacent Waters: Family Eunicidae. Zoologica 

Scripta. 18(4):483-500. 

415

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Andrews, E.A. 

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Anonymous 

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Benham, W.B. 

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Berthold, Arnold Adolph 

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Blanchard, E. 

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Buchanan, Florence 

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Pomona College Journal of Entomology and Zoology, 11


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