West African Brachyuran Crabs - Smithsonian Institution Libraries

West African Brachyuran Crabs 

(Crustacea: Decapoda) 

RAYMOND B. MANNING 

and 

L. B. HOLTHUIS 

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 306 

;

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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 3 0 6 



Raymond B. Manning 

and L. B. Holthuis 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1981

ABSTRACT 

Manning, Raymond B., and L. B. Holthuis. West African Brachyuran Crabs 

(Crustacea: Decapoda). Smithsonian Contributions to Zoology, number 306, 379 

pages, 88 figures, 1981.—The West African marine brachyuran crab fauna, 

comprising 218 named species in 120 genera and 26 familes, is surveyed. 

Sixteen new genera and 24 new species are recognized. Synonymies are 

updated for the tropical species, and all 300 + Eastern Atlantic species are 

listed. Original references and synonymies are provided for all 146 Eastern 

Atlantic genera. Synonymies have been compiled for all 36 currently recognized 

families of marine crabs. Twenty-nine families are represented in the 

Eastern Atlantic fauna. One family, Hexapodidae Miers, 1886, and one 

subfamily, Camptandriinae Stimpson, 1858 (Ocypodidae) are revised at the 

generic level. The genera Liocarcinus Stimpson, 1871 (Portunidae), Machaerus 

Leach, 1818 (Goneplacidae), and Lambdophallus Alcock, 1900, Paeduma Rathbun, 

1897, Parahexapus Balss, 1922, Pseudohexapus Monod, 1956, and Thaumastoplax 

Miers, 1881 (all Hexapodidae), are defined and recognized. It is 

suggested that the family Geryonidae Colosi, 1923, shows closest affinities 

with the family Portunidae Rafinesque, 1815. 

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 

cavemosa (Linnaeus). 

Library of Congress Cataloging in Publication Data 

Manning, Raymond B. 1934- 

West Africa brachyuran crabs (Crustacea: Decapoda) 

(Smithsonian contributions to zoology ; no. 306) 

Bibliography: p. 

Supt. of Docs, no.: SI 1.27:306 

1. Crabs—Africa, West—Classification, 2. Crustacea—Classification. 3. Crustacea—Africa, 

West—Classification. I. Holthuis, L. B., joint author. II. Title. III. Series: Smithsonian 

Institution. Smithsonian contributions to zoology ; no. 306. 

QL1.S54 no. 306 (QL444.M33] 591s [595.3'842] 80-607105

Contents 

Page 

Introduction 1 

Collections Studied 3 

Format Considerations 4 

Repositories and Abbreviations 5 

Acknowledgments 6 

Family RANINIDAE de Haan, 1839 6 

Subfamily NOTOPODINAE Serene and Umali, 1972 7 

Genus Ranilia H. Milne Edwards, 1837 7 

* Ranilia constricta (A. Milne Edwards, 1880) 7 

Subfamily RANININAE de Haan, 1839 9 

Genus Cyrtorhina Monod, 1956 9 

Cyrtorhina granulosa Monod, 1956 9 

Genus Raninoides H. Milne Edwards, 1837 10 

* Raninoides bouvieri Capart, 1951 10 

Family HOMOLODROMIIDAE Alcock, 1899 11 

Family DROMIIDAE de Haan, 1833 11 

Genus Dromia Weber, 1795 11 

Dromia bollorei Forest, 1974 11 

Dromia marmorea Forest, 1974 11 

* Dromia monodi Forest and Guinot, 1966 15 

Dromia nodosa A. Milne Edwards and Bouvier, 1898 19 

Genus Sternodromia Forest, 1974 19 

* Sternodromia spinirostris (Miers, 1881) 19 

Family DYNOMENIDAE Ortmann, 1892 23 

Genus Dynomene Desmarest, 1823 23 

* Dynomene filholi Bouvier, 1894 23 

Family LATREILLIIDAE Stimpson, 1858 24 

Genus Latreillia Roux, 1830 24 

Latreillia elegans Roux, 1830 25 

Family HOMOLIDAE de Haan, 1839 25 

Genus Homola Leach, 1815 25 

* Homola barbata (Fabricius, 1793) 25 

Genus Paromola Wood-Mason and Alcock, 1891 27 

Paromola cuvieri (Risso, 1816) 27 

Family CYCLODORIPPIDAE Ortmann, 1892 28 

Family CYMONOMIDAE Bouvier, 1898 28 

Family DORIPPIDAE MacLeay, 1838 28 

Key to Subfamilies and Genera of Dorippidae 29 

Subfamily DORIPPINAE MacLeay, 1838 30 

Genus Medorippe, new genus 31 

iii

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

* Medorippe lanata (Linnaeus, 1767), new combination 31 

Genus Phyllodorippe, new genus 35 

* Phyllodorippe armata (Miers, 1881), new combination 36 

Subfamily ETHUSINAE Guinot, 1977 38 

Genus Ethusa Roux, 1830 38 

* Ethusa rosacea A. Milne Edwards and Bouvier, 1897 38 

* Ethusa rugulosa A. Milne Edwards and Bouvier, 1897 39 

* Ethusa vossi, new species 39 

Genus Ethusina Smith, 1884 42 

Key to Eastern Atlantic Species of Ethusina 43 

Ethusina alba (Filhol, 1884) 43 

* Ethusina beninia, new species 46 

Family CALAPPIDAE de Haan, 1833 49 

Subfamily CALAPPINAE de Haan, 1833 49 

Genus Acanthocarpus Stimpson, 1871 49 

* Acanthocarpus brevispinis Monod, 1946 50 

Genus Calappa Weber, 1795 51 

Calappa gallus (Herbst, 1803) 51 

Calappa granulata (Linnaeus, 1758) 51 

* Calappa pelii Herklots, 1851 52 

* Calappa rubroguttata Herklots, 1851 54 

Genus Cycloes de Haan, 1837 56 

Cycloes cristata (Brulle, 1837) 56 

Subfamily MATUTINAE de Haan, 1835 56 

Genus Matuta Weber, 1795 56 

Matuta michaelseni Balss, 1921 56 

Family LEUCOSIIDAE Samouelle, 1819 57 

Subfamily EBALIINAE Stimpson, 1871 59 

Genus Atlantotlos Doflein, 1904 59 

* Atlantotlos rhombifer Doflein, 1904 59 

Genus Ebalia Leach, 1817 60 

* Ebalia qffinis Miers, 1881 60 

Ebalia cranchii Leach, 1817 61 

Ebalia nux A. Milne Edwards, 1883 61 

* Ebalia tuberculata Miers, 1881 61 

Ebalia tuberosa (Pennant, 1777) 63 

Genus Merocryptus A. Milne Edwards, 1873 64 

Merocryptus obsoletus A. Milne Edwards and Bouvier, 1898 ... 64 

Subfamily ILIINAE Stimpson, 1871 64 

Genus Ilia Leach, 1817 64 

Ilia nucleus (Linnaeus, 1758) 64 

* Ilia spinosa Miers, 1881 64 

Subfamily LEUCOSIINAE Samouelle, 1819 65 

Genus Philyra Leach, 1817 65 

Philyra cristata Miers, 1881 66 

* Philyra laevidorsalis Miers, 1881 66 

Page

NUMBER 306 

Genus Pseudomyra Capart, 1951 66 

* Pseudomyra mbizi Capart, 1951 66 

Family BELLIIDAE Dana, 1852 67 

Family ATELECYCLIDAE Ortmann, 1893 68 

Genus Atelecyclus Leach, 1814 68 

Atelecydus rotundatus (Olivi, 1792) 68 

Atelecyclus undecimdentatus (Herbst, 1783) 69 

Family THIIDAE Dana, 1852 69 

Genus Thia Leach, 1815 69 

Thia scutellata (Fabricius, 1793) 69 

Family CANCRIDAE Latreille, 1803 69 

Family PIRIMELIDAE Alcock, 1899 69 

Genus Pirimela Leach, 1816 70 

Pirimela denticulata (Montagu, 1808) 70 

Genus Sirpus Gordon, 1953 70 

Sirpus monodi Gordon, 1953 70 

* Sirpus gordonae, new species 70 

Family CORYSTIDAE Samouelle, 1819 72 

Genus Nautilocorystes H. Milne Edwards, 1837 72 

Nautilocorystes ocellatus (Gray, 1831) 72 

Family BYTHOGRAEIDAE Williams, 1980 72 

Family PORTUNIDAE Rafinesque, 1815 72 

Subfamily CARCININAE MacLeay, 1838 75 

Genus Carcinus Leach, 1814 75 

Carcinus maenas (Linnaeus, 1758) 75 

Genus Xaiva MacLeay, 1838 75 

Xaiva biguttata (Risso, 1816) 75 

Xaiva mcleayi (Barnard, 1947) 76 

Subfamily POLYBIINAE Ortmann, 1893 76 

Genus Bathynectes Stimpson, 1871 76 

Bathynectes maravigna (Prestandrea, 1839), new combination .. 76 

* Bathynectes piperitus, new species 77 

Genus Liocarcinus Stimpson, 1871 83 

Liocarcinus arcuatus (Leach, 1814), new combination 84 

Liocarcinus corrugatus (Pennant, 1777), new combination 84 

Genus Macropipus Prestandrea, 1833 85 

Macropipus australis Guinot, 1961 85 

* Macropipus rugosus (Doflein, 1904) 86 

Subfamily PORTUNINAE Rafinesque, 1815 87 

Genus Callinectes Stimpson, 1860 87 

Key to West African Species of Callinectes 88 

* Callinectes amnicola (De Rochebrune, 1883), new combination 88 

* Callinectes marginatus (A. Milne Edwards, 1861) 92 

* Callinectes pallidus (De Rochebrune, 1883), new combination 95 

Genus Cronius Stimpson, 1860 98 

* Cronius ruber (Lamarck, 1818) 98 

Page

vj SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

Genus Portunus Weber, 1795 100 

Portunus hastatus (Linnaeus, 1767) 101 

* Portunus inaequalis (Miers, 1881) 102 

* Portunus validus Herklots, 1851 103 

Portunus vocans (A. Milne Edwards, 1878) 107 

Genus Thalamita Latreille, 1829 107 

Thalamita poissonii (Audouin, 1826) 107 

Family GERYONIDAE Colosi, 1923 108 

Genus Geryon Kr0yer, 1837 109 

Geryon affinis A. Milne Edwards and Bouvier, 1894 110 

* Geryon maritae, new species 112 

Family PLATYXANTHIDAE Guinot, 1977 118 

Family XANTHIDAE MacLeay, 1838 118 

Genus Cataleptodius Guinot, 1968 120 

* Cataleptodius floridanus (Gibbes, 1850) 120 

Genus Coralliope Guinot, 1967 121 

Coralliope parvula (A. Milne Edwards, 1869) 121 

Genus Cycloxanthops Rathbun, 1897 122 

Cycloxanthops occidentalis (A. Milne Edwards, 1867) 122 

Genus Domecia Eydoux and Souleyet, 1842 122 

* Domecia acanthophora africana Guinot, 1964 122 

Genus Epixanthus Heller, 1861 123 

* Epixanthus hellerii A. Milne Edwards, 1867 123 

Genus Eriphia Latreille, 1817 124 

Eriphia verrucosa (Forskal, 1775) 124 

Genus Euryozius Miers, 1886 124 

Euryozius bouvieri (A. Milne Edwards, 1869) 125 

* Euryozius pagalu, new species 126 

Genus Eurypanopeus A. Milne Edwards, 1878 130 

* Eurypanopeus blanchardi (A. Milne Edwards, 1881) 130 

Genus Globopilumnus Balss, 1933 133 

Globopilumnus africanus (A. Milne Edwards, 1867) 133 

* Globopilumnus stridulans Monod, 1956 134 

Genus Glyptoxanthus A. Milne Edwards, 1879 135 

* Glyptoxanthus angolensis (De Brito Capello, 1866) 135 

Glyptoxanthus cavernosus (A. Milne Edwards, 1878) 135 

Glyptoxanthus corrosus (A. Milne Edwards, 1869) 135 

Genus Heteropanope Stimpson, 1858 136 

Heteropanope acanthocarpus Crosnier, 1967 136 

Heteropanope tuberculidens Monod, 1956 136 

Genus Leopoldius Serene, 1971 136 

* Leopoldius pisifer (MacLeay, 1838), new combination 136 

Genus Menippe de Haan, 1833 137 

Menippe nodifrons Stimpson, 1859 137 

Genus Microcassiope Guinot, 1967 138 

* Microcassiope minor (Dana, 1852) 138 

Page

NUMBER 306 Vll 

Genus Monodaeus Guinot, 1967 140 

Monodaeus couchii (Couch, 1851) 141 

Monodaeus rectifrons (Crosnier, 1967) 141 

* Monodaeus rouxi (Gapart, 1951) 142 

Genus Nanocassiope Guinot, 1967 142 

* Nanocassiope melanodactyla (A. Milne Edwards, 1867) 143 

Genus Nanopilumnus Takeda, 1974 145 

* Nanopilumnus boletifer (Monod, 1956) 145 

Genus Panopeus H. Milne Edwards, 1834 145 

* Panopeus africanus A. Milne Edwards, 1867 146 

Genus Paractaea Guinot, 1969 148 

* Paractaea margaritaria (A. Milne Edwards, 1867) 148 

Paractaea monodi Guinot, 1969 149 

* Paractaea rufopunctata africana Guinot, 1976 150 

Genus Paraxanthias Odhner, 1925 151 

Paraxanthias eriphioides (A. Milne Edwards, 1867) 151 

Genus Pilumnopeus A. Milne Edwards, 1863 151 

* Pilumnopeus africanus (De Man, 1902) 151 

Pilumnopeus caparti (Monod, 1956) 151 

Genus Pilumnus Leach, 1815 152 

Pilumnus hirtellus (Linnaeus, 1761) 152 

Pilumnus inermis A. Milne Edwards and Bouvier, 1894 152 

* Pilumnus perrieri A. Milne Edwards and Bouvier, 1898 153 

Pilumnus spinifer H. Milne Edwards, 1834 154 

* Pilumnus stebbingi Capart, 1951 154 

Genus Platychelonion Crosnier and Guinot, 1969 155 

Platychelonion planissimum Crosnier and Guinot, 1969 155 

Genus Platypodiella Guinot, 1967 155 

Platypodiella picta (A. Milne Edwards, 1869) 155 

Genus Pseudomedaeus Guinot, 1968 155 

* Pseudomedaeus africanus (Monod, 1956) 155 

Genus Xantho Leach, 1814 156 

Xantho incisus (Leach, 1814) 156 

Xantho pilipes A. Milne Edwards, 1867 156 

Xantho sexdentatus (Miers, 1881) 157 

Genus Xanthodius Stimpson, 1859 157 

* Xanthodius denticulatus (White, 1848) 157 

Xanthodius inaequalis faba (Dana, 1852) 158 

* Xanthodius inaequalis inaequalis (Olivier, 1791) 158 

Family GONEPLACIDAE MacLeay, 1838 159 

Subfamily CARCINOPLACINAE H. Milne Edwards, 1852 160 

Genus Carcinoplax H. Milne Edwards, 1852 160 

* Carcinoplax barnardi Capart, 1951 160 

Subfamily EURYPLACINAE Stimpson, 1871 161 

Genus Machaerus Leach, 1818 161 

* Machaerus atlanticus (Miers, 1881), new combination 162 

Page

vlll 


* Machaerus oxyacantha (Monod, 1956), new combination 163 

Subfamily GONEPLACINAE MacLeay, 1838 163 

Genus Goneplax Leach, 1814 163 

Goneplax rhomboides (Linnaeus, 1758) 164 

Subfamily RHIZOPINAE Stimpson, 1858 165 

Genus Acidops Stimpson, 1871 165 

Acidops cessacii (A. Milne Edwards, 1878) 165 

Subfamily TYPHLOCARCINOPINAE Rathbun, 1909 165 

Genus Typhlocarcinodes Alcock, 1900 165 

* Typhlocarcinodes integrifrons (Miers, 1881) 165 

Family HEXAPODIDAE Miers, 1886 166 

Key to Genera of Hexapodidae 168 

Genus Hexapinus, new genus 169 

Hexapinus buchanani (Monod, 1956), new combination 171 

Genus Hexaplax Doflein, 1904 171 

Genus Hexapus de Haan, 1833 171 

Genus Lambdophallus Alcock, 1900 173 

Genus Paeduma Rathbun, 1897 173 

Genus Parahexapus Balss, 1922 175 

Parahexapus africanus Balss, 1922 175 

Genus Pseudohexapus Monod, 1956 176 

Pseudohexapus platydactylus Monod, 1956 176 

Genus Spiroplax, new genus 176 

Genus Stevea, new genus 177 

Genus Thaumastoplax Miers, 1881 177 

Thaumastoplax anomalipes Miers, 1881 179 

Genus Tritoplax, new genus 180 

Family PINNOTHERIDAE de Haan, 1833 181 

Subfamily ASTHENOGNATHINAE Stimpson, 1858 181 

Genus Asthenognathus Stimpson, 1858 181 

Asthenognathus atlanticus Monod, 1933 181 

Subfamily PINNOTHERINAE de Haan, 1833 182 

Genus Pinnotheres Bosc, 1802 182 

Pinnotheres conicola, new species 182 

Pinnotheres leloeuffi Crosnier, 1969 185 

Pinnotheres mccainae Schmitt, 1973 185 

Pinnotheres pinnotheres (Linnaeus, 1758) 187 

Pinnotheres pisum (Linnaeus, 1767) 187 

Pinnotheres tellinae, new species 187 

Pinnotheres sp. A 190 

Pinnotheres sp. B 190 

Pinnotheres sp. D 191 

Pinnotheres sp 191 

Family RETROPLUMIDAE Gill, 1894 191 

Family MICTYRIDAE Dana, 1851 191 

Family PALICIDAE Rathbun, 1898 191 

Page

NUMBER 306 

Genus Palicus Philippi, 1838 191 

* Palicus caromi (P. Roux, 1830) 191 

Family OCYPODIDAE Rafinesque, 1815 192 

Subfamily CAMPTANDRIINAE Stimpson, 1858 193 

Key to Genera of Camptandriinae 193 

Genus Calabarium, new genus 195 

Calabarium crinodytes, new species 196 

Genus Camptandrium Stimpson, 1858 199 

Genus Cleistostoma de Haan, 1833 200 

Genus Deiratonotus, new genus 201 

Genus Ecphantor, new genus 202 

Ecphantor modestus, new species 203 

Genus Ilyogynnis, new genus 206 

Genus Leipocten Kemp, 1915 207 

Genus Paracleistostoma De Man, 1895 208 

Genus Paratylodiplax Serene, 1974 209 

Genus Serenella, new genus 211 

Genus Telmatothrix, new genus 212 

Telmatothrix powelli, new species 213 

Genus Tylodiplax De Man, 1895 217 

Subfamily OCYPODINAE Rafinesque, 1815 217 

Genus Ocypode Weber, 1795 217 

Ocypode africana De Man, 1881 218 

* Ocypode cursor (Linnaeus, 1758) 219 

Genus Uca Leach, 1814 220 

* Uca tangeri (Eydoux, 1835) 221 

Family GRAPSIDAE MacLeay, 1838 225 

Subfamily GRAPSINAE MacLeay, 1838 226 

Genus Geograpsus Stimpson, 1858 226 

* Geograpsus lividus (H. Milne Edwards, 1837) 226 

Genus Goniopsis de Haan, 1833 227 

* Goniopsis pelii (Herklots, 1851) 227 

Genus Grapsus Lamarck, 1801 232 

* Grapsus grapsus (Linnaeus, 1758) 232 

Genus Pachygrapsus Randall, 1840 233 

* Pachygrapsus gracilis (De Saussure, 1858) 233 

* Pachygrapsus transversus (Gibbes, 1850) 234 

Genus Planes Bowdich, 1825 235 

Planes cyaneus Dana, 1851 235 

Planes minutus (Linnaeus, 1758) 236 

Subfamily PLAGUSIINAE Dana, 1851 237 

Genus Pennon Gistel, 1848 237 

* Pennon gibbesi (H. Milne Edwards, 1853) 238 

Genus Plagusia Latreille, 1804 238 

* Plagusia depressa (Fabricius, 1775) 239 

Subfamily SESARMINAE Dana, 1851 239 

Page


Genus Cyclograpsus H. Milne Edwards, 1837 239 

* Cyclograpsus integer H. Milne Edwards, 1837 239 

Genus Metagrapsus H. Milne Edwards, 1853 240 

* Metagrapsus curvatus (H. Milne Edwards, 1837) 240 

Genus Sesarma Say, 1817 .' 241 

Subgenus Chiromantes Gistel, 1848 242 

Sesarma (Chiromantes) angolense De Brito Capello, 1864 243 

* Sesarma (Chiromantes) buettikoferi De Man, 1883 243 

Sesarma (Chiromantes) elegans Herklots, 1851 244 

Subgenus Perisesarma De Man, 1895 245 

Sesarma (Perisesarma) alberti Rathbun, 1921 245 

Sesarma (Perisesarma) huzardi (Desmarest, 1825) 245 

Sesarma (Perisesarma) kamermani De Man, 1883 247 

Subfamily VARUNINAE H. Milne Edwards, 1853 247 

Genus Brachynotus de Haan, 1833 247 

Brachynotus atlanticus Forest, 1957 247 

Genus Euchirograpsus H. Milne Edwards, 1853 247 

Euchirograpsus liguricus H. Milne Edwards, 1853 247 

Family GECARCINIDAE MacLeay, 1838 248 

Genus Cardisoma Latreille, 1828 249 

Cardisoma armatum Herklots, 1851 249 

Genus Gecarcinus Leach, 1814 250 

Gecarcinus weileri (Sendler, 1912) 250 

Family HAPALOCARCINIDAE Caiman, 1900 250 

Genus Neotroglocarcinus Fize and Serene, 1957 250 

Neotroglocarcinus balssi (Monod, 1956) 251 

Family HYMENOSOMATIDAE MacLeay, 1838 251 

Genus Elamena H. Milne Edwards, 1837 252 

Elamena (Trigonoplax) gordonae Monod, 1956 252 

Genus Hymenosoma Desmarest, 1825 252 

Hymenosoma orbiculare Desmarest, 1825 252 

Family MAJIDAE Samouelle, 1819 252 

Subfamily EPIALTINAE MacLeay, 1838 255 

Genus Acanthonyx Latreille, 1828 255 

Key to Eastern Atlantic Species of Acanthonyx 255 

Acanthonyx brevijrons A. Milne Edwards, 1869 256 

Acanthonyx depressifrons, new species 258 

Acanthonyx lunulatus (Risso, 1816) 260 

* Acanthonyx minor, new species 261 

Subfamily INACHINAE MacLeay, 1838 263 

Key to Eastern Atlantic Genera of Inachinae 265 

Genus Achaeus Leach, 1817 266 

Key to Eastern Atlantic Species of Achaeus 268 

* Achaeus buderes, new species 269 

Achaeus cranchii Leach, 1817 271 

* Achaeus foresti Monod, 1956 271 

Page

NUMBER 306 XI 

Achaeus monodi (Capart, 1951) 272 

Achaeus trifalcatus Forest and Guinot, 1966 272 

* Achaeus turbator, new species 272 

Genus Calypsachaeus, new genus 275 

* Calypsachaeus calypso (Forest and Guinot, 1966), new combination 

275 

Genus Capartiella, new genus 277 

* Capartiella longipes (Capart, 1951), new combination 278 

Genus Dorhynchus Thomson, 1873 279 

Dorhynchus thomsoni Thomson, 1873 281 

Genus Ergasticus Studer, 1883 281 

Ergasticus clouei Studer, 1883 281 

Genus Inachus Weber, 1795 282 

Key to Tropical West African Species of Inachus 282 

Inachus aguiarii De Brito Capello, 1876 283 

* Inachus angolensis Capart, 1951 283 

* Inachus biceps, new species 285 

Inachus dorsettensis (Pennant, 1777) 287 

* Inachus grallator, new species 287 

Inachus guentheri (Miers, 1879) 291 

Inachus leptochirus Leach, 1817 291 

* Inachus nanus, new species 291 

Inachus phalangium (Fabricius, 1775) 293 

Inachus thoracicus Roux, 1830 294 

Genus Macropodia Leach, 1814 294 

Key to Tropical West African Species of Macropodia 295 

Macropodia doracis, new species 295 

* Macropodia gilsoni (Capart, 1951) 297 

* Macropodia hesperiae, new species 298 

Macropodia intermedia Bouvier, 1940 300 

Macropodia longicornis (A. Milne Edwards and Bouvier, 1899) 300 

Macropodia longipes (A. Milne Edwards and Bouvier, 1899) ... 300 

* Macropodia macrocheles (A. Milne Edwards and Bouvier, 1898) 301 

* Macropodia spinulosa (Miers, 1881) 301 

* Macropodia straeleni Capart, 1951 303 

Genus Stenorhynchus Lamarck, 1818 304 

* Stenorhynchus lanceolatus (Brulle, 1837) 304 

Subfamily MAJINAE Samouelle, 1819 307 

Genus Maja Lamarck, 1801 307 

Maja crispata Risso, 1827 307 

Maja goltziana d'Oliveira, 1888 307 

Maja squinado (Herbst, 1788) 307 

Subfamily PISINAE Dana, 1851 307 

Genus Apiomithrax Rathbun, 1897 307 

Apiomithrax bocagei (Osorio, 1887) 308 

* Apiomithrax violaceus (A. Milne Edwards, 1867) 309 

Page

xjj SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

Genus Eurynome Leach, 1814 311 

* Eurynome aspera (Pennant, 1777) 311 

* Eurynome parvirostris Forest and Guinot, 1966 312 

Genus Herbstia H. Milne Edwards, 1834 312 

Key to Species of Adult Herbstia from West Africa 312 

* Herbstia condyliata (Fabricius, 1787) 312 

* Herbstia nitida, new species 315 

Herbstia rubra A. Milne Edwards, 1869 317 

Genus Micropisa Stimpson, 1858 318 

Micropisa ovata Stimpson, 1858 318 

Genus Pisa Leach, 1814 318 

* Pisa armata (Latreille, 1803) 318 

* Pisa calva Forest and Guinot, 1966 319 

* Pisa carinimana Miers, 1879 320 

Pisa nodipes (Leach, 1815) 321 

Pisa tetraodon (Pennant, 1777) 321 

Family MIMILAMBRIDAE Williams, 1979 322 

Family PARTHENOPIDAE MacLeay, 1838 322 

Subfamily AETHRINAE Dana, 1851 322 

Genus Heterocrypta Stimpson, 1871 322 

* Heterocrypta maltzami Miers, 1881 322 

Genus Sakaila, new genus 324 

* Sakaila africana, new species 325 

Subfamily PARTHENOPINAE MacLeay, 1838 327 

Genus Daldorfia Rathbun, 1904 327 

Daldorfia bouvieri (A. Milne Edwards, 1869) 327 

Genus Parthenope Weber, 1795 327 

* Parthenope expansa (Miers, 1879) 328 

* Parthenope massena (Roux, 1830) 330 

Parthenope miersii (A. Milne Edwards and Bouvier, 1898) 331 

* Parthenope notialis, new species 331 

Genus Solenolambrus Stimpson, 1871 336 

* Solenolambrus noordendei (Capart, 1951) 336 

Appendix I: Station Data 337 

Appendix II: Gazetteer 342 

Addendum 348 

Genus Lillyanella, new genus 348 

Lillyanella plumipes, new species 349 

Addendum to Key 352 

Literature Cited 353 

Page



Introduction 

Raymond B. Manning 

and L. B. Holthuis 

This report is based primarily on collections 

made in the Gulf of Guinea in 1964 and 1965 

during two cruises of the University of Miami 

Research Vessel John Elliott Pilhbury. Material 

from the Pillsbury collections has been supplemented 

by other brachyuran crabs from West 

Africa in the collections of the Rijksmuseum van 

Natuurlijke Historie, Leiden, and the National 

Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 

Our original intent was to prepare a list of the 

Brachyura obtained during the two Pillsbury 

cruises, a job that we anticipated would be quite 

simple, as the West African Brachyuran fauna 

has been investigated in detail in the last 30 years. 

In 1951, A. Capart published a beautifully illustrated 

volume on the West African crabs collected 

by the Belgian Oceanographic Expedition, 1948- 

1949, aboard the Noordende III. Capart's study was 

overshadowed by the publication in 1956 by Th. 

Monod of a review of West African crabs. 

Monod's work, a monumental compilation of 

information on West African Brachyura, was illustrated 

with almost 900 figures. It remains the 

basic tool in the study of West African crabs. 

Raymond B. Manning, Department of Invertebrate Zoology, National 

Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. L. B. Holthuis, Rijksmuseum van Natuurlijke Historie, 

Postbus 9517, 2300 RA Leiden, The Netherlands. 

These reports were soon followed by others 

based on smaller collections, those by M. Rossignol 

(1957 and 1962) on decapods from the Congo 

region and by D. Guinot and A. Ribeiro (1962) 

on collections from the Cape Verde Islands and 

Angola, as well as several other smaller reports. 

Material collected by the Calypso in 1956 from 

West African localities between Spanish Sahara 

and Gabon on the mainland, as well as from the 

offshore islands of the Gulf of Guinea, Principe, 

Sao Tome, and Annobon, formed the basis of an 

important report by J. Forest and D. Guinot in 

1966. 

Realizing that the West African tropical brachyuran 

fauna was one of the best known in the 

world, we anticipated few difficulties in preparing 

a report on the materials collected by the Pillsbury. 

As our study progressed, however, we continually 

encountered problems that demonstrated that 

our original optimism was largely unfounded. 

Whenever comparative study material was 

available in our collections, we compared specimens 

from West African localities with those from 

other areas. Several of the common species in the 

Gulf of Guinea have in the past been identified 

with European-Mediterranean species. Thus 

Ethusa mascarone, Inachus dorsettensis, Macropodia rostrata, 

and Parthenope macrochelos, among others, all 

have been included in the tropical fauna by 

Monod (1956), as well as other authors. In the

case of these four species, direct comparison of 

material from northern and southern areas has 

demonstrated that the southern forms are distinct; 

southern counterparts of three of these are 

described as new, and a name is resurrected from 

synonymy for the fourth species. 

Similarly, direct comparison of material from 

both sides of the Atlantic yielded surprising results. 

In one case, that of one species of Ranilia, 

such a comparison showed that populations on 

each side of the Atlantic were conspecific whereas 

they had previously been considered to be distinct. 

Other comparisons, based on species in 

Eurypanopeus, Goniopsis, Callinectes, and Bathynectes, 

resulted in our recognition of distinct species on 

each side of the Atlantic. 

Two family-group taxa, the family Hexapodidae 

and the subfamily Camptandriinae of the 

family Ocypodidae, had to be surveyed at the 

generic level in order to determine the status of 

the West African species. These revisions are incorporated 

herein. In addition, a key to the 

subfamilies and genera of family Dorippidae is 

presented. 

We also encountered unanticipated problems 

in various aspects of nomenclature: Citations of 

brachyuran family and subfamily names vary 

widely in the existing literature and often are 

incorrect; information on type-species and original 

citations of genera is widely scattered and has 

never been compiled for the Eastern Atlantic 

species; and numerous older names in the literature 

have never been properly identified. 

Finally, there is no single reference work containing 

at least a list of all of the Eastern Atlantic 

Brachyura. The Scandinavian species have been 

dealt with by M. Christiansen (1969) in a wellprepared 

handbook; the British species have been 

studied by R. W. Ingle (1980) of the British 

Museum (Natural History); the Spanish and Portuguese 

species were the subject of the most upto-date 

review of European-Mediterranean crabs 

by the late R. Zariquiey Alvarez (1968); the Black 

Sea species were studied by Bacescu (1967); the 

French species were studied by Bouvier (1940) in 

a work in which the nomenclature is now com- 


pletely out of date; the Adriatic species were 

reported by O. Pesta in 1918, but, again, all the 

names are not current; and the eastern Mediterranean 

species were listed by L. B. Holthuis and 

E. Gottlieb in 1958. The latter fauna contains 

several species that have entered and colonized 

the region from the Red Sea via the Suez Canal. 

About 300 species of Brachyura are known from 

the Eastern Atlantic in literature that is widely 

scattered. 

Thus our simple list of species taken off West 

Africa by the Pillsbury changed into a guide to the 

Eastern Atlantic crab fauna, with major emphasis 

on that of tropical West Africa. We have listed 

all Eastern Atlantic species of Brachyura known 

to us, and we have included all references to West 

African crabs published since 1956 that have 

come to our attention. This work, then, is basically 

an update of Monod's monumental work, 

based primarily on the collections made by the 

Pillsbury. 

In our accounts, the term "West African" or 

"tropical" is used to indicate tropical West Africa: 

the Cape Verde Islands and the African mainland 

from Mauritania southward to Angola, including 

the offshore islands of the Gulf of Guinea. 

"Extralimital" is generally used to describe more 

northern forms, including those that live in the 

Mediterranean. We have not attempted to deal 

with the poorly known fauna of South-West 

Africa. 

In the systematic account herein, we include 

original citations for all 36 families of marine 

Brachyura, as well as references to synonyms at 

the family-group level. Twenty-nine families are 

represented in the Eastern Atlantic fauna and 26 

of these have representatives in the tropical fauna. 

The seven families of crabs not occurring in the 

Eastern Atlantic are the Belliidae, the Bythograeidae, 

the Mimilambridae, the Cyclodorippidae, 

the Mictyridae, the Platyranthidae, and the 

Retroplumidae. The three families of crabs occurring 

in the Eastern Atlantic but not represented 

in the tropical fauna are the Cancridae, 

the Cymonomidae, and the Homolodromiidae. 

Original citations are provided for all 146 genera

NUMBER 306 

now known from the Eastern Atlantic, along with 

an indication of their type-species and gender; 

there are representatives of 120 genera in the 

tropical fauna. The Eastern Atlantic fauna comprises 

about 300 species, 218 of which have been 

recorded from localities between Mauritania and 

Angola. 

Sixteen new genera, six of which are extralimital, 

and 24 new species from West Africa are 

recognized. 

Most of the 300+ Eastern Atlantic species are 

indigenous to the Eastern Atlantic. Fifteen species 

have been introduced into the Eastern Mediterranean 

through the Suez Canal and apparently 

have become established there: Atergatis rosens, 

Charybdis helleri, C. longicollis, Eucrate crenata, Heteropanope 

laevis, Hyastenus hilgendorfi, Ixa monodi, 

Leucosia signata, Myra fugax, Notopus dorsipes, Pilumnopeus 

vauquelini, Pilumnus hirsutus, Portunus pelagicus, 

Sphaerozius nitidus, and Thalamita poissonii. Five 

other species have been introduced from other 

areas: Callinectes sapidus, Eriocheir sinensis, Neopanope 

sayi, Pilumnoides perlatus, and Rhithropanopeus 

harrisii. 

Several other species have been introduced into 

the Eastern Atlantic but apparently have not 

become established there. Catta (1876) reported 

the following species taken from a vessel in Marseilles 

harbor: Pachygrapsus advena, new species (= 

P. transversus), Nautilograpsus minutus (= Planes minutus), 

Plagusia squamosa (= P. depressd), and P. 

tomentosa (= P. chabrus (Linnaeus, 1758)); the first 

three are included in the West African fauna. De 

Man (1913) found Menippe convexa Rathbun, 

1893, Leptodius voeltzkowi Lenz, 1905, Pilumnus longicomis 

Hilgendorf, 1878, Pilumnus malardi De 

Man, 1913 (= Parapilumnus malardi), and Pilumnus 

truncatospinosus De Man, 1913 (= Parapilumnus truncatospinosus) 

in the harbor of St. Vaast-la-Hougue, 

Normandy, France, where they were obtained 

from barnacles attached to a ship coming from 

Madagascar. 

The following species have been erroneously 

recorded from West Africa: Chlorodiella longimana, 

Hepatus princeps, Libinia erinacea, Metopograpsus mes- 

sor, Notolopas brasiliensis, Ocypode ceratophthalmus, O. 

quadrata, Pilumnoides hassleri, Platychirograpsus spectabilis, 

Pseudograpsus elongatus, Rochinia gracilipes, 

Sesarma roberti, and Uca burgersi. 

In addition, there are several species described 

or recorded from Eastern Atlantic localities based 

on incorrectly labeled material, such as Portunus 

sanguinolentus (Herbst, 1783), reported from the 

Adriatic by Pesta (1918:458). These species are 

not included in our lists of extralimital species 

under each family. 

COLLECTIONS STUDIED.—Most of the specimens 

reported here were collected during two cruises 

of the research vessel John Elliot Pillsbury of the 

Rosenstiel School of Marine and Atmospheric 

Science, University of Miami, in 1964 and 1965. 

A narrative of the cruise was given by Voss (1966) 

and Bayer (1966), who summarized dredging and 

trawling records. Cruise tracks of the Pillsbury off 

West Africa are shown in Figure 87. Species taken 

by the Pillsbury are marked with an asterisk (*) in 

the discussion sections under each family heading 

and in the headings of the species accounts. The 

Pillsbury collections have been deposited in the 

Rijksmuseum van Natuurlijke Historie, Leiden 

(L) and the National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. (W). 

The Pillsbury collections are quite important. 

They are slightly smaller than those taken by the 

Calypso, containing the representatives of 98 

rather than 108 species (Forest and Guinot, 1966: 

27). Although this represents less than half of the 

known West African crab fauna, 16 of the 24 new 

species recognized here are based on specimens 

taken by the Pillsbury. A smaller collection from 

off Gabon, taken during an exploratory cruise of 

the U. S. Fish and Wildlife Service vessel Geronimo 

also is reported on here. All of the material from 

the Geronimo collections has been deposited at 

Washington. The bulk of the brachyuran collection 

made off southern Angola and northern 

South-West Africa by the U. S. Fish and Wildlife 

Service vessel Undaunted and reported upon by 

Crosnier (1970) has been deposited at Leiden and 

is reported here. 

In the species accounts, station data for the


collections made by the Pillsbury, Geronimo, andcitations, 

not an exhaustive list of references. 

Undaunted are abbreviated to station number, Where appropriate, we have indicated which 

depth in meters, and observed bottom type. Com- family-group names are on the Official List of 

plete station data and a list of brachyuran species Family Group Names in Zoology established by the 

taken at each station are given in Appendix I. International Commission on Zoological Nomen- 

Smaller collections from the northwestern coast clature (ICZN) by stating "name 00 on Official 

of Africa by the Rijksmuseum van Natuurlijke List." 

Historie aboard the research vessel Onversaagd also In each family account we have included a 

are reported under "Material Examined" sections paragraph entitled "Eastern Atlantic Genera," in 

of the appropriate species discussion. Many of the which we state how many genera of that family 

species collected during the Onversaagd cruises do occur in the area. Extralimital or nontropical 

not occur in the tropical fauna and will be studied 

and possibly reported upon separately. In addition, 

the collections at Washington include series 

of decapods from Liberia donated by G. C. 

Miller, then with the U. S. Fish and Wildlife 

Service; material from Ghana donated by G. W. 

Bane, then with Cornell University; and several 

collections from Gulf of Guinea, donated by A. 

Crosnier, then with the Centre ORSTOM, 

Pointe-Noire. 

genera are then listed along with their original 

citations, an indication of their type-species and 

genders, and a statement that they are on the 

Official List, as appropriate. We have not tried to 

cite synonyms of extralimital genera. 

That paragraph is followed by one entitled 

"Eastern Atlantic Species," in which the numbers 

of species occurring in that region are given. This 

is followed by a comparison of the names of 

tropical species, as cited by Monod (1956), com- 

In both Washington and Leiden are a few lots pared to the current names; in several families, 

from the Travailleur and Talisman collections, ac- notably the Xanthidae and the Majidae, there 

quired on exchange from the Museum national have been numerous name changes since 1956. In 

d'Histoire naturelle, Paris. The Smithsonian re- those lists, the names used by Monod are given 

ceived representatives of 80 species from those in the same order in which he cited them, to 

collections in 1899 through E.-L. Bouvier. make it easier to work from our paper back to 

FORMAT CONSIDERATIONS.—We have adopted a Monod's. Also in those lists, species taken by the 

fairly rigid format in order to ensure that our Pillsbury are marked with an asterisk (*). Those 

accounts are comparable at the family, genus, lists are then followed by a list, in alphabetical 

and species levels. 

order by genus and species, of extralimital species, 

At the family level, we have first given the along with a brief comment on their range and 

original citation for each family of marine brach- one or more pertinent references. These two inyuran 

crabs; the freshwater families have been troductory paragraphs may be followed by a 

excluded. In addition to the original citation we 

have included original citations to all synonyms 

that we could find at the family-group level, 

without indicating necessarily which of these are 

now in current use as family-group categories. 

Thus in the Portunidae, for example, we have 

cited 20 different family-group taxa that have 

been recognized in the literature; only a few of 

these are now in current use. The list of synonyms 

for each family does not include erroneous spellings 

or subsequent uses of the family names; each 

of the family synonymies is a guide to original 

third entitled "Remarks" or by a key to genera. 

The families usually are arranged in the order 

in which they were treated in Monod. Within 

families, subfamilies, if recognized, are arranged 

alphabetically except in the Xanthidae where 

taxa are arranged alphabetically by genus and by 

species within each genus, genera are arranged 

alphabetically within subfamilies, and species are 

arranged alphabetically within genera. 

For genera represented in the tropical fauna 

we have included not only the original citation 

but references to all synonyms known to us. In

NUMBER 306 

the cases of some genera, such as Dorippe sensu abbreviated; full data and a list of species taken 

lato, Sesarma, and Uca, all of which are in need of at each station are in Appendix 1. Geographic 

revision, we have given a complete list of original localities in the sections on material and distri- 

citations of genus-group names in these groups as bution are corrected to approved spellings by the 

a guide to their future revision. We do not mean U. S. Board on Geographic Names as published 

to imply that the names so cited are to be consid- in their gazetteers. A list of localities, by country, 

ered as synonyms. 

with coordinates, is given in Appendix 2. Locali- 

Tropical species are treated in two ways. If ties in the literature accompanied by coordinates 

material from the tropical region was studied, we are not repeated in Appendix 2. Alternate spell- 

give an expanded account, with a synonymy, a ings often are given in the text. The names of 

list of synonyms, material examined (with a sep- most countries (e.g., Senegal, Guinea) have been 

arate paragraph for Pillsbury, Geronimo, and Un- anglicized. In the sections of ecology we have 

daunted collections), citations to a description, to summarized available information on habitat, 

a good illustration, and to an illustration of the depth distribution, and occurrences of ovigerous 

gonopod (used interchangeably with male pleo- females; much of this information has been taken 

pod and first male pleopod), an expanded section from the recent literature, that is since 1956. 

on biology, and an expanded section on distri- In the text, author and date citations accombution. 

If no tropical material is available, we panying a name (e.g., Dehaanius MacLeay, 1838) 

have given an abbreviated account, with refer- are considered to be part of the name, not a 

ences to Capart (1951) and Monod (1956), and bibliographic citation. References to Monod 

subsequent references, with parenthetic remarks without a date always refer to the monograph of 

on origin of the material reported in the litera- West African crabs published by Monod in 1956. 

ture, followed by a brief statement on distri- Although we have tried to make our lists of 

bution. 

family citations, generic names and synonyms, 

In our synonymies we have tried to include and species synonyms as complete as possible, we 

references to Capart (1951) and Monod (1956), realize that these lists are by no means exhaustive. 

all subsequent references based on West African We would appreciate having any omissions or 

specimens, and a few minor references overlooked errors brought to our attention. 

by Monod. We have not duplicated the extensive In some accounts in the literature based on 

synonymies given by Monod, except in the case expeditions sponsored by the French in the latter 

of new species, where complete synonymies are half of the 19th century (A. Milne Edwards and 

given and in cases where the name has been Bouvier, 1894, 1899, 1900), coordinates were 

changed since 1956. Original citations for all based on the Paris observatory, not Greenwich, 

species then reported were given by Monod in his which is 2° 20' West of Paris. Later (Bouvier, 

synonymies. The separate list of synonyms may 1922), these coordinates were based on Green- 

seem redundant, but even though the nomenclawich. In our text we have corrected the earlier 

ture of the Eastern Atlantic brachyurans is rela- observations from Paris to Greenwich; to avoid 

tively stable, current studies are demonstrating errors, we often have given both, so that a coor- 

the existence of previously unrecognized species dinate may be cited as: 05°10'N, 05°05'W of 

there, and many older names are available in the Paris (= 02°45'W of Greenwich). 

literature dealing with the European-Mediterra- REPOSITORIES AND ABBREVIATIONS.—The bulk 

nean fauna. In this region older names must be of the materials reported herein have been depos- 

searched for carefully before new species are ited in two institutions: The Rijksmuseum van 

named. 

Natuurlijke Historie, Leiden, The Netherlands, 

In the sections on material based on the Pills- abbreviated to "L" in the material sections and 

bury, Geronimo, and Undaunted collections, data are often referred to as "Leiden" in the text. Registry

numbers (Crust. D. 000) are for the Decapod 

collection at Leiden. The National Museum of 

Natural History, Smithsonian Institution, Washington, 

D. C, abbreviated to "W" in the material 

and referred to as "Washington" or "USNM" in 

the text. Catalog numbers for material at Washington 

are given under the acronym USNM, 

referring to catalogs established under the former 

U. S. National Museum. For new species deposited 

in these collections, registry or catalog numbers 

are given only for holotypes. 

A few specimens from other museums also have 

been studied. These are cited as follows: 

BM British Museum (Natural History), London 

MP Museum national d'Histoire naturelle, Paris 

(Paris Museum) 

ZMB Zoologisches Museum, Berlin 

A few abbreviations have been used throughout 

the text: 

cb 

cl 

cm 

fm 

juv 

m 

mm 

ov 

carapace breadth 

carapace length 

centimeter 

fathom 

juvenile 

meter 

millimeter 

ovigerous 

ACKNOWLEDGMENTS.—We are indebted to Gilbert 

L. Voss, Rosenstiel School of Marine and 

Atmospheric Science, University of Miami, for 

inviting us to participate in the Pillsbury cruises 

and for making the material available for study. 

Participation in the cruises was supported by the 

National Geographic Society under grants to the 

University of Miami. Part of this study was supported 

by the National Geographic Society under 

grant NGS 1042 to one of us (RBM). 

R. W. Ingle, British Museum (Natural History), 

London, and J. Forest, Museum national 

d'Histoire naturelle, Paris, provided working 

space on several occasions; Ingle also read and 

commented on part of the manuscript. H. Zibrowius, 

Station Marine d'Endoume, Marseille, 

and H.-E. Gruner, Zoologisches Museum, Berlin, 

sent material to us. C. B. Powell, University of 

Port Harcourt, Nigeria, collected and donated 


specimens to the Rijksmuseum van Natuurlijke 

Historie, Leiden and the National Museum of 


D. C. 

C. Froglia, Laboratorio di Tecnologia della 

Pesca, Ancona, brought an obscure paper by N. 

Prestandrea and the account of Portunus Maravigna 

to our attention. P. Le Loeuff, Antenne OR- 

STOM, Centre de Bretagne, Brest, provided us 

with a copy of a paper by C. A. Dias and J. F. 

Seita Machado that we were unable to obtain 

elsewhere. Angelo A. DiMauro, University of 

Connecticut, Torrington, provided information 

on the rediscovery of the type of Amorphopus cylindraceus 

in the Bell collection at Oxford University, 

England. C. F. Cowan, Cumbria, England, provided 

information on the dates of publication of 

Guerin's Iconographie. We thank M. Tiirkay, Natur-Museum 

Senckenberg, for bringing to our 

attention the account ofLambrus spinosissimus Osorio, 

1923, which we otherwise would have overlooked, 

and R. W. Ingle, British Museum (Natural 

History), for pointing out that Verany (1846) 

had introduced a generic and specific name for 

Brachynotus sexdentatus (Risso) (see synonymy for 

Brachynotus, p. 247). 

We thank two staff members of the Department 

of Invertebrate Zoology (Crustacea), National 

Museum of Natural History, Smithsonian 

Institution, for helping with various aspects of 

preparation of the manuscript: Anne Cohen for 

help in checking the bibliography and Cynthia J. 

Hemmings for proofreading parts of the manuscript. 

The manuscript was read by our colleague 

at the Smithsonian, Fenner A. Chace, Jr., with 

his usual and welcome attention to detail. Lilly 

King Manning executed most of the original 

illustrations and prepared all of the illustrations 

for publication. Much of her work was done as a 

volunteer. Carolyn S. Hahn and Jack F. Marquardt 

of the Smithsonian Library helped to 

locate many obscure references. 

Family RANINIDAE de Haan, 1839 

RANINOIDEA de Haan, 1839:102 [emended to Raninidae by 

White, 1847a:56].

NUMBER 306 

NOTOPINAE Serene and Umali, 1972:22, 25, 29 [herein corrected 

to Notopodinae]. 

EASTERN ATLANTIC GENERA.—Four, of which 

three, Cyrtorhina, Ranilia, and Raninoides, are represented 

by tropical species. The fourth genus, 

which occurs in the eastern Mediterranean, is 

Notopus de Haan (1841:138). Type-species: Cancer 

dorsipes Linnaeus, 1758, by monotypy; gender: 

masculine; name 1570 on Official List. 

EASTERN ATLANTIC SPECIES.—Four, three of 

which were recorded by Monod (1956) as follows: 

Name in Monod 

Ranilia atlantica 

Cyrtorhina granulosa 

Raninoides bouvieri 

Current Name 

Ranilia constrieta* 

Cyrtorhina granulosa 

Raninoides bouvieri* 

The fourth raninid occurring in the eastern 

Atlantic is Notopus dorsipes (Linnaeus, 1758): Eastern 

Mediterranean; an Indo-West Pacific species 

that has entered the Mediterranean via the Suez 

Canal, first recorded from the Mediterranean by 

Lewinsohn and Holthuis (1964). 

Subfamily NOTOPODINAE Serene and Umali, 

1972 

Genus Ranilia H. Milne Edwards, 1837 

Ranilia H. Milne Edwards, 1837:195 [type-species: Ranilia 

muricata H. Milne Edwards, 1837, by monotypy; gender: 

feminine]. 

Raninops A. Milne Edwards, 1880:34 [type-species: Raninops 

constrictus A. Milne Edwards, 1880, by subsequent designation 

by Rathbun, 1937:17; gender: masculine]. 

* Ranilia constricta (A. Milne Edwards, 1880) 

FIGURES 1, 2 

Raninops constrictus A. Milne Edwards, 1880:35. 

Notopus (Raninoides?) atlanticus Studer, 1883:17, pi. 1: figs. 

5a, b. 

Ranilia constricta.—A. Milne Edwards and Bouvier, 1923:302, 

pi. 1: figs. 11-13, pi. 3: figs. 2-5.—Rathbun, 1937:20, pi. 

4: fig. 5, pi. 5: figs. 1, 2.—Gomes Correa, 1970:2, pis. 1, 2, 

7: figs. 56-58.—Pequegnat, 1970:180. 

Notopus atlanticus.—Gurney, 1939:103 [listed]. 

Ranilia atlantica.—Monod, 1956:47, 631, figs. 17, 18.— 

Longhurst, 1958:87.—Rossignol, 1962:113 [listed].— 

Gomes Correa, 1970:5 [discussion]. 

Raninids.—Voss, 1966:50. 

Lyndeus.—Bayer, 1966:102. 

MATERIAL EXAMINED.—Pillsbury Material: Annobon: 

Sta 275, 9-69 m, rubble of coralline algae, 3

a 


FIGURE 1.—Ranilia constricta (A. Milne Edwards), male cl ca. 15 mm, Pillsbury Sta 275: a, front; 

b, cheliped; c, fourth pereiopod; d, e, first pleopod of male;/, g, second pleopod of male. 

The largest specimen recorded in the literature is 

a male 41 mm long (Gomes Correa, 1970). 

REMARKS.—Both Monod (1956) and Gomes 

Correa (1970) have suggested that Ranilia atlantica 

(Studer, 1883) from Ascension Island and West 

Africa was conspecific with R. constricta (A. Milne 

Edwards, 1880) from the western Atlantic, but, 

so far as we can determine, specimens from both 

sides of the Atlantic have not been compared 

directly. We have been able to compare the four 

specimens taken by the Pillsbury off Annobon with 

a female (cl 22.2 mm) from Bahia Honda, Cuba 

(USNM 48642), a female (cl 19.2 mm) from 

Barbados, 92-366 m (USNM 110223), a male (cl 

11.6 mm) and two females (cl 11.0-11.3 mm) 

from off Palm Beach, Florida, 55-73 m (USNM 

169698), and a male (cl 17.2 mm) and a female 

(cl 18.0 mm) from off Sombrero Light, Florida, 

92-110 m (USNM 169699), as well as with the 

syntypes of/?, atlantica from Ascension Island. We 

can find no significant differences between these 

specimens. 

The males from Annobon differ from the female 

in having slightly more divergent anterolateral 

spines on the carapace. Our female from 

Annobon resembles those from other localities in 

that the anterolateral spines do not diverge perceptibly 

from the lateral margins of the carapace. 

The illustrations published by A. Milne Edwards 

and Bouvier (1923) and the figures given 

by Rathbun (1937) differ in several respects, as 

pointed out by Monod (1956), who questioned 

whether the two specimens previously identified 

with R. constricta were conspecific. Gomes Correa 

(1970:5) commented on the resemblance of the 

figures given by Studer (1883) and Rathbun 

(1937). 

In all of our specimens, the rostral spine extends 

to or exceeds the adjacent frontal teeth by less 

than half its length; in no specimen is the rostrum 

so long as shown by A. Milne Edwards and 

Bouvier (1923, pi. 3: fig. 2) (Figure 2a). In addition, 

all of the specimens seen by us have a short 

but well-marked median carina on the rostrum. 

This carina is shown in the figures of A. Milne 

Edwards and Bouvier (1923, pi. 3: fig. 2), Rathbun 

(1937, pi. 5: fig. 1), and Studer (1883, pi. 1: 

fig 5a), but not by Monod (1956, figs. 17,18) or 

by Gomes Correa (1970, pi. 1: figs. 1,2). 

We have reproduced here the illustrations published 

by Studer (1883) and A. Milne Edwards 

and Bouvier (1923) (Figure 2). These figures, 

especially those by Studer, may be inaccessible to 

most workers. In addition, we have added 

sketches from our specimens (Figure 1), including 

a figure of the male pleopods.

NUMBER 306 

FIGURE 2.—Ranilia constricta (A. Milne Edwards): a, dorsal view (from A. Milne Edwards and 

Bouvier, 1923, pi. 3: fig. 2); b, cheliped (from A. Milne Edwards and Bouvier, 1923, pi. 1: fig. 

11); c, dorsal view; d, cheliped (from Studer, 1883, pi. 1: fig. 5a,b). 

BIOLOGY.—This species has been taken in West 

African waters in depths between 40 and 69 m; 

off Ascension Island it occurs in 110 m. In the 

western Atlantic it has been recorded from the 

littoral zone to 366 or 481 m, usually in less than 

100 m. The record of 100-200 fm (183-366 m) in 

Pequegnat (1970) is somewhat puzzling. Pequegnat 

(1970:180) gave this depth in the species 

account, but in the station list (1970:6), published 

in the same volume, the station at which the 

species was collected, 65-A-9-15, is indicated as 

having a depth of 263 or 330 fm (481 or 604 m). 

The Pillsbury specimens were taken on rubble 

of coralline algae, whereas those from Sierra 

Leone were taken on muddy sand (Longhurst, 

1958). In the western Atlantic this species has 

been collected on a reef (Rathbun, 1937) and on 

a rocky reef off Palm Beach, Florida. Monod 

(1956) reported this species from the stomach 

contents of the fishes Syacium micrurum Ranzani 

and Trygon marmorata Steindachner. Nothing is 

known of the biology of this species. Ovigerous 

females have not been observed off West Africa. 

DISTRIBUTION.—Tropical Atlantic Ocean. 

Monod (1956), who reported material from Senegal, 

SW of Goree, 40 m, and between Joal and 

Sangomar, 42 m, as well as from Moyen Congo 

d 

[Congo], was the first to record the species from 

West Africa. Other records in the literature include: 

Eastern Atlantic. Sierra Leone: 07°43'N, 13°43'W, 40 m 

(Longhurst, 1958). 

Annobon Island: 01°24'S, O5°37'E to 01°24'S, 05°38'E, 

55-69 m (Bayer, 1966; Voss, 1966). 

Central Atlantic. Ascension Island: 60 fm (110 m) (Studer, 

1883). 

Western Atlantic. Florida: Off Sombrero [reef], 47 fm 

(86 m) (A. Milne Edwards, 1880; A. Milne Edwards and 

Bouvier, 1923). 

Cuba: Bahia Honda (Rathbun, 1937). 

Gulf of Mexico: 23°OO'N, 86°48'W, 100-200 fm (183-366 

m) (or 481-604 m?) (Pequegnat, 1970). 

Brazil: Cabo Frio Island, littoral; Pai Island, Rio de 

Janeiro State, 20-30 m; off Guaritiba, Guanabara State, 35- 

40 m (all Gomes Correa, 1970). 

Subfamily RANININAE de Haan, 1839 

Genus Cyrtorhina Monod, 1936 

Cyrtorhina Monod, 1956:49 [type-species: Cyrtorhina granulosa 

Monod, 1956, by monotypy; gender: feminine]. 

Cyrtorhina granulosa Monod, 1956 

Cyrtorhina granulosa Monod, 1956:49, figs. 19-31 [Ghana].— 

Gauld, 1960:68 [Ghana].—Monod, 1963, fig. 31 [no locality].—Forest 

and Guinot, 1966:42 [Principe].

10 

Cyrthorina granulosa.—Forest, 1959:23 [Principe; erroneous 

spelling]. 

Cyrtorhyna granulosa.—Forest, 1959, pi. 2: fig. 1 [Principe; 

erroneous spelling]. 

DISTRIBUTION.—Gulf of Guinea, from off 

Ghana and Ilha do Principe, in 5-6 and 12 m. 

Genus Raninoides H. Milne Edwards, 1837 

Raninoides H. Milne Edwards, 1837:196 [type-species: Ranina 

loevis Latreille, 1825, by monotypy; gender: masculine]. 

* Raninoides bouvieri Capart, 1951 

Raninoides bouvieri Capart, 1951:59, fig. 17.—Monod, 1956: 

54, figs. 32-34.—Longhurst, 1958:87.—Forest, 1959:15.— 

Gauld, 1960:68.—Rossignol, 1%2:113.—Crosnier, 1964: 

35._Forest and Guinot, 1966:42.—Le LoeufT and Intes, 

1968, table 1. 

MATERIAL EXAMINED.—Pillsbury Material: Liberia: 

Sta 68, 70 m, broken shell, 4d, 2$ (1 ov), 2 juv (L). 

Ivory Coast: Sta 62, 46 m, brown, branched and foliate 

Foraminifera, 1$ (L). Sta 64, 68 m, 16* (L). 

Ghana: Sta 28, 49-53 m, 1$ (L). 

Nigeria: Sta 241, 59-63 m, mud and shell, 2

NUMBER 306 11 

Liberia: 04°34.5'N, 08°31'W, 64 m (Forest and Guinot, 

1966). 

Ivory Coast: Off Fresco, 40 m (Le Loeuff and Intes, 1968). 

Ghana: Off Accra, 28-80 m (Gauld, 1960). 

Cameroon: No specific locality (Crosnier, 1964). 

Gabon: 00°38'25"S, 08°46'E, 5 m (Forest and Guinot, 

1966). 

Cabinda: W of Landana, 45 m (Rossignol, 1962). 

It has not previously been recorded from Nigeria 

or Dahomey, although both of these localities 

are well within its known range. 

Family HOMOLODROMIIDAE Alcock, 1899 

HOMOLODROMIDAE Alcock, 1899b: 127, 130 [corrected to 

Homolodromiidae by Stebbing, 1905:58; considered a 

subfamily of family Prosopidae von Meyer, 1860, by 

Glaessner, 1969:R486]. 

EASTERN ATLANTIC GENERA.—One, which has 

not been recorded from tropical waters, is Dicranodromia 

A. Milne Edwards (1880:31). Typespecies: 

Dicranodromia ovata A. Milne Edwards, 

1880, by monotypy; gender: feminine. 

EASTERN ATLANTIC SPECIES.—One, Dicranodromia 

mahieuxii A. Milne Edwards, 1883. Bay of 

Biscay, Azores, and off the Sahara coast in depths 

between 454 and 1330 m (Zariquiey Alvarez, 

1968). 

Family DROMIIDAE de Haan, 1833 

DROMIACEA de Haan, 1833:ix [corrected to Dromiidae by 

Ortmann, 1892:541, 543; name 356 on Official List}. 

EASTERN ATLANTIC GENERA.—Two, Dromia and 

Sternodromia, both represented by species occurring 

off tropical West Africa. 

EASTERN ATLANTIC SPECIES.—Six, of which all 

but one are tropical. Three species were recorded 

by Monod (1956): 

Name in Monod Current Name 

Dromia caputmortuum 

Dromia nodosa [part] 

Dromia nodosa [part] 

Dromidiopsis spinirostris 

Dromia marmorea 

Dromia nodosa 

Dromia monodi* 

Sternodromia spinirostris* 

The fifth tropical species, D. bollorei, was named 

in 1974. 

The extralimital species is Dromia per sonata (Lin- 

naeus, 1758). Southern North Sea southward 

to Spanish Sahara, Mediterranean; sublittoral 

(Christiansen, 1969; Forest, 1974). 

Genus Dromia Weber, 1795 

Dromia Weber, 1795:92 [type-species: Cancer personatus Linnaeus, 

1758, by subsequent designation under the Plenary 

Powers of the International Commission on Zoological 

Nomenclature (Opinion 688 in Bulletin of Zoological Nomenclature, 

21:16-19); gender: feminine; name 1568 on Official 

List]. 

Dromia bollorei Forest, 1974 

Dromia bollorei Forest, 1974:91, figs. Id, 2, 3d, 5, 6b, 7c,d, pi. 

2: figs. 1, 2, pi. 3: fig. 4, pi. 6: fig 1 [Mauritania and Ivory 

Coast]. 

DISTRIBUTION.—Known only from off Mauritania 

and off the Ivory Coast in 100 m. 

Dromia marmorea Forest, 1974 

Dromia vulgaris.—Osorio, 1889:135, 139; 1898:193.—A. 

Milne Edwards and Bouvier, 1900:17, pi. 9: figs. 12-14 

[not fig. 15].—Rathbun, 1900a:300.—Balss, 1921:47.— 

Gordon, 1950:246 [part], figs. 24, 25.—Longhurst, 1958: 

87.—Buchanan, 1958:20. [Not Dromia vulgaris H. Milne 

Edwards, 1837 = D. personata (Linnaeus, 1758).] 

Hairy Brown Sea Crab.—Irvine, 1932:13, fig. 16. 

Dromia caput-mortuum.—Irvine, 1947:301, fig. 205.—Gauld, 

1960:68. [Not Cancer caputmortuum Linnaeus, 1767 = Dromia 

personata (Linnaeus, 1758).] 

Dromia caputmortuum.—Monod, 1956:59, figs. 35-51, 83a.— 

Rossignol, 1962:113. [Not Cancer caputmortuum Linnaeus, 

1767 = Dromia personata (Linnaeus, 1758).] 

Dromia nodosa.—Monod, 1956:65 [part]. [Not Dromia nodosa 

A. Milne Edwards and Bouvier, 1898.] 

Dromia personata.—Crosnier, 1967:321 [not Cancer personatus 

Linnaeus, 1758]. 

Dromia marmorea Forest, 1974:79, 81, figs, lc, 2, 3b, 4d-f, j,k, 

5, pi. 1: figs. 2,4, pi. 3: fig. 2, pi. 4: fig. 7, pi. 5: figs. 3,4, pi. 

8: figs. 3,4.—Tiirkay, 1976b:61 [listed], 62. 

MATERIAL EXAMINED.—Pillsbury Material: None. 

Other Material: Senegal: Dakar, 1949, R. Mauny, paratypes, 

1

12 

Ghana: No specific locality, 1966, F. R. Irvine, 1(5 (BM). 

Accra, 1938, F. R. Irvine, paratypes, 2? (1 ov) (BM). 

Gabon: Cap Lopez, 20 m, Dec 1956, A. Crosnier, paratype, 

19 ov (MP). 

Congo: Baie de Pointe-Noire, beach seine, Oct 1955, M. 

Rossignol, paratypes, 26 (MP). Baie de Pointe-Noire, widemouthed 

nets, Aug 1967, 1

NUMBER 306 

cellent figures of the present species, which show 

all the important details, and to which we have 

little to add. The only discrepancy that we can 

find is in the shape of the dactyli of the walking 

legs, which in Monod's figures 48 and 49 show 

minute ventral teeth, while in most of our specimens 

they bear very distinct spiniform teeth. The 

shape of these dactyli is very similar to that found 

in D. erythropus and D. personate 

Osorio (1889) reported "Dromia vulgaris" from 

Sao Tome and Ilha do Principe without giving 

morphological details of his material. In the same 

paper he also reported material of "Dromia spinirostris" 

from the islands; this latter material, as 

shown below, in all probability belongs to Dromia 

monodi. Therefore, it seems most likely that Osorio's 

Dromia vulgaris belongs to the other common 

shallow-water Dromia of the area, D. marmorea, the 

moreso as the latter species has often been confused 

with D. personata (= D. vulgaris and D. 

caputmortuum). 

Osorio (1898), Rathbun (1900a), and Balss 

(1921) just listed Osorio's 1889 records under 

Dromia vulgaris, and obviously did not see any new 

material. The references by Rathbun (1900a) and 

Balss (1921) to the occurrence of the species at St. 

Helena in all probability do not refer to the 

present species (see "Distribution," p. 14). 

The first original record of this species subsequent 

to that by Osorio (1889) is that by A. Milne 

Edwards and Bouvier (1900:17, pi. 9: figs. 12- 

14), who reported "Dromia vulgaris" from Senegal 

(16*) and from Porto da Praia (as La Praya), Cape 

Verde Islands (1 6 juv). Forest (1974), who examined 

both specimens, identified them with D. 

marmorea. A Milne-Edwards and Bouvier's description 

and illustrations of these West African 

specimens of "Dromia vulgaris" contain some mistakes 

that have caused considerable confusion. 

The two French authors, in their account of the 

Talisman and Travailleur Brachyura reported two 

species of Dromia: Dromia vulgaris (1900:17) and D. 

nodosa (1900:18). The latter species was said in the 

text (1900:18) and in the explanation of the 

figures (1900:369) to be illustrated on plate 9: 

figs. 12-14, whereas Dromia vulgaris was said to be 

figured on plate 9: fig. 15. Actually, however, the 

reverse is true: plate 9: figs. 12-14 shows one of 

the two specimens of D. vulgaris, probably the 

larger one from Senegal, and figure 15 is based 

on the syntype of D. nodosa, which was figured 

again by Forest and Guinot (1966, fig. 2a). This 

mixup was the primary reason that Monod (1956) 

incorrectly interpreted D. nodosa. 

Irvine (1932) gave a short account and a recognizable 

figure of the present species, which he 

indicated as "Hairy Brown Sea Crab," based on 

material from the Gold Coast (Ghana), presumably 

from Accra. In a later publication, Irvine 

(1947) repeated this information and listed the 

species under the name Dromia caput-mortuum. 

According to Irvine (1947), this species is not 

considered to be edible in Ghana. Irvine also 

noted that the vernacular name of this crab in 

the Ga language of Ghana is "Tsitsikuntu," a 

name derived from the word "kuntu" for blanket 

and obviously referring to the wooly hair cover of 

the body. 

Gordon (1950:244-251), in her study of the 

morphology of the spermatheca in Dromiidae, 

figured and described the thoracic sternum of a 

female of "Dromia vulgaris" from Madeira. Forest 

(1974), who later examined the specimen, identified 

it with D. marmorea. 

Capart (1951:21, figs. 1, 3a) also reported 

"Dromia vulgaris" from West Africa. He mentioned 

a young female from Pulpito Bay, Rio de Oro 

(Spanish Sahara). Forest (1974), who examined 

the specimen, identified it with D. personata. Capart's 

illustrations were based on a specimen of 

D. personata from the English Channel, so that 

both his text and illustrations pertain to D. personata, 

not to the present species. 

The material reported upon by Sourie (1954a), 

Buchanan (1958), and Longhurst (1958) as 

Dromia vulgaris had been identified as such by 

Monod, and thus must be considered to be D. 

marmorea. As shown by the superb figures of 

"Dromia caputmortuum" in Monod's (1956) great 

work, the specimens that he considered to be that 

species (= D. vulgaris — D. personata) actually were 

D. marmorea. 

13

14 

It was Forest (1974) who first pointed out that 

the West African form is distinct from the Mediterranean 

and NE Atlantic D. personata, and Forest 

proposed the name Dromia marmorea for it, at 

the same time providing an excellent, well-illustrated 

description. 

BIOLOGY.—This species has been reported from 

depths between 0 and 96 m, with one doubtful 

record from 100 m, but more than 90% of the 

records are from depths of less than 42 m, and 

about 60% are from 20 m or less. Irvine (1947: 

302) remarked that the species "lives in deep 

water and is sometimes caught in bottom-nets a 

mile or more from land." It has been reported 

from the following types of bottom: mud (Crosnier, 

1967); very fine sand (Buchanan, 1958); 

sand and shells (Forest, 1974); muddy sand 

(Longhurst, 1958); rocks (Sourie, 1954a; Monod, 

1956; Forest, 1974). Monod (1956) reported a 

juvenile specimen from the hull of a ship. Sourie 

(1954a) classed this species as characteristic of the 

"hypobioses lapidicoles du sous-etage," while Buchanan 

considered it to belong to the active 

epifauna of the inshore fine sand community off 

Ghana. 

Osorio (1889) is the only author to report that 

this species carries a sponge. Sourie (1954a: 247) 

stated that it mostly bears didemnid or polycitorid 

ascidians. Two specimens from Bel-Air, 

Dakar, in the collection of the Smithsonian Institution 

(USNM 152646) have balanids all over 

the central and anterior part of the carapace; if 

these specimens carried a sponge or ascidian it 

must have covered a small area on the posterior 

part of the body. 

Ovigerous females have been taken in September, 

November, and December (Monod, 1956; 

Forest, 1974). 

DISTRIBUTION.—Dromia marmorea is a West African 

species, reported from the Azores, Madeira, 

and the Canary Islands south to Cabinda, and 

Sao Tome and Principe islands in the Gulf of 

Guinea. Monod (1956:62) reported two males 

and a female from "Cap Blanc, M. H. Routh 

coll., 1951 (B.M.)"; an examination of this lot in 

the British Museum revealed that Monod's local- 


ity indication was erroneous, for the specimens 

actually originate from Gambia (see "Other Material" 

p. 11). The records of the species in the 

literature are as follows: 

Azores: Ilha do Muda [?], off Ilha das Flores, 22-30 m; 

Caldeira Inferno, Ilha do Faial, 10 m; Ponta Sao Diego [?], 

Ilha Terceira, 0-30 m; Ponta Delgada, 15-20 m, Caloura, 3 

m, and Ponta da Galera, 10-12 m, Ilha de Sao Miguel (all 

Forest, 1974). 

Madeira: No specific locality (Gordon, 1950; Forest, 1974; 

Tiirkay, 1976b). Funchal harbor (Tiirkay, 1976b). 

Canary Islands: Santa Cruz de La Palma, Isla de La 

Palma, 15 m; Playa de los Abrigos, Las Caletillas, 100 m 

(?), Puerto de la Cruz, intertidal, and Ensenada de Cristianos, 

intertidal, Isla de Tenerife; Playa Quemada, 2-3 m, 

and Arrecife, Isla de Lanzarote (all Forest, 1974). 

Cape Verde Islands: Porto da Praia (as La Praya), Sao 

Tiago, 10-30 m (A. Milne Edwards and Bouvier, 1900; 

Forest, 1974); Porto da Praia, Sao Tiago (as Porto Praia (I. 

Santiago)), the type-locality (Forest, 1974). 

Mauritania: No specific locality (Monod, 1956; Forest, 

1974). 

Senegal: No specific locality, in 80 m (A. Milne Edwards 

and Bouvier, 1900; Forest, 1974). Dakar (Monod, 1956; 

Forest, 1974). Goree Island, near Dakar, beach, 2-3 m and 

96 m; S of Goree, 33-35 m and 40 m (Monod, 1956; Forest, 

1974). Bel-Air, near Dakar, 5-10 m (Forest, 1974). Anse 

Bernard, Dakar; Goree, 23 m; Mbour, 25 m; Joal (Monod, 

1956). 

Gambia: Erroneously cited as Cap Blanc, Mauritania 

(Monod, 1956). 

Guinea-Bissau: 10°22'N, 16°22'W, 41 m (Forest, 1974). 

Guinea: 09°22'N, 13°42'W, 20-35 m; Guinea, 15-20 m; 

09°N, 13°50'W, 30 m (Monod, 1956). 09°20'N, 14°15'W, 

32 m (Forest, 1974). 

Sierra Leone: Off Freetown, 15 m (Forest, 1974). 

Ghana: Gold Coast (Irvine, 1932); same, 23-45 m 

(Longhurst, 1958). Near Accra, in deep water (Irvine, 1947); 

same (Monod, 1956); same, in 5.5-14.6 m (Buchanan, 1958); 

same, in shallow water to 25 m (Gauld, 1960); same (Forest, 

1974). Chorkor, near Accra, beach seine (Monod, 1956). 

Principe: No specific locality (Osorio, 1889, 1898). 

Sao Tome: Praia Lagarto (Osorio, 1889, 1898). 

Gabon: Cap Lopez, 20 m (Crosnier, 1967; Forest, 1974). 

Congo: Baie de Pointe-Noire, beach seine (Rossignol, 

1962; Crosnier, 1967; Forest, 1974). Off Pointe-Noire, 50 m 

(Crosnier, 1967; Forest, 1974). 

According to Forest (1974) the present species 

also occurs in Saint Helena, and he assigned 

material from there identified as Dromia vulgaris 

by Melliss (1875) and Cunningham (1910), as 

well as that reported upon as Dromia species? by

NUMBER 306 

Colman (1946) and as Dromia erythropus by Chace 

(1966), to the present species. The records by 

Rathbun (1900a) and Balss (1921) of Dromia vulgaris 

from St. Helena are not original, but based 

on Melliss' (1875) record. We have examined 

Chace's (1966) St. Helena material and found it 

impossible to confidently assign it to the present 

species, although it indeed is very similar, but so 

is it similar to Dromia erythropus and D. personata. 

Therefore, we have not included St. Helena in 

the range of D. marmorea. It seems possible that 

the St. Helena Dromia belongs to a species distinct 

from either D. marmorea and D. erythropus, but 

more material is needed to decide the status of 

that form; it also is possible, although less likely, 

that more than one species of Dromia occurs in St. 

Helena waters. 

* Dromia monodi Forest and Guinot, 1966 

FIGURE 3a, b 

Dromia spinirostris Osorio, 1889:136, 139; 1898:193—Balss, 

1921:47—Capart, 1951:23 [part, only specimens from A.S. 

141 and Mercator]. [Not Dromia spinirostris Miers, 1881 = 

Stemodromia spinirostris.] 

Dromia atlantica.—Rathhun, 1921:393, fig. 1, pi. 18: fig. 3 

[not Dromia atlantica Dofiein, 1904 = Stemodromia spinirostris 

(Miers, 1881).] 

Dromia nodosa.—Monod, 1956:65 [part], figs, 52-71, 83b.— 

Longhurst, 1958:87.—Gauld, 1960:68. [Not Dromia nodosa 

A. Milne Edwards and Bouvier, 1898.] 

Dromia monodi Forest and Guinot, 1966:43, fig. 1.—Crosnier, 

1967:321 [part, only the male specimens].—Le Loeuff and 

Intes, 1968:38, tables 1,5,9.—Forest, 1974:96, figs, le, 2, 

3f,g, 5, 7a,b, pi. 4: figs. 1-3, pi. 6: fig. 2. 

Dromia sp.—Forest and Guinot, 1966:46.—Forest, 1974:99. 

PDromia.—Maurin, 1968b, fig. 4. 

PDromia nodosa.—Maurin, 1968b:484, 489. 

Not Dromia nodosa.—Rossignol, 1962:113 [= Stemodromia spinirostris 

(Miers, 1881)]. 

MATERIAL EXAMINED.—Pillsbury Material: Ghana: 

Sta 26, 27 m, shell bottom (scallops), 1$ ov (L). 

Nigeria: Sta 248, 33 m, 5c5, 5$ (2 ov) (L, W). Sta 250, 

24 m, brackish water, 19 (L). 

Other Material: Senegal: Dakar, 10 m, Dec 1951, E. Postel, 

1

16 


FIGURE 3—Sternal grooves of females of various sizes. Dromia monodi Forest and Guinot, Pillsbury 

Sta 248: a, cb 14 mm, b, cb 38 mm. Sternodromia spinirostris (Miers), Pillsbury Sta 47: c, cb 21 mm. 

two ovigerous females, cl 14 and 18 mm, bridging 

nicely the existing gap. The smallest of these 

specimens has the sternal grooves rather straight 

and wide apart distally and each ends at the base 

of a tubercle; these tubercles are connected by a 

low wide ridge. The structure of the two tubercles 

and the ridge could very well be described as a 

strong transverse swelling, excavated in the middle. 

The specimen of 18 mm shows about the 

same situation, only the tubercles are less distinct 

and the ridge is absent, the surface between the 

tubercles being flush with the rest of the surface. 

In the ovigerous female of cl 47 mm the situation 

resembles that in the specimen of 18 mm, but 

here the tubercles are even less distinct as shown 

for a specimen 38 mm long in Figure 3b. We have 

not been able to find any consistent difference in 

the sharpness of the tubercles of the carpus of the 

cheliped in large and small, ovigerous and nonovigerous 

specimens, and believe that this character, 

to which Forest and Guinot themselves 

attached very little importance, indeed is of no 

taxonomic value here. We have come to the 

conclusion therefore that the specimens indicated 

by Forest and Guinot as Dromia sp. can safely be 

considered to belong to D. monodi. Forest (1974: 

100) considered it most likely that the specimens 

are precocious D. monodi, but did not exclude the 

possibility that they are a new species. 

An interesting feature of this species is the 

arrangement of the sternal grooves in the female. 

As already stated above, in the ovigerous females, 

whether large or small, these grooves end wide 

apart, each at the base of a low rounded tubercle, 

which in the small specimens is somewhat more 

distinct than in the large. From the genital opening 

the groove extends straight backward, parallel 

and rather close to the lateral margins of the 

thoracic sternum; the anterior ends of these 

grooves are definitely not turned inward. In the 

non-ovigerous females, again whether small or 

large, the tips of the grooves are distinctly curved 

inward towards each other, and no tubercles are 

visible; in these specimens the tubercle is replaced 

by a depression (Figure 3a, $ cb 14 mm; 3b, 9 cb 

38 mm). An excellent figure of the latter situation 

also is given by Rathbun (1921:393, fig. le). 

Forest and Guinot (1966) showed that the 

Dromia species from West Africa indicated by 

Monod (1956) as Dromia nodosa is specifically distinct 

from the type-specimen of the true Dromia 

nodosa A. Milne Edwards and Bouvier, 1898, and 

gave it the new name Dromia monodi. The source 

of the confusion was given by Forest and Guinot 

(1966:45) as "l'inexactitude du dessin ensemble 

(A. Milne Edwards et Bouvier, pi. IX; fig. 14)," 

evidently meaning figure 12, not 14; also Forest 

(1974:97) mentioned "le peu de fidelite du dessin 

publie par A. Milne Edwards et Bouvier (1900; 

pi. 9, fig. 12)." Examination of the text and

NUMBER 306 

figures published by A. Milne Edwards and Bouvier 

shows, however, that figure 12 is not inexact, 

but has only been given the wrong name. Comparing 

Forest and Guinot's figure 2a of the type 

of Dromia nodosa with A. Milne Edwards and 

Bouvier's (1900, pi. 9: fig. 15) of "Dromia vulgaris" 

it is clear that these two are made after the same 

specimen. What has happened, therefore, obviously 

is that A. Milne Edwards and Bouvier 

(1900) on their plate 9 gave as figures 12 to 14 

the entire animal and details of what in the text 

they called "Dromia vulgaris," and on figure 15 the 

type of Dromia nodosa in dorsal view, but that in 

the explanation of these figures (1900:369 and pi. 

9) they switched the names, and also in the text 

referred to the wrong figures. This is confirmed 

in the explanation (1900:369) of plate 9: figure 

15 ("Dromia vulgaris"), which is said to be of "un 

des petits exemplaires des lies du Cap-Vert," 

which evidently refers to Dromia nodosa of which 

the authors had 5 specimens from the Cape Verde 

Islands, while in their material "Dromia vulgaris" 

was represented by only a single Cape Verde 

Islands specimen. A Milne Edwards and Bouvier's 

(1900, pi. 9: figs. 12-14) illustrations are 

actually made after a specimen they named 

Dromia vulgaris, probably the large male from 

Senegal, which was shown by Forest (1974:79) to 

belong to Dromia marmorea. It is not surprising 

therefore that Monod (1956) arrived at the wrong 

conclusion about the identity of Dromia nodosa, 

especially since A. Milne Edwards and Bouvier's 

(1900) figure 12 shows the second and third anterolateral 

tooth of the carapace quite close together 

as in Dromia monodi. 

As Forest and Guinot (1966:46) indicate, it is 

more likely that Osorio's (1889:136, 139) Dromia 

spinirostris from Sao Tome and Principe islands 

belongs to Dromia monodi rather than to Sternodromia 

spinirostris, because the latter species usually 

is found at greater depths than the 4 or 5 m from 

which Osorio reported his male. Monod (1956) 

referred the specimens listed by Balss (1921) under 

Dromia spinirostris to the present species, evidently 

because Balss did not find a median tubercle 

between the ends of the sternal grooves of 

his female. As Balss' specimens were collected at 

depths of 10 and 20 m, it seems highly likely that 

Monod's surmise is correct, even though the character 

of the sternal tubercle does not always hold 

good. Monod (1956) also referred the material 

that Odhner (1923) identified as Dromia spinirostris 

to the present species, but there are reasons to 

believe that Odhner's identification was correct 

(see p. 19). 

Rathbun's (1921) description and illustrations 

of what she thought to be Dromia atlantica Doflein 

leave no doubt that her material was actually D. 

monodi; already Monod (1956) pointed this out. 

Monod (1956) very extensively figured the 

present species and identified it with Dromia nodosa 

A. Milne Edwards and Bouvier, being followed 

in this by Longhurst (1958), Gauld (1960), Rossignol 

(1962), and possibly Maurin (1968b). Forest 

and Guinot (1966) pointed out that the present 

species is different from the true Dromia nodosa, 

and proposed the new name Dromia monodi for it, 

which name has been adopted by subsequent 

authors. 

Some of the specimens, mostly juveniles, that 

Monod (1956) identified as Dromia nodosa proved 

on examination to belong to neither that species 

nor to D. monodi. Forest (1974:79) found that the 

specimens indicated by Monod with the numbers 

1-2 (Mauritania), 5, 7, 12(1), 15, 16(4) (all from 

Goree), actually belong to Dromia marmorea. Examination 

of the specimen from Murray Town, 

Sierra Leone (Monod, 1956:69, no. 52), proved 

that it belongs to Stemodromia spinirostris. 

Crosnier (1967:321) assigned two males and 

two females to the present species. The females 

are both juvenile, one is from Dahomey, the other 

from "Pte Ste Clara" (= Cap Santa Clara), Gabon. 

The Gabon female had been reported on 

before by Rossignol (1962) under the name 

Dromia nodosa. The two females are now in the 

collection of the Museum national d'Histoire naturelle, 

Paris, and were examined by us. Although 

juveniles of Dromia monodi and Stemodromia spinirostris 

are usually difficult to distinguish, we believe 

that both specimens should be assigned to 

Stemodromia spinirostris rather than to Dromia mon- 

17

18 

odi. Forest (1974:101) also assigned the Dahomey 

specimen to Sternodromia spinirostris, but did not 

mention the Cap Santa Clara specimen. The two 

male specimens listed by Crosnier (1967), from 

Cap Lopez (Gabon) and Cabinda, are true Dromia 

monodi (Forest, 1974:96, and p. 15). 

Maurin (1968b:484, 489) reported Dromia nodosa 

from 200 m depth, dredged between Cabo 

Corbeiro and Cabo Blanco (= Cap Blanc), Spanish 

Sahara, and also from 60 and 70 m depth on 

the Bane d'Arguin, Mauritania. The species is 

not noted in Maurin's (1968a) more extensive 

treatise of the same area. As no morphological 

data are provided of the material, it is impossible 

to conclude whether the true Dromia nodosa or D. 

monodi is the species obtained by Maurin; the 

depth indications would indicate the former. 

Pechiiel-Loesche (1882) reported a specimen of 

Dromia from the "Banya" (= Crique Banjia, 00° 

14'N, 09°40'E), Gabon. As insufficient morphological 

details are provided by Pechiiel-Loesche, 

the identity of the species is uncertain, the description 

of its habits make it more likely that not 

Dromia but Phyllodorippe was meant (see p. 35). 

BIOLOGY.—As Forest and Guinot (1966) 

pointed out, this species occurs mostly between 0 

and 25 m depth. This is fully confirmed by the 

records below, all but one (60 m) of which are 

from less than 45 m and about 80% from 25 m or 

less. 

As to the bottom on which the species lives, 

this is given variously as green and black mud 

(Capart, 1951; Forest, 1974), "sable" (Osorio, 

1889), "sable vaseuse" (Le Loeuff and Intes, 

1968), "between sponges that surrounded Pinna" 

(Rathbun, 1921), sand, shelly sand and shelly 

mud (Longhurst, 1958), "roche, coraux" and 

"vase, coquilles" (Forest and Guinot, 1966), 

"fonds a Palythoa et Moigula, sable," "sable dur" 

(Monod, 1956). Specimens have been reported as 

collected with beach seines on sandy beaches 

(Monod, 1956; and p. 15). 

Like other species of Dromia the present species 

has been noted to carry sponges (Monod, 1956) 

and composite ascidians (Monod, 1956). Osorio 

(1889:136) reported on a specimen that carried a 


piece of decaying wood, by which it was entirely 

covered, but "il ne marchait pas tout a son aise"; 

it is possible that the collector mistook a sponge 

or ascidians for a piece of decaying wood. 

Ovigerous females have been collected in February, 

March, May, July, and October (Monod, 

1956; Pillsbury). 

DISTRIBUTION.—Dromia monodi has been reported 

from the west coast of Africa between 

Mauritania and Angola. The records from between 

Cabo Corbeiro and Cabo Blanco, Spanish 

Sahara, 200 m and Bane d'Arguin, Mauritania, 

60-70 m (both Maurin, 1968b) and Crique Banjia, 

Gabon, on the shore (Pechiiel-Loesche, 1882), 

discussed above, are excluded. Other records include 

the following: 

Mauritania: No specific locality (Forest, 1974). 

Senegal: Near Dakar, 15-20 m (type-locality) (Monod, 

1956; Forest and Guinot, 1966; Forest, 1974). SE of lie de la 

Madeleine, near Dakar, 35 m; near Dakar, 10 m and 34 m; 

Joal; Lagoba [?], 10-11 m (Monod, 1956; Forest, 1974). Bel- 

Air, near Dakar, 5-10 m (Forest, 1974). Near Goree, in 20 m 

(Balss, 1921); same, in 15-20 m (Monod, 1956). Anse Bernard, 

near Dakar, 13-15 m; between Goree and Rufisque, 

25 m; E of Bel-Air, 14 m; Ngazobil, 4 m; Joal, 4 m, 

10-11 m and 15 m (Monod, 1956). 

Guinea: 09°25'N, 13°55'W, 15 m; 09°16'N, 13°34'W, 

10 m; between Tamara and lie de Corail, 10-12 m (all 

Monod, 1956). Near Conakry, 20 m (Monod, 1956). 

09°51'N, 15°30'W (Capart, 1951; Forest, 1974). 

Sierra Leone: No specific locality, 10-22 m (Longhurst, 

1958). 08°38'-08°42'N, 8-12 m; Murray Town; Freetown 

(Monod, 1956). Off Freetown, 15 m (Forest, 1974). 

Ivory Coast: Grand-Bassam, 30 m, 60 m (Le Loeuff and 

Intes, 1968). Off San-Pedro, 20 m (Forest, 1974). 

Ghana: Near Accra, in 14, 35, 37, and 44 m (Monod, 

1956); same, in 0-45 m (Gauld, 1960). Chorkor, near Accra; 

Christiansborg; Ningo [?] (all Monod, 1956). 

Togo: No specific locality (Monod, 1956); Lome (Monod, 

1956). 

Cameroon: Douala (Balss, 1921). 

Principe: No specific locality (Osorio, 1889, 1898). 

Sao Tome: No specific locality (Balss, 1921). Praia de 

Santa Catarina, 00°16'N, 06°29'E, 3-10 m (Forest and 

Guinot, 1966). Sao Joao dos Angolares, 4-5 m (Osorio, 1889, 

1898). 

Gabon: No specific locality (Monod, 1956). Cap Lopez, 

20 m (Crosnier, 1967; Forest, 1974). Near Cap Lopez, 

OO°53'S, 08°40'E, in 33-55 m (Capart, 1951; Forest, 1974). 

Cabinda: No specific locality, in 25 m (Crosnier, 1967; 

Forest, 1974).

NUMBER 306 

Angola: Ambrizete (as Ambrizette), 10 m; Quicembo (as 

Kinsembo) (Balss, 1921). Luanda (as Sao Paulo de Loanda), 

a few feet deep (Rathbun, 1921). 

Dromia nodosa A. Milne Edwards and Bouvier, 

1898 

Dromia nodosa.—Monod, 1956:65 [part].—Forest, 1974:94, 

figs. 2, 3e, 5, 7e,f, pi. 2: figs. 3, 4, pi. 5: fig. 6 [references]. 

REMARKS.—There is a male syntype of this 

species (USNM 22937), carapace length 7.2 mm, 

taken by the Talisman in the Cape Verde Islands 

in 75 m, 29 July 1883, in the collection of the 

Smithsonian Institution. Forest (1974:95) noted 

that only three of the original five specimens 

could then be located. 

DISTRIBUTION.—Cape Verde Islands, in 75 m, 

and off the Atlantic coast of Morocco. 

Genus Sternodromia Forest, 1974 

Stemodromia Forest, 1974:100 [type-species: Dromia spinirostris 

Miers, 1881, by original designation and monotypy; gender: 

feminine]. 

* Sternodromia spinirostris (Miers, 1881) 

FIGURE 3C 

Dromia spinirostris Miers, 1881a:271, pi. 16: fig. 2.—Rathbun, 

1900a:300.—Odhner, 1923:15.—Capart, 1951:23 [part, 

not the specimens from A.S. 141 and Mercator], figs. 2, 

3b.—Rossignol, 1957:75. 

Dromia fulvo-hispida Miers, 1881a:270, pi. 16: fig. 1. 

Dromia fulvohispida.— Rathbun, 1900a: 300—Balss, 1921:47. 

Dromia atlantica Doflein, 1904:10, 189, 252, 253, pi. 7: figs. 3, 

4. 

Dromidtopsis spinirostris.—Monod, 1956:72, figs. 72-82, 83c— 

Longhurst, 1958:87.—Rossignol, 1962:113.—Forest and 

Guinot, 1966:47.—Crosnier, 1967:322.—Le Loeuff and 

Intes, 1968:38, 65, 67, 68, 71, fig. 62, tables 1, 6, 7, 8, 9; 

1969:64, 65.—Maurin, 1968a:48.—Maurin, 1968b:484, 

489, 491, fig. 7.—Tiirkay, 1975a: 71 [listed], 72.—Pastore, 

1976:114, fig. 1 [Mediterranean].—Lewinsohn, 1977:7 

[discussion]. 

Dromia nodosa.—Monod, 1956:69 [part, specimen from Murray 

Town].—Rossignol, 1962:113. [Not Dromia nodosa A. 

Milne Edwards and Bouvier, 1898.] 

Dromia.— Voss, 1966:22. 

Dromia monodi.—Crosnier, 1967:321 [part, the two female 

specimens]. 

Dromidiopsis.—Maurin, 1968b, figs. 4, 9. 

Dromidopsis spinifrons.—Maurin, 1968b:486 [erroneous spelling]- 

Stemodromia spinostris.—Forest, 1974:100, figs, If, 2, 3h, 5, 

6c,d, 7g-i, pi. 4: figs. 4,5, pi. 6: fig. 3, pi. 7: figs. 2, 4 

[erroneous spelling]. 

Not Dromia spinirostris.—Osorio, 1889:136, 139; 1898:193.— 

Balss, 1921:47. [= Dromia monodi Forest and Guinot, 

1966.] 

Not Dromia atlantica.—Rathbun, 1921:393, fig. 1, pi. 18: fig. 

3 [= Dromia monodi Forest and Guinot, 1966]. 


Sta 68, 70 m, broken shell, 2 juv (L). 

Ivory Coast: Sta 46, 38-42 m, mud bottom with dense 

Jullienella, 6

20 

and a carapace width of 72 mm; in the largest 

female these values are 58 mm and 68 mm, 

respectively. Ovigerous (and impregnated) females 

have cl 48 to 58 mm and cb 55 to 58 mm. 

In juveniles the carapace width is slightly less (0.9 

times) than the carapace length, and the larger 

specimens become relatively wider. Even in the 

largest specimen however, the ratio between carapace 

width and carapace length does not go 

beyond 1.19, the females as a rule being on the 

average somewhat wider than the males. Our 

finds agree with those by Crosnier (1967) who 

found that this species is slightly wider on the 

average than Dromia monodi, although the difference 

is slight. 

REMARKS.—The present species greatly resembles 

Dromia monodi. Differences between adult 

specimens of the two species, some of which have 

already been pointed out by previous authors 

(Monod, 1956; Forest and Guinot, 1966; Crosnier, 

1967; Forest, 1974) are the following: 

1. The pubescence in Stemodromia spinirostris 

consists of a very dense cover of short hairs of 

equal length giving the animal a smooth and 

shiny, almost sealskin-like surface. In Dromia monodi 

the hairs are of two sizes, the shorter are less 

closely placed than in S. spinirostris and therefore 

do not produce the skin-like effect, while groups 

of longer hairs, especially in the anterior and 

lateral portions of the carapace give the pubescence 

a still more shaggy appearance. 

2. The interantennular spine in S. spinirostris 

is hardly visible in dorsal view; in Dromia monodi 

it is distinct and reaches beyond the middle of the 

rostral teeth. 

3. The anterolateral teeth of the carapace in 

D. monodi are far stronger and more conspicuous 

than in Stemodromia spinirostris; moreover, they are 

directed more outwards and are less appressed. In 

5. spinirostris the teeth are inconspicuous, especially 

the second and third, which are often hidden 

by the pubescence. Some of the teeth, especially 

the fourth and fifth are often continued 

backward as a marginal carina. The first anterolateral 

tooth is far more conspicuous than the 

second. 


4. Between the first anterolateral tooth and 

the suprasutural tooth (Ihle, 1913:9, fig. 5) there 

is a marked groove in Dromia monodi; in Stemodromia 

spinirostris such a groove is hardly noticeable, 

while the teeth are less distinct. 

5. The chelipeds in Dromia monodi have strong 

tubercles: one on the outer surface of the palm at 

the base of the dactylus, and two near the anterior 

margin of the carpus. These tubercles are hardly 

noticeable in Stemodromia spinirostris being quite 

obscured by the pubescence. 

6. The outer surface of the carpus of the 

second and third pereiopods (= first and second 

walking legs) is evenly rounded in Stemodromia 

spinirostris, while in Dromia monodi it shows a distinct 

longitudinal ridge in the pubescence. 

7. The dactyli of the second and third pereiopods 

in D. monodi are shorter than in S. spinirostris 

and have stronger spines on the lower margin. 

8. The upper margin of the propodus of the 

fourth pereiopod is longer than the height of this 

segment in S. spinirostris, much shorter than the 

height in D. monodi. The dactylus is twisted in the 

latter species, lying in the same plane as the 

propodus in the former. 

9. The fifth pereiopod is more slender in D. 

monodi than in S. spinirostris. 

10. The female abdomen in non-ovigerous females 

is triangular with a rounded tip in D. 

monodi, more trapezoidal with a truncated tip in 

S. spinirostris. 

11. In the same non-ovigerous females the sternal 

grooves in S. spinirostris end somewhat closer 

together than in D. monodi. 

12. In our material of S. spinirostris the pubescence 

is grayish brown and the finger tips of the 

chelae "are white; in D. monodi the pubescence is 

more yellowish brown and the finger tips are 

pink. 

Young specimens of Stemodromia spinirostris differ 

from the adults in the following points: 

1. The pubescence of the carapace shows 

some longer hairs in the anterior part, by which 

the smooth appearance is lost and becomes more 

shaggy like in D. monodi.

NUMBER 306 21 

2. The interantennular spine is relatively 

longer and more distinct. 

3. The anterolateral teeth of the carapace are 

even less conspicuous than in the adults. 

These young specimens still can be distinguished 

from young of Dromia monodi by the character 

of the smaller anterolateral teeth and the 

more slender dactyli of the second and third 

pereiopods, in which the lower margin bears 

smaller spinules. 

The sternal grooves in the females collected by 

the Pillsbury Expedition, all of which are nonovigerous, 

very closely resemble those of non-ovigerous 

females of Dromia monodi, the only difference 

being that the anterior ends of the two 

grooves are placed somewhat closer together. In 

the Museum national d'Histoire naturelle, Paris, 

we examined five large females (cl 48 to 58 mm) 

of the present species, some of which are impregnated 

or ovigerous. In these specimens the sternal 

grooves converge anteriorly and end at a distinct 

median tubercle, as shown in Monod's (1956) 

figures 78 and 79. In a juvenile female in the 

Paris Museum (cl 34 mm) no tubercle was seen, 

but the grooves resemble those found in the Pillsbury 

specimens (Figure 3c, 9 cb 21 mm) in which 

the tubercle is visible. Evidently the arrangement 

of the sternal grooves changes drastically when 

sexual maturity is reached. It is debatable 

whether the sole character of the placement of 

the female sternal grooves is sufficient to distinguish 

genera. This seems the more doubtful when 

one considers the fact that Dromia monodi and 

Sternodromia spinirostris resemble each other so 

closely that they are difficult to separate, while 

both differ less from one another than from the 

other West African Dromiidae. Monod (1956:75) 

already commented that it does not seem logical 

that these two species should be placed in different 

genera. For the time being, however, we 

followed Forest (1974), who separated the present 

species from the other Dromidiopsis and Dromia 

species and erected a new genus, Sternodromia, for 

it. Lewinsohn (1977:7), when dealing with Dromiidae 

of the Red Sea, commented that in his view 

too much importance has been attached to the 

female sternal grooves as a generic character and 

that a revision of the family on a generic level is 

highly desirable. 

There can be little doubt that the type material 

of Miers' Dromia spinirostris belongs to the present 

species, and in the modern literature the epithet 

spinirostris has practically always consistently and 

correctly been used for it. A quite different matter 

is the identity of Dromia fulvo-hispida Miers, 1881. 

This species was based on a single juvenile specimen 

(8X11 mm), taken together with the types 

of Dromia spinirostris. The specimen was reexamined 

by Monod (1956:59), who gave additional 

details of it but did not arrive at a conclusion 

concerning its identity. We have also examined 

the specimen, which is in a poor condition; it is 

a juvenile female, which is partly damaged; therefore, 

the measurements 8X11 mm cannot be 

relied upon. We decided after studying this specimen 

that it is a juvenile specimen of Sternodromia 

spinirostris. The fact that the anterolateral margins 

show hardly any teeth (Miers described these 

margins as entire, but as Monod correctly pointed 

out there is at least one visible tooth) points 

strongly in this direction. Furthermore that it was 

taken at a depth of 9 to 15 fathoms (16.5-27 m) 

together with the types of Dromia spinirostris make 

the identity of the two highly likely. As the type 

specimen of Dromia fulvohispida shows no characters 

that make it impossible to identify it with D. 

spinirostris and as there are several indications that 

the two species are synonymous, we confidently 

sink the epithet Julvohispida as a synonym of spinirostris. 

Under Dromia monodi (p. 17) we have given the 

reasons why the specimens that Osoria (1889, 

1898) and Balss (1921) considered to be Dromia 

spinirostris actually should be referred to Dromia 

monodi. Also we point out that the female specimens 

from Dahomey and Cap Santa Clara, identified 

by Rossignol (1962) and Crosnier (1967) as 

Dromia monodi, be better referred to the present 

species. All this again clearly shows the close 

resemblance of the two species. 

The juvenile specimens from Angola that 

Odhner (1923) identified as Dromia spinirostris

22 

were referred by Monod (1956:72) to Dromia nodosa 

(= D. monodi). Although Monod examined 

one of Odhner's specimens, we are not convinced 

that his conclusion is correct. Odhner himself was 

convinced that his specimens were conspecific 

with Dromia atlantica Doflein, after he compared 

them with Doflein's description and figures; 

moreover Odhner sent his material to W. T. 

Caiman, who did not find any characters that 

would make them specifically different from 

Miers' types of Dromia spinirostris. Also the fact 

that Odhner's specimens came from depths of 72 

and 108 m makes it more likely that they belong 

to Sternodromia spinirostris than to Dromia monodi. 

As already suggested by Balss (1921), Odhner 

(1923), Monod (1956), and others, Dromia atlantica 

of Doflein (1904) is identical with the present 

species, as is clearly shown by Doflein's account. 

The juvenile male from Sta A.S. 141 and the 

juvenile male collected by the Mercator expedition, 

both of which were identified by Capart (1951) 

with the present species, were examined by Forest 

(1974:96) and found to be Dromia monodi. 

The specimen from Murray Town, Sierra 

Leone, assigned by Monod (1956:69) to Dromia 

nodosa, was examined by us and proved to belong 

to the present species. 

The frequent misidentification, especially of 

juvenile specimens, clearly shows the difficulty of 

distinguishing the various species of West African 

Dromiidae. 

BIOLOGY.—The species has been reported from 

depths between 7.5-9.5 and 108 m. As already 

pointed out by Forest and Guinot (1966) it has 

its optimum occurrence in water deeper than 25 

m, this in contrast to Dromia monodi, which is 

hardly ever found in water deeper than 25 m. 

About 90% of the published records for Sternodromia 

spinirostris are from depths of 25 m or over 

and about 80% from depths between 25 and 75 

m. Le Loeuff and Intes (1968) indicate that the 

species "vit de 35 a 60 m et surtout a 40 et 50 m." 

The bottom on which the species has been taken 

has variously been described as "partly shelly and 

partly muddy" (Miers, 1881a); sand and shell 

fragments, sand and muddy sand with broken 


shells and stones (Odhner, 1923); sand, mud, 

coral and rock, black mud, mud and brown sand, 

green mud and sand, brown mud, mud (Capart, 

1951); mud (Rossignol, 1957); muddy sand 

(Longhurst, 1958); mud, rock, calcareous algae, 

sand and Foraminifera, mud and sand, mud and 

shells, muddy sand and Foraminifera, mud with 

Area shells (Forest and Guinot, 1966); sandy mud 

and shells (Voss, 1966; and p. 19 herein); sandy 

mud with Foraminifera (Crosnier, 1967); "fonds 

detritiques en vases" (Maurin, 1968a, 1968b); 

sand or sand with shells and Suberites, liquid mud 

sometimes mixed with very fine sand (Maurin, 

1968b); mud, Foraminifera, dark red bryozoans, 

mud with Jullienella (p. 19). Le Loeuff and Intes 

(1968), who obviously carefully studied the species, 

stated that in Ivory Coast waters they only 

found it on sandy mud bottoms, and they ranged 

it among the "vasicoles." 

None of our specimens carried a sponge or 

ascidians, and we know of no record in the literature 

indicating such an association. 


April, May, and December (Capart, 1951; 

Forest and Guinot, 1966; Forest, 1974; and p. 

19). 

DISTRIBUTION.—Sternodromia spinirostris is a West 

African species which occurs from the Spanish 

Sahara to Angola. Pastore (1976) recently recorded 

the species from the Gulf of Taranto in 

the Mediterranean. The West African records in 

the literature are as follows: 

Spanish Sahara: Islote Virginia (as Hot Virginie) (Forest, 

1974). Between Cabo Barbas and Cabo Blanco, 50-90 m 

(Maurin, 1968b). Between Cabo Corbeiro and Cabo Blanco, 

60-80 m (Maurin, 1968a). Cabo Blanco, 2O°37.7'N, 

17°24.4'W, 57 m (Tiirkay, 1975). 

Mauritania: 20 c 55'N, 17°22'W, 60 m; and 20°19.5'N, 

17°13.8'W, 20 m (Forest, 1974). Bane d'Arguin, 30-40 m 

(Maurin, 1968b). 

Cape Verde Islands: 15°16'28"N, 23°27'24"W, 40-45 m 

(Forest, 1974). 

Senegal: PSenegal, 50-62 m (Forest, 1974). Saint-Louis, 

35-40 m (Maurin, 1968b). Baie de Goree, 18-28 m (Miers, 

1881a). Between Goree and Cap Manuel, 7.5-9.5 m; Ngaparou 

(Forest, 1974). 13°01'N, 17°24'W, 51-55 m; and 

12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 1966; 

Forest, 1974).

NUMBER 306 

Gambia: 13°12'N, 17°O3.5'W, 21 m (Forest, 1974). 

Guinea-Bissau: 10°19'N, 16°24'W, 60-73 m (Forest and 

Guinot, 1966; Forest, 1974). 

Guinea: 09°28'N, 14°58'W, 45 m (Forest, 1974). 09°15'N, 

14°50'W, 45 m (Monod, 1956). 

Sierra Leone: No specific locality, in 39 m (Longhurst, 

1958). Murray Town (Monod, 1956). 

Ivory Coast: No specific locality (Le Loeuff and Intes, 

1969). 04°35'N, 06°40'W to 04°35'N, 06°41'W, 64 m (Voss, 

1966). O5°O2.5'N, 05°25'W, 21-27 m (Forest and Guinot, 

1966; Forest, 1974). 05°11.5'N, 04°09'W, 40 m; 05°12.5'N, 

04°05'W, 40 m; and 05°04'N, 03°22.5'W, 50 m (Forest, 

1974). Fresco; Grand-Lahou, 40 m; Jacqueville; Grand-Bassam, 

35-60 m (overall depth range 15-60 m) (all Le Loeuff 

and Intes, 1968). 

Ghana: 05°18'N, 00°24'W, 35-37 m (Forest, 1974). 

Dahomey: 06°04'N, 01°38.5'E, 48 m (Forest, 1974). 06° 

10'N, 02°02'E, 45 m (Crosnier, 1967; Forest, 1974). 

Nigeria: 04°03'N, 06°12'E, 32 m (Forest and Guinot, 

1966; Forest, 1974). 

Cameroon: 03°54'N, 08°50'E, 65-70 m (Forest, 1974). 

Gabon: W of Cap Santa Clara, 20-40 m (Rossignol, 1962; 

Crosnier, 1967). Off Pointe Banda, 03°57.5'S, 10°36.5'E, 

85 m (Capart, 1951). 

Congo: Pointe-Noire, 100 m (Forest, 1974). Off Pointe- 

Noire, 50 m (Rossignol, 1957, 1962). Off Pointe-Noire, 04° 

23'S, 1 l°07'E, 88 m, and 05°S, 11°36'E, 50 m (Forest, 1974). 

Zaire: Off Banana, 05°54'S, 11°58.5'E, 50 m; 05°55'S, 

12°01'E, 25-30 m; and 05°56'S, 12°E, 50-60 m (Capart, 

1951; Forest, 1974). 

Angola: Off Moita Seca, 06°06'S, 12°14'E, 12-15 m 

(Capart, 1951; Forest, 1974), and 06°16'S, 12°07'E, 50 m 

(Capart, 1951). Off the mouth of the Congo River, 06°18'S, 

12°02'E, 44 m (Doflein, 1904). Near Ambriz, O7°57'S, 13° 

05'E, 40-50 m (Capart, 1951; Forest, 1974). Ambriz, 70 m 

(Forest, 1974). Porto Alexandre, 72 and 108 m (Odhner, 

1923). 

Family DYNOMENIDAE Ortmann, 1892 

DYNOMENIDAE Ortmann, 1892:538, 541, 543. 

EASTERN ATLANTIC GENERA.—One, Dynomene, 

represented in the tropical fauna. 

EASTERN ATLANTIC SPECIES.—One, Dynomene Jilholi, 

material of which was collected by the Pillsbury. 

Genus Dynomene Desmarest, 1823 

Dynomene Desmarest, 1823:422 [index] [a genus without included 

nominal species; type-species: Dynomene hispida 

Guerin-Meneville, 1832, by subsequent monotypy by 

Guerin-Meneville, 1832, in 1829-1844, pi. 14; gender: 

feminine]. 

Maxillothrix Stebbing, 1921:456 [type-species: Maxillothrix actaeiformis 

Stebbing, 1921, a subjective junior synonym of 

Dynomene pilumnoides Alcock, 1899, by monotypy; gender: 

feminine]. 

REMARKS.—So far as we can determine, the 

first latinized use of the generic name Dynomene 

was by Desmarest (1823:422) in the index to his 

article in the Dictionnaire des Sciences Naturelles. In 

the text of this article (182S:249, footnote) the 

name is used in the vernacular (DYNOMENE). Desmarest 

again used the vernacular form in his book 

Considerations Generates sur la Classe des Crustaces 

(1825:133, footnote), using DYNOMENE in the text 

(p. 133), DYNOMENE HISPIDE in the list of 

figures (p. 432), and Dynomene hispide on the 

plate itself (plate 18); in this work he gave the 

Latin name again in the index (p. 442). The 

specific epithet hispida, often attributed to Latreille 

or Desmarest, apparently was first used by 

Guerin-Meneville (14 July 1832: pi. 14). Guerin 

also used the Latin name in the text of his Iconographie 

which, however, was published as late as 

September 1844. We are much indebted to Col. 

C. F. Cowan (in litt., 29 September 1971), who 

provided the information that livraison 22 of 

Guerin's Iconographie, which was published on 14 

July 1832 (Cowan, 1971:29), contained Crustacea 

plates 12 and 14. 

* Dynomene filholi Bouvier, 1894 

Dynomene filholi.—Monod, 1956:76, figs. 84-88, 873.—Forest 

and Guinot, 1966:48. 


Sta 275, 9-69 m, rubble of coralline algae, 26, 1? (L). Sta 

282, 18-37 m, nodular coralline algae, 16, 59, 6 juv (L). Sta 

283, 51-55 m, nodular coralline algae, 25

24 

circular, about IV4 times broader than long, depressed, 

moderately convex. Fronto-orbital region 

forming on obtuse triangle, frontal margin thickened, 

fronto-orbital width about 2/3 carapace 

width. Regions well marked but obscured by coat 

of low setae. Several rounded prominences on 

surface, each with long tuft of setae: usually 1 

epigastric, 2 or 3 protogastric, 1 metagastric, 1 

intestinal, and 5 branchial. Anterolateral margins 

with 5 blunt teeth, each tufted. 

Chelipeds stout, subequal, larger in males, covered 

with low setae, ornamented with rounded, 

tufted prominences. Merus with subdistal prominence 

on posterior margin, smaller, lower naked 

tubercles on inner margin. Carpus with 3 tufted 

prominences on posterior margin, 1 dorsally, 1 

anteriorly; inner angle produced into slender, 

flattened lobe, almost spatulate. Propodus stout, 

high, slightly longer than movable finger, all with 

almost naked ventral surface ornamented with 

rounded prominences. Upper and outer surfaces 

of chela with longer setae, denser on inner face. 

Fingers gaping, movable finger with prominent 

tooth near midlength, naked apically, apices 

spatulate. 

Walking legs decreasing in length posteriorly, 

surface densely tomentose, dorsal surfaces with 

rounded, tufted prominences and some scattered 

longer setae. Dactyli curved, apices corneous, ventral 

margins toothed. Fifth legs very short, not 

extending much beyond midlength of merus of 

fourth, dactylus very small, subterminal, leg appearing 

chelate. 

Abdomen of 7 somites in both sexes, broadening 

posteriorly in males. 

Figures: A. Milne Edwards and Bouvier, 1900, 

pi. 3: fig. 3 (color); Monod, 1956, fig. 873. 

Male Pleopod: A. Milne Edwards and Bouvier, 

1900, pi. 8: fig. 13; Monod, 1956, figs. 84-88 

(both Cape Verde Islands). 

MEASUREMENTS.—Our specimens have carapace 

widths of 3 to 16 mm; the carapace widths 

of ovigerous females are 7 to 12 mm. 

REMARKS.—The male pleopods of our specimens 

have a large, triangular apical lobe, resembling 

that of the pleopod figured by A. Milne 


Edwards and Bouvier (1900, pi. 8: fig. 13) rather 

than that figured by Monod (1956, figs. 84-88). 

BIOLOGY.—The depth range of the species is 

from 23 to 1477 m (with one record of 9-47 m), 

but 80% of the records are from between 23 and 

75 m. As the Talisman and Travailleur records are 

notoriously unreliable, it is possible that the records 

from 110-180 and 150-275 m are incorrect. 

Also the depth of 1477 m (Monaco Expedition 

Sta 1209) needs confirmation. 

The Pillsbury specimens all were taken from a 

bottom of nodular coralline algae (diameter of 

the nodules about 100-150 mm; see Voss, (1966: 

50, fig. 14)). In the literature the species is reported 

from "fond dur" (Bouvier, 1922:90), 

"banes de corail rouge" (A. Milne Edwards and 

Bouvier, 1900:9) and from bottoms with calcareous 

algae or sand, rock, corals and calcareous 

algae (Forest and Guinot, 1966). Off West Africa, 

ovigerous females have been collected in May 

{Pillsbury). 

DISTRIBUTION.—This is an insular West African 

species, known so far from the Cape Verde islands 

(the type-locality) and the Gulf of Guinea Islands, 

Principe, and Annobon; it has not been recorded 

from the mainland. Monod (1956) reported material 

from the Cape Verde Islands and Principe. 

Since 1956 Dynomene filholi has been recorded from 

the following localities: 

Principe: 01°35'N, O7°28'E, 45 m; and 01°38'35"N, 

07°21'35"E, 35 m (Forest and Guinot, 1966). 

Annobon: O1°27.5'S, 05°36.5'E, 35 m; N of San Antonio, 

23 m; 01°26'15"S, 05°35'40"E, 60 m (Forest and Guinot, 

1966). 

Family LATREILLIIDAE Stimpson, 1858 

LATREILLIDEA Stimpson, 1858c:226 [corrected to Latreilliidae 

by Stebbing, 1902:23; name 373 on Official List]. 

EASTERN ATLANTIC GENERA.—One, Latreillia, 

represented in the tropical fauna. 

EASTERN ATLANTIC SPECIES.—One, Latreillia elegans, 

not represented in the Pillsbury collections. 

Genus Latreillia Roux, 1830 

Latreillia P. Roux, 1830, pi. 22 [type-species: Latreillia elegans

NUMBER 306 

Roux, 1830, by monotypy; gender: feminine; name 1630 

on Official List]. 

Praetor Gistel, 1848:ix [erroneously substituted for Latreillia 

(see Rathbun, 1937:73); type-species: Latreillia elegans 

Roux, 1830; gender: masculine]. 

Latreillia elegans Roux, 1830 

Latreillia elegans.—Monod, 1956:78 [references].—Zariquiey 

Alvarez, 1968:307 [Spain; references]. 

Latreilla elegans.—Tiirkay, 1976a:25 [listed], 36, fig. 16 [Morocco; 

erroneous spelling]. 

DISTRIBUTION.—Mediterranean Sea and adjacent 

eastern Atlantic, from Portugal, the Azores, 

off the Sahara coast, and the Cape Verde Islands, 

in depths between 100 and 475 m. Ascension 

Island (Stebbing, 1914); western Atlantic (Rathbun, 

1937). 

Family HOMOLIDAE de Haan, 1839 

HOMOLIDEA de Haan, 1839:102 [corrected to Homolidae by 

White, 1847a:55; name 243 on Official List, there attributed 

to White, 1847, in error]. 

THELXIOPEIDAE Rathbun, 1937:62 [name 278 on Official 

Index]. 

EASTERN ATLANTIC GENERA.—Three, two of 

which, Homola and Paromola, are represented by 

species occurring off tropical West Africa. The 

other genus is Homologenus A. Milne Edwards (in 

Henderson, 1888:20): A substitute name for Homolopsis 

A. Milne Edwards (1880:34), an invalid 

junior homonym of Homolopsis Bell, 1862 (typespecies: 

Homolopsis rostratus A. Milne Edwards, 

1880, by monotypy; gender: masculine). 

EASTERN ATLANTIC SPECIES.—Three, two of 

which were recorded by Monod (1956), as follows: 


Thelxiope barbata Homola barbata* 

Paromola cuvieri Paromola cuvieri 

The extralimital species is Homologenus rostratus 

(A. Milne Edwards, 1880): Eastern Atlantic from 

the Azores, Madeira, and off Morocco, 1435 to 

2195 m (Tiirkay, 1976a); western Atlantic (Rathbun, 

1937). 

Genus Homola Leach, 1815 

Thelxiope Rafinesque, 1814:21 [suppressed by the International 

Commission on Zoological Nomenclature, Opinion 

522 in Bulletin of Zoological Nomenclature, 19:211; type-species: 

Thelxiope palpigera Rafinesque, 1814, by monotypy; 

gender: feminine; name 1190 on Official Index]. 

Homola Leach, 1815a:324 [type-species: Homola spinifrons 

Leach 1815, a subjective junior synonym of Cancer barbatus 

Fabricius, 1793, by monotypy; gender: feminine; name 

1301 on Official List]. 

Homolax Alcock, 1899b: 124, 129, 156 [type-species: Homola 

megalops Alcock, 1894, by monotypy; gender: feminine]. 

Moloha Barnard, 1947:371 [type-species: Latreillopsis alcocki 

Stebbing, 1920, by monotypy; gender: feminine]. 

* Homola barbata (Fabricius, 1793) 

Thelxiope barbata.—Gordon, 1950:221, 230, 232, 239, 250, 

figs. 19, 26b-d.—Monod, 1956:79.—Maurin, 1968b: 

486.—Le Loeuff and Intes, 1968:31, table 1. 

Homola barbata.—Figueira, 1960:7.—Guinot and Ribeiro, 

1962:23.—Peres, 1964:20.—Forest and Guinot, 1966: 

48—Crosnier, 1967:322.—Maurin, 1968a: 19, 30, 43, 61, 

107 [p. 107 Balearic Isles]; 1968b:480, 484.—Zariquiey 

Alvarez, 1968:304, figs. 12g, 106c [Spain; references].— 

Rice and Provenzano, 1970:446 ff., figs. 1-15 [larvae].— 

Tiirkay, 1976a:25 [listed], 36 [Portugal, in part]; 1976b: 

61 [listed], 62. 

SYNONYMS.—Cancer cubicus Forskal, 1775 (suppressed 

by ICZN); Cancer novemdecos Sulzer, 1776 

(suppressed by ICZN); Thelxiope palpigera Rafinesque, 

1814; Homola spinifrons Leach, 1815; Dorippe 

spinosus Risso, 1816. 


Sta 68, 70 m, broken shell, 19 ov (L). 

Nigeria: Sta 241, 59-63 m, mud and shell, 19 (W). 

Annobon: Sta 283, 51-55 m, nodular coralline algae, 1(5 

(W). 

Other Material: Congo: Off Pointe-Noire, 04°56'S, 11° 

31'E, 95-97 m, trawl, 21-22 Sep 1965, 2

26 

Lateral margin of carapace with line of spinules 

decreasing in size posteriorly. Linea anomurica 

indistinct. Front, under orbit, with 3 spines forming 

triangle. Pterygostomian region with curved 

row of 6 spines dorsally, lower margin with curved 

row of 5-6 spines, and between these, 1-2 spines; 

below and posterior to these larger spines are 

numerous small spines. Raised anterior margin of 

epistome terminating before meeting in midline, 

leaving smooth median gap. Eyes elongate, stalk 

greatly enlarged proximal to cornea. Basal segment 

of antennal peduncle with outer spine. 

Chelae slender, equal. Merus with 2 lines of 

spines below, 1 above, lower, outer spines largest. 

Carpus with 1 large and 1 small inner spines, 

outer surface with 6-8 spines arranged in 2 irregular 

rows and 2 distal spines. Palm with low, 

blunt spine on outer margin. Fingers shorter than 

palm, cutting edges corneous, tips of fingers crossing 

when closed, movable finger with low, obtuse 

prominence proximally on cutting edge. 

Walking legs compressed, thin, elongate. Meri 

with row of fixed spines above. Propodi elongate, 

each shorter than respective merus, with row of 

4 slender, movable spines below and 2 distal 

spines. Dactyli elongate, each equal to or slightly 

longer than respective carpus, more than half as 

long as respective propodus, with 11-13 spines 

below. Fifth leg with dorsal spine on coxa, 1 or 2 

spines on ischium; merus with 4 large posterior 

spines and 1 terminal anterior spine; carpus with 

numerous small spines anteriorly; propodus with 

numerous spines on opposable margin; dactylus 

with 4 spines on opposable margin. 

Abdomen of 7 somites in both sexes. Second 

segment of abdomen with prominent conical 

tooth. 

Figures: Rathbun, 1937, fig. 16, pi. 15: figs. 1, 

2; Rice and Provenzano, 1970, fig. 2. 

Male Pleopod: Gordon, 1950, figs. 26b-d (Madeira). 


lengths of 16 to 28 mm; the carapace length 

of the ovigerous female is 22 mm. 

REMARKS.—In addition to the specimens collected 

by Pillsbury, we have been able to examine 


two larger males (cl 29 and 30 mm) reported off 

Pointe-Noire in 95-97 meters by Crosnier (1967: 

322). The male pleopods of these specimens and 

that from a male (cl 32 mm, Leiden Crust. 

D.I2989) from Naples, the type-locality, closely 

resemble that illustrated from a small male, cl 14 

mm, from Madeira by Gordon (1950). The Mediterranean 

and eastern Atlantic populations of 

this species apparently are conspecific. 

On the basis of their study of larvae identified 

with this species, Rice and Provenzano (1970) 

reinforced earlier suggestions by Rice (1964) and 

Rice and Von Levetzow (1967) that there might 

be several taxa, each with a distinctive larval 

form, currently placed in H. barbata. The recognized 

three groups of larvae are the western Atlantic, 

eastern Atlantic (including the Mediterranean 

and some specimens from South Africa), 

and a group found only off South Africa. Rice 

and Provenzano examined postlarval specimens 

from the western North Atlantic, Brazil, the Mediterranean, 

and South Africa but could find no 

consistent differences between specimens from 

these areas. However, they suggested that eastern 

Atlantic representatives of H. barbata matured 

faster and grew larger than specimens from other 

areas. 

BIOLOGY.—Homola barbata inhabits moderate 

depths, having been taken in 55-679 m in the 

western Atlantic (Rathbun, 1937) and between 

50 and 400 m in the eastern Atlantic (Monod, 

1956); records since 1956 indicate that the species 

lives in depths between 10-30 and 470-500 m in 

the eastern Atlantic. The Pillsbury specimens were 

taken on broken shell in 70 m, on mud and shell 

in 59-63 m, and in nodular coralline algae in 51- 

55 m. This species also has been taken on mud in 

215 m (Guinot and Ribeiro, 1962); reddish gravel 

with shell debris in 210 m (Peres, 1964); calcareous 

algae in 73 m (Forest and Guinot, 1966); 

muddy sand in 95-98 m (Crosnier, 1967); muddy 

sand or sandy detritus in 10-30 m, slightly sandy 

mud with funiculines in 200-250 m, alcyonarians 

in mud in 90-150 m, and muddy sand in 470- 

500 m (all Maurin, 1968a); and on muddy shell 

in 50 m, mud in 40-60 m, muddy detritus in 200

NUMBER 306 

m, and mud or sandy mud in 90-100 m (all 

Maurin, 1968b). 

Off West Africa, ovigerous females have been 

collected in June and September (Crosnier, 1967; 

Pillsbury). 

DISTRIBUTION.—Mediterranean, eastern and 

western Atlantic, and off South Africa, in depths 

between 10-30 m and 679 m. In the western 

Atlantic it has been recorded from localities between 

Massachusetts and Brazil (Rathbun, 1937; 

Rice and Provenzano, 1970). In the eastern Atlantic 

it is known from scattered localities in the 

Mediterranean and in the Atlantic from Portugal 

and Spain southward to South Africa, including 

the Azores, the Cape Verde Islands, and Madeira 

(Barnard, 1950; Gordon, 1950; Monod, 1956; 

Zariquiey Alvarez, 1968). Since 1956 it has been 

recorded from the following: 

Spain: Off Guadiana, 470-500 m (Maurin, 1968a). 

Azores: Off Horta, Ilha do Faial, in deep water (Figueira, 

1960). 

Madeira: Funchal, 100-ca. 250 m; Ilhas Desertas (Tiirkay, 

1976b). 

Morocco: Bane de Spartel, 35°54'N, 06°14'W, 210 m 

(Peres, 1964). Between Cap Rhir and Cap Draa, 10-30 m 

(Maurin, 1968a). Off Cap Juby, 50 m (Maurin, 1968b). 

31°01'N, 10°16'W, 360-375 m (Tiirkay, 1976a). 

Spanish Sahara: Off Medano de Aaiun and Cabo Bojador, 

200-300 m; Morro Garnet, 300-350 m (Maurin, 1968a); 

between Cabo Corbeiro and Cabo Blanco, 200 m (Maurin, 

1968b). 

Mauritania: S Bane d'Arguin, 90-150 m (Maurin, 

1968a). Bane d'Arguin, 40-60 and 90-100 m (Maurin, 

1968b). 

Ivory Coast: 05°06'N, 03°49'W, 50 m (Crosnier, 1967). 

In 40-50 m (Le Loeuff and Intes, 1968). 

Principe: 01°43'10"N, O7°28'2O"E, 73 m (Forest and 

Guinot, 1966). 

Congo: Off Pointe-Noire, 05°00'S, 11°32'E, 98 m, and 

04°56'S, 11°31'E, 95-97 m (Crosnier, 1967). 

Angola: Baia dos Tigros, 215 m (Guinot and Ribeiro, 

1962). 

It has not been recorded previously from Liberia, 

Nigeria, or Annobon, although these localities 

are within the known range of the species. 

Genus Paromola Wood-Mason and Alcock, 

1891 

Paromola Wood-Mason and Alcock, 1891:267 [type-species: 

Dorippe cuvieri Risso, 1816, by monotypy; gender: feminine; 

name 1641 on Official List]. 

Paromola cuvieri (Risso, 1816) 

Paromola cuvieri.—Capart, 1951:25, fig. 4.—Monod, 1956:79, 

fig. 89.—Guinot and Ribeiro, 1962:23.—Rossignol, 1962: 

113.—Forest, 1963:628.—Peres, 1964:27, 29.—Figueira, 

1964:69, pis. 1,2.—Zariquiey Alvarez, 1968:301, fig. 106b 

[Spain; references].—Maurin, 1968a:29, 33, 50, 64, 106, 

114, 121 [p. 106, 114, 121 Mediterranean], fig. 22; 1968b: 

479, 480, 482, 484, 489, 491, 492, figs. 3, 6.—Christiansen, 

1969:24, fig. 8, map 2 [Scandinavia].—Crosnier, 1969: 

529.—Tiirkay, 1976a:25 [listed], 36; I976b:61 [listed], 

62.—Intes and Le Loeuff, 1976:103. 

Paromola.—Maurin, 1968a, fig. 23; 1968b, figs. 1, 9. 

Parhomola.—Maurin, 1968a, fig. 29 [erroneous spelling]. 

Paramola.—Maurin, 1968b, fig. 4 [erroneous spelling]. 

SYNONYM.—Maia dumerili Risso, 1816. 


Other Material: Morocco: Off Cap de Mazagan, 33°40'N, 

08°45'W, 570 m, Agassiz trawl, 28 Mar 1976, Onversaagd Sta 

159, 19 (L). 

DESCRIPTION.—Capart, 1951:27. 

Figures: Capart, 1951, fig. 4; Monod, 1956, fig. 

89; Christiansen, 1969, fig. 8. 

Male Pleopod: Sankarankutty, 1968, fig. la,b 

(North Sea). 

Color: "Couleur generate jaune orange, plus 

rouge a l'avant de la carapace et sur les epines 

anterieures; les pattes un peu plus foncees" (Capart, 

1951:27). 

BIOLOGY.—Paromola cuvieri is a deep water species, 

generally occurring on soft bottoms in depths 

between 150 and about 1000 m; there is one 

record from a depth of 952-1038 m (Tiirkay, 

1976a), and another from a depth of 10 m (Doflein, 

1904), but generally the species occurs beyond 

shelf depths. It has been recorded from mud 

or sandy mud. 


recorded in March and October (Capart, 1951; 

Monod, 1956). 

DISTRIBUTION.—Eastern Atlantic, from the Hebrides 

and southern Scandinavia southward to 

Angola, including the Azores, the Cape Verde 

Islands, and the Mediterranean, in depths between 

10 and more than 1000 m, usually deeper 

27

28 

than 150 m. Monod (1956) summarized earlier 

West African records and reported material from 

Senegal. Since 1956 the species has been recorded 

from: 

Azores: Ilha Terceira (Figueira, 1964). 

Madeira: No specific locality; fish market, Madeira (Tiirkay, 

1976b). 

Morocco: Foum Agouitir (as Puerto-Cansado), 350-450 

m (Maurin, 1968a). 35°19'N, O6°32'W to 35°28.8'N, 

06°39.2'W, 333-360 m, and 35°17.5'N, 06°10.3'W to 

35°13.9'N, 06°36.2'W, 295-340 m (PeVes, 1964). 33°37.5'N, 

09°02.2'W, 952-1038 m (Tiirkay, 1976a). Asilah (as Arzila) 

to Larache, 300-350 m (Maurin, 1968a). 

Spanish Sahara: Off Medano de Aaiiin and W of Cabo 

Bojador, 300-500 m; off Cabo Bojador and Morro Garnet, 

530-720 m (Maurin, 1968b). Off Villa Cisneros, 300-500 m 

(Maurin, 1968a,b). 

Mauritania: Off Nouakchott, 350-400 to 600 m (Maurin, 

1968a). Bane d'Arguin, 200-300 m, and Tamzak (as 

Tamxat), 350-600 m (Maurin, 1968b). 

Senegal: Fosse de Kayar, 300-350 to 600 m (Maurin, 

1968b). 

Ivory Coast: No specific locality (Intes and LeLoeuff, 

1976). 04°32'30"N, 06°31'W, 300-455 m, and 04°54'N, 

03°23'W, 380-400 m (Forest, 1963). 


Congo: W of Banga (as Banda), 250 m (Rossignol, 1962). 

Off Pointe-Noire, 04°54'S, 11°19'E, 300 m (Crosnier, 1969). 

Angola: Baia dos Tigres, 453-478 m (Guinot and Ribeiro, 

1962). 06°24'N, 11°34'E, 325 m (Crosnier, 1969). 

Family CYCLODORIPPIDAE Ortmann, 1892 

CYCLODORIPPIDAE Ortmann, 1892:552. 

TYMOLINAE Alcock, 1896:273, 274. 

REMARKS.—This family is not represented in 

the eastern Atlantic fauna. It was considered by 

Glaessner (1969:R492, as Tymolinae) to be a 

subfamily of the Dorippidae. 

Family CYMONOMIDAE Bouvier, 1898 

CYMONOMAE Bouvier, 1898:55, 59 [corrected to Cymonomidae 

by Glaessner, 1969:R627]. 

EASTERN ATLANTIC GENERA.—One, occurring 

to the north of the tropical region, Cymonomus A. 

Milne Edwards (1880:26). Type-species: Cymonomus 

quadratus A. Milne Edwards, 1880, by mono- 


typy; gender: masculine; name 1618 on Official 

List. 

EASTERN ATLANTIC SPECIES.—Two, both occurring 

north of the tropical region: 

Cymonomus granulatus (Thomson, 1873). Ireland 

southward to Sahara coast, Mediterranean; sublittoral, 

300-1350 m (Zariquiey Alvarez, 1968). 

Cymonomus normani Lankester, 1903. Between 

Iceland and the Faroes, 875-1269 m, and off 

Portugal, 1370-1430 m (see Turkay, 1976a:36, 

for references). 

Family DORIPPIDAE MacLeay, 1838 

DORIPPINA MacLeay, 1838:69 [corrected to Dorippidae by 

White, 1847a:53; name 355 on Official List, there attributed 

to de Haan, 1841, in error.] 

ETHUSINAE Guinot, 1977:1052. 

EASTERN ATLANTIC GENERA.—Four, Ethusa, 

Ethusina, Medorippe, new genus, and Phyllodorippe, 

new genus, each represented by tropical species. 

EASTERN ATLANTIC SPECIES.—Ten, six of which 

were recorded by Monod (1956), as follows: 

Name in Monod 

[Cymonomus granulatus 

(Dorippidae)] 

Ethusina abyssicola 

Ethusa mascarone 

Ethusa rosacea 

Ethusa rugulosa 

Dorippe lanata 

Dorippe armata 

Current Name 

[Cymonomus granulatus 

(Cymonomidae)] 

Ethusina alba 

Ethusa vossi, new species* 

Ethusa rosacea* 

Ethusa rugulosa* 

Medorippe lanata* 

Phyllodorippe armata* 

A seventh West African species, Ethusina beninia, 

new species, is represented in the Pillsbury collections. 

One species assigned to the Dorippidae by 

Monod (1956), Cymonomus granulatus (Thomson, 

1873), is included under the Cymonomidae 

(above), following Garth and Haig (1971:6.7). 

The extralimital species are as follows: 

Ethusa mascarone (Herbst, 1785). Mediterranean 

and adjacent Atlantic, including Canary Islands 

(Miers, 1881a; A. Milne Edwards and Bouvier, 

1900), Portugal (Nobre, 1936), and Bay of Biscay 

(Zariquiey Alvarez, 1968). For possible references

NUMBER 306 

to E. mascarone from the NW African coast, see 44 off Cap Rhir, Morocco, 29°52'N, 14°07'W of 

account of E. vossi, new species (p. 39). 

Paris (11°47'W of Greenwich), 2083 m, 25 June 

Ethusa microphthalma Smith, 1881. Azores, in 1883. 

1000 fm (1830 m) (Miers, 1886:329); western REMARKS.—Usually the genera Dorippe sensu 

Atlantic (Rathbun, 1937). 

lato, Ethusa, and Ethusina are placed in the nom- 

Ethusina talismani A. Milne Edwards and Bouinate subfamily of the Dorippidae. Recently 

vier, 1897. Off the Azores and Cap Rhir, Mo- Guinot (1977:1052) recognized a second subfamrocco, 

in 2075 to 2235 m (Monod, 1956). A male ily, Ethusinae (p. 38), for Ethusa and Ethusina, an 

syntype (USNM 22940) in the collections of the 

action with which we concur. 

Smithsonian Institution was taken at Talisman sta 

Key to Subfamilies and Genera of Dorippidae 

1. Afferent branchial orifices narrow and placed somewhat before bases of 

chelipeds, being separated from chelipeds by narrow process of carapace 

reaching down to sternum. Last two pereiopods distinctly subchelate, 

dactylus closing against tubercle on propodus (Dorippinae) 2 

Afferent branchial orifices wide and placed immediately against bases of 

chelipeds. Last two pereiopods not subchelate; dactylus short and not 

closing against tubercle on propodus (Ethusinae) 8 

2. Carapace with distinct epibranchial tooth 3 

Carapace without epibranchial tooth 5 

3. Male gonopod short, broad, and straight. Gastric region of carapace with 

V-shaped ridge. Dactylus of second and third pereiopod with upper 

margin naked or with few hairs in extreme basal part (less than VA of 

length) 4 

Male gonopod slender, S-shapedly curved. Gastric region without V-shaped 

ridge. Dactylus of second and third pereiopod with fringe of short hairs 

over more than basal third of dorsal margin [Carapace broader than 

long.] Phyllodorippe, new genus 

4. Meri of second and third pereiopods with dorsal row of spines or spinules; 

dactyli of these legs with few hairs and spinules in extreme basal part. 

Carapace broader than long with some low elevations. Abdomen of male 

and female without teeth Medorippe, new genus 

Meri of second and third pereiopods without dorsal spines or spinules; 

dactyli of these legs naked and smooth. Carapace longer than wide, with 

distinct tubercles. Abdomen of male and female with distinct teeth on 

third and fourth somites (also on second in male) Dorippe 

5. Male gonopod short, stubby, with one or more narrow appendages, one or 

more of which are T-, hammer-, or cross-shaped. Carapace distinctly 

wider than long. Front reaching about as far forward as outer orbital 

teeth 6 

Male gonopod very slender and curved, ending in 3 or 4 rounded or 

subacute lobes. Carapace longer than wide or almost as long as wide. 

Front reaching beyond outer orbital teeth 7 

29

30 


6. Male gonopod straight, with single twisted L- or T-shaped appendage at 

apex, and lobe at base Dorippoides 

Male gonopod with sharp angular bend in middle, with several appendages 

at top, without lobe at base Paradorippe 

7. Carapace distinctly longer than wide, flattened dorsally, without tubercles, 

but with grooves. Front reaching far beyond outer orbital teeth. Dactyli 

of second and third pereiopods with fringe of long hairs along both 

upper and lower margins Neodorippe 

Carapace slightly wider than long, convex, smooth or with tubercles. Front 

reaching only slightly beyond outer orbital teeth. Dactyli of second and 

third pereiopods with dorsal and ventral fringes of short hairs. Nobilum 

8. Eyes movable. Basal segment of antennules normal Ethusa 

Eyes immovable. Basal segment of antennules very large and swollen 

Ethusina 

Subfamily DORIPPINAE MacLeay, 1838 

REMARKS.—The two eastern Atlantic species 

belonging to this subfamily are present in the 

Pillsbury collections. Until recently both would 

have been placed in the genus Dorippe, but in 

1969, Serene and Romimohtarto in a paper dealing 

with Indo-West Pacific species, recognized 

instead of the old genus Dorippe three genera 

{Dorippe, Neodorippe, and Paradorippe) and two new 

subgenera, Dorippoides (in Dorippe) and Nobilum (in 

Neodorippe). 

We tried to accommodate the Atlantic species 

of Dorippe in the genera and subgenera recognized 

by Serene and Romimohtarto, but encountered 

several difficulties. Therefore a study first had to 

be made of the Indo-West Pacific species of Dorippinae. 

During this study we could only confirm 

the correctness of the taxonomic views of Serene 

and Romimohtarto, and even had to raise some 

of their subgenera to the rank of full genera. 

Furthermore, we found it necessary to establish 

two new genera, one each for the Atlantic species. 

It is our intention to discuss the Indo-West Pacific 

species of Dorippe, sensu lato, in a future paper, 

but our main conclusions can be found in the 

following list of the genera of Dorippinae that we 

at present recognize: 

Dorippe Weber (1795:93). Type-species: Cancer 

quadridens Fabricius, 1793, a subjective junior synonym 

of Cancer frascone Herbst, 1785, by subse- 

quent selection by Latreille, 1810:96, 422; gender: 

feminine. So far Dorippe contains only the typespecies. 

Dorippoides Serene and Romimohtarto (1969:3, 

4, 8). Type-species: Cancer facchino Herbst, 1785, 

by original designation and monotypy; gender: 

masculine. So far only containing the type-species. 

Medorippe, new genus. Type-species: Cancer lanatus 

Linnaeus, 1767, by present designation and 

monotypy; gender: feminine. 

Neodorippe Serene and Romimohtarto (1969:3, 

4, 11). Type-species: Dorippe astuta Fabricius, 

1798, a subjective junior synonym of Cancer facchino 

Herbst, 1785, by original designation; gender: 

feminine. An examination of three syntypes 

of Dorippe astuta Fabricius, 1798, in the collection 

of the Leiden Museum, and photographs of two 

of the four syntypes in the Zoological Museum, 

Copenhagen, showed that Dorippe astuta Fabricius, 

1798, is a subjective junior synonym of Cancer 

facchino Herbst, 1785, as defined by the lectotype 

selection for the latter species by Serene and 

Romimohtarto (1969:9). The specimens usually 

assigned by authors to Dorippe astuta and also 

described and figured under that name by Serene 

and Romimohtarto (1969:11, figs. 3, 7, 12, 17, pi. 

Id, pi. 4a,b), prove to be Dorippe callida Fabricius, 

1798. Photographs of the two type-specimens of 

Dorippe callida in the collection of the Copenhagen 

Museum, which could be examined by us, clearly

NUMBER 306 

showed the identity of the species. Thus, Neodorippe 

is a genus based on a misidentified typespecies, 

and in order to comply with Article 70(a) 

of the International Code of Zoological Nomenclature, 

we will submit this case to the International 

Commission on Zoological Nomenclature, requesting 

that under their plenary powers Dorippe 

callida Fabricius, 1798, be designated the typespecies 

of the genus Neodorippe. 

Although Serene and Romimohtarto assigned 

two species and a subspecies (the type-species, 

plus Dorippe japonica Von Siebold and D. japonica 

taiwanensis Serene and Romimohtarto) to the 

nominate subgenus of Neodorippe, we prefer to 

consider Neodorippe a genus with a single species, 

Neodorippe callida (Fabricius, 1798). 

Nobilum Serene and Romimohtarto (1969:3, 5, 

14). Type-species: Dorippe histrio Nobili, 1903, by 

original designation and monotypy; gender: neuter. 

We prefer to place Dorippe japonica in this 

genus rather than in Neodorippe. The gonopods of 

the two species, Dorippe japonica and D. histrio, in 

our opinion do not differ in essential details, while 

the general morphology of the body shows the 

species to be much closer to one another than to 

Neodorippe callida. Therefore we consider the genus 

Nobilum at present to consist of Nobilum histrio 

(Nobili, 1903), N. japonicum japonicum (Von Siebold, 

1824) and N. japonicum taiwanense (Serene 

and Romimohtarto, 1969). 

Paradorippe Serene and Romimohtarto (1969:3, 

5, 15). Type-species: Dorippe granulata de Haan, 

1841, by original designation; gender: feminine. 

This genus contains three species: Paradorippe granulata 

(de Haan, 1841), P. australiensis (Miers, 

1884), and P. polita (Alcock and Anderson, 1894). 

Phyllodorippe, new genus. Type-species: Dorippe 

armata Miers, 1881, by present designation and 

monotypy; gender: feminine. 

The new genera, Medorippe and Phyllodorippe, 

are restricted to the eastern Atlantic. The other 

five genera inhabit the Indo-West Pacific region. 

The subfamily, so far, has not been found in 

American waters (neither in the western Atlantic 

nor in the eastern Pacific). 

Genus Medorippe, new genus 

TYPE-SPECIES.—Cancer lanatus Linnaeus, 1767. 

ETYMOLOGY.—The name is formed from the 

Greek prefix me- (not) and the feminine generic 

name Dorippe. 

DIAGNOSIS.—Carapace broader than long, pubescent, 

with distinct epibranchial spine on either 

side. Extra-orbital teeth slightly surpassing front. 

Tubercles on dorsal surface of carapace few and 

low, grooves conspicuous; V-shaped ridge present 

on cardiac region. Eyes not reaching beyond 

extra-orbital teeth. Lower margin of orbit, between 

extra- and infra-orbital teeth, smooth. Second 

and third pereiopods with dorsal margin of 

merus carrying row of spines. Dactylus of these 

pereiopods narrow and somewhat twisted, not 

fringed with hairs (at most with very short row of 

hairs in extreme basal part of dorsal margin). 

Abdomen of male with blunt and low elevations, 

but without teeth or spines. First pleopod of male 

short, stubby and straight, with lobe at outer 

margin of base; apex acute, turned inward 

abruptly (almost at right angle), without distal 

appendages. 

REMARKS.—In the genus Medorippe, the shape 

of the male gonopod is closest to those of Dorippe 

and Dorippoides, but differs from that in both 

genera in the sharp straight apex which is turned 

inward. From the other genera, Medorippe can be 

distinguished by the short, straight gonopod without 

distal appendages. Like Dorippe and Phyllodorippe, 

Medorippe has a distinct epibranchial spine, 

which in Dorippe, however, is preceded by a row 

of granules. The presence of spines on the dorsal 

margin of the meri of pereiopods 2 and 3 distinguishes 

Medorippe from the others. 

* Medorippe lanata (Linnaeus, 1767), 

new combination 

FIGURE Aa-h 

Dorippe lanata.—Capart, 1951:30, fig. 6.—Monod, 1956:90, 

fig. 103 [as Dorippe armata; not fig. 102 = Phyllodorippe 

armata].—Rossignol, 1957:75, pi. 3: fig. 3.—Forest and 

Gantes, 1960: 350—Gauld, 1960:68.—Guinot and Ri- 

31

32 

beiro, 1962:25.—Rossignol, 1962:114.—Crosnier, 1964: 

34.—Forest and Guinot, 1966:50.—Guinot, 1967a:244 

[listed; Indian Ocean].—Zariquiey Alvarez, 1968:312, 

figs. 2f, 14b, 105a,b, 106b [Spain; references].—Maurin, 

1968a:30,41; 1968b:480, 486.—Le Loeuffand Intes, 1968: 

38, table 1, figs. 47, 62; 1969:63, 65.—Serene and Romimohtarto, 

1969:6 [discussion].—Bas, Arias, and Guerra, 

1976, table 3.—Tiirkay, 1976a:25 [listed], 37. 

Dorippe armata.—Monod, 1956, fig. 103.—Crosnier, 1964, fig. 

on pi. A.—Maurin, 1968b, figs. 5, 7. [Not Phyllodorippe 

armata (Miers, 1881).] 

Dorripe lanata.—Rossignol, 1957:126 [key].—Crosnier, 1970: 

1215. [Erroneous spelling.] 

SYNONYM.—Dorippe qffinis Desmarest, 1823. 


Sta 68, 70 m, broken shell, 16\ 1$ ov, 6 juv (L). 

Ivory Coast: Sta 42, 62-75 m, mud with brown, branched 

Foraminifera, 2 juv (W). Sta 46, 38-42 m, mud with dense 

Jullienella, lie?, 11$, 27 juv (L). Sta 47, 37 m, bottom with 

Jullienella, 46*, 19 (W). Sta 60, 79-82 m, coral or rock, 1(5, 1 

juv (L). Sta 62, 46 m, brown, branching and foliate Foraminifera, 

19(5, 89 (W). Sta 64, 68 m, 19 (L). Sta 65, 46-49 m, 

26, 49 (W). 

Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 1(5 

(W). Sta 23, 42 m, foliate brown to orange bryozoans, 36, 29 

(L). Sta 24, 35-37 m, dark red bryozoans, 1(5 (L). Sta 28, 49- 

53 m, 26, 19 (W). Sta 30, 61-64 m, coral, 1(5, 19 (L). 

Nigeria: Sta 241, 59-63 m, mud and shell, 4c?, 19 (W). Sta 

248, 33 m, 26 (L). 

Cameroon: Sta 260, 46 m, 1(5, 19 (W). 

Undaunted Material: Angola: Sta 102, 54 m, 19 (L). Sta 

103, 90 m, 16 (L). 

Other Material: Morocco: Off Cap Hadid, 31°55'N, 09°- 

52'W, 78 m, muddy sand, 5m beam trawl, 25 Mar 1976, 

Onversaagd Sta 127, 1(5 (L). 

DESCRIPTION.—Monod, 1933b:490-494; Capart, 

1951:30. 


103. 

Male Pleopod: Monod, 1933b, fig. 3f,g (Morocco, 

Syria) (Figure 4g,h). 

Color: Capart (1951:31) noted that his material 

was "uniforme ocre rose, la carapace souvent 

encrassee de vase; les doigts de la pince roses." 

The Pillsbury specimens also were uniform pinkish 

in color, without obvious bands on the pereiopods. 

MEASUREMENTS.—Carapace lengths of our 

specimens range from 4 to 28 mm; the single 

ovigerous female has a carapace length of 28 mm. 


REMARKS.—The West African specimens of 

Medorippe lanata closely resemble those from localities 

in the Mediterranean as well as those from 

the Atlantic coasts of Spain and Portugal with 

which they have been compared. The male pleopods 

of our specimens apparently are identical 

with those of Mediterranean specimens, resembling 

those illustrated by Monod (1933b, fig. 3f, 

6* from Morocco) (Figure 4g); the pleopods of 

immature males are like that figured by Monod 

(1933b: fig. 3g) (Figure Ah) from a specimen from 

Syria; Monod questioned whether his specimens 

from the eastern Mediterranean actually were 

mature. 

In spite of their close resemblance, some differences 

can be observed between our material from 

the Gulf of Guinea and specimens from more 

northern localities, but these observed differences 

overlap so broadly that we believe that the Gulf 

of Guinea and the Mediterranean populations of 

this species cannot be separated nomenclaturally 

at this time. Perhaps with additional material, 

distinct northern and southern subspecies can be 

recognized. In general, the Mediterranean specimens 

appear to be more pubescent, the spines on 

the merus of the walking legs are larger and give 

the impression of being less numerous, the teeth 

on the cutting edges of the chelae are sharper, 

and the walking legs appear to be longer and 

slenderer. The second and third pereiopods of a 

specimen from Tunisia (Figure Ac,d) and one 

from the Ivory Coast (Figure AeJ) are shown 

here. Of these apparent differences, the first three 

vary widely and cannot be used to distinguish 

representatives of the two populations. Mediterranean 

specimens have 7-15 (usually 10-11) 

spines on the dorsal edge of the merus, whereas 

the Gulf of Guinea specimens have 7 to approximately 

20 (usually 12) spines. The most striking 

difference is in the proportion of length to width 

of the merus of the walking legs (Table 1). The 

Mediterranean specimens appear to have much 

slenderer and longer legs, and this is in general 

supported by measurements, although for each 

size range the ratios overlap to some extent. 

The Gulf of Guinea population may mature at

NUMBER 306 

FIGURE 4.—Medorippe lanata (Linnaeus): a, dorsal view (from Monod, 1956, fig. 103); b, dorsal 

view of carapace (from Monod, 1933b, fig. 4c); c, second pereiopod, male, cl 23 mm, Tunis; d, 

third pereiopod, same specimen; e, second pereiopod, male, cl 19 mm, Pillsbury Sta 47;/, third 

pereiopod, same specimen; g, h, male, first pleopods (from Monod, 1933b, fig. 3f,g). Phyllodorippt 

armata (Miers): i, dorsal view (from Monod, 1956, fig. 102);^, dorsal view of carapace (from 

Monod, 1933b, fig. 4a); k, I, male first pleopods and apexes (from Monod, 1933b, fig. 3h). 

b 

33

34 

TABLE 1.—Length-width ratios of the propodus of the third 

pereiopods for M. lanala from the Gulf of Guinea and the 

Mediterranean 

Carapace 

length 

(mm) 

8-10 

11-13 

14-16 

17-19 

20-22 

23-25 

26-28 

Number 

of 

specimens 

3 

11 

12 

1 

3 

- 

- 

Gulf of Guinea 

Range 

(mm) 

4.2-4.9 

3.8-4.9 

3.7-4.7 

- 

3.3-3.8 

- 

- 

Mean 

(mm) 

4.6 

4.4 

4.1 

3.9 

3.5 

- 

- 

Number 

of 

specimens 

1 

- 

4 

2 

13 

11 

8 

Mediterranean 

Range 

(mm) 

_ 

- 

4.5-5.5 

4.3-4.6 

3.8-5.1 

3.8-4.9 

4.1-4.8 

Mean 

(mm) 

4.6 

- 

5.1 

4.5 

4.5 

4.4 

4.3 

a smaller size. A 22 mm long male from Angola 

has the right cheliped greatly inflated, whereas in 

specimens from Spain males with carapace 

lengths of 23-26 mm show only slight inflation of 

the chela and smaller ones, 18-22 mm long, have 

chelae similar to those of females. 

Monod (1933b:492) suggested that the eastern 

Mediterranean population of this species might 

be subspecifically distinct from those in the western 

Mediterranean and the Atlantic. One of the 

differences he noted was in the shape of the male 

pleopod, but, as already pointed out, his Syrian 

specimens were juveniles and the pleopods were 

not fully developed. He also noted that the Syrian 

specimens differed from others he examined in 

having a more spinulose sternal border; this feature 

varies widely in our material, too. 

As indicated by Forest and Guinot (1966:50), 

figures 102 and 103 of Monod's (1956) monograph 

are interchanged, figure 102 showing Phyllodorippe 

armata, figure 103 Medorippe lanata. Crosnier's 

(1964) figure of "Dorippe armata" on his 

plate A, as well as Maurin's (1968b) figures 5D.a. 

and 7D.a., all are copies of Monod's (1956) figure 

103, and thus do not represent Phyllodorippe armata, 

but Medorippe lanata. 

BIOLOGY.—The depth at which this species 

occurs is usually indicated as sublittoral to about 

100 m (Bouvier, 1940:200; Monod, 1956:92). The 

depth of 952-1038 m off Morocco given by Tiirkay 

(1976a) is far in excess of the depths at which 


the species normally is found and is probably 

erroneous. Zariquiey Alvarez (1968:313) indicated 

that this species is usually taken in Spanish 

waters in depths of 40 or 50 to 100 m; and Pesta 

(1918:287) gave these depths for the Adriatic 

population, with the reservation that the species 

may occur in depths of 10 m. So far as West 

Africa is concerned, evidently the same situation 

is found: Crosnier (1964) listed the species as 

occurring in cold water in depths greater than 50 

m off Cameroon, whereas Le Loeuff and Intes 

(1968) stated that although Medorippe lanata is 

sometimes taken in Ivory Coast waters in depths 

between 25 and 30 m (in their table they give 15 

m as least depth; this may be an error for 25), the 

majority of the specimens there were found in 

depths between 35 and 50 m and as deep as 100 

m. Of the specimens examined by us only one 

was found at a depth less than 35 m (viz. 33 m), 

10 specimens (55%) were found between 35 and 

55 m, and the remainder were from deeper water, 

55-90 m. Also, more than 80% of the material 

listed by Monod (1956) for which depth is indicated 

was from depths of 35 to 55 m, whereas 

only two specimens, the remainder, were taken at 

96 and 100 m. Capart's West African material for 

the larger part (14 of the 17 lots for which depth 

was recorded) came from between 30 and 92 m, 

and only two lots were from less than 30 m (12- 

15 m and 20-24 m). Most other West African 

records also conform to this pattern: the far 

greater part is from deeper than 35 m. Only 

Maurin (1968a) reported the species from a depth 

of 10-30 m, and only Tiirkay (1976a) has reported 

it from much deeper water, as noted 

above. 

Capart (1951) reported the occurrence of this 

species on coastal bottoms of mud or muddy sand 

generally in depths between 25 and 100 m, with 

one capture at 15 (12-15) m. It was taken only 

on one occasion with the shallower Phyllodorippe 

armata. 

The bottom types on which this species was 

collected by the Pillsbury include: mud with Foraminifera, 

46 m; foliate brown to orange bryozoans 

in 42 m and dark red bryozoans in 35-37

NUMBER 306 

m; coral in 61-64 m and coral and rock in 79-82 

m; broken shell in 70 m; and mud and shell in 

59-63 m. Most of the Pillsbury specimens were 

taken on mud bottom with foliate Foraminifera, 

probably Jullienella, in depths between 37 and 

75 m. 

Other bottom types on which the species has 

been collected include: mud and sand in 65- 

75 m, mud in 50-64 m, mud, stones, calcareous 

algae, sand, and Foraminifera in 51 to 55 m and 

gravel, shell, and Foraminifera in 50 m (all Forest 

and Guinot, 1966); sand and muddy sand in 10- 

30 m and muddy sand in 40-50 m (Maurin, 

1968a); shelly mud, in 50 m (Maurin, 1968b); 

and mud and sandy mud in 15[25?]-100 m (Le 

Loeuff and Intes, 1968). Le Loeuff and Intes 

(1969:63) indicated that this species was a mud 

dweller off the Ivory Coast. It seems that this 

species, like E. vossi, prefers a soft bottom with 

hard particles in it. 


collected in March, April, May, June, August, 

and October (Capart, 1951; Monod, 1956; 

Guinot and Ribeiro, 1962; Pillsbury). 

DISTRIBUTION.—Eastern Atlantic, from the 

Mediterranean Sea and Portugal to South Africa 

and Mozambique (Barnard, 1955). Monod 

(1956) enumerated earlier records from West Africa; 

since 1956 M. lanata has been recorded from 

the following localities: 

Morocco: 33°37.5'N, 09°02.2'W, 952-1038 m (?) (Tiirkay, 

1976a). Agadir, 60-130 m (Forest and Games, 1960); 

Agadir (Maurin, 1968a). Between Cap Rhir (as Cap Ghir) 

and Cap Draa (as Cap Noun), 10-30 m (Maurin, 1968a). 

Off Essaouira, 32°08'N, 09°02'W, 33 m, and 31°37'N, 09°- 

54'W, 70 m (Forest and Games, 1960). SW of Cap Juby, 

40-50 m (Maurin, 1968a). Cap Juby, 50 m (Maurin, 1968b). 

Spanish Sahara: 21°44'N, 17°03'W to 21°46.5'N, 17°- 

04'W, 40 m, and 21°3O'N, 17°24'W to 21°27'N, 17°24'W, 

95-137 m (Bas, Arias, and Guerra, 1976). 

Mauritania: Bane d'Arguin, 40-60 m and 90-100 m 


Senegal: 13°01'N, 17°24'W, 51-55 m, and 12°55.5'N, 

17°33'W, 65-75 m (Forest and Guinot, 1966). 

Sierra Leone: 07°15.5'N, 12°51'W, 64 m (Forest and 



1969). Off Sassandra, off Fresco, off Grand-Lahou, off Jac- 

queville, and off Grand-Bassam, 15[25?]-100 m (Le Loeuff 


Ghana: Off Accra, 37-51 m (Gauld, 1960). 04°40'N, 

02°08'W to 04°39'N, 02°05'W, 50 m, and 04°36.5'N, 01°- 

31'W, 50 m (Forest and Guinot, 1966). 

Cameroon: No specific locality, more than 50 m (Crosnier, 

1964). 

Congo: Near Pointe-Noire (Rossignol, 1957, 1962). 

Angola: 17°O2'S, ll°40'E, 54 m, and 17°06'S, 11°35'E, 

90 m (Crosnier, 1970). 

Genus Phyllodorippe, new genus 

TYPE-SPECIES.—Dorippe armata Miers, 1881. 

ETYMOLOGY.—The name is formed by a combination 

of the Greek word phyllon (leaf) and the 

feminine name Dorippe. 

DIAGNOSIS.—Carapace distinctly broader than 

long, with sparse pubescence dorsally and distinct 

epibranchial spine on either side. Extra-orbital 

teeth reaching slightly farther forwards than 

front. Tubercles on dorsal surface low, grooves 

distinct. Cardiac region without V-shaped ridge. 

Eyes not reaching beyond extra-orbital teeth. 

Lower margin of orbit between extra- and infraorbital 

teeth smooth. Dorsal margin of pereiopods 

2 and 3 without spines on dorsal margin of merus. 

Dactyli of these pereiopods narrow and slightly 

twisted; with fringe of short hairs present in basal 

half of upper margin. Abdomen of male with 

blunt low elevations, but without teeth or spines. 

First pleopod of male long, slender, S-shapedly 

curved, with 2 short appendages in distal part, 

lacking lobe in proximal part. 

REMARKS.—In the shape of the male gonopod, 

Phyllodorippe is closest to the genera Neodorippe and 

Nobilum, in both of which the male gonopod is 

also slender and curved; the gonopod of Phyllodorippe 

differs from those of the other genera 

because it ends in a narrow acute point and 

carries two lobiform appendages. The presence of 

epibranchial spines distinguishes the present genus 

from all other dorippine genera except Dorippe 

and Medorippe. Furthermore, in Neodorippe and 

Nobilum the front reaches farther forward than the 

anterolateral teeth. So far the genus is represented 

by a single species. 

35

36 

* Phyllodorippe armata (Miers, 1881), 


FIGURE 4/-/ 

?Dromia Pechiiel-Loesche, 1882:288. 

Dorippe Senegalensis Monod, 1933b:548 [footnote; nomen nudum, 

published in synonymy]. 

Dorippe armato.—Monod, 1933b:548 [footnote; erroneous 

spelling]. 

Dorippe armata.—Capart, 1951:33, fig. 7.—Monod, 1956:92, 

figs. 102 [as Dorippe lanata], 104-107 [not fig. 103 = 

Medorippe lanata].—Rossignol, 1957:75.—Longhurst, 1958: 

87— Lebour, 1959:131, 132, 134-136, fig. 19 [larvae].— 

Gauld, 1960:68—Rossignol, 1962:114.—Guinot and Ribeiro, 

1962:24.—Crosnier, 1964:32, 38 [not fig. on pi. A 

= Medorippe lanata].—Forest and Guinot, 1966:50.—Le 

Loeuff and Intes, 1968:38, figs. 47, 61, tables 1, 3-6, 8; 

1969:63-65.—Maurin, 1968a:48, 59, 62; 1968b:484, 486, 

489, 491 [not figs. 5D.a. and 7D.a. = Medorippe lanata].— 

Serene and Romimohtarto, 1966:6—Monod, 1970:66. 

Dorrippe armata.—Rossignol, 1957:126 [key; erroneous spelling] 

Dorippe armate.—Lebour, 1959:136 [larvae; erroneous spelling] 

Dorippe.—Voss, 1966:19.—Maurin, 1968b, figs. 4, 9. 

MATERIAL EXAMINED.—Pillsbury Material: Ivory 

Coast: Sta 46, 38-42 m, mud with dense Jullienella, 3$ (L). 

Sta 47, 37 m, bottom with Jullienella, 96, 4$ ov (L). Sta 48, 

22 m, 126\ 16$ (13 ov) (W). Sta. 49, 73-77 m, 1

NUMBER 306 

28 specimens of the species were obtained) and 

accidentally remained in the net. 

Of the post-1956 records listed under "Distribution," 

those by Maurin (1968a, 1968b) are the 

deepest (1 record of 35-40 m, 6 records between 

50 and 100 m, and 1 of about 200 m). Forest and 

Guinot (1966) reported the species from depths 

between 4 and 50 m (all but one of these 9 records 

from 34 m or less). Le Loeuff and Intes (1968:38) 

remarked that "Z). armata est la plus cotiere [of 

the two Dorippe species] et ne depasse pas les 35 

m," although in their table 1 they give the depth 

of the species as 15-50 m. Guinot and Ribeiro 

(1962) reported the species from very shallow 

localities (5 and 8 m), but also from 42 m. Other 

authors indicated depth ranges up to 43 m 

(Gauld, 1960) or even 60 m (Longhurst, 1958). It 

seems likely that the species has its greatest concentration 

below 35 m, but also occurs occasionally 

as deep as 60 m. The very deep records of 

the species need verification. 

Inasmuch as Maurin (1968b) followed Monod 

(1956) and reversed the figures of M. lanata and 

P. armata and Maurin's records for the shallow 

water species, P. armata, are deeper than usual 

and his records for the deeper living species, M. 

lanata, are shallower than usual, it seems likely 

that the two species were reversed throughout his 

accounts (1968a,b). 

Phyllodorippe armata is usually found on a sand 

or mud bottom, with shell, calcareous algae, or 

Foraminifera. It seems to be able to tolerate 

low salinities. Balss (1921:48) reported it from 

"Flusswasser" (in Nigeria), from the "Nyanga- 

Fluss, Salzwasser" (Gabon), and "Siisswasser des 

Kongoflusses" (near Boma [05°51'S, 13°03'E], 

Zaire). Voss (1966:37) remarked that Pillsbury Sta 

250, at which P. armata was taken, had "a typically 

brackish water or estuarine fauna due to our 

proximity to the mouths of the Niger, or Kwara, 

which strongly influence this section of the coast." 

Sourie (1954b) found it on coarse shelly sand 

bottom with Area and Pyura, in 10-12 m in the 

Baie de Dakar. 

Crosnier (1964) found this species on mud or 

sandy mud off Cameroon and characterized it as 

a warm water species, living in 0-30 m. Longhurst 

(1958) reported if from muddy sand, shelly sand, 

and shelly mud in 10-60 m off Sierra Leone. Le 

Loeuff and Intes (1969:63) characterized this species 

as a coastal species indifferent to the nature 

of the substrate. 

Ovigerous females have been collected in January, 

February, March, April, May, June, August, 

September, November, and December (Capart, 

1951; Monod, 1956; Forest and Guinot, 

1966; and p. 36). 

Pechiiel-Loesche (1882:288) described the behavior 

of a crab that he observed at the Crique 

Banjia, Gabon, as follows: "Urn vieles komischer 

nimmt sich aber eine andere Krabbenart 

(Dromia?) aus, die ich am Banya beobachtete; 

beim Spazierengehen pflegt sie mittelst der hinteren, 

am Riicken entspringenden Beinpaare einen 

Sonnenschirm iiber sich zu halten, welcher 

gewohnlich aus einem halben Mangrovenblatte 

besteht. Fliichtet sie eilig, oder geht sie ins Wasser, 

so lasst sie das wunderliche Schutzdach fallen." 

As no morphological details of this crab are provided 

by Pechiiel-Loesche, it is impossible to state 

with certainty which species was observed by 

him. We know of no Dromia carrying leaves, but 

Serene and Romimohtarto (1969:12), when dealing 

with Neodorippe callida, remarked that "in 

Singapore the species is very common on the 

mangrove swamps and generally all specimens 

hook a leaf of a mangrove tree on their backs." If 

Pechiiel-Loesche's specimen actually is a dorippid, 

it is more likely to be Phyllodorippe armata than 

Medorippe lanata, as the former is known to live in 

shallower depths than the latter. New field observations 

may solve the problem and could add 

much of interest to the knowledge of the biology 

of these little known animals. 

DISTRIBUTION.—Off West Africa, where it has 

been recorded from many localities between Cabo 

Corbeiro, Spanish Sahara and Angola. Monod 

(1956) summarized earlier records and added 

many new ones. Since 1956, P. armata has been 

recorded from the following localities: 

Spanish Sahara: Between Cabo Corbeiro and Cabo 

Blanco, 60-80 m (Maurin, 1968a), in about 200 m (Maurin, 

37

38 

1968b). Between Cabo Barbas and Cabo Blanco, 50-90 m 


Mauritania: N Bane d'Arguin, 20°20'N-20°40'N, 90-100 

m (Maurin, 1968a); same, less than 45 m, 40-60 m, 60-70 

m, and 90-100 m (Maurin, 1968b). Tamzak (as Tamxat), 

70-75 m (Maurin, 1968a), in 70 m (Maurin, 1968b). 

Senegal: Mboro, 35-40 m (Maurin, 1968b). 


1958). O7°2O'N, 12°39'W, 30-34 m (Forest and Guinot, 

1966). 

Ivory Coast: No specific locality (Le LoeufT and Intes, 

1969). 05°00'N, 05°28.5'W, 27 m, and 05°03'N, 05°25'W, 

20-25 m (Forest and Guinot, 1966). Off Sassandra, off 

Fresco, off Grand-Lahou, and off Grand-Bassam, 15-50 m 

(Le Loeuff and Intes, 1968). 

Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m 

(Forest and Guinot, 1966). Off Accra and Takoradi, 10-43 

m (Gauld, 1960). 

Cameroon: No specific locality, 0-30 m (Crosnier, 1964). 

Principe: Between Ponta da Mina and Ilheu Santana, 

10-12 m; off (Cais de) Santana, 11 m; between Ponta da 

Mina and Ponta Novo Destino, 6 m (Forest and Guinot, 

1966). 

Sao Tome: Off Ponta Diogo Nunes, 4 m (Forest and 


Gabon: ?Crique Banjia (as Banya) (Pechiiel-Loesche, 

1882). 

Congo: Off Pointe-Noire (Rossignol, 1957, 1962). 

Angola: Porto Amboim, 42 m; Baia de Benguela, 5 m; 

Baia da Caota, Benguela, 8 m (Guinot and Ribeiro, 1962). 


* Ethusa rosacea A. Milne Edwards and 

Bouvier, 1897 

Ethusa rosacea.—Capart, 1951:28, fig. 5.—Monod, 1956:88.— 

Rossignol, 1957:126 [key].—Crosnier, 1967:323; 1969:530, 

figs. 20-22. 


Sta 74, 641-733 m, l?ov(L). 

Ivory Coast: Sta 41, 641-842 m, 26* (W). Sta 44, 403-586 

m, hard, dark gray mud, lc? (L). 


Figure: Capart, 1951, fig. 5. 

Male Pleopod: Crosnier, 1969, figs. 20-22 (Angola). 

Color: Capart (1951:28) gave the following account 

of color pattern: "Couleur brun clair, sauf 

la face ventrale et les pinces, qui sont blanc-rose." 

MEASUREMENTS.—Our male specimens have 

carapace lengths of 8 to 11 mm; the ovigerous 

female has a carapace length of 12 mm. 

REMARKS.—Crosnier (1969) figured the first 

male pleopod of this species and commented on 

its distinctness from that of E. rugulosa, figured in 

Monod (1956:140). Some of the apparent differences 

between the first pleopods of these two 

species may be the result of their having been 

Lebour (1959:136) recorded larvae of this spe- illustrated from slightly different angles. 

cies from several localities in the Gulf of Guinea: BIOLOGY.—Ethusa rosacea is a deep-water spe- 

Ivory Coast, 04°29'N, 06°41'W; Ghana, 05°29'N, cies, occurring generally in depths below 100 m. 

00°20'E and 05°37'N, 00°38'E; and Nigeria, off One of the Pillsbury samples was taken on hard, 

Lagos, off Bonny River, and 05°58'N, 04°38'E. dark gray mud in 403-586 m. Material of this 

species reported by Crosnier (1969) was taken on 

mud in depths between 545 and 602 m. Capart 

Subfamily ETHUSINAE Guinot, 1977 (1951) reported the species from brown muddy 

sand in 125 m; brown, black mud in 100 m; 

Genus Ethusa Roux, 1830 

sandy mud in 290-390 and 500 m; and green 

sandy mud in 280-420 m. 

Ethusa P. Roux, 1830, pi. 18 [type-species: Cancer mascarone Ovigerous females have been recorded in 

Herbst, 1785, by subsequent designation by Fowler, 1912: March, April, and June (Capart, 1951; Crosnier, 

590; gender: feminine; name 1622 on Official List]. 1969; Pillsbury). 

?Pndope Nardo, 1847a:2 [nomen nudum; without included 

DISTRIBUTION.—Eastern Atlantic, where it has 

species). 

Pridope Nardo, 1869:307 [type-species: Pridope typica Nardo, 

been recorded from localities between the Canary 

1869, a subjective junior synonym of Cancer mascarone Islands and Luanda, Angola, in depths between 

Herbst, 1785, by original designation by use of typica and 100 and 1113 m (Monod, 1956). Since 1956 it has 

monotypy; gender: feminine). 

been recorded from the following localities.

NUMBER 306 39 

Congo: Off Pointe-Noire, 05°04'S, 1 l°20'E, 500 m (Crosnier, 

1967). 

Angola: 08°35'S, 12°51'E, 545-555 m; 09°27'S, 12°38'E, 

545-555 m; 11°55'S, 13°20'E, 545-552 m; and 11°57'S, 

13°29'E, 595-602 m (all Crosnier, 1969). 

It has not been recorded previously from the 

Ivory Coast or Liberia. 

* Ethusa rugulosa A. Milne Edwards and 

Bouvier, 1897 

Ethusa rugulosa.—Monod, 1956:89, figs. 99-101.—Longhurst, 

1958:87.—Rossignol, 1962:114.—Forest and Guinot, 

1966:49.—Crosnier, 1970:1215. 


Coast: Sta 49, 73-77 m, 1$ (L). Sta 50, 128-192 m, 29 (W). 

Sta 60, 79-82 m, coral or rock, 1 juv (L). 

Other Material: Ivory Coast: Off Grand-Lahou, 20 m, 

dredge, 31 Mar 1964, Guinean Trawling Survey, Tr 24, Sta 

1, 1

40 


FIGURE 5.—Ethusa vossi, new species, dorsal view (from Monod, 1956, fig. 90). 

56, 17$ (L). Sta 47, 37 m, bottom with Jullienella, 1(5, 1$ ov 

(W). 

Ghana: Sta 22, 51 m, rough bottom, 1$ ov (W). Sta 23, 

42 m, foliate brown to orange bryozoans, 1

NUMBER 306 41 

FIGURE 6.—Ethxtsa vossi, new species, male paratype, cl 9 mm, Pillsbury Sta 248: a, dorsal view; 

b, terminal two somites of abdomen; c, palm of chela. Ethusa mascarone (Herbst), male, cl 10 mm, 

Naples: d, dorsal view; e, terminal two somites of abdomen; /, palm of chela. 

and merus of third and fourth legs. Terminal 

somite of male abdomen shorter than penultimate 

(Figure 6b). 

Male Pleopod: Illustrated in Figure la,b. 

Color: Our specimens had the legs banded with 

red or orange and white. In males the enlarged 

right cheliped is white, with the tips of the fingers 

brown. Monod (1956) recorded one specimen 

with irregular white spots on the branchial regions. 


lengths of 4 to 11 mm. The ovigerous females 

have carapace lengths of 10 mm. 

REMARKS.—Ethusa vossi closely resembles E. 

mascarone (Herbst) from the Mediterranean and 

adjacent Atlantic; this new species had been identified 

with E. mascarone in the past, but it differs 

in several important respects. In E. vossi the terminal 

somite of the male abdomen is shorter than 

the penultimate somite (Figure 6b), whereas in E. 

mascarone the terminal somite is the longer (Figure 

6

42 

the orbit is smoother in E. mascarone (Figure 6d) 

than in E. vossi (Figure 6a), and females of the 

two species can be distinguished by this feature. 

Miers (1881a:270), in identifying material of 

this new species from Goree, Senegal, with E. 

mascarone, noted that "the larger chelipede in the 

male has the palm deeper and externally somewhat 

more convex than the male from the Mediterranean 

in the [British] Museum collection." 

In addition to the tropical West African records 

cited below (and assigned to E. vossi), E. mascarone 

has been recorded from the following localities 

outside of the Mediterranean: Canary Islands (A. 

Milne Edwards and Bouvier, 1900; Miers, 

1881a); Cape Verde Islands (A. Milne Edwards 

and Bouvier, 1900); and Cabo Blanco, Spanish 

Sahara (Cap Blanc, Mauritania) (Monod, 1933b; 

Tiirkay, 1975a). These materials will have to be 

reexamined to determine whether E. mascarone or 

E. vossi is involved. 

TYPE-LOCALITY.—Gulf of Guinea, off Nigeria, 

04°03'N, 05°41'E to 04°07'N, 05°40'E, Pillsbury 

Sta 248. 

DISPOSITION OF TYPES.—The holotype (Crust. 

D. 31538), a male, cl 8 mm, is in the Rijksmuseum 

van Natuurlijke Historie, Leiden. The other specimens 

are paratypes; they have been divided 

between the Rijksmuseum van Natuurlijke Historie 

in Leiden and the National Museum of 

Natural History, Smithsonian Institution, Washington. 

ETYMOLOGY.—It is a pleasure to name this 

species for Gilbert L. Voss of the Rosenstiel 

School of Marine and Atmospheric Science, University 

of Miami, Florida, who organized and led 

the Pillsbury expeditions to West Africa in 1964 

and 1965. 

BIOLOGY.—This species has been recorded from 

depths between 6 and 96 m, but 80% of the 

records are from depths of less than 45 m, and 

somewhat more than half from between 30 and 

45 m. The bottoms on which the species has been 

found have been described as mud with shells 

(Miers, 1881a); coarse shelly sand, bottom with 

Area and Pyura (Sourie, 1954b); partly shelly and 

partly muddy and shelly sand (Longhurst, 1958); 

mud and shells, mud, calcareous algae and shells, 


algae and calcareous algae, gravel, shells and 

Foraminifera, and mud, calcareous algae and 

shells (all Forest and Guinot, 1966). The Pillsbury 

specimens were taken on bottoms with JullieneIla 

(Sta 46, 47) (Voss, 1966), broken shell (Sta 68), 

and branched Foraminifera (Sta 70). It seems, 

therefore, that the species prefers a muddy bottom 

on which solid particles (shells, Foraminifera, 

etc.) occur. 

Ovigerous females have been recorded in January, 

February, April, May, August, and September 

(Monod, 1956; Forest and Guinot, 1966; Pillsbury). 

DISTRIBUTION.—Off tropical West Africa, 

where it has been recorded from localities between 

Senegal and Angola; there is one record 

from Mauritania, which requires corroboration. 

Records in the literature include: 

Mauritania: Cap Blanc (?) (Monod, 1933b). 

Senegal: S of Mbao (Monod, 1956). Near Dakar, 

14°38.5'-14°41'N, 17°20.5'-17°23.5'W, 22-34 m (Monod, 

1956). Anse de Hann (as Baie de Hann) (Monod, 1956). 

Between Goree and Thiaroye-sur-Mer, ca. 15 m (Monod, 

1956). Near Goree, 25 m (Balss, 1921); same, 17-% m 

(Monod, 1956). Baie de Goree, 18-28 m (Miers, 1881a). 

Near Bane du Seminole, Baie de Goree, 38 m (Monod, 

1956). Near lie de la Madeleine, 35 and 40 m (Monod, 

1956). 

Guinea: 09°40'N, 14°05'W, 18 m (Forest and Guinot, 

1966). Near Conakry, 09°N, 13°5O'W, 30 m; 09°22'N, 13°- 

37^, 10 m; 3 mi NW of Tamara, 12 m; between Tamara 

and lie de Corail, 10 m (all Monod, 1956). 

Sierra Leone: No specific locality, 44 m (Monod, 1956); 

6-44 m (Longhurst, 1958). 

Ivory Coast: 05°07'N, 04°32'W to O5°O7'N, 04°36'W, 

43-38 m (Voss, 1966). 

Ghana: Off Accra, 16-44 m (Monod, 1956; Gauld, 1960). 

04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m, and 

04°36.5'N, 01°31'W, 50 m (Forest and Guinot, 1966). 

Principe: O1°37'N, 07°22'E, 30 m; 01°38'25"N, 07°- 

22'05"E, 31 m; and 01°43'10"N, 07°28'20"E, 73 m (Forest 

and Guinot, 1966). 

Sao Tome: 00°25'40"N, 06°40'10"E, 50 m (Forest and 


Cabinda: No specific locality, 38-39 m (Guinot and 

Ribeiro, 1962). 

Angola: Off the mouth of the Congo River, 06°18'S, 

12°O2'E, 44 m (Doflein, 1904). 

Genus Ethusina Smith, 1884 

Ethusitu Smith, 1884:349 [type-species: Ethusina abyssicola 

Smith, 1884, by monotypy; gender: feminine].

NUMBER 306 43 

Key to Eastern Atlantic Species of Ethusina 

1. Outer orbital spine very small, directed laterally, not overreaching eye 

K alba 

Outer orbital spine large, directed anteriorly, overreaching eye 2 

2. Outer orbital spine extending to base of inner orbital spine . E. talismani 

Outer orbital spine extending to or beyond apex of inner orbital spine ... 

E. beninia, new species 

Ethusina alba (Filhol, 1884) 

FIGURES 8, 9d, 10 

Ethusa alba Filhol, 1884:199 [23 February; nomen nudum], 

230 [8 March; nomen nudum], 232 [8 March; available 

name].—Anonymous, 1884:7 [20 April].—Filhol, 1885b: 

129 [1 December]. 

Ethusina abyssicola.—Bouvier, 1898:66 [listed].—A. Milne Ed- 

wards and Bouvier, 1899:18;-1900:29, pi. 1: fig. 6 [color; 

outline based on E. abyssicola Smith from Smith, 1884: pi. 

2: fig. 1].—Bouvier, 1922:53, pi. 2: fig. 1 [color].—Monod, 

1956:85[references only]. [Not Ethusina abyssicola Smith, 

1884.] 

Ethusa abyssicola.—Bouvier, 1922:91 [not Ethusina abyssicola 

Smith, 1884]. 


Other Material: North Atlantic: Azores to France, 42° 15'N, 

23°37'W (of Paris, = 21°17'W of Greenwich), 3975 m, to 

42°19'N, 23°36'W (of Paris, = 21°16'W of Greenwich), 4060 

m, soft white mud, Talisman Sta 133, 134, 24 Aug 1883, Id, 

(holotype) 1$ (L,W). 

DESCRIPTION.—Carapace (Figures 8a, 10) longer 

than broad in male (width = 0.95 X length), 

broader than long in female (width = 1.03 X 

length), noticeably narrowing anteriorly. Carapace 

coarsely granular, at most lightly pubescent 

dorsally, few longer hairs present laterally. Cervical 

and cardiobranchial grooves distinct. Branchial 

regions moderately inflated. Front with 4 

upturned spines, submedians broad, blunt, laterals 

narrower, sharper, falling short of submedians 

in male (Figure 8a), overreaching submedians 

in female (Figure 8*). Front less than half greatest 

width of carapace. Outer orbital spine small, 

triangular, scarcely overreaching eye, not extending 

to front. Eyes short, stout, fixed, with terminal 

black cornea, narrower than stalk, visible in dorsal 

view. Basal segment of antennule inflated, 

surface irregular, not tuberculate. Basal article of 

antenna short, not extending to front. Antennal 

flagellum elongate, extending to posterior third 

of carapace. Ventral surface of body less granular 

but more pubescent than dorsal. 

Outer maxillipeds slender, merus narrower 

than, but more than half as long as, ischium. 

Chelipeds similar, equal, small, extending with 

merus to cornea in both sexes; propodus slightly 

larger in male than in female but not markedly 

inflated. Fingers (Figure 8b) longer than palm, 

slender, flattened, with shallow longitudinal 

groove, cutting edges crenulate. 

Second and third pereiopods similar, third 

longer, more than twice as long as chela, about 

2 1 A times as long as carapace. On each leg dactylus 

longer than propodus but shorter than car- 

FIGURE 8.—Ethusina alba (Filhol). Male, cl 10.4 mm, North 

Atlantic: a, front, dorsal view; b, chela; c, propodus and 

dactylus of third pereiopod, posterior view; d, abdomen. 

Female, cl 9.3 mm, North Atlantic: e, front, dorsal view; /, 

front, lateral view; g, abdomen.

44 

FIGURE 9.—Male pleopods: a, Ethusina beninia, new species, 

holotype, cl 14.0 mm, Pillsbury Sta 18; b, Ethusina robusta 

(Miers), cl 13.9 mm, Galapagos Islands; c, Ethusina talismani 

A. Milne Edwards and Bouvier, syntype, carapace broken, 

Morocco; d, Ethusina alba (Filhol), cl 10.4 mm, North Atlantic; 

e, Ethusina abyssicola Smith, cl 11.9 mm, Massachusetts. 

apace, subequal to or slightly shorter than merus, 

flattened dorsoventrally, grooved longitudinally, 

spatulate, slightly twisted, apex corneous; dactyli 

of both legs shorter than carapace in our material. 

Fourth and fifth pereiopods dissimilar, much 

shorter than second and third, both overreaching 

merus of third by length of dactylus; dactyli of 

last 2 legs very short, curved, apices corneous, 

with row of spinules on ventral margin; ischium 

of fifth pereiopod longer than that of fourth. 

Pereiopods with light surface pubescence, especially 

on fourth and fifth, latter also with some 

longer hairs. 

Male abdomen with 5 somites, that of female 

with 7 (Figure 8dg). 

Male Pleopod: Illustrated in Figure 9d. 

Color: "Cephalothorax bleuatre sur le vivant, 

avec une legere teinte violacee; pattes et abdomen 

d'un blanc jaunatre, avec les doigts de couleur 

rose" (A. Milne Edwards and Bouvier, 1900:30). 

Bouvier (1922:53) noted that material from the 

Cape Verde Islands was "jaune d'ocre pale avec 


les doigts et les antennules roses" and that in 

material collected earlier by the Talisman the 

carapace was "uniformement bleuatre." The 

color of this species has been illustrated by A 

Milne Edwards and Bouvier (1900, pi. 1: fig. 6) 

and Bouvier (1922, pi. 2: fig. 1). 

MEASUREMENTS.—The single male examined 

has a carapace length of 10.4 mm, a carapace 

width of 9.9 mm; the only female examined 

measures 9.3 X 9.6 mm. 

REMARKS.—The present species was identified 

by A. Milne Edwards and Bouvier (1900:29) with 

Ethusina abyssicola Smith. Originally, however, A. 

Milne Edwards must have thought that the species 

was new and gave it provisionally the manuscript 

name Ethusa alba. Filhol (1884:199, 230, 

232), in a popular article on the Travailleur and 

Talisman expeditions, introduced, obviously unintentionally, 

the name Ethusa alba into the literature; 

A. Milne Edwards was cited as the author. 

In that paper, Filhol used the name on three 

occasions; in two of these (pp. 199, 230) the name 

was not accompanied by sufficient data to make 

it an available name. On page 232, however, 

Filhol remarked that Ethusa alba differed from 

other species of the genus by the presence of eyes 

("dans un meme genre Ton trouve des especes 

aveugles et d'autres qui ne le sont pas, ainsi 

YEthusa granulata [= Cymonomus granulatus] qui vit 

dans les mers du Nord entre 200 et 1300 metres 

FIGURE 10.—Ethusina alba (Filhol) (from Bouvier, 1922, pi. 1: 

fig. 1).

NUMBER 306 45 

est aveugle, alors que YEthusa alba que nous avons 

pris dans 1'Ocean par 5000 metres ne Test pas"). 

Hereby the name Ethusa alba becomes available 

as from 8 March 1884 and Filhol has to be cited 

as the author. Since the publication of Filhol's 

paper, the name Ethusa alba has only been mentioned 

a few times in the literature. First in an 

anonymous guide to the "exposition sous-marine 

du Travailleur et du Talisman" issued for the 

"visiteurs de l'Exposition organisee sous la direction 

de M. Alph. Milne-Edwards, president de la 

Commission des dragages". This guide was a 

"publication du journal La Nature" and consists 

of an abstract of Filhol's 1884 paper, probably 

made by Gaston Tissandier, chief editor of the 

journal. The dates of the exposition are given on 

page 16 of the guide as follows: 

Cette exposition des collections du Travailleur et du Talisman 

a ete ouverte le 26 Janvier 1884 et fermee le 16 mars. 

On 1'a ouverte de nouveau le 7 avril, a l'occasion de la 

reunion des societes savantes a la Sorbonne, et elle sera 

fermee le 20 du meme mois. 

The fact that the last verb is written in the 

future tense shows that the catalog was written 

and probably published before 20 April 1884. 

The remark on Ethusa alba in the guide is verbally 

the same as the one made by Filhol (1884:199), 

viz., "les Ethuses dont une espece nouvelle, 

YEthusa alba, a ete prise a 5000 metres de profondeur." 

Filhol (1885b: 129) in his book La vie au 

fonds des mers, does not add anything new to the 

knowledge of the species. We have been unable 

to find any later reference to Ethusa alba in the 

literature, probably because A. Milne Edwards 

and Bouvier (1899) identified their specimens 

with Ethusina abyssicola S. I. Smith and used that 

name ever since. 

Apart from the fact that the species has eyes, 

the only information on it provided by Filhol is 

that it was taken by the Talisman at 5000 m 

(Filhol, 1884:199, 232; 1885b: 129) or "entre 

quatre et cinq mille metres" (Filhol, 1884:230). 

A. Milne Edwards and Bouvier (1900) in their 

account of the Talisman and Travailleur Brachyura, 

listed the following Ethusinae collected by the 

Talisman: Ethusa mascarone (Herbst) in 38 and 60 

m (Sta 56); E. rugulosa A. Milne Edwards and 

Bouvier in 150-275 m (Sta 103); E. rosacea A. 

Milne Edwards and Bouvier in 930 and 1013— 

1113m (Stas 85,87); Ethusina abyssicola S. I. Smith 

in 2995, 3655, 3975, and 4060 m (Stas 102, 131, 

133, 134); and E. talismani A. Milne Edwards and 

Bouvier in 2075, 2083, 2115, 2212, and 2235 m 

(Stas 40, 41, 43, 44, 130). Of Ethusina abyssicola 

they remarked (1900:30): "C'est, de tous les 

Crabes, le plus abyssal." Although none of the 

crabs of the Talisman expedition was collected at 

5000 m, as stated for Ethusa alba, Ethusina abyssicola 

is the only species then taken in a depth of more 

than 4000 m and would fit Filhol's statement that 

Ethusa alba was taken "entre quatre et cinq mille 

metres." The only Talisman station of 5000 m 

depth is no. 137 of 27 August 1883 (4975-5005 

m), but no crabs are recorded from it. Filhol's 

statement that Ethusa alba was taken at 5000 m 

either is an exaggeration or an error (he may have 

confused stations 134 and 137), or a careless 

abbreviation of 4000-5000 m. Confusion in the 

station numbers of the Talisman expedition occurred 

very frequently (the expedition is rather 

notorious because of it). So de Folin (1887:332) 

when dealing with station 136 (26 August 1883, 

depth 4255 m) mentioned from it "des Ethuses 

differentes de celles deja connues," although A. 

Milne Edwards and Bouvier (1900) do not mention 

any Ethusinae from that station. However 

this may be, there can be little doubt that the 

present species at first was thought by A. Milne 

Edwards to be a new species, Ethusa alba, and that 

he later changed his views and considered it 

identical with S. I. Smith's Ethusina abyssicola. 

Filhol's use of the name Ethusa alba in print made 

that name available and it has to be used, now 

that the specific distinctness of the Talisman specimens 

from S. I. Smith's Ethusina abyssicola has 

been demonstrated. 

In identifying material of this species collected 

by the Talisman at several different localities, A. 

Milne Edwards and Bouvier (1900:29) noted: 

"Nos specimens ressemblent completement a ceux 

qu'a decrits ou figures S.-I. Smith. Nous ferons 

remarquer, toutefois, que les doigts de leurs

46 

grandes pattes ambulatoires sont manifestement 

tordus, que leurs saillies rostrales sont tantot obtuses, 

tantot acuminees, enfin que leurs epines 

susorbitaires ont une longueur extremement variable. 

L'aire cardiaque est toujours assez distincte 

et les dents laterales du 3° segment abdominal 

sont peu saillantes." 

Our material ofE. alba differs from E. abyssicola 

in having much shorter frontal spines with their 

apices only slightly reflected dorsally. In E. abyssicola 

the frontal spines are much more prominent 

and their apices, especially of the outer pair, are 

strongly reflected dorsally, being situated at an 

angle of almost 45° from the longitudinal axis of 

the carapace. The most important difference between 

the two species is in the structure of the 

male pleopod, which in E. alba (Figure 9d) is 

slenderer, not sharply bent basally, with a distinct, 

corneous angled prominence apically; the 

apex of the pleopod in E. abyssicola is rounded or 

flattened distally (Figure 9*). 

As in E. abyssicola, the female of E. alba is 

comparatively broader and has much shorter 

frontal spines than the male. 

The figure of this species given by A. Milne 

Edwards and Bouvier (1900, pi. 1: fig. 6) is based 

on an outline of E. abyssicola published by Smith 

(1884). 

Coordinates for Talisman station 133 given by 

A. Milne Edwards and Bouvier (1900:30), 42°- 

45'N, 23°37 / W of Paris (- 21 O 17'W of Greenwich), 

may be in error, for that station was made 

on the same day as station 134 and the latitude 

for station 133 is given as 42°15'N in a list of 

Talisman stations prepared by the Paris Museum. 

These stations were so close to each other that the 

material from them may have been combined. 

The male specimen from Leiden has the following 

label data: 42°19 / N, 23°37'W [Paris], 4060 m, 

Talisman Sta. 134 [(it should read longitude 

23°36'W (Paris)], whereas that from Washington 

bears the coordinates 42°15'N, 23°37'W (Paris), 

3975-4060 m; both sets of data were originally 

based on longitude of Paris and are corrected to 

Greenwich (see "Distribution"). 

BIOLOGY.—Ethusina alba, like E. abyssicola, is a 


true abyssal species, living in depths exceeding 

1000 m. It has been collected on white mud with 

Foraminifera in 4261 m (A. Milne Edwards and 

Bouvier, 1899); soft white mud in 2995, 3975 and 

4060 m, and gray mud in 3655 m (A. Milne 

Edwards and Bouvier, 1900); and on muddy 

volcanic sand with Foraminifera in 3890 m (Bouvier, 

1922). 

Ovigerous females have not been recorded. 


North Atlantic N of the Azores to the Cape Verde 

Islands, in depths between 2995 and 4261 m. 

Records in the literature include: 

North Atlantic between the Azores and France: 42°15'N, 

23°37'W of Paris (= 21°17'W of Greenwich), 3975 m, and 

42°19'N, 23°37'W of Paris (= 21°17'W of Greenwich), 4060 

m (A. Milne Edwards and Bouvier, 1900). 

Azores: 38°38'N, 27°26'W of Paris (= 25°06'W of Greenwich), 

2995 m (A. Milne Edwards and Bouvier, 1900). 

36°55'N, 24°43'W of Paris (= 22°23'W of Greenwich), 4261 

m (A. Milne Edwards and Bouvier, 1899). 

Cape Verde Islands: 16° 12'N, 24°43'45"W, 3890 m (Bouvier, 

1922). 15°48'N, 22°43'W of Paris (= 20°23'W of Greenwich), 


* Ethusina beninia, new species 

FIGURES 9a, 11 


Sta 18, 3047-3129 m, soft, dark gray clay, 16* (holotype), 19 

(L). Sta 34, 1948-1984 m, mud, 19 (W). 

DESCRIPTION.—Carapace (Figure 1 la) slightly 

broader than long (1.05 times broader in male, 

1.01 to 1.05 times in females), noticeably narrowing 

anteriorly. Surface finely granular, lightly 

pubescent. Cervical and cardiobranchial grooves 

distinct, latter deeper, better defined. Branchial 

regions inflated. Intestinal region with pair of 

rounded prominences. Front (Figure 1 \a-d) with 

4 upturned spines, inner pair broader and shorter 

than outer, with sinus between inner spines 

broader and deeper than that between inner and 

outer spines. Inner frontal spines shorter, blunter 

in female than in male. Front less than half 

greatest width of carapace. Outer orbital spine 

strongly developed, extending to or beyond base 

of frontal sinus, apex curved upward, directed

NUMBER 306 47 

a d 

FIGURE 11.—Ethusina beninia, new species. Holotype, male, cl 14.0 mm, Pillsbury Sta 18: a, dorsal 

view; b, front, dorsal view; c, front, lateral view; g, abdomen. Paratype, female, cl 16.1 mm, 

Pillsbury Sta 18: d, front, ventral view; e, chela;/, dactylus of third pereiopod; h, abdomen. 

anterolaterally, broader and shorter in female 

than in male. Eyes (Figure 1 lc,d) short, stout, 

fixed, with terminal black cornea; eyes tapering 

distally, scarcely or not at all visible in dorsal 

view. Basal segment of antennule inflated, with 

1 or 2 anterior tubercles. Basal article of antenna 

short, not extending to front. Antennal flagellum 

elongate, extending to posterior fourth of carapace. 

Ventral surface of body more granular than 

dorsal. 

Outer maxillipeds slender, merus narrower 

than, but more than half as long as, ischium. 

Chelipeds similar, equal, small, slender, with 

propodus slightly stouter than carpus. Palm 1.44

48 

times longer than high in male, 1.33-1.34 times 

in female. Left chela broken on only intact male, 

meri of chelipeds on that male subsimilar in size 

and shape, intact cheliped slightly larger than 

those of female (Figure 11

NUMBER 306 49 

30°03'N, 14°02'W of Paris (= 11°42'W of Greenwich), 

in 2212 m, was shown on plate 10: figure 

9. Apparently the same male was figured in color 

on plate 3: figure 6, although according to the 

text a female was figured there (only one female 

was reported in the collection). According to the 

figure and to the account given in the text (A. 

Milne Edwards and Bouvier, 1900:31), "les femelles 

se distinguent des males par leur carapace 

plus large, par leurs pointes frontales plus courtes 

et plus obtuses . . . ," the female differed from the 

males in having the frontal spines shorter and 

broader, the lateral spines originating more anteriorly 

and extending further forward, and the 

eyes were not visible in dorsal view. In these 

features the female more closely resembles E. 

beninia, differing mainly in having shorter exorbital 

spines which are directed anteriorly. The 

female reported by A. Milne Edwards and Bouvier 

and E. beninia both differ from males of E. 

talismani in having eyes that are hidden in dorsal 

view and in having the exorbital spines originating 

nearer the anterior margin of the carapace. 

There is no indication of dimorphism to such an 

extent in our material of E. beninia, so it seems 

likely that the female from the Talisman collection 

may represent a distinct species. 

TYPE-LOCALITY.—Bight of Benin, off Ghana, 

05°01'N, 00° 12'E, in 3047-3129 m. 

DISPOSITION OF TYPES.—The male holotype 

(Crust. D. 31781) and female paratype from Pillsbury 

Sta 18 are in the Rijksmuseum van Natuurlijke 

Historie, Leiden. A female paratype from 

Pillsbury Sta 34 is in the National Museum of 

Natural History, Smithsonian Institution, Washington. 

ETYMOLOGY.—The specific name, beninia, alludes 

to its occurrence in the Bight of Benin. 

BIOLOGY.—This is a deep water species occurring 

between 1984 and 3047 m (the two stations 

at which it was collected being from 1948-1984 

m and 3047-3129 m). The bottom where the 

species was obtained in 3047-3129 m was soft, 

dark gray clay. 

DISTRIBUTION.—This species is known only 

from the Bight of Benin, off Ghana, at 05°01'N, 

00°12'E and 03°53'N, 02°33'W. 

Family CALAPPIDAE de Haan, 1833 

CALAPPIDEA de Haan, 1833:ix [corrected to Calappidae by 

White, 1847a:44; name 371 on Official List]. 

MATUTOIDEA de Haan, 1835:36. 

ORITHYINAE Dana, 1852b:391. 

EASTERN ATLANTIC GENERA.—Four, Acanthocarpus, 

Calappa, Cycloes, and Matuta, all of which have 

representatives in tropical waters. 

EASTERN ATLANTIC SPECIES.—Seven, all of 

which were recorded by Monod (1956), as follows: 

Name in Monod 

Matuta michaelseni 

[Osachila stimpsoni 

(Calappidae)] 

Calappa gallus 

Calappa peli 

Calappa granulata 

Calappa rubroguttata 

Acanthocarpus brevispinis 

Cryptosoma cristatum 

Current Name 

Matuta michaelseni 

[Sakaila africana, 

new genus, new species* 

(Part henopi dae) ] 

Calappa gallus 

Calappa pelii* 

Calappa granulata 

Calappa rubroguttata* 

Acanthocarpus brevispinis* 

Cycloes cristata 

As noted above, we have placed West African 

material identified with Osachila stimpsonii in Sakaila 

africana, new genus, new species, which has 

been assigned to the Subfamily Aethrinae, Family 

Parthenopidae (p. 322). 

We believe that one other species reported in 

the West African fauna by Monod (1956:151) is 

based on an erroneously labeled specimen. It is 

Hepatus princeps (Herbst, 1794): a western Atlantic 

species reported from "Guinea" by Rathbun 

(1937:237) based upon a single specimen in the 

Zoological Museum, Copenhagen. 

Subfamily CALAPPINAE de Haan, 1833 

Genus Acanthocarpus Stimpson, 1871 

Acanthocarpus Stimpson, 1871a: 152 [type-species:- Acanthocarpus 

alexandri Stimpson, 1871, by monotypy; gender: masculine].

50 

* Acanthocarpus brevispinis Monod, 1946 

Acanthocarpus africanus Capart, 1951:35, fig. 8.—Rossignol, 

1957:127 [key]. 

Acanthocarpus brevispinis Monod, 1956:109, figs. 125-132.— 

Guinot-Dumortier and Dumortier, 1960:129 [stridulation].—Guinot 

and Ribeiro, 1962:27.—Forest, 1963: 

628.—Monod, 1963, fig. 32.—Maurin, 1968b:484, 492, 

fig. 6— Le Loeuffand Intes, 1968, table 1; 1969:66. 

Acanthocarpus bispinosus.—Longhurst, 1958:87 [not Acanthocarpus 

bispinosus A. Milne Edwards, 1880]. 

Acanthocarpus brevispinnis.—Maurin, 1968b:489 [erroneous 

spelling]. 


Coast: Sta 51, 329-494 m, 1

NUMBER 306 51 

March and October (Capart, 1951; Monod, 

1956). 

DISTRIBUTION.—West Africa, where it has been 

recorded from a few localities between Cap Juby, 

Morocco [the type-locality] and Angola, in depths 

between 100 and 500 m. Monod (1956) studied 

material from Senegal, Gambia, and Guinea. 

Since 1956 it has been recorded from the 

following: 

Spanish Sahara: Off Villa Cisneros, 300-500 m; off 

Tamzak (as Tamxat), 200-400 m (both Maurin, 1968b). 

Mauritania: Bane d'Arguin, 200-300 m (Maurin, *968b). 

Senegal: Off Saint-Louis, 300 m; Fosse de Kayar, 250 to 

300-350 m (both Maurin, 1968b). 


1958). 


1969); Off Grand-Bassam, 200 m (Le Loeuff and Intes, 

1968). 04°32'30"N, 06°31'W, 300-455 m (Forest, 1963). 


Angola: Ponta de Sao Jose, Baia Farta, Benguela, 235- 

310 m (Guinot and Ribeiro, 1962). 

Genus Calappa Weber, 1795 

Calappa Weber, 1795:92 [type-species: Cancer granulatus Linnaeus, 

1758, by subsequent designation by Latreille, 1810: 

422; gender: feminine; name 1611 on Official List]. 

Camara de Haan, 1837:67 [type-species: Calappa fomicata Fabricius, 

1798, a subjective junior synonym of Calappa 

calappa Linnaeus, 1758, by monotypy; gender: feminine]. 

Lophos de Haan, 1837:67 [type-species: Cancer lophos Herbst, 

1785, by tautonymy; gender: masculine]. 

Gallus de Haan, 1837:67 [type-species: Cancer gallus Herbst, 

1803, by tautonymy and monotypy; gender: masculine]. 

Pistor Gistel, 1848:ix [substitute name for Gallus de Haan, 

1837; type-species: Cancer gallus Herbst, 1803; gender: 

masculine]. 

Calappa gallus (Herbst, 1803) 

Calappa gallus.—Monod, 1956:100, figs. 115, 116 [Cape 

Verde Islands, Senegal, Sierra Leone, Ghana].—Rossignol, 

1957:76, 127 [key], fig. 1 [Congo].—Longhurst, 1958: 

87 [Sierra Leone]—Gauld, 1960:68 [Ghana].—Guinot and 

Ribeiro, 1962:26 [Cape Verde Islands].—Rossignol, 1962: 

114 [Congo].—Ribeiro, 1964:4 [Cape Verde Islands].— 

Forest and Guinot, 1966:51 [Principe, Sao Tome, Annobon].—Guinot, 

1967a:245 [Indian Ocean, listed].— 

Monod, 1967:178 [no locality].—Ribeiro, 1973:5 [Cape 

Verde Islands]. 

SYNONYMS.—Calappa galloides Stimpson, 1859; 

Calappa squamosa Desbonne and Schramm, 1867; 

Calappa gallus var. capellonis Laurie, 1906. 

DISTRIBUTION.—Tropical Atlantic and Indo- 

West Pacific. In the eastern Atlantic it has been 

recorded from scattered localities between the 

Cape Verde Islands and Senegal to Angola, in- 

cluding the offshore islands of the Gulf of Guinea; 

from shore to a depth of about 50 m. 

Calappa granulata (Linnaeus, 1758) 

Calappa granulata.—Gruvel, 1913:168 [listed].—Chapman 

and Santler, 1955:374.—Monod, 1956:105 [references 

only].—Figueira, 1960:8.—Peres, 1964:28.—Zariquiey Alvarez, 

1968:315, figs. 105c, 107a [Spain; references].— 

Maurin, 1968a: 18 [Portugal], 43, 107 [Mediterranean], 

fig. 4; 1968b:479, 480, 482, 484, fig. 2.—Ribeiro, 1973:4, 

fig. 1.—Turkay, 1976a:25 [listed], 36, fig. 17 [Portugal in 

part]; 1976b:61 [listed], 62.—Bas, Arias, and Guerra, 1976, 

table 3. 

Calappa.—Maurin, 1968b, fig. 1. 


Other Material: Morocco: Off Cap Hadid, 31°54'N, 09° 

55^, 85 m, muddy sand, 5 m beam trawl, 25 Mar 1976, 

Onversaagd Sta 126, 19 (L). Casablanca fishmarket, 31 Mar 

1976, 1

52 

Geer ex Algiria mihi," showing that De Geer's 

specimen(s), originating from Algeria, were examined 

by Linnaeus himself. Linnaeus' (1758) 

description evidently is entirely or partly based 

on this Algerian material. In this description, 

namely, Linnaeus described "Rostrum bilobum, obtusum" 

while Catesby's figure shows a narrowly 

rounded truncate rostrum, which certainly cannot 

be called bilobed. As Cancer granulatus is a 

composite species, its identity can only be fixed 

by a lectotype selection. As this, so far as is known 

to us, has so far not been done, we now select as 

the lectotype of Cancer granulatus Linnaeus, 1758, 

the specimen (or if more than one specimen is 

represented in the material, the largest specimen) 

from Algeria in the De Geer collection. 

Monod (1956:105) noted that "la seule mention 

possible de l'espece dans notre dition est celle 

d'A. MILNE-EDWARDS et BOUVIER [1900:59-60]. 

Mais comme ces auteurs considerent rubroguttata 

comme probablement synonyme de granulata, rien 

ne prouve qu'ils aient bien eu cette derniere 

espece sous les yeux." 

The confirmation of the early record of this 

species from the Cape Verde Islands (A. Milne 

Edwards and Bouvier, 1900) by Ribeiro (1973), 

the records of Maurin (1968a,b) from Morocco, 

Spanish Sahara, and Mauritania, and the present 

records from Morocco suggest that this species is 

more widely distributed in the Atlantic than previously 

believed. 

BIOLOGY—Calappa granulata is a sublittoral species 

which, according to Bouvier (1940) and Zariquiey 

Alvarez (1968), lives in depths between 30 

and 150 m. If the recent records in the literature 

are accurate, the species occurs in much deeper 

water off the northwestern coast of Africa. Tiirkay 

(1976a) recorded it from depths of 160-250 m off 

Morocco and (1976b) in 13 m and 20-30 m off 

Madeira. Peres (1964) found it off Morocco in 

260-500 m on compact bathyal mud. Maurin 

(1968a,b) recorded it from the following depths 

and bottom types off Spanish Sahara: 200-300 m 

and 200-350 m, slightly sandy mud with Pennatulacea; 

400-700 m, sandy mud; 50-90 m, on 

muddy detritus; and 60 m, shelly, muddy sand, 


as well as on bathyal mud with funiculines and 

Isidella in 350-450 m off Morocco. 


Mediterranean, the Atlantic coasts of Portugal 

and Spain, southward to Cap Blanc, Mauritania, 

including the Azores, Madeira, and the Cape 

Verde Islands, in depths between 13 and 400-700 

m (Zariquiey Alvarez, 1968). Monod (1956) summarized 

the literature but reported no material. 

In addition, the following species localities have 

been recorded. 

North Atlantic: Conception Bank, 200 and 237 m 


Azores: Faial (Figueira, 1960). Horta Harbor, Faial 

(Chapman and Santler, 1955). 

Madeira: No specific locality; Funchal harbor, 13 m; 

Ponta dos Reis Magos, 20-30 m (Tiirkay, 1976b). 

Morocco: Foum Agoiiitir (as Puerto-Cansado), 350-450 

m (Maurin, 1968b). 35°05.5'N, 09°18'W, 160-250 m (Turkay, 

1976a). 34°39.6'N, 06°54.5'W to 34°33.5'N, 06°56'W, 

260-500 m (Peres, 1964). 

Spanish Sahara: 23°30'N, 16°06'W to 23°28.5'N, 

16°09.5'W, 24-29 m (Bas, Arias and Guerra, 1976). Off 

Medano de Aaiun and Cabo Bojador, 200-300 m (Maurin, 

1968a,b). Between Cabo Bojador and Morro Garnet, 200- 

350 m; off Morro Garnet, 400-700 m; between Cabo Corbeiro 

and Cabo Blanco, 50-90 m; and off peninsula of Cabo 

Blanco, 60 m (all Maurin, 1968b). 

Cape Verde Islands: Boavista, 15°58'N, 22°43'W, 80 m 

(Ribeiro, 1973). 

* Calappa pelii Herklots, 1851 

FIGURE 12a 

Calappa pelt.— Capart, 1951:39, figs. 9, 10.—Monod, 1956: 

102, figs. 117-121.—Rossignol, 1957:75, 127 [key].— 

Longhurst, 1958:87.—Gauld, 1960:69.—Guinot and Ribeiro, 

1962:26.—Rossignol, 1962:114.—Crosnier, 1964:34, 

35, fig. on pi. B.—Forest and Guinot, 1966:52—Monod, 

1967:178 [no material].—Maurin, 1968a:48, 59, 64, fig. 

23; 1968b:484, 486, 489, 491, figs. 5, 9.—Le Loeuff and 

Intes, 1968:40, table 1, figs. 48, 61, 63; 1969:63, 64.— 

Crosnier, 1970:1215 [listed], 1216. 

Calappa.—Voss, 1966:19.—Maurin, 1968b, figs. 1, 4[?]. 

Calappa Pelii.—Holthuis, 1968:29 [listed]. 

SYNONYM.—Calappa piscatorum Caiman, 1914. 


Sta 68, 70 m, broken shell, 39, 16 juv (L). 

Ivory Coast: Sta 42,62-75 m, mud with brown, branched

NUMBER 306 

Foraminifera, 2 juv (L). Sta 46, 38-42 m, mud with dense 

Jullienella, 3 juv (W). Sta 47, 37 m, bottom with Jullienella, 

1 juv (L). Sta 49, 73-77 m, 2 juv (W). Sta 62, 46 m, brown, 

branching and foliate Foraminifera, 7 juv (L). Sta. 64, 68 m, 

1$, 1 juv (W). 

Ghana: Sta 17, 48 m, fine sand and green mud, 1? (L). 

Sta 22, 51 m, rough bottom, Id (W). Sta 23, 42 m, foliate 

brown to orange bryozoans, 3 juv (L). 

Nigeria: Sta 237, 101 m, 16* (W). Sta 241, 59-63 m, mud 

and shell, 8 juv (L). Sta 253, 33-40 m, mud, 1 juv (L). 

Cameroon: Sta 259, 59 m, mud and broken shell, 2 juv 

(W). Sta 260, 46 m, 7 juv (L). 

Gerommo Material: Gabon: Sta 212, 200 m, 1

54 

The deepest records for this species that we 

have found are those of Maurin (1968a,b), who 

reported material from depths of 200 m off Spanish 

Sahara and 200-350 and 200-400 m off Mauritania. 

The species generally is taken in depths of 

50 to 150 m. 

Ovigerous females have been collected in 

March, May, August, October, and December 

(Capart, 1951; Guinot and Ribeiro, 1962). 

DISTRIBUTION.—Off West Africa, from localities 

between Spanish Sahara and Angola, in 

depths between 8-20 and 400 m, usually between 

50 and 150 m. Monod (1956) recorded material 

from localities between Port-Etienne, Mauritania, 

and Luanda, Angola. Since 1956 this species 

has been recorded from the following localities. 

West Africa: No specific locality (Monod, 1967). 


Blanco, 60-80 m (Maurin, 1968a); in 200 m (Maurin, 

1968b). Between Cabo Barbas and Cabo Blanco, 50-90 m 


Mauritania: N of Bane d'Arguin, 20°20'N to 20°40'N, 

90-100 m (Maurin, 1968a). Bane d'Arguin, 40-60 m, 60-70 

m, 90-100 m (Maurin, 1968b). Between Cap Timiris and 

Tamzak (as Tamxat), 200-350 m (Maurin, 1968a). Off 

Tamzak (as Tamxat), 200-400 m (Maurin, 1968b). 

Senegal: Saint-Louis, 35-40 m (Maurin, 1968b). Off 

Mboro, 35-40 m (Maurin, 1968b). 13°01'N, 17°24'W, 5U 

55 m, and 12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 

1966). 



Sierra Leone: No specific locality, in 12-120 m (Longhurst, 

1958). 


1969). Off Sassandra, off Fresco, off Grand-Lahou, off Jacqueville, 

and off Grand-Bassam, 22-200 m (Le Loeuff and 

Intes, 1968). 05°07'N, 04°32'W to O5°O7'N, 04°36'W, 38- 

42 m (Voss, 1966). 

Ghana: Off Accra, 37 m (Gauld, 1960). 04°36.5'N, 01° 

31'W, 50 m (Forest and Guinot, 1966). 

Cameroon: No specific locality, in more than 50 m (Crosnier, 

1964). 

Principe: 01°38'25"N, 07°22'05"E, 31 m (Forest and 


Gabon: 00°25'N, 09°00'E, 73 m (Forest and Guinot, 

1966). 

Congo: Pointe-Noire, 40-75 m (Rossignol, 1957). W of 

Pointe-Noire, 80-100 m (Rossignol, 1962). 


Angola: 16°41'S, 11°21'E, 162 m (see "Remarks"); 16° 

37'S, 11°22'E, 122 m; 17°O2'S, 1 l°40'E, 54 m; and 17°06'S, 

11°35'E, 90 m (all Crosnier, 1970). Luanda, 110-113 m; 

Mussulo Grande, Luanda, 86-90 m; Porto Amboim, 85 m; 

Baia da Caota, Benguela, 8-20 m, 13 m, 17 m, 30 m; 

Sombreiro, 13 m; between Ponta da Caruita and Sombreiro, 

29 m; Baia Farta, 22 m, 100-144 m; Baia de Mocamedes; 

Baia dos Tigres, 107 m (all Guinot and Ribeiro, 1962). 

*Calappa rubroguttata Herklots, 1851 

FIGURES 126, 13 

Calappa rubroguttata.—Biiuikofer, 1890:466, 487.—Johnston, 

1906: 862.—Postel, 1950:25, 26.—Monod, 1956:106, figs. 

122-124.—Rossignol, 1957:76.—Longhurst, 1958:87.— 

Buchanan, 1958:23.—Gauld, 1960:69.—Rossignol, 1962: 

144.—Guinot and Ribeiro, 1962:27.—Crosnier, 1964:32, 

35, fig. on pi. B.—Monod, 1967:178, pi. 16: fig. 1 [no 

material].—Forest and Guinot, 1966:53.—Maurin, 1968b: 

491, fig. 9.—Le Loeuff and Intes, 1968:40, table 1, figs. 

48, 61, 63; 1969:63, 64, 65.—Uschakov, 1970:439, 455 

[listed]. 

?Calappa granulata.—Bouvier, 1911:226 [not C. granulata (Linnaeus, 

1767)]. 

Calappa rubroguttatus.—Capan, 1951:43, figs. 10, 11.—-Rossignol, 

1957:127 [key]. [Erroneous spelling.] 

Calappa.— Bassindale, 1961:499. 

Callappa rubroguttata.—Bott, 1968:170 [erroneous spelling]. 

SYNONYM.—Calappa bocagei De Brito Capello, 

1871. 

MATERIAL EXAMINED.—Pilhbury Material: Ivory 

Coast: Sta 47, 37 m, bottom with Jullienella, 1? ov (L). 

Ghana: Sta 24, 35-37 m, dark red bryozoans, 2 juv (W). 

Nigeria: Sta 248, 33 m, 2 juv (W). 

Other Material: Liberia: No specific locality, 1879-1882, J. 

Biittikofer, 3cJ, 1? (L) (smallest male with huge Chelonibia 

patula (Ranzani) on posterior margin of carapace). 

Ghana: Butre (04°49'N, 01°55'W), 1841-1851, H. S. Pel, 

lectotype, \6 (L); paralectotypes, 8

NUMBER 306 

pattern in this species: "Coloration typique: la 

carapace beige clair, marquee des bandes et 

taches rondes carminees. La partie anterieure a 

fond carmin clair. La pince marquee de trois 

taches rondes disposees en triangle; le carpe de 

deux taches. Les pattes de couleur claire." 


widths of 14 to 104 mm; the carapace width of 

the single ovigerous female examined is 95 mm. 

REMARKS.—The lectotype of this species, selected 

here, is a male, cb 104 mm, from Butre 

(= Boutry), east of Dixcove, Ghana, collected 

between 1841 and 1851 by H. S. Pel (Crust. D. 

772). There are nine paralectotypes, a female, cb 

51 mm, and eight males, cb 82 to 99 mm, two of 

which are preserved dry (Crust. D. 771). 

A sketch of the carapace of a juvenile is shown 

in Figure \2b along with that of a young C. pelii 

(Figure 12a). We also reproduce here an original 

illustration of this species prepared by Herklots 

(Figure 13). 

BIOLOGY.—This species lives in shallower water 

than does C. pelii, from shore to a maximum 

depth of about 90 m. It apparently can live on all 

level substrates, but may prefer sand. Gauld 

(1960:69) reported that off Ghana this species was 

"very common on sand from shallow water to 30 

m. Catches of up to 250 have been taken in a 

single net; large catches have been taken only 

beyond 10 m." Crosnier (1964) characterized this 

species as a warm water crustacean, living in 0- 

30 m off Cameroon; there it replaces C. pelii in 

FIGURE 13.—Calappa ruboguttata Herklots (sketch by J. A. 

Herklots). 

shallow water. Off the Ivory Coast, Le Loeuff and 

Intes (1968:40) found this species to be very 

abundant all year on all sediments in depths less 

than 40 m, usually between 20 and 35 m. In 

depths between 30 and 40 m it occurred together 

with C. pelii. In 1969 they noted that it appeared 

to be indifferent to the nature of the substrate 

and that it was eurythermic. 

More than 85% of the occurrences reported in 

the literature for which depth information is 

available are from depths of 40 m or less. There 

are about equal numbers of records of the species 

in depths below 20 m and between 20 and 40 m. 

Ovigerous females have been collected in May, 

September, and October (Capart, 1951; Forest 

and Guinot, 1966; Pillsbury). 

DISTRIBUTION.—Off West Africa, between Senegal 

and Angola, usually in depths of less than 50 

m. Monod (1956) summarized the literature prior 

to 1956 and reported numerous specimens from 

localities between Senegal and Luanda, Angola. 

In addition this species has been recorded from 

the following localities. 


Senegal: No specific locality, 30-80 m (Postel, 1950). Off 

Saint-Louis, 35-40 m; off Mboro, 35-40 m (Maurin, 1968b). 

13°01'N, 17°24'W, 51-55 m (Forest and Guinot, 1966). Baie 

de Rufisque (Bouvier, 1911). 

Guinea: No specific locality, depths greater than 20 m 

(Uschakov, 1970). 


1958). 

Liberia: No specific locality (Biittikofer, 1890; Johnston, 

1906). 


1969). 05°00'N, 05°28.5'W, 27 m; 05°03.5'N, 05°25'W, 20- 

25 m; 05°02.5'N, 05°25'W, 21-27 m (Forest and Guinot, 


and off Grand-Bassam, 8-40 m (Le Loeuff and 

Intes, 1968). 

Ghana: Off Accra (Bassindale, 1961). In 3-8 fm (5-15 

m) (Buchanan, 1958). In shallow water to 30 m (Gauld, 

1960). 


Principe: Between Ponta da Mina and Ponta Novo Destino, 

6 m (Forest and Guinot, 1966). 

Gabon: 00°38'25"N, 08°46'E, 5 m (Forest and Guinot, 

1966). 

55

56 

Congo: Baie de Pointe-Noire, beach (Rossignol, 1957). 

Pointe-Noire, beach (Rossignol, 1962). 

Angola: Luanda (Bott, 1968). Bai'a de Benguela, shore 

(Guinot and Ribeiro, 1962). 

Genus Cycloes de Haan, 1837 

Cydoes de Haan, 1837:67, 68, 69 [type-species: Cycloes granulosa 

de Haan, 1837, by monotypy; gender: feminine]. 

Cryptosoma Brulle, 1837:16 [an invalid junior homonym of 

Cryptosoma Bert hold, 1827 (Coleoptera); type-species: Cryptosoma 

cristatwn Brulle, 1837, by monotypy; gender: neuter]. 

REMARKS.—Chace (1968:610) discussed the use 

of the two names previously applied to this genus 

and gave convincing reasons for using Cycloes 

rather than Cryptosoma. He also provided a key to 

the species of Cycloes. 

Cycloes cristata (Brulle, 1837) 

Cryptosoma cristatum.—Monod, 1956:114, fig. 133 [Cape 

Verde Islands; references].—Tiirkay, 1976b:61 [listed]. 

Cycloes cristata.—Guinot-Dumortier and Dumortier, 1961: 

561, figs. 1-4 [Cape Verde Islands].—Guinot and Ribeiro, 

1962:27, figs. 3, 4 [Cape Verde Islands]. 

Cycloes cristata.—Chace, 1968:610 [key].—Guinot, 1968b, fig. 

13 [morphology]. 

Cycloes cristata.—Ribeiro, 1964:4 [Cape Verde Islands]. 

Crytosoma cristatwn.—Tiirkay, 1976b:62 [Madeira, Porto 

Santo, Ilhas Desertas; erroneous spelling]. 

SYNONYM.—Cryptosoma dentatum Brulle, 1839. 

DISTRIBUTION.—Eastern Atlantic, from Madeira, 

the Canary Islands, and the Cape Verde 

Islands; sublittoral (Monod, 1956). 

Subfamily MATUTINAE de Haan, 1835 

Genus Matuta Weber, 1795 

Matuta Weber, 1795:92 [type-species: Cancer victor Fabricius, 

1786, a subjective junior synonym of Cancer lunaris Forskal, 

1775, by subsequent designation by Latreille, 1810:422; 

gender: feminine]. 

Matutinus MacLeay, 1838:70 [type-species: Cancer victor Fabricius, 

1786, a subjective junior synonym of Cancer lunaris 

Forskal, 1775, by monotypy; gender: masculine]. 


Matuta michaelseni Balss, 1921 

Matuta michaelseni.—Capart, 1951:45, fig. 12.—Monod, 1956: 

98, figs. 108-114.—Rossignol, 1957:77, 127 [key].—Buchanan, 

1958:20.—Longhurst, 1958:87.—Gauld, 1960: 

68.—Guinot and Ribeiro, 1962:25.—Rossignol, 1962: 

114.—Forest and Guinot, 1966:51.—Le Loeuff and Intes, 

1968, table 1. 


Other Material: Liberia: Off St. Paul River, Monrovia, 

trawl, 6 Jan 1953, G. C. Miller, \6 (W). 

Ivory Coast: Off Sassandra, 11 m, 3 Apr 1969, Guinean 

Trawling Survey, Tr 22, Sta 1, 19 (L). 

Ghana: Takoradi, 14 Aug 1961, Amegah, 16 (W). 

Congo: Beach at Pointe-Noire, beach seine, Jul 1963, A. 

Crosnier, 26, 79 (W). 


Figures: Monod, 1956, figs. 108-114. 

Male Pleopod: Monod, 1956, figs. 110-114 (Senegal, 

Togo). 

Color: "Blanc-rose uniforme" (Rossignol, 1957: 

77). 


lengths of 9 to 14 mm. 

BIOLOGY.—This species inhabits primarily 

sandy bottoms in shallow water, from the shore 

to a depth of about 30 m. Buchanan (1958) found 

it in the inshore fine sand community, in 3-8 fm 

(5-15 m) off Accra, Ghana, and Longhurst 

(1958) found it on sand and muddy sand, but 

noted that it occurred mostly on shelly sand in 5- 

30 m off Sierra Leone. Guinot and Ribeiro (1962) 

reported material collected on a beach at low 

tide. 


March, April, and September (Capart, 

1951; Monod, 1956; Guinot and Ribeiro, 1962). 

DISTRIBUTION.—West Africa, from Senegal to 

Angola, in shallow water, shore to about 30 m. 

Monod (1956) summarized earlier records and 

reported material from localities between Senegal 

and Togo. Since 1956 the species has been recorded 

from: 


1958). 

Ivory Coast: Off Sassandra, 15 m (Le Loeuff and Intes, 

1968). 05°03'N, 05°25'W, 20-25 m (Forest and Guinot, 

1966).

NUMBER 306 57 

Ghana: No specific locality, shore to 15 m (Gauld, 1960). 

Accra, 3-8 fm (5-15 m) (Buchanan, 1958). 

Congo: Pointe-Noire, beach (Rossignol, 1957). Baie de 

Pointe-Noire, beach (Rossignol, 1962). 

Angola: Bala de Benguela, beach at low tide (Guinot and 


Family LEUCOSIIDAE Samouelle, 1819 

LEUCOSIADAE Samouelle, 1819:91 [corrected to Leucosiidae 

by Miers, 1879a:671; name 374 on Official List}. 

ILIINAE Stimpson, 1871a: 155 [name 372 on Official List]. 

EBALIINAE Stimpson, 1871a: 159. 

MYRODINAE Miers, 1886:297. 

OREOPHORINAE Miers, 1886:297. 

MYROIDA Alcock, 1896:167. 

IPHICULOIDA Alcock, 1896:167. 

NURSILIOIDA Alcock, 1896:167. 

NURSIOIDA Alcock, 1896:166. 

NUCIOIDA Alcock, 1896:167. 

CRYPTOCNEMIDAE Stimpson, 1907:161. 

PHILYRINAE Rathbun, 1937:151. 

EASTERN ATLANTIC GENERA.—Nine, of which 

six, Atlantotlos, Ebalia, Ilia, Merocryptus, Philyra, and 

Pseudomyra, are represented by tropical species 

occurring off West Africa. Indo-West Pacific species 

belonging to three genera have become established 

in the eastern Mediterranean, having colonized 

that area via the Suez Canal. These genera 

are as follows: 

Ixa Leach (1815a:334). Type-species: Cancer cylindricus 

Fabricius, 1777, by monotypy; gender: 

feminine; name 161 on Official List. 

Leucosia Weber (1795:92). Type-species: Cancer 

craniolaris Linnaeus, 1758, by subsequent designation 

by Holthuis, 1959a: 106; gender: feminine; 

name 1631 on Official List. 

Myra Leach (1817:19, 23). Type-species: Leucosia 

fugax Fabricius, 1798, by monotypy; gender: 


EASTERN ATLANTIC SPECIES.—Twenty, of which 

12 occur in tropical waters. The extralimital species 

are as follows. 

Ebalia deshayesi Lucas, 1846. Canary Islands, 

Madeira (Tiirkay, 1976b), and Mediterranean; 

sublittoral, to 100 m (Zariquiey Alvarez, 1968). 

Ebalia edwardsii Costa, 1838. Mediterranean; 

littoral and sublittoral, to about 200 m (Zariquiey 

Alvarez, 1968). 

Ebalia granulosa H. Milne Edwards, 1837. Eastern 

Atlantic, from England to the Mediterranean; 

littoral and sublittoral, to 445 m (Zariquiey Alvarez, 

1968). 

Ebalia tumefacta (Montagu, 1808). Norway and 

Shetland Islands to Spanish Sahara; sublittoral, 

to 130 m (Zariquiey Alvarez, 1968; Christiansen, 

1969). 

Ixa monodi Holthuis and Gottlieb, 1956. An 

Indo-West Pacific immigrant into the eastern 

Mediterranean, Turkey; Red Sea (Holthuis and 

Gottlieb, 1956, 1958). 

Leucosia signata Paulson, 1875. An Indo-West 

Pacific immigrant into the eastern Mediterranean; 

Israel and Egypt (Holthuis and Gottlieb, 

1958; Lewinsohn and Holthuis, 1964; Ramadan 

and Dowidar, 1976). 

Merocryptus boletifer A. Milne Edwards and Bouvier, 

1894. Azores, Seine Seamount, Mediterranean; 

sublittoral, 100-600 m (Zariquiey Alvarez, 

1968). 

Myra fugax (Fabricius, 1798). An Indo-West 

Pacific immigrant into the eastern Mediterranean; 

Egypt, Israel, and possibly Turkey (Holthuis 

and Gottlieb, 1958; Ramadan and Dowidar, 

1976). 

The status of Ebalia fragifera Miers (1881a:268) 

from the Canary Islands, which Monod (1956: 

131) considered to be distinct from E. tuberculata, 

remains to be determined. 

The names used by Monod (1956) have not 

been changed, so the species therein are not listed 

here. 

REMARKS.—Glaessner's (1969:R496) observation 

that "this family is commonly divided into 

subfamilies which are constituted and defined 

differently by different authors (Miers, Alcock, 

Ihle, Rathbun, Balss) and are not considered 

helpful to paleontologists at the present stage of 

our knowledge of the family" is still quite pertinent. 

Of the four subfamilies currently recognized 

within the Leucosiidae, two, Ebaliinae and Cryptocneminae, 

do not offer too many difficulties 

and most authors agree in the limits of the two. 

The confusion that exists concerning the size and

58 

composition of the two other subfamilies, however, 

is very great. 

The root of all the difficulties in the subfamily 

classification of the Leucosiidae is to be found in 

a nomenclatural mix-up concerning the type genus 

of the family, Leucosia. Until 1897 the name 

Leucosia Fabricius, 1798, was used for a genus of 

Indo-West Pacific crabs, the type of which is 

Cancer craniolaris Linnaeus, 1758. In 1897(b), however, 

Rathbun showed this usage to be incorrect 

as the first type selection for Leucosia Fabricius, 

1798, is that by Latreille (1810:97, 422), who 

selected Cancer nucleus Linnaeus, 1758, as the type 

of the genus. As Cancer nucleus Linnaeus is also the 

type-species of the genus Ilia Leach, 1817, the 

name Leucosia Fabricius, 1798, had to be substituted 

for Ilia Leach, a name until then generally 

accepted by carcinologists. For the genus Leucosia 

auctt. (non Fabricius, 1798), Rathbun (1897b: 

160) proposed the new name Leucosides. There 

was considerable opposition to Rathbun's action, 

although this action was nomenclaturally entirely 

justified; most non-American authors ignored the 

changes proposed. In 1959 Holthuis (1959a: 106) 

showed that Leucosia Fabricius, 1798, is a junior 

homonym of Leucosia Weber, 1795, and thus is 

invalid. For Leucosia Weber, 1795, Holthuis selected 

as type-species Cancer craniolaris Linnaeus, 

1758, thereby making this name a senior synonym 

of Leucosides Rathbun, 1897, and Leucosia auctt. 

(non Fabricius, 1798). In this way the former, 

pre-1897, usage of Leucosia was restored, albeit 

with a different author's name. In 1964 the situation 

was further consolidated by the International 

Commission on Zoological Nomenclature 

(1964, in Opinion 712, Bulletin of Zoological Nomenclature, 

21(4): 336-351), who placed both 

names Leucosia Weber, 1795, and Ilia Leach, 1817, 

on the Official List of Generic Names in Zoology. 

At the generic level Rathbun's action had 

caused hardly any disturbance because (1) non- 

American authors had, incorrectly, ignored her 

changes, and (2) the genera Ilia Leach, 1817, and 

Leucosia Weber, 1795, do not occur in American 

waters, the former being restricted to the eastern 

Atlantic, the latter to the Indo-West Pacific (ex- 


cept for L. signata which has entered the Mediterranean 

via the Suez Canal). 

Rathbun's (1937:194) record of "Leucosia planata 

(Fabricius)" from Tierra del Fuego rests upon 

an uncharacteristic mistake on her part. Fabricius' 

species is not a leucosiid but is Halicarcinus 

planatus (Fabricius), a species of Hymenosomatidae, 

already dealt with by Rathbun (1925:563) 

in an earlier work. 

At the subfamily level Rathbun's (1897b) 

transfer of the name Leucosia from one genus 

(Leucosia Weber, 1795) to another (Ilia Leach, 

1817) caused much confusion as these two genera 

were generally considered to form part of two 

different subfamilies. These subfamilies were indicated 

by Rathbun (1937:122, 151, and 123, 

183) with the names Philyrinae and Leucosiinae, 

respectively, while most other authors used the 

names Leucosiinae and Iliinae, respectively, for 

them. The fact that the name Leucosiinae could 

stand for two entirely different groups was not 

realized by several authors. 

A year before Rathbun (1897b) published her 

controversial paper, Alcock (1896) gave an important 

classification of the Leucosiidae, which 

he divided into two subfamilies: Leucosiinae 

(containing 5 subdivisions, named by him Alliances 

i-v) and Iliinae (with Alliances I-IV). 

Ihle (1918) recognized three subfamilies within 

the Leucosiidae: Ebaliinae (with 12 genera, most 

of which were placed in Alcock's Alliances I-III of 

his Leucosiinae), Iliinae (with 19 genera including 

all of those placed by Alcock in his Iliinae, those 

of his Alliance iv of Leucosiinae and part of those 

of Alliances n and m of the latter subfamily), and 

Leucosiinae (with 7 genera including all those of 

Alcock's Alliance v of the Leucosiinae and the 

genera Cryptocnemus and Carcinaspis which Alcock 

had placed (the latter with some doubt) in Alliance 

i). 

Rathbun (1937) followed Ihle in recognizing 

three subfamilies of Leucosiidae, and in his delimitation 

of the subfamily Ebaliinae. Rathbun's 

Philyrinae contained all the genera that Ihle 

assigned to the Leucosiinae plus several that he 

considered to belong to the Iliinae (viz., all those

NUMBER 306 59 

of Alcock's Leucosiinae Alliance iv, plus Randallia 

of Alliance m). Rathbun's Leucosiinae consisted 

of the other Iliinae sensu Ihle. As Rathbun dealt 

only with the American representatives, it is difficult 

to know the exact limits of her subfamilies. 

Balss (1957:1612-1615), in his authoritative 

treatment of the Decapoda in Bronn's Klassen und 

Ordnungen des Tierreichs, which of necessity is 

mostly based on study of the literature, evidently 

became severely confused. He recognized, like 

Rathbun, the two subfamilies Philyrinae and 

Leucosiinae, but unlike Rathbun used the generic 

names Leucosia and Ilia for the genera that she 

named Leucosides and Leucosia, respectively. In the 

subfamily Philyrinae Balss placed all the genera 

that Ihle had assigned to the Iliinae with the 

exception of two, viz., Calhdactylus and Iliacantha, 

the only two that Rathbun (1937) mentioned by 

name and which she placed in the Leucosiinae. 

Balss' Leucosiinae contained all genera placed by 

Ihle in that subfamily with the exception of Philyra 

(which Rathbun, 1937, had placed in the 

Philyrinae) and Pseudophilyra. Balss' definition of 

the Philyrinae is based almost exclusively on that 

by Rathbun (1937), but his definition of the 

Leucosiinae has the first five sentences adapted 

from Ihle's (1918:207) account of the Leucosiinae 

(sometimes in the identical wording), while the 

last sentence (dealing with the third maxilliped) 

is taken from Rathbun's (1937:183) definition of 

her Leucosiinae. Balss' action was taken in spite 

of the fact that Rathbun (1937:151, 183) made 

clear that her subfamily Philyrinae corresponds 

with all of Ihle's Leucosiinae plus part of his 

Iliinae, while her Leucosiinae are formed by the 

rest of Ihle's Iliinae. Unfortunately Balss's (1957: 

1612-1615) garbled version of the classification 

of the Leucosiinae provides the most recent complete 

list of all the genera of the family and has 

been accepted as the basis for the work of several 

subsequent authors (Guinot, 1967a:246-251; Serene, 

1968:41-48; Sakai, 1976:76-126). Serene 

(1965:11-17) separated the subfamily Cryptocneminae 

from the Leucosiinae, placing the genera 

Cryptocnemus, Leucisca (= Carcinaspis), Onychomorpha 

and Lissomorpha in the former subfamily, 

leaving only Leucosia in the latter (probably Callidactylus 

and Iliacantha are also left in the Leucosiinae 

by Serene, but as these genera do not occur 

in the Indo-West Pacific region they are not 

mentioned by him). For the time being we follow 

the system of Ihle (1918), which seems to be more 

logical than the one by Rathbun (1937) or than 

the garbled version of the latter produced by 

Balss (1957). A closer comparative study of the 

various genera of Leucosiidae is, however, badly 

needed. 

Subfamily EBALIINAE Stimpson, 1871 

Genus Atlantotlos Doflein, 1904 

Atlantotlos Doflein, 1904:49 [type-species: Atlantotlos rhombifer 

Doflein, 1904, by monotypy; gender: masculine]. 

* Atlantotlos rhombifer Doflein, 1904 

Atlantotlos rhombifer.—Capart, 1951:57, fig. 16, pi. 2: fig. 1.— 

Monod, 1956:134, figs. 162, 163.—Longhurst, 1958:87.— 

Gauld, 1960:69. 

Atlantotlus rhombifer.—Rossignol, 1962:114 [erroneous spelling]- 

Atlantotlos.—Voss, 1966:35. 

Merocryptus rhombifer.—Forest and Guinot, 1966:54. 


Sta 68, 70 m, broken shell, 56, 4$ (W). 


Foraminifera, let (L). Sta 63, 64 m, sandy mud with shells, 

1$ (L). 

Nigeria: Sta 239, 73 m, 1$ (L). Sta 241, 59-63 m, mud 

and shell, Id, 1$ ov (L). Sta 248, 33 m, 29 (W). 

Cameroon: Sta 260, 46 m, 3d, 19, 1 juv (L). 

Other Material: Ghana: 05°13'42"N, 03°59'48"W, 50-55 

m, dredge, Guinean Trawling Survey, La Rafale, Tr 12, Id 

(W). 


Figures: Capart, 1951, fig. 16, pi. 2: fig. 1. 

Male Pleopod: Capart, 1951, pi. 2: fig. 1 (Gabon); 

Monod, 1956, fig. 163 (Senegal). 

Color: According to Capart (1951:58), in this 

species "Les males et les femelles sont orange rose, 

mais les sillons entre les saillies sont bruns avec 

des tubercules presque blancs." 

MEASUREMENTS.—Our specimens have cara-

60 

pace widths of 7 to 16 mm. The carapace width 

of the ovigerous female is 15 mm. 

REMARKS.—Forest and Guinot (1966) synonymized 

Atlantotlos Doflein, 1904 with Merocryptus A. 

Milne Edwards, 1873, noting Monod's (1956: 

136) comment: "Je ne suis pas certain du tout 

que le genre Atlantotlos soit distinct de Merocryptus." 

However, the male pleopod of A. rhombifer, 

figured in Monod, differs from that of two species 

of Merocryptus figured in Guinot and Ribeiro 

(1962, fig. 5a, b (M obsoletus), and fig. 6 (M. 

boletifer)), in having a subterminal bulbous enlargement 

rather than tapering evenly to the 

apex. Until all the representatives of these genera 

can be studied in more detail, we prefer to retain 

Atlantotlos for Doflein's species. 

BIOLOGY.—This species occurs in depths between 

44 and 115m, but most specimens were 

taken in depths between 70 and 100 m. The 

Pillsbury specimens were collected on mud with 

brown, branched Foraminifera in 62-75 m, sandy 

mud with shells in 64 m, and on mud and shell 

in 59-63 m. The material reported by Capart 

(1951) was taken on brown muddy sand and 

coral, brown sandy mud and sandy brown mud, 

and on muddy sand or muddy sand and rock in 

85 to 100 m. Longhurst (1958) found it on shelly 

mud in 106 m, and Rossignol (1962:114) found 

it in 115 m, "en bordure des affleurements rocheux." 

Forest and Guinot (1966) found the species 

on mud, rocks, calcareous algae, sand and 

Foraminifera in 51-55 m and on mud and shell 

in 73-80 m. 

Ovigerous females have been recorded in February, 

May, September, and December (Capart, 



recorded from a few localities between Senegal 

and Angola, in depths between 44 and 115 m. 


recorded the species from Senegal. Since 1956 it 


Senegal: 13°01'N, 17°24'W, 51-55 m (Forest and Guinot, 

1966). 

Liberia: 05°21.5'N, 09°54.5'W, 73-80 m (Forest and 



Sierra Leone: No specific locality, 106 m (Longhurst, 

1958). 

Ghana: Off Accra, 80 m (Gauld, 1960). 

Nigeria: 04°56'N, 05°00'E, to 04°54'N, 05°05'E, 73 m 

(Voss, 1966). 

Congo: W of Pointe-Noire, 115 m (Rossignol, 1962). 

Genus Ebalia Leach, 1817 

Ebalta Leach, 1817, in 1815-1875, pi. 25 [type-species: Ebalia 

bryerii Leach, 1817, a subjective junior synonym of Cancer 

tumefactus Montagu, 1808, by subsequent designation by 

H. Milne Edwards, 1837, in 1836-1844, pi. 24: fig. 3 (as 

Ebalia brayerii); gender: feminine; name 145 on Official 

List, type-species there given as Cancer tuberosus Pennant, 

1777, in error]. 

Phlyxia Bell, 1855:303 [type-species: Phlyxia crassipes Bell, 

1855, by selection by Bell, 1855:304; gender: feminine]. 

Bellidilia Kinahan, 1856:115, 117, 128 [type-species: Bellidilia 

undecimspinosa Kinahan, 1856, by present designation; gender: 

feminine]. 

* Ebalia affinis Miers, 1881 

Ebalia atlantica.—Capart, 1951:54. 

Ebalia affinis.—Monod, 1956:117, figs. 134-144.—Longhurst, 

1958:87.—Gauld, 1960:69.—Forest and Guinot, 1966:53. 

SYNONYM.—Ebalia atlantica A. Milne Edwards 

and Bouvier, 1898. 


Sta 68, 70 m, broken shell, 3c? (W). 

Ivory Coast: Sta 47, 37 m, bottom with Jullienella, 3c5, 1$ 

(L). 

Ghana: Sta 17, 48 m, fine sand and green mud, Ic5 (W). 

Sta 23, 42 m, foliate brown to orange bryozoans, 1

NUMBER 306 61 

BIOLOGY.—This species has been recorded from 

depths between 4 and 140 m, but more than 85% 

of the records are between 4 and 45 m. Capart 

(1951) reported it on brown muddy sand in 100 

m. Longhurst (1958) found it on muddy sand, 

shelly sand, and shelly mud in depths between 8 

and 140 m. Forest and Guinot (1966) reported it 

from mud or shelly mud in 18-30 m, mud, calcareous 

algae, and shell in 31 m, rocks and coral 

in 3-10 m, sand, algae, and calcareous algae in 

8-30 m, and mud and calcareous algae in 4-5 m. 

The Pillsbury specimens were taken on fine sand 

and green mud in 48 m, on bottom with bryozoans 

in 35-37 and 42 m, on bottom with Jullienella 

in 37 m, and on broken shell in 70 m. 

Apparently the species prefers soft bottom mixed 

with shell or other hard substances. 


February, March, April, May, and June 

(Monod, 1956; Forest and Guinot, 1966). 

DISTRIBUTION.—Off West Africa, from localities 

between Senegal and Angola; it also has been 

reported from the Seine Seamount north of Madeira. 

Monod (1956) summarized earlier records 

and reported material from Senegal, Guinea, 

Sierra Leone, and Ghana. Since 1956 the species 

has been reported from the following localities. 

Guinea: 09°40'N, 14°05'W, 18 m, andO9°36'N, 13°57'W, 

18-30 m (Forest and Guinot, 1966). 


1958). 

Ghana: Accra, 32 m (Gauld, 1960). 

Principe: O1°37'N, 07°22'E, 30 m, and 01°38'25"N, 


Sao Tome: Praia de Santa Catarina, W coast, 3-10 m; 

00°25'15"N, 06°43'05"E, 8-30 m; Baia de Ana de Chaves, 

5 m; and in front of Ponta Diogo Nunes, 4 m (Forest and 


Ebalia cranchii Leach, 1817 

Ebalia cranchi. —Monod, 1956:122, figs. 145, 146 [Senegal; 

references].—Zariquiey Alvarez, 1968:329, figs. 108a,b, 

lllb,d, lllAa, lllCb [Spain; references].—Tiirkay, 

1976a:25 [listed], 37, fig. 19 [Morocco]. 

Ebalia cranchii.—Christiansen, 1969:31, fig. 11, map 5 [Scandinavia]. 

DISTRIBUTION.—Eastern Atlantic, from Norway 

to Senegal, Mediterranean, in depths between 20 

and 550 m (Zariquiey Alvarez, 1968). 

Ebalia nux A. Milne Edwards, 1883 

Ebalia nux A. Milne Edwards, 1883, pi. 5.—Monod, 1956: 

121 [references].—Zariquiey Alvarez, 1968:328, figs. 108c, 

111 Ab [Spain; references].—Turkay, 1976a:25 [listed], 37, 

fig. 18 [Morocco]. 




150, 1

62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

SYNONYM.—Lithadia barnardi Stebbing, 1920. 

MATERIAL EXAMINED.—Pillsbuty Material: Ghana: 

Sta 22, 51 m, rough bottom, 29 (L). Sta 23, 42 m, foliate 

brown to orange bryozoans, Ic5 (L). Sta 24, 35-37 m, dark 

red bryozoans, 7

NUMBER 306 63 

Burger (1902, 1904) described a gynandromorph 

specimen of Jasus frontalis (H. Milne Edwards), 

and Hay (1905) mentioned the case of such a 

specimen in the freshwater crayfish Orconectes propinquus 

(Girard). According to Balss (1944:634, 

635), apart from a case in Lucifer, no gynandromorphs 

have been reported in Decapoda other 

than the Macrura Reptantia. However, a gynandromorph 

of the Mediterranean crab, Brachynotus 

gemmellari (Rizza), was reported by Froglia 

and Manning (1978:700, fig. 5). In Homarus and 

Jasus the body of the gynandromorph usually is 

skewed because of the different longitudinal proportions 

of males and females. 

Nothing irregular was found in the shape of 

the carapace or the thoracic appendages of this 

specimen of E. tuberculata, which in this case may 

be an indication that in this species the sexes are 

not different in size. 

Both Capart (1951:56) and Guinot (1967a:247, 

footnote) suggest that material from South Africa 

referred to this species by Barnard (1950) probably 

belongs to a distinct species. Barnard (1955: 

4) proposed Ebalia pondoensis for South African 

specimens previously identified with E. tuberculata. 

COMMENSALS AND PARASITES.—The carapace of 

the gynandromorph is covered dorsally and ventrally 

by Bryozoa, but the appendages, sternum, 

and abdomen are entirely free from this growth. 

In the space enclosed by the abdomen and the 

thoracic sternum a number of sausage-shaped 

organisms, resembling Rhizocephala of the genus 

Thompsonia, can be seen. One such organism is 

fastened to the male half of the thoracic sternite 

of the fifth pereiopod, whereas on the female half 

of the abdomen seven similar organisms are present, 

two attached to a pleopod, the other five to 

the abdominal sternites. 

BIOLOGY.—The depths at which this species 

has been taken range from 10-12 m to 135-250 

m. Of the known depth records, more than 85% 

are from depths of less than 75 m, the optimal 

depth apparently being between 30 and 50 m. 

Little is known of the habitat preferences of the 

species. The Pillsbury specimens were taken on 

rough bottom in 51 m, on bottom with bryozoans 

in 35-37 and 42 m, and on bottom with nodular 

coralline algae in 51-55 m. The species was reported 

from mud, sand, and compacted sand 

[sable construit] in 65-75 m and on mud with 

Area in 32 m by Forest and Guinot (1966). 

Apparently the species spawns all year. Ovigerous 

females have been recorded in January, 

March, April, May, and September through December 

(Monod, 1956; Pillsbury). 

DISTRIBUTION.—Off West Africa, where it occurs 

from localities between Morocco and Angola 

and possibly South Africa. Monod (1956), who 

recorded material from Morocco, the Cape Verde 

Islands, Senegal, Ghana, and Angola, summarized 

the earlier literature. Since 1956 this species 


Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest and 


Ghana: Accra, 37-51 m (Gauld, 1960). 

Nigeria: Off the mouths of the Niger River, 04°03'N, 

06°12'E, 32 m (Forest and Guinot, 1966). 

This species has not been recorded previously 

from Annobon. 

Ebalia tuberosa (Pennant, 1777) 

Ebalia tuberosa.—Lebour, 1954:236.—Monod, 1956:124, figs. 

150, 151 [England, Algeria, Bonifacio, Ilhas Desertas, 

Canary Islands, Mauritania; references].—Zariquiey Alvarez, 

1968:326, figs. 109d, HOa-c, Ilia [Spain; references].—Christiansen, 

1969:27, fig. 9, map 3 [Scandinavia].— 

Tiirkay, 1976a:25 [listed], 37, fig. 20 [Morocco]; 

1976b:61 [listed], 63 [Madeira, Ilhas Desertas]. 

SYNONYMS.—Ebalia pennantii Leach, 1817; Ebalia 

insignis Lucas, 1849; PEbalia madeirensis Stimpson, 

1858. 

REMARKS.—This species is included in this list 

of tropical species on the basis of the single juvenile 

female reported by Monod (1956:1Z6) from 

the Bane d'Arguin, Mauritania, 21°51'N, 

19° 48'W. Other specimens from the same station 

off the Bane d'Arguin were referred to E. tuberculata 

by A. Milne Edwards and Bouvier (1900:50). 

Its occurrence northward, off Spanish Sahara 

and Morocco, is not questioned. Christiansen 

(1969) did not accept Monod's record, noting

64 

that the species occurred southward to Spanish 

Sahara. Lebour (1954:236) reported on some larval 

and postlarval specimens of Ebaha from the 

Benguela Current that she believed might be 

identified with this species. 


and the Hebrides southward to Mauritania (?), 

including the Azores and the Canary Islands, 

Mediterranean; sublittoral to about 138 m (Zariquiey 

Alvarez, 1968; Christiansen, 1969). 

Genus Merocryptus A. Milne Edwards, 1873 

Merocryptus A. Milne Edwards, 1873a: 78 [type-species: Merocryptus 

lambriformis A. Milne Edwards, 1873, by monotypy; 

gender: masculine; name 166 on Official List]. 

Merocryptus obsoletus A. Milne Edwards and 

Bouvier, 1898 

Merocryptus obsoletus.— Monod, 1956:132, figs. 161, 161 bis 

[Morocco; Senegal].—Guinot and Ribeiro, 1962:28, figs. 

5a, b, pi. 1: figs. 1, 2, 4 [Cape Verde Islands, Senegal, 

Angola]. 

DISTRIBUTION.—Eastern Atlantic, from Morocco, 

the Cape Verde Islands, Senegal, and Angola, 

in depths between 75 and 132 m. 

Subfamily ILIINAE Stimpson, 1871 

Genus Ilia Leach, 1817 

Leucosia Fabricius, 1798:313, 349 [an invalid junior homonym 

of Leucosia Weber, 1795 (Decapoda); type-species: 

Cancer nucleus Linnaeus, 1758, by subsequent designation 

by Latreille, 1810:97, 422; gender: feminine; name 1732 

on Official Index]. 

Ilia Leach, 1817:19, 24 [type-species: Cancer nucleus Linnaeus, 

1758, by monotypy; gender: feminine; name 1628 on 

Official List]. 

Thaumasta Gistel, 1848:ix [substitute name for Leucosia Fabricius, 

1798; type-species: Cancer nucleus Linnaeus, 1758; 


Ilia nucleus (Linnaeus, 1758) 

Ilia nucleus.—Monod, 1956:139 [references].—Guinot and 

Ribeiro, 1962:30 [Cape Verde Islands].—Zariquiey Alvarez, 

1968:322, figs. 1 lc, 94f [Spain; references]. 


PIlia nucleus spinosa.— Tiirkay, 1975a:71 [listed], 72 [Spanish 

Sahara]. [Not Ilia spinosa Miers, 1881.] 

SYNONYMS.—Cancer orbicularis Olivi, 1792; Leucosia 

leachii Risso, 1822; Ilia laevigala Risso, 1827; 

Ilia rugulosa Risso, 1827; Ilia parvicauda Costa, 

1853. 

REMARKS.—Tiirkay's (1975a) record of Ilia 

from off Cabo Blanco, Spanish Sahara, may be 

referable to this species rather than to /. spinosa. 

Monod (1933b) recorded /. nucleus from the same 

locality, and this apparently is the southernmost 

limit of the species along the African mainland. 

Ilia spinosa is not known with certainty to occur 

north of Mauritania. 


Cape Verde Islands (where it occurs together with 

/. spinosa), Spanish Sahara, and from the Mediterranean, 

in depths between 4 and 162 m (Zariquiey 


* Ilia spinosa Miers, 1881 

PIlia nucleus.—Bouvier, 1911:226 [not Ilia nucleus (Linnaeus, 

1758)]. 

Leucosia spinosa.—Capart, 1951:52, fig. 15. 

Ilia spinosa.—Monod, 1956:136, 632, figs. 164-166.—Buchanan, 

1958:28, 54.—Longhurst, 1958:87.—Lebour, 

1959:133, 135, 136, 137, fig. 21 [larvae].—Gauld, 1960: 

69.—Rossignol, 1962:114.—Guinot and Ribeiro, 1962:30, 

pi. 4: fig. 1.—Crosnier, 1964:38.—Ribeiro, 1964:4.—Forest 

and Guinot, 1966:55.—Crosnier, 1967:323.—LeLoeuff 

andlntes, 1968:31, table 1. 


Sta 68, 70 m, broken shell, 1 juv (L). 


Foraminifera, 1(5, 1 juv (W). Sta 46, 38-42 m, mud bottom 

with dense Julltenella, 26, 19 (W). Sta 48, 22 m, 1(5 (W). Sta 

62, 46 m, brown, branching and foliate Foraminifera, 3

NUMBER 306 65 

mud, stones, calcareous algae, sand, Foraminifera, 15 May 

1956, Calypso Sta 3, 1$ ov (W). 

Cameroon: 02°39'N, 09°40'E, 50-65 m, mud, 22 Aug 

1963, A. Crosnier, 29 (1 ov) (W). 



Male Pleopod: Monod, 1956, figs. 165, 166 (Senegal). 

Color: Capart (1951:53) noted that in this species 

"Le carapace est orange clair, les epines et 

appendices presque blancs." 


lengths of 5 to 17 mm; the carapace lengths 


REMARKS.—In none of our specimens are the 

posterolateral spines of the carapace so well developed 

as shown for a specimen from Angola by 

Guinot and Ribeiro (1962, pi. 4: fig. 1). 

It seems likely that Tiirkay's (1975a) record of 

Ilia nucleus spinosa from Spanish Sahara is based 

on material of/, nucleus (Linnaeus) rather than /. 

spinosa. For that reason we have placed it in the 

synonymy of the latter species. 

Guinot and Ribeiro (1962) noted that one of 

their specimens from the Cape Verde Islands was 

similar to /. nucleus in having relatively short 

posterolateral spines on the carapace. Apparently 

both species occur in the Cape Verde Islands. 

Bouvier (1911:226) reported /. nucleus from Baie 

de l'Ouest, Mauritania, and noted "avec passage 

a la spinosa, Miers." 

BIOLOGY.—This species lives in relatively shallow 

water, in depths between 6 and 132 m. Of 49 

depth records in Monod (1956), 1 was at 132 m, 

1 at 96 m, 1 at 75 m, and 46 were at 50 m or less. 

Apparently, like many of the leucosiids, it prefers 

soft bottoms with harder (larger?) substances in 

it. Sourie (1954b) found it on coarse shelly sand, 

bottom with Area and Pyura, in 10-12 m in the 

Baie de Dakar, and Buchanan (1958) took it in 

the silty sand community with Jullienella off 

Ghana. Crosnier (1964) reported that it was 

found on mud or sandy mud in cold water off 

Cameroon. Forest and Guinot (1966) reported it 

from a variety of substrates in depths between 18 

and 90-100 m. Most of the Pillsbury specimens 

were taken on bottom with shell, bryozoans, or 

Foraminifera. 


June, July, August, October, November, and December 

(Capart, 1951; Monod, 1956; Forest and 

Guinot, 1966; Crosnier, 1967; Pillsbury). 

DISTRIBUTION.—Off West Africa, from the Canary 

Islands, the Cape Verde Islands, and Mauritania 

southward to Angola, in depths between 4 

and 132 m, generally in less than 50 m. Monod 

(1956) reported material from Mauritania, Senegal, 

Guinea, Sierra Leone, Ghana, and Principe. 

In addition, the species has been recorded from 

the following localities. 

Cape Verde Islands: Bafa de Porto Grande, Sao Vicente, 

8 m (Guinot and Ribeiro, 1962; Ribeiro, 1964). 

Mauritania: Baie de l'Ouest (Bouvier, 1911). 

Senegal: 13°01'N, 17°24'W, 51-55 m, and 12°55.5'N, 


Guinea-Bissau: 10°19'N, 16°34'W, 18 m (Forest and 


Guinea: 09°36'N, 13°57'W, 18-30 m (Forest and Guinot, 

1966). 


1958). 

Ivory Coast: Off Jacqueville and Grand-Bassam, 40- 

100 m (Le Loeuff and Intes, 1968). 

Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m; 

04°36.5'N, 01°31'W, 50 m; 04°37'N, 00°50'W, 90-100 m 

(all Forest and Guinot, 1966). Off Accra (Buchanan, 1958); 

in 20-55 m (Gauld, 1960). 


Principe: 01°38'25"N, 07°22'05"E, 31 m (Forest and 


Annobon: 01°25'30"S, 05°39'E, 52 m (Crosnier, 1967). 

Gabon: 00°40'S, 08°46'25"E, 18 m, and 00°38'20"S, 

08°48'30"E, 35 m (Forest and Guinot, 1966).—W of Mayumba, 

20 m (Rossignol, 1962). 

Congo: W of Pointe-Noire, 20-30 m (Rossignol, 1962). 

Angola: Baia Farta, Benguela, 40 m (Guinot and Ribeiro, 

1962). 

Lebour (1959) recorded larvae of this species 

from the following localities: Guinea, 10°22'N, 

16°34'W; Sierra Leone, 08°22'N, 14°08'W; 

Ghana, 05°44'N, 01°02'E; Cameroon, 04°01'N, 

07°23'E; and Angola, 07°35'S, 12°46'E. 

Subfamily LEUCOSIINAE Samouelle, 1819 

Genus Philyra Leach, 1817 

Philyra Leach, 1817:18, 22 [type-species: Leucosia globulosus

66 

Bosc, 1802, a subjective junior synonym of Cancer globus 

Fabricius, 1775, by subsequent designation by H. Milne 

Edwards, 1837, in 1836-1844, pi. 24: fig. 4 (as Philyra 

globulosa); gender: feminine; name 1642 on Official List.] 

Philyra cristata Miers, 1881 

Philyra cristata.—Monod, 1956:144, figs. 177-183 [Senegal, 

Guinea, Sierra Leone; references].—Longhurst, 1958:87 

[Sierra Leone].—Rossignol, 1962:115 [Congo].—Forest 

and Guinot, 1966:56 [Guinea; Sao Tome]. 

DISTRIBUTION.—West Africa, from Senegal to 

the Congo, in depths between 4 and 25 m. 

* Philyra lacvidorsalis Miers, 1881 

Philyra laevidorsalis.—Capart, 1951:47, fig. 13.—Monod, 

1956:141, figs. 169-176.—Rossignol, 1957:77.—Longhurst, 

1958:87.—Buchanan, 1958:20.—Gauld, 1960: 

69.—Rossignol, 1962:115.—Forest and Guinot, 1966:56. 


Coast: Sta 47, 37 m, bottom with Jullienella, \6 (W). 

Other Material: Liberia: Off St. Paul River, Monrovia, 

22-29 m, trawl, 4 Nov 1953, G. C. Miller, \6 (W). 

Ivory Coast: Off Sassandra, 11 m, 3 Apr 1964, Guinean 

Trawling Survey, Tr 22, Sta 1, 46\ 69 (L). 


Figures. Monod, 1956, figs. 169-176. 

Male Pleopod: Monod, 1956, figs. 173-176 (Senegal). 

Color: Rossignol (1957) noted that this species 

is a uniform gray beige, with the ventral surface 

lighter. Capart (1951:47) reported that this species 

has "couleur uniforme gris brunatre, plus 

claire sur la face inferieure; telson blanc ivoire." 


lengths of 6 to 11 mm. 

BIOLOGY.—Philyra laevidorsalis is a coastal species, 

generally found in shallow water, in depths 

between 4 and 18-30 m. Longhurst (1958) found 

it in 9-86 m off Sierra Leone, but all other depth 

records in the literature are from 20-25 m or less. 

Sourie (1954b) found it on coarse shelly sand, 

bottom with Area and Pyura, in the Baie de Dakar, 

and Buchanan (1958) characterized it as a member 

of the active epifauna, inshore fine sand 

community, in 3-8 fm (5-15 m) off Ghana. Off 


Sierra Leone, Longhurst (1958) found it on shelly 

sand. Forest and Guinot (1966) reported it from 

mud in 18-30 m, shells in 20-25 m, calcareous 

algae in 10-12 m, and calcareous algae, sand and 

mud or shells in 6 to 11 m. 

Ovigerous females have been collected in April 

and May (Monod, 1956; Forest and Guinot, 

1966). 

DISTRIBUTION.—West coast of Africa, from Cap 

Blanc, Mauritania, southward to Angola, including 

the Cape Verde Islands and Principe, in 

shallow water, from a depth of 4 m to about 30 

m. Monod (1956), who summarized the earlier 

literature, reported material from Senegal and 

Ghana. Since 1956 it has been recorded from the 

following localities. 

Guinea: 09°36'N, 13°57'W, 18-30 m (Forest and Guinot, 

1966). 


1958). 

Ivory Coast: 05°03'N, 05°25'W, 20-25 m (Forest and 


Ghana: Accra, 3-8 fm (5-15 m) (Buchanan, 1958). Off 

Accra and Takoradi, 15-20 m (Gauld, 1960). 


10-12 m; in front of (Cais de) Santana, 11 m; and between 

Ponta da Mina and Ponta Novo Destino, 6 m (all Forest 


Congo: Pointe-Noire, beach (Rossignol, 1957). Baie de 

Pointe-Noire, 7-15 m (Rossignol, 1962). 

Genus Pseudomyra Capart, 1951 

Pseudomyra Capart, 1951:48 [type-species: Pseudomyra mbizi 

Capart, 1951, by original designation and monotypy; 


* Pseudomyra mbizi Capart, 1951 

Pseudomyra mbizi Capart, 1951:49, fig. 14, pi. 2: fig. 24.— 

Monod, 1956:140, figs. 167, 168.—Rossignol, 1962:115.— 

Guinot and Ribeiro, 1962:30.—Crosnier, 1964:35.—Forest 

and Guinot, 1966:56.—Voss, 1966:35.—Le Loeuff and 

Intes, 1968:40.—Maurin, 1968b:491, 492.—Crosnier, 

1970:1215, 1216. 

Pseudomyra.—Voss, 1966:33, 36. 

Pseudomyra m'bizi.—Le Loeuff and Intes, 1968, table 1 [erroneous 

spelling]; 1969:66. 

MATERIAL EXAMINED.—Pillsbury Material: Liberia:

NUMBER 306 67 

Sta 68, 70 m, broken shell, 196", 32$ (23 ov), 7 juv (W). Sta 

69, 29 m, coral or rock, lcj (W). 


Foraminifera, 174c5, 9, juv (L,W). Sta 45, 73-97 m, 96\ 3$ 

(W). Sta 49, 73-77 m, lie?, 39, 7juv (L). Sta 50, 128-192 m, 

16, 29 (W). Sta 59, 55-64 m, mud with dense, branched 

Foraminifera, 5ct, 49 (W). Sta 60, 79-82 m, coral or rock, 3

68 

the eastern Atlantic. Guinot (1976) provided a 

recent review of the family. 

Family ATELECYCLIDAE Ortmann, 1893 

ATELECYCLIDAE Ortmann, 1893a:27 [name 369 on Official 

List]. 

CHEIRAGONIDAE Ortmann, 1893b:413, 419. 

EASTERN ATLANTIC GENERA.—One, Atelecyclus, 

represented in the tropical fauna of West Africa. 

EASTERN ATLANTIC SPECIES.—Two, both occurring 

in Europe as well as off tropical West Africa. 

Monod (1956) included accounts of both species, 

as follows, but had material of only one: 


Atelecyclus septemdentatus Atelecyclus rotundatus 

Atelecyclus undecimdentatus Atelecyclus undecimdentatus 

REMARKS.—Neither species was taken by the 

Pillsbury. 

Genus Atelecyclus Leach, 1814 

Atelecyclus Leach, 1814:430 [type-species: Cancer {Hippa) septemdentatus 

Montagu, 1813, a subjective junior synonym of 

Cancer rotundatus Olivi, 1792, by monotypy; gender: masculine; 


Atelecyclus rotundatus (Olivi, 1792) 

Cancer rotundatus Olivi, 1792, pi. 2: fig. 2. 

Atelecyclus rotundatus.— Sourie, 1954b: 150.—Forest, 1958:472, 

fig. 2 [references].—Peres, 1964:20.—Forest and Guinot, 

1966:57.—Maurin, 1968a:30.—Zariquiey Alvarez, 1968: 

342, figs. Id, 112b [Spain; references].—Christiansen, 

1969:37, fig. 13, map 7 [Scandinavia].—Turkay, 1976a:25 

[listed], 37, fig. 21 [Portugal]. 

Atelecyclus septemdentatus.—Bouvier, 1911:236.—Monod, 

1956:148.—Longhurst, 1958:87. 

Atelecyclus.— Maurin, 1968a, fig. 13. 

SYNONYMS.—Cancer septemdentatus Montagu, 

1813; A telecyclus heterodon Leach, 1815. 


Gtrommo Material: Gabon: Sta 211, 100 m, 1$ (W). 

DESCRIPTION.—Christiansen, 1969:37. 

Figure: Christiansen, 1969, fig. 13. 

MEASUREMENTS.—Our specimen, a non-oviger- 


ous female, has a carapace length and a carapace 

width of 15 mm. 

REMARKS.—Our specimen agrees well with the 

accounts of this species given by Christiansen 

(1969) and Forest (1958). The latter author established 

the identity of A. rotundatus (Olivi, 1792) 

with A. septemdentatus (Montagu, 1813). 

A comparison of our young specimen with 

material of this species from the Mediterranean 

shows that the West African specimen has a 

considerably rougher carapace; no other differences 

were observed. 

BIOLOGY.—Little is known about the habitat 

requirements of this species, which, as Forest 

(1958) noted, lives from shallow water to a depth 

of 200-300 m; Forest questioned the validity of 

some early records from very deep water. Off 

Scandinavia, Christiansen (1969) found it on 

sand and soft bottom, often with gravel and small 

stones, in depths between 15-40 and 190-324 m. 

Longhurst (1958) collected it on shelly sand in 

121-160 m. Peres (1964) recorded it from reddish 

gravel with shell debris in 210 m off the Bane de 

Spartel, and Maurin (1968a) reported it from 

muddy sand and sandy detritus in 10-30 m. It 

was collected on mud, rocks, calcareous algae, 

sand, and Foraminifera by the Calypso (Forest 


DISTRIBUTION.—Eastern Atlantic, from Scandinavia 

and the Hebrides southward to South 

Africa, including the Mediterranean, and, off the 

African Coast, a few localities between Morocco 

and Gabon, as well as the Cape Verde Islands; 

littoral to about 300 m. Monod (1956) summarized 

the literature but recorded no material of 

this species. In addition, the species has been 

recorded from the following localities. 

Morocco: Bane de Spartel, 35°54'N, 06°14'W, 210 m 

(Peres, 1964). Between Cap Rhir (as Cap Ghir) and Cap 

Draa (as Cap Noun), 10-30 m (Maurin, 1968a). 

Mauritania: Baie de l'Ouest (Bouvier, 1911). 

Senegal: Baie de Dakar, 0-17 m (Sourie, 1954b). 13°01'N, 



1958). 

Although Gabon is well within the known

NUMBER 306 69 

range of the species, it is the southernmost locality 

on the tropical mainland. 

Atelecyclus undecimdentatus (Herbst, 1783) 

Atelecyclus cruentatus.—Capart, 1951:136, pi. 2: fig. 7 [Spanish 

Sahara]. 

Atelecyclus undecimdentatus.—Monod, 1956:148, figs. 184-186 

[Mauritania, Senegal, Gambia, Gabon; references].—Forest, 

1958:472, fig. 1 [references].—Longhurst, 1958:87 

[Sierra Leone].—Forest and Gantes, 1960:350 [Morocco]. 

—Zariquiey Alvarez, 1968:342, fig. 112d [Spain; references].—Maurin, 

1968b:486, 489, fig. 4 [Mauritania].— 

Bas, Arias, and Guerra, 1976, table 3 [Spanish Sahara]. 

SYNONYMS.—Atelecyclus cruentatus Desmarest, 

1825; Atelecyclus omiodon Risso, 1827. 

DISTRIBUTION.—Eastern Atlantic, from France 

to Gabon, Mediterranean, from shore to about 

30 m. 

Family THIIDAE Dana, 1852 

THIIDAE Dana, 1852a: 120 [name 361 on Official List, dated 

1862 in error]. 

EASTERN ATLANTIC GENERA.—One, Thia, represented 

in the fauna of tropical West Africa. 

EASTERN ATLANTIC SPECIES.—One (possibly 

two), occurring off tropical West Africa. The 

species reported as Thia residua by Monod (1956) 

is now known as Thia scutellata. 

REMARKS.—This family was not represented in 

the Pillsbury collections. 

Genus Thia Leach, 1815 

Thia Leach, 1815a:312 [type-species: Thiapolila Leach, 1815, 

a subjective junior synonym of Hippa scutellatus Fabricius, 



Thia scutellata (Fabricius, 1793) 

Hippa scutellata Fabricius, 1793:474. 

Thia residua.—Monod, 1956:153, figs. 186 bis, 186 ter [Sierra 

Leone; references].—Longhurst, 1958:87 [Sierra Leone]. 

Thia scutellata.—Zariquiey Alvarez, 1968:343, fig. 11 If 

[Spain; references].—Christiansen, 1969:40, fig. 14, map 

8 [Scandinavia]. 

Thia sp. aff. residua.—Forest and Guinot, 1966:57, fig. 3 [Sao 

Tome]. 

SYNONYMS.—Cancer residuus Herbst, 1799; Thia 

polita Leach, 1815; Thia blainvillii Risso, 1822. 

REMARKS.—Forest and Guinot (1966) pointed 

out several differences between Monod's and their 

specimens from West Africa and European specimens 

and suggested that a direct comparison of 

material from the two areas might result in the 

recognition of a small, distinct West African species. 


British Isles and Sweden southward to Portugal, 

Mediterranean, and off Sierra Leone and Sao 

Tome in the Gulf of Guinea; usually in depths 

between 10 and 25 m. 

Family CANCRIDAE Latreille, 1803 

Cancerides Latreille, 18O3b:35O [corrected to Cancridae by 

MacLeay, 1838:59]. 

TRICHOCERIDAE Dana, 1852a: 120. 

EASTERN ATLANTIC GENERA.—One, not represented 

in the tropical fauna, is Cancer Linnaeus, 

1758:625. Type-species: Cancer pagurus Linnaeus, 

1758, by subsequent designation by Latreille, 

1810:422; gender: masculine; name 491 on Official 

List. 

EASTERN ATLANTIC SPECIES.—Two, both extralimital: 

Cancer bellianus Johnson, 1861. Shetland Islands 

and Iceland southward to Canary Islands, Madeira 

(Tiirkay, 1976b), and Spanish Sahara 

(Maurin, 1968b); sublittoral, to 620 m (Zariquiey 


Cancer pagurus Linnaeus, 1758. Northern Norway 

to the Mediterranean; sublittoral, between 

6 and 100 m (Zariquiey Alvarez, 1968; Christiansen, 

1969). 

Family PIRIMELIDAE Alcock, 1899 

Pirimelides A. Milne Edwards, 1862a:41; 1865:181, 204. 

PIRIMELINAE Alcock, 1899a:5, 95. [Elevated to Pirimelidae 

by Bouvier, 1940:217.]

70 

PERIMELIDAE Monod, 1956:147, 157. [Unjustified emendation.] 

EASTERN ATLANTIC GENERA.—Two, Pirimela 

and Sirpus, both represented in the fauna of tropical 

West Africa. 

EASTERN ATLANTIC SPECIES.—Four, three of 

which occur in tropical waters. The extralimital 

species, Sirpus zariquieyi Gordon, 1953, occurs in 

the Mediterranean (Lewinsohn and Holthuis, 

1964; Zariquiey Alvarez, 1968). 

REMARKS.—Monod (1956) recorded two species 

from West Africa, as listed below, and we 

add a previously undescribed species of Sirpus (S. 

gordonae), the only representative of the family 

collected by the Pillsbury: 

Name in Monod Current "Name 

Perimela denticulata Pirimela denticulata 

Sirpus monodi Sirpus monodi 

Genus Pirimela Leach, 1816 

Pirimela Leach, 1816, in 1815-1875, pi. 3 [type-species: Cancer 

denticulatus Montagu, 1808, by monotypy; gender: feminine; 


Perimela Agassiz, 1846:280, 293 [unjustified emendation of 

Pirimela Leach, 1816; type-species: Cancer denticulatus Montagu, 

1808; gender: feminine]. 

Pirimela denticulata (Montagu, 1808) 

Pirimela denticulata.—Bouvier, 1911:226 [PMauritania; listed]. 

—Zariquiey Alvarez, 1968:350, figs. 7a, lid, 112a, 113a 


11 [Scandinavia].—Tiirkay, 1976b:61 [listed], 64 [Madeira]. 

Perimela denticulata.—Capart, 1951:137 [Spanish Sahara].— 

Monod, 1956:157, figs. 187-190 [Mauritania, Senegal, 

Gabon(?)]. 

SYNONYM.—Pirimela princeps Hope, 1851. 


to Senegal, Mediterranean, usually in shallow 

water, intertidal to 40-50 m. There is one questionable 

record from Gabon (Monod, 1956); 

Bouvier (1940) recorded material to a depth of 

200 m. 


Genus Sirpus Gordon, 1953 

Sirpus Gordon, 1953a:304 [type-species: Sirpus zariquieyi Gordon, 

1953, by original designation and monotypy; gender: 

masculine]. 

Sirpus monodi Gordon, 1953 

Sirpus monodi.— Monod, 1956:159, figs. 191-193 [Mauritania, 

Senegal; references].—Rossignol, 1962:115 [Congo].— 

Monod, 1963, fig. 33 [no locality], 

Zirpus Monodi.—Monod, 1963:124 [erroneous spelling]. 

DISTRIBUTION.—Off West Africa, from Mauritania, 

Senegal, and the Congo, intertidal to a 

depth of 12 m. 

* Sirpus gordonae, new species 

FIGURE 15 


Sta 271, shore, 3c5 (includes holotype), 39 (L,W). 

DESCRIPTION.—Carapace (Figure I5a,b) about 

as long as wide, greatest width behind postorbital 

tooth, at level of first anterolateral tooth. Front 

(Figure I5a,b) 3-lobed rather than 3-toothed as 

in Sirpus monodi Gordon and S. zariquieyi Gordon, 

median lobe bluntly triangular, extending farthest 

anteriorly, submedian lobes shorter, 

rounded triangular in shape, pointed or with 

small, sharp, almost granuliform apex. Anterior 

supraorbital angle triangularly pointed or 

rounded, with small granuliform apex. Dorsal 

orbital margin straight, sloping posteriorly, terminating 

in distinct incision. Posterior orbital 

margin, between incision and postorbital tooth, 

rounded or angular. Postorbital tooth large, 

sharply pointed, curved anteriorly. Lateral margin 

(Figure I5a,b), posterior to postorbital tooth, 

with 3 spiniform teeth, anteriormost larger than 

postorbital, with margins, particularly anterior, 

crenulate. Posterior 2 lateral teeth short, small, 

contrasting greatly with anterior tooth, terminating 

in sharp apices, often dorsally directed or 

twisted. Surface of carapace strongly areolated, 

in this resembling other Sirpus species, with elevations 

as follows: 2 protogastric, 1 mesogastric,

NUMBER 306 71 

FIGURE 15.—Sirpus gordonae, new species, paratypes. Female, cl 2.8 mm, Pillsbury Sta 271: a, 

carapace. Male, cl 2.8 mm, Pillsbury Sta 271: b, carapace; c, abdomen; d, first pleopod; e, apex 

of first pleopod;/, second pleood. 

1 on each branchial region, and 2 submedian on 

cardiac region. No elevation present at base of 

posterior lateral tooth, but small spinule or tooth 

there on dorsal surface of carapace in some specimens. 

Posterior part of carapace rugose, with 

short transverse striae, as in other species of genus. 

Lower border of orbit terminating in blunt 

tooth. Antepenultimate segment of antennular 

peduncle, forming part of orbital floor, terminating 

in distinct anterior spine and spinules. Antennal 

peduncle extending beyond front. 

Male abdomen (Figure 15

72 

TYPE-LOCALITY.—Annobon Island, 01°25'S, 

05°38'E, Gulf of Guinea. 


D. 31543), a male specimen with carapace length 

and width 2.6 mm, is in the Rijksmuseum van 

Natuurlijke Historie, Leiden. Two paratypes, a 

male and a female, also are in that collection and 

a male and two female paratypes (USNM 

169534) are in the National Museum of Natural 

History, Smithsonian Institution, Washington, 

D. C. 

ETYMOLOGY.—It is a great pleasure to dedicate 

this new species to Dr. Isabella Gordon, British 

Museum (Natural History), who was the first to 

make the existence of the genus Sirpus known to 

the world and who exhaustively treated the first 

two species assigned to it. 

DISTRIBUTION.—Our material of Sirpus gordonae 

was obtained at Annobon during a fish poison 

station at a sandy beach in a rocky cove. Sirpus 

zariquieyi is a Mediterranean species, known from 

scattered localities between Spain and Israel; it 

has been found in depths between 1 to about 40 

m. Sirpus monodi is known from NW Africa, from 

Mauritania and Senegal, and has been taken in 

depths between 0 and 12 m. This represents the 

first find of the genus in the southern hemisphere. 

Family CORYSTIDAE Samouelle, 1819 

CORYSTIDAE Samouelle, 1819:82 [name 357 on Official List). 

NAUTILOCORYSTIDAE Ortmann, 1893a:26, 28. 

EURYALIDAE Rathbun, 1930:10. 

EASTERN ATLANTIC GENERA.—Two, only one of 

which, Nautilocorystes, is represented in the tropical 

fauna. The other genus is Corystes Bosc (1802:65). 

Type-species: Hippa dentata Fabricius, 1793, by 

monotypy; gender: masculine; name 1571 on Official 

List. 

EASTERN ATLANTIC SPECIES.—Two, the first of 

which is the only species of the family occurring 

in tropical waters: 

Nautilocorystes ocellatus, was first discovered there 

after Monod's account of the tropical species; 

Monod remarked (1956:155) that it occurred as 


far north as 17°S. No representatives of the family 

were taken by the Pillsbury. 

Corystes cassivelaunus (Pennant, 1777). Southern 

Norway and Sweden southward to Gibraltar and 

the Mediterranean, in depths between 7 and 90 

m; both Zariquiey Alvarez (1968) and Christiansen 

(1969) gave accounts of this species. 

Genus Nautilocorystes H. Milne Edwards, 1837 

Dicera de Haan, 1833:14 [invalid junior homonym of Dicera 

Germar, 1817 (Hymenoptera); type-species: Corystes (Dicera) 

octodentata de Haan, 1833, a subjective junior synonym 

of Corystes ocellatus Gray, 1831, by monotypy; gender: 

feminine]. 

Nautilocorystes H. Milne Edwards,, 1837:149 [type-species: 

Nautilocorystes ocellatus H. Milne Edwards, 1837, an invalid 

junior homonym of Corystes ocellatus Gray, 1831, by 

monotypy; gender: masculine]. 

Alyptes Gistel, 1848:ix [substitute name for Dicera de Haan, 

1833; type-species: Corystes (Dicera) octodentata de Haan, 

1833, a subjective junior synonym of Corystes ocellatus Gray, 

1831; gender: masculine]. 

Nautilocorystes ocellatus (Gray, 1831) 

Corystes ocellata Gray, 1831:39. 

Nautilocorystes ocellata.—Barnard, 1950:303, figs. 57a-c [South 

Africa].—Monod, 1956:155 [Great Fish Bay (= Baia dos 

Tigres), Angola].—Rossignol, 1962:115 [listed].—Crosnier, 

1967:324 [Congo]. 

Nautilocorystes ocellatus.—Capart, 1951:110, fig. 38 [Waivis 

Bay, Southwest Africa]. 

SYNONYMS.—Corystes {Dicera) octodentata de 

Haan, 1833; Nautilocorystes ocellatus H. Milne Edwards, 

1837. 

DISTRIBUTION.—South Africa and South-West 

Africa northward to Angola and off the Congo, 


Family BYTHOGRAEIDAE Williams, 1980 

BYTHOGRAEIDAE Williams, 1980:444. 

This family, comprising a single genus and 

species from the Galapagos Rift in the eastern 

Pacific, is not represented in the study area. 

Family PORTUNIDAE Rafinesque, 1815 

PORTUNIDIA Rafinesque, 1815:97 [corrected to Portunidae 

by Samouelle, 1819:83; name 69 on Official List]. 

MEGALOPIDAE Haworth, 1825:184.

NUMBER 306 73 

CARCINIDAE MacLeay, 1838:59. 

LUPINAE Dana, 1851b: 129. 

ARENAEINAE Dana, 1851b: 129. 

PLATYONYCHIDAE Dana, 1851b: 130. 

PODOPHTHALMIDAE Dana, 1851b: 130. 

Neptuniden Nauck, 1880:65. 

THALAMITINAE Miers, 1886:170, 193. 

CAPHYRINAE Miers, 1886:170. 

Cronius ruber 

Neptunus vocans 

Neptunus validus 

Neptunus inaequalis 

Callinectes gladiator 

Callinectes marginatus 

Callinectes latimanus 

Cronius ruber* 

Portunus vocans 

Portunus validus* 

Portunus inaequalis * 

Callinectes pallidus * 

Callinectes marginatus * 

Callinectes amnicola * 

POLYBIIDAE Ortmann, 1893a:66, 68. 

The extralimital species are as follows: 

CARUPIDAE Ortmann, 1893a:66, 68. 

Bathynectes longipes (Risso, 1816). England to 

LISSOCARCINIDAE Ortmann, 1893a:67, 87. 

LUPOCYCLOIDA Alcock, 1899a:22. 

PORTUMNINAE Ortmann, 1899:1170. 

GONIOCAPHYRINAE Borradaile, 1900:577. 

XAIVIDAE Berg, 1900:224. 

Portugal, Madeira, Mediterranean; sublittoral, 

20-90 m (Zariquiey Alvarez, 1968; Tiirkay, 

1976b). 

Bathynectes maravigna (Prestandrea, 1839). See 

CATOPTRINAE Borradaile, 1902a: 200. 

page 76. 

LIOCARCININAE Rathbun, 1930:18. 

MACROPIPINAE Stephenson and Campbell, 1960:75, 76, 88. 

Callinectes sapidus Rathbun, 1896. A. Milne Edwards 

and Bouvier (1900:71, pi. 4: fig. 5) reported 

EASTERN ATLANTIC GENERA.—Twelve, of which 

three specimens (2 6, 1 9) of "Callinectes diacanthus, 

nine, Bathynectes, Callinectes, Carcinus, Cronius, Lio- 

Latr., var. africanus, A. M.-Edw." from Porto da 

carcinus, Macropipus, Portunus, Thalamita, and Xaiva, 

Praia (as Praya), Cape Verde Islands. The colored 

are represented by tropical species. Three genera 

figure that they published shows definitely Calli- 

do not occur in the tropical region: 

nectes sapidus, having only two frontal teeth. As 

Charybdis de Haan (1833:3, 10). Type-species: 

Monod (1956:204) pointed out, the figure is noth- 

Cancer sexdentatus Herbst, 1783, a subjective junior 

ing but a colored copy of the figure that A. Milne 

synonym of Cancer feriatus Linnaeus, 1758, by Edwards (1861:425, pi. 30: fig. 1) published of 

subsequent designation by Glaessner (1929:113); "Neptunus diacanthus. Individu male de gran- 

gender: feminine; name 1616 on Official List. deur naturelle, rapporte des cotes de l'Amerique 

Polybius Leach (1820, in 1815-1875, pi. 9b: figs. 

septentrionale. (Variete a front quadrilobe)." As 

1-4). Type-species: Polybius henslowii Leach, 1820, to the identity of the three specimens taken by 

by monotypy; gender: masculine; name 184 on the Talisman, Monod (1956:204) stated: "J'ai vu 

Official List. 

un des 6 d'africanus (M.P.): c'est un C. marginatus." 

Portumnus Leach (1814:391, 429-430). Type- This identification is confirmed by Williams 

species: Cancer latipes Pennant, 1777, by mono- (1974:729), who likewise examined the male spectypy; 

gender: masculine; name 185 on Official imen in the collection of the Paris museum. The 

List. 

female specimen of the Talisman set now forms 

EASTERN ATLANTIC SPECIES.—36, of which 16 part of the collection of the Museum of Compar- 

occur in tropical waters. The following species ative Zoology, Harvard University (no. 6530) and 

were recorded by Monod (1956): 

was likewise examined by Williams (1974:729); it 

also is C. marginatus. The third specimen of the lot, 


a male (USNM 23950), is now in the collection 

Carcinus maenas Carcinus mamas 

of the National Museum of Natural History, 

Xaiva biguttata Xaiva biguttata 

Xaiva mcleayi Xaiva mcleayi 

Smithsonian Institution, Washington, D.C.; it 

Portunus arcuatus Liocarcinus arcuatus 

was examined by us, and like the two other 

Portunus corrugatus Liocarcinus corrugatus 

specimens belongs to C. marginatus. A. Milne Ed- 

Portunus tuberculatus 

Bathynectes superbus 

Thalamita africana 

Macropipus rugosus * 

Bathynectes piperitus, 

new species* 

Thalamita poissonii 

wards and Bouvier (1900:71) thus did not correctly 

report C. sapidus from West Africa. 

A second dubious record of C. sapidus from

74 

West Africa is the one by Gruvel (1912, pi. 2: fig. 

1), who in an account of the edible Crustacea of 

West Africa figured as Callinectes africanus a specimen 

that clearly represents C. sapidus. The explanation 

of Gruvel's figure does not give any information 

about the locality or origin of the specimen 

figured. As Callinectes sapidus otherwise has 

not been reported from West Africa, notwithstanding 

intensive collecting, we agree with 

Monod (1956:204) who thinks that "on est bien 

oblige de penser- qu'ici encore le specimen represente 

est un sapidus (americain)." 

Although so far there has not been a single 

reliable record of Callinectes sapidus from West 

Africa, and the species certainly is not native 

there, the possibility exists that it will be accidentally 

introduced in West African waters as it was 

in Europe, the Mediterranean, and Japan; for an 

enumeration of the European and eastern Mediterranean 

localities of the species see Holthuis 

(1969a:34, fig. 1) and Christiansen (1969:72, fig. 

29, map 23). Since 1969, introduced specimens 

have been reported from the North Sea off the 

English coast (Dr. R. W. Ingle, in litt.), Normandy, 

France (Maury, 1975:25), possibly from 

Nice (Tiirkay, 1971:129), the central Adriatic Sea 

coast of Italy (Froglia, 1972:48), Strait of Messina 

(Cavaliere and Berdar, 1977), and Japan (Dr. T. 

Sakai, in litt.). In view of what is said in the 

previous paragraphs, it is unlikely that Rathbun's 

(1921:384) suggestion that the European species 

came from a West African population is correct. 

Carcinus aestuarii Nardo, 1847. Mediterranean, 

also in Canary Islands, possibly in Atlantic adjacent 

to Mediterranean and possibly in portions 

of the Suez Canal; intertidal (Zariquiey Alvarez, 

1968). Until now this species was known as Carcinus 

mediterraneus Czerniavsky, 1884. Nardo's 

(1847b) account, however, based on material 

from the lagoon at Venice, clearly provides the 

oldest available name for the Mediterranean species 

of Carcinus. 

Charybdis helleri (A. Milne Edwards, 1867). Eastern 

Mediterranean, Egypt and Israel (from the 

Red Sea); sublittoral (Holthuis and Gottlieb, 

1958; Ramadan and Dowidar, 1976). 


Charybdis longicollis Leene, 1938. Eastern Mediterranean, 

Egypt to Turkey (from the Red Sea); 

sublittoral (Holthuis, 1961; Lewinsohn and Holthuis, 

1964; Ramadan and Dowidar, 1976). 

Liocarcinus bolivari (Zariquiey Alvarez, 1948). 

Mediterranean, 8-60 m (Zariquiey Alvarez, 

1968). 

Liocarcinus depurator (Linnaeus, 1758). Norway 

to Spanish Sahara, Mediterranean; in depths to 

450 m, usually shallower than 100 m (Zariquiey 


Liocarcinus holsatus (Fabricius, 1798). Hebrides 

southward to Portugal, Canary Islands, possibly 

also Cap Blanc, Mauritania (Monod, 1956, recorded 

a juvenile 8 X 10 mm from there); in 

depths to 350 m, generally in less than 100 m 

(Zariquiey Alvarez, 1968; Christiansen, 1969). 

Liocarcinus maculatus (Risso, 1827). Mediterranean 

records of L. pusillus are referable to this 

species, according to a study by Manning and C. 

Froglia, Laboratorio di Tecnologia della Pesca, 

Ancona, to be published in the proceedings of II 

Colloquium Crustacea Decapoda Mediterranea 

held in Ancona, Italy, in May 1979. 

Liocarcinus marmoreus (Leach, 1814). Southern 

North Sea and British Isles southward to Spain, 

Azores, and Madeira; in shallow water to about 

85 m (Zariquiey Alvarez, 1968; Christiansen, 

1969; Tiirkay, 1976b). 

Liocarcinus puber (Linnaeus, 1767). Norway to 

Spanish Sahara, Mediterranean; littoral to 70 m 

(Zariquiey Alvarez, 1968; Christiansen, 1969). 

Liocarcinus pusillus (Leach, 1815). Norway to 

Portugal, possibly on northwest African coast (see 

Christiansen, 1969, for comments); in depths 

from 6 to 200 m, usually less than 50 m (Zariquiey 


Liocarcinus vernalis (Risso, 1816). Mediterranean, 

and possibly (doubtfully) off Cabo Blanco, 

Spanish Sahara (Monod, 1956); shallow water 

(Zariquiey Alvarez, 1968). 

Liocarcinus zariquieyi Gordon, 1968. England to 

Canary Islands, Mediterranean; 5 to 30 m (Zariquiey 


Macropipus tuberculatus (Roux, 1830). Norway to 

the Azores, Morocco, Mediterranean; 20-30 to

NUMBER 306 75 

834 m (Zariquiey Alvarez, 1968; Christiansen, 

1969). 

Polybius henslowii Leach, 1820. North Sea and 

British Isles southward to Morocco, Mediterranean; 

pelagic (Zariquiey Alvarez, 1968; Christiansen, 

1969). 

Portumnus latipes (Pennant, 1777). North Sea 

and British Isles to northern Morocco (there is 

one doubtful record from Mauritania), Mediterranean; 

intertidal and subtidal to a depth of 28 

m (Zariquiey Alvarez, 1968; Christiansen, 1969). 

Portumnus pestai Forest, 1967. Mediterranean; 

shallow water (Forest, 1967). 

Portunus pelagicus (Linnaeus, 1758). Mediterranean 

(from the Red Sea, Egypt to Turkey, Italy); 

sublittoral (Holthuis and Gottlieb, 1958; Ramadan 


Portunus sayi (Gibbes, 1850). Canary Islands (see 

P. hastatus, p. 101); Cap Spartel, Morocco and 

Balearic Islands, from drifting Sargassum (Bouvier, 

1922; Zariquiey Alvarez, 1968). Bouvier (1940) 

does not refer to his earlier record of this species 

from the Balearic Islands, so there is some question 

as to its authenticity. There are no other 

records of this species from the Mediterranean. 

Subfamily CARCININAE MacLeay, 1838 

Genus Carcinus Leach, 1814 

Ligia Weber, 1795:92 [type-species: Cancer granarius Herbst, 

1783, a subjective junior synonym of Cancer maenas Linnaeus, 

1758, by monotypy; gender: feminine; name suppressed 

by the International Commission on Zoological 

Nomenclature in Opinion 330, 1955, and placed on the 

Official Index as name 207]. 

Carcinus Leach, 1814:390 [type-species: Cancer maenas Linnaeus, 

1758, by monotypy; gender: masculine; name 798 


Megalopa Leach, 1814:431 [type-species: Cancer granarius 

Herbst, 1783, a subjective junior synonym of Cancer maenas 

Linnaeus, 1758, by present selection; gender: feminine]. 

Macropa Latreille, 1822:9 [type-species: Megalopa montagui 

Leach, 1817, an objective synonym of Cancer rhomboidalis 

Montagu, 1804, a subjective junior synonym of Cancer 

maenas Linnaeus, 1758, by monotypy; gender: feminine]. 

Megalops H. Milne Edwards, 1837:260 [erroneous spelling of 

Megalopa Leach, 1814]. 

Sympractor Gistel, 1848:ix [replacement name for Megalopa 

Leach, 1814 (as Megalopus); type-species: Cancer granarius 

Herbst, 1783, a subjective junior synonym of Cancer maenas 

Linnaeus, 1758; gender: masculine]. 

Carcinides Rathbun, 1897b: 164 [replacement name for Carcinus 

Leach, 1814; type-species: Cancer maenas Linnaeus, 

1758; gender: masculine; name 209 on Official Index]. 

Carcinus maenas (Linnaeus, 1758) 

Carcinus maenas.—Monod, 1956:165, fig. 194 [Mauritania; 

references].—Forest and Games, 1960:351 [Morocco].— 

Guinot, 1967a:253 [Indian Ocean localities; listed].— 

Monod, 1967:178 [no locality].—Zariquiey Alvarez, 1968: 

354, figs. 115a, c [Spain; references].—Christiansen, 1969: 

49, fig. 18, map 12 [Scandinavia]. 

SYNONYMS.—Cancer viridis Herbst, 1783; Cancer 

granarius Herbst, 1783; Cancer pygmaeus Fabricius, 

1787; Cancer rhomboidalis Montagu, 1804; Megalopa 

montagui Leach, 1817; Portunus menoides Rafinesque, 

1817; Cancer granulatus Say, 1817. 

DISTRIBUTION.—Eastern Atlantic, from Norway, 

N of 70° N latitude, S to Mauritania, about 

20° S latitude; western Atlantic; eastern Pacific; 

Australia. In the Mediterranean it is replaced by 

the allied C. aestuarii Nardo, 1847 (= C. mediterraneus 

Czerniavsky, 1884) (see Bacescu, 1967, and 

Zariquiey Alvarez, 1968, for differences between 

the two species). 

Genus Xaiva MacLeay, 1838 

Xaiva MacLeay, 1838:62 [type-species: Xaiva pulchella Mac- 

Leay, 1838 (? = Portunus biguttatus Risso, 1816), by monotypy; 

gender: feminine; name 1649 on Official List]. 

Portumnoides Bohn, 1901:270, 271 [type-species: Portumnoides 

garstangi Bohn, 1901, a subjective junior synonym of Portunus 

biguttatus Risso, 1816, by monotypy; gender: masculine]. 

Xaiva biguttata (Risso, 1816) 

Xaiva biguttata.—Monod, 1956:168, fig. 195 [Cape Verde 

Islands].—Guinot, 1967a:253 [Indian Ocean; listed].— 

Zariquiey Alvarez, 1968:359, figs. 8d, 116c, 122b [Spain; 

references].—Kensley, 1970:182 [South-West Africa].— 

PenrithandKensley, 1970b:248,261 [South-West Africa]. 

SYNONYMS.—Platyonichus nasutus Latreille, 1828; 

Portumnoides garstangi Bohn, 1901.

76 

REMARKS.—Material of this species from the 

northern and southern parts of its reported range 

(England to Cape Verde Islands versus South- 

West Africa and South Africa) should be studied 

to determine whether or not two species might be 

recognized. If the southern form proves to be 

distinct, the name Xaiva pulchella MacLeay, 1838, 

is available. 

DISTRIBUTION.—Eastern Atlantic, from England 

southward to the Cape Verde Islands, 

South-West Africa, South Africa, and the Mediterranean, 

in shallow water. 

Xaiva mcleayi (Barnard, 1947) 

Portumnus mcleayi.—Capart, 1951:114, fig. 40, pi. 2: fig. 9 

[Angola].—Rossignol, 1957:123 [key]. 

Xaiva mcleayt.—Monod, 1956:169, 632, figs. 196-201 [Mauritania, 

Senegal, Sierra Leone; references].—Longhurst, 

1958:87 [Sierra Leone].—Guinot and Ribeiro, 1962:31 

[Angola].—Forest and Guinot, 1966:59 [Principe, Sao 

Tome].—Guinot, 1967a:253 [Indian Ocean; listed].— 

Crosnier, 1967:324 [Dahomey, Annobon]. 

?Xaiva biguttata.—Gauld, 1960:69 [Ghana] [not Xaiva biguttata 

(Risso, 1816)]. 

REMARKS.—We suspect that Gauld's record 

from Ghana of X. biguttata, a species not known 

to occur south of the Cape Verde Islands on the 

tropical West African Coast, may have been 

based on this species, now known to occur at least 

as far north as Mauritania on the West African 

coast (Monod, 1956). 

DISTRIBUTION.—Eastern Atlantic, from scattered 

localities between Mauritania and Angola, 

South Africa, in depths between 8-30 and 73 m. 

Subfamily POLYBIINAE Ortmann, 1893 

Genus Bathynectes Stimpson, 1871 

Bathynectes Stimpson, 1871a: 145 [type-species: Bathynectes longispina 

Stimpson, 1871, by subsequent designation by 

Rathbun, 1930:27; gender: masculine; name 127 on Official 

List]. 

Thranites Bovallius, 1876:60, 61 [type-species: Thranites velox 

Bovallius, 1876, a subjective junior synonym of Portunus 

maravtgna Prestandrea, 1839, by monotypy; gender: masculine]. 


Bathynectes maravigna (Prestandrea, 1839), 


Portunus Maravigna Prestandrea, 1839:132. 

Portunus superbus Costa, in Costa and Costa, 1838-1871: 19, 

pi. 5 [color]. 

Bathynectes.—Filhol, 1885a:56.—Maurin, 1968a, fig. 29; 

1968b, figs. 1,4. 

Bathynectes superba.—Forest and Gantes, 1960:351 [Morocco].—Peres, 

1964:20, 26, 29 [Morocco]. 

Bathynectes superbus.—Maurin, 1968a: 19, 45, 50, 64 [Spain, 

Spanish Sahara, Mauritania]; 1968b:482, 484, 489, 491, 

fig. 6 [Spanish Sahara, Mauritania].—Zariquiey Alvarez, 

1968:382, fig. 127g [Spain; references].—Christiansen, 

1969:70, fig. 28, map 22 [Scandinavia].—Tiirkay, 1976b: 

61 [listed], 64 [Madeira].—Lewis and Haefner, 1978:164 

[part; Spain]. 

SYNONYM.—Thranites velox Bovallius, 1876. 

MATERIAL EXAMINED.—France: Bane de la Chapelle, 

off Brittany, 47°58'N, 08°00'W, 9 Sep 1921, L. Fage, 16\ 1$ 

(MP). 

Spain: NE of Santander, (43°28'N, 03°48'W), 564 m, 

sand and shells, 6 Jul 1882, Travailleur, 6 juv (MP, W). Off 

western Galicia, 43°41.2'N, 08°57.6'W, 995 m, rocky bottom, 

8 Aug 1967, Thalassa Sta T.491, 1 juv (MP). 

Madeira: SE of Madeira, 32°43'N, 16°43'W, 420 m, 16 

(L). 

Morocco: Fosse de Rabat (approx. 34°N, 07°W), 300- 

400 m, N. Pigeault, 1

NUMBER 306 77 

delle articolazioni dei piedi camminatori. . . . Gli occhi sono m, 3 juv (W). Sta 220, 300 m, 1 carapace (W). Sta 247, 400 

sferici nerastri. [The anterolateral teeth] sono tutti del colore m, 1 9(W). 

del torace con l'estremita color caffe cotto. . . . Tutte le spine Undaunted Material: Angola: Sta 111, ca. 366 m, 2 6* 

dei piedi-mani [= chelipeds] sono bianche colla estremita (includes holotype), 39 (1 ov) (L). 

color caffe cotto; pero quella del lato interno del carpo a Other Material: Cape Verde Islands: 16°53'N, 27°30'W to 

una banda bianca; quindi una rosso-corallo chiara e la 16°54'N, 27°30'W of Paris [= 25° 10'W of Greenwich], 410estremita 

come le altre. [The fingers are] verso l'estremita 460 m, sand and gravel, 29 Jul 1883, Talisman Sta 110, 16 

brunastre. 

(MP). 15°14'N, 23°03'45"W, 628 m, muddy sand, fish trap, 

In listing material of this species that does not 

occur off tropical West Africa, we have departed 

14-15 Aug 1901, Princesse Alice Sta 1189, 19 (MP). 

Senegal: NW of Pointedes Almadies (14°45'N, 17°32'W), 

250-300 m, 21 Oct 1952, Ghard Tre'ca, 26, 19 (MP). Off 

from our format. However, the material listed Pointe des Almadies, ca. 300 m, 16 Oct 1952, Cremoux, leg., 

above was used to compare this species with the Gerard Tre'ca, 19 (MP). 

new species from West Africa described below. DESCRIPTION—Front (Figures 16, 17a) with 4 

Material from Spanish Sahara and Mauritania rounded teeth, inner 2 about half as wide as 

identified with this species by Maurin (1968a,b) outer, extending about as far forward as outer. 

might well be referable to B. piperitus, new species. Latter broadly rounded, more semicircular than 

This cannot be determined without examining triangular. Inner orbital teeth low, rectangular, 

his material. 

wide, with blunt apex and short dorsal carina. 

DISTRIBUTION.—Eastern Atlantic, from Norway Upper orbital margin with 2 distinct narrow, 

and the Faroes southward to NW Morocco, pos- open fissures. Exorbital tooth triangular, blunt, 

sibly to Spanish Sahara and Mauritania, Medi- especially in larger specimens. Lower orbital marterranean; 

in moderately deep water, from 100 to gin with small, triangular, often blunt, narrow 

1455 m. 

tooth next to exorbital tooth. Third tooth present 

medially, wider, blunter than second, separated 

*Bathynectes piperitus, new species from second by U-shaped incision. Inner, lower 


orbital tooth triangular, with narrowly rounded 

apex, latter occasionally curved inward. First 3 

Bathynectes superba.—A. Milne Edwards and Bouvier, 1899, anterolateral teeth of carapace, excluding exor- 

pi. 2: figs. 16-18 [part, not p. 25, pi. 2: figs. 1-15, bital tooth, sharply triangular, curved anteriorly, 

19-24]; 1900:65 [part].—Bouvier, 1922:59 [part, Cape 

Verde Islands only].—Monod, 1933b:510 [listed].—Ca- third smallest. Fourth (= last) tooth large, spinepart, 

1951:121, fig. 42. [Not Portunus superbus Costa, 1853.] like, curved anteriorly, especially apically. Dorsal 

Bathynectes superbus.—Monod, 1956:183, figs. 210-212.— surface of carapace with 4 transverse ridges: (1) 

Longhurst, 1958:87.—Guinot and Ribeiro, 1962:45.— indistinct postfrontal ridge, widely interrupted in 

Forest, 1963:628.—Monod, 1967, pi. 15: fig. 4 [no records].—Maurin, 

1968b:492.—Intes and Le Loeuff, 1976: 

103.—Lewis and Haefner, 1978:164, pi. la [part]. [Not 

Portunus superbus Costa, 1853.]. 

Bathynectes suberbus.—Gauld, 1960:69 [erroneous spelling]. 

Bathynectes.—Voss, 1966:19, 25.—Maurin, 1968a, fig. 29; 

1968b, fig. 9 [part, Senegal only]. 

midline; (2) mesogastric ridge, divided by median 

MATERIAL EXAMINED.—Pilbbury Material: Liberia: 

Sta 73, 311-366 m, 13 6, 4 9 ov (L, W). 

Ivory Coast: Sta 44, 403-586 m, hard dark gray mud, 

5 juv (L). Sta 51, 329-494 m, 31 6\ 16 9 (1 ov), 13 juv (L, 

W). 

Geronimo Material: Gabon: Sta 179, 293 m, 1 9 (W). Sta 

191, 300 m, 2 6 (W). Sta 198, 300 m, 3 6 (W). Sta 199, 400 

m, 1 6, 1 9 (W). Sta 203, 200 m, 3 6\ 2 9 (W). Sta 206, 455- 

610 m, 1 9, 1 juv (W). Sta 213, 300 m, 1 9 (W). Sta 214, 546 

FIGURE 16.—Bathynectes piperitus, new species (from Capart, 

1951, fig. 42).

78 

FIGURE 17.—Bathynectes piperitus, new species: a, dorsal view; 

b, c, first pleopod of male and apex; d, second pleopod of 

male. (All from Monod, 1956, figs. 210-212.) 

interruption, each half again divided into 2 short, 

slightly convex ridges; (3) long lateral spines connected 

by third transverse ridge, sinuous and 

tuberculate; third ridge convex anteriorly, with 

posteriorly directed indentations (incurvations) at 

branchiocardiac grooves; and (4) short transverse 

carina placed between ends of branchiocardiac 

grooves on posterior half of carapace. Carapace 

finely and closely pubescent. 

Mouthparts not significantly different from 

those of Bathynectes longispina Stimpson. 

First pereiopods unequal, larger chela higher 

and heavier than smaller, with teeth on cutting 

edges of fingers larger, blunter. Otherwise chelipeds 

very similar. Fingers about as long as palm. 

Dactylus with usual 5 ridges, dorsalmost tuberculated 

basally. Upper ridge on inner surface of 

dactylus divided into 2 branches proximally, 


branches united distally; upper branch with 

blunt tubercle. In larger chela, basal tooth of 

cutting edge of dactylus large, swollen, directed 

somewhat posteriorly. Opposite tooth of fixed 

finger molar-like, with 2 small teeth proximally 

at base. Fixed finger, like dactylus, with laterally 

compressed teeth, usually alternately large and 

small. In smaller chela all teeth on cutting edges 

of compressed type. Fixed finger also with 5 

ridges. Upper carina of palm with large, subdistal 

sharp tooth, lacking spinule adjacent to tooth; 

carina tuberculate posterior to tooth. Upper outer 

carina of palm with 6 large, triangular, blunt 

teeth, several smaller tuberculiform denticles 

present between larger teeth. Carina below upper, 

outer carina low, often partly absent, with row of 

small granules, latter often inconspicuous or absent, 

especially in middle of carina. Next lower 

carina strong, high and sharp, granular, terminating 

before reaching base of fingers; row of 

granules below distal end of this carina extending 

to first tooth of fixed finger. Lower part of palm 

with carina extending onto fixed fingers, more 

distinct proximally than distally, ornamented 

with numerous granules. All of lower surface of 

palm covered by transverse rows of granules; 

anteriorly this area terminates on ventral carina 

of fixed finger. Inner surface of palm with high, 

distinct longitudinal carina terminating somewhat 

above base of upper carina of inner surface 

of fixed finger. Carpus with very large, sharply 

pointed tooth on inner margin, anterior margin 

of tooth with 1 to 3, rarely 4, spinules, remainder 

of margin smooth or granular. Upper surface of 

carpus with 2 spines placed in line with upper 

articulation of palm, inconspicuous row of tubercles 

extending from posterior spine towards inner 

tooth, with more distinct row of tubercles extending 

obliquely proximally. Some tubercles scattered 

over remainder of upper surface of carpus. 

Outer anterior angle of carpus with sharp tooth. 

Anterior margin of carpus, between sharp tooth 

and upper articulation with propodus, smooth or 

evenly granular. Merus slender, more than twice 

as long as high, with strong spiniform tooth on 

anterior third of upper margin, second tooth

NUMBER 306 79 

somewhat before middle of inner, lower margin. 

Remainder of merus smooth or slightly tuberculate. 

Walking legs (pereiopods 2 to 4) very slender. 

Dactylus narrow, evenly curved, upper and lower 

surfaces with narrow, longitudinal groove over 

entire length; lateral surfaces with broad groove, 

distally divided in two by longitudinal carina. 

Propodus as long as dactylus, measured dorsally, 

and more than 5 to more than 6 times as long as 

high. Upper and lower margins with deep longitudinal 

groove, flanked by 2 sharp carinae. Outer 

surfaces with 2 longitudinal carinae separating 3 

grooves, middle groove particularly wide; upper 

carina placed close to outer carina of upper margin. 

On inner surface of propodus upper carina 

absent or very reduced, almost entirely covered 

by pubescence, thus only lower carina visible on 

surface. Carpus 2/3 to 4/7 length of propodus, 

with blunt longitudinal dorsal ridge, flanked on 

either side by distinct pubescent groove. Merus 2 

to 3V3 times as long as carpus and 4-5 times as 

long as wide, upper part with transverse subdistal 

groove, occasionally flanked posteriorly by blunt 

tubercle. Upper surface of merus sometimes 

slightly tuberculate. Dactylus of last pereiopod 3 

times as long as high, oval, with lower margin 

straight for most of length, merging with distal 

tooth in straight line. Upper margin regularly 

curved, merging with distal tooth in concave line. 

Propodus about 2/3 as long as dactylus. Carpus 

less than half as long as dactylus. Merus about as 

long as propodus and 2 l k times as long as high. 

Male abdomen triangular, third to fifth somites 

fused, sixth and seventh somites free, latter triangular, 

apex broadly rounded, appearing 

broader than that of B. longispina. 

Male gonopods similar to those of B. longispina, 

with apex less slender, less strongly curved laterally, 

as illustrated by Monod (1956, fig. 211) 

(Figure \lb-d). 

Color: The first color note published of this 

species is the one by Capart (1951:122): "Couleur 

rouge orange, les epines generalement plus 

rouges; les pattes marquees de zones rouges et 

orange alternees." Monod (1956:185) described 

the color as follows: "gris a rouge-brique, parfois 

rose vif; chelipedes en partie rouges, un anneau 

rouge sur les merus, carpe, propode et base du 

dactyle des pattes." The following color description 

is made after notes and photographs taken 

of specimens of Pillsbury Station 73, immediately 

after they came aboard: The carapace is orangered 

with a white spot at the front, one such spot 

just behind either of the large lateral teeth, one 

in each posterolateral angle, and one in the middle 

of the posterior margin. The apex of the 

lateral spine is very dark, almost black. The chelipeds 

are orange with the main spines and teeth 

white. Also the distal part of the palm and the 

fingers are white, except for an orange spot at the 

base of the dactylus. The tips of the fingers and 

a large part of the cutting edges are of a dark, 

almost black color. The walking legs have a broad 

orange band over the distal part of the merus, 

one over the carpus and one over the proximal 

half of the propodus, sometimes the proximal 

part of the dactylus also is orange. All the rest of 

the leg is white, viz., the base of the merus, all the 

articulations, the distal part of the propodus and 

the entire dactylus or the larger part of it. In the 

fifth leg the orange color is somewhat more extensive 

than in the other legs, here the basal part of 

the merus is white, from there on the leg is orange 

up to the middle of the propodus or slightly 

beyond (with the exception of small white articulations), 

and also the basal part of the dactylus 

is orange, the distal part white. The lower surfaces 

of body and chelipeds are white, that of the 

second to fifth pereiopods is of the same color as 

the dorsal surface. The eggs are carmine. 

MEASUREMENTS.—The specimens examined 

had the carapace width from 15 to 86 mm (inclusive 

of the lateral spine); the largest specimen is 

a male. The ovigerous females have carapace 

widths between 55 and 72 mm; the largest of the 

females is ovigerous. In the literature, male specimens 

with carapace lengths from 17 to 57 mm 

and carapace widths 32 to 80 mm (including the 

lateral spines) are recorded; for the non-ovigerous 

females these values are 22 to 49 mm and 38 to 

68 mm, respectively, and for the ovigerous fe-

80 

males 40 to 54 mm and 62 to 78 mm. The 

diameter of the eggs is 0.42 mm (Monod, 1956). 

REMARKS.—Most authors have considered all 

Atlantic Bathynectes, other than Bathynectes longipes 

(Risso, 1816), to belong to a single species, B. 

superbus (Costa, 1853), the type-locality for which 

is Naples, Italy. As stated (p. 76), that species 

should be known as Bathynectes maravigna (Prestan- 

drea, 1839), type-locality "Mare di Messina," 

Sicily, Italy. However, a comparison of the present 

West African material with West Atlantic 

specimens and with material from the Mediterranean, 

European, and NW African waters, 

showed constant differences between the three 

groups, which leads to the conclusion that three 

species are involved. For the West Atlantic species 

the name Bathynectes longispina Stimpson, 1871, is 

available (B. brevispina Stimpson, 1871, is a synonym). 

The name Bathynectes maravigna (Prestandrea, 

1839) has to be used for the species from 

the Mediterranean and NE Atlantic. A new 

name, Bathynectes piperitus, is proposed here for the 

West African species. Bathynectes piperitus differs 

from B. maravigna in the following respects: 

1. The frontal teeth of the carapace in B. 

piperitus are always blunt, even in the large specimens, 

and the median teeth are always distinctly 

narrower than the outer. In B. maravigna the large 

specimens have the frontal teeth sharply pointed 

and triangular, while the median teeth are hardly 

narrower than the outer. In juveniles of B. maravigna 

the situation is much like in the large B. 

piperitus. 

2. The lateral spine of the carapace in B. maravigna 

is long and straight, in B. piperitus it is 

much shorter than in B. maravigna and is curved 

forward in the adults. Although in juveniles of B. 

piperitus the lateral spines are straighter and relatively 

longer than in the adults, they are still 

shorter than in the corresponding stages of B. 

maravigna. 

3. The postfrontal ridges in B. maravigna as a 

rule are more distinct than in B. piperitus, and 

each of the two mesogastric carinae is mostly 

entire and not divided in two short ridges. 

4. The last somite of the male abdomen is 


wider and flatter in B. piperitus, being relatively 

longer and narrower, with a convex upper surface, 

in B. maravigna. The fourth and fifth abdom- 

inal somites in males of B. piperitus each show a 

distinct transverse carina, which are absent or 

very vague in B. maravigna. 

5. The outer half of the anterior margin of the 

carpus of the cheliped (between the outer spine 

and the upper articulation with the palm) in B. 

piperitus usually is straight or finely granular; in 

B. maravigna it carries some spinules. 

6. The inner surface of the propodus of the 

second pereiopod in B. maravigna shows two distinct 

longitudinal carinae separating three longitudinal 

pubescent grooves. In B. piperitus the upper 

of these carinae is absent or hardly indicated, 

so that the lower carina is separated from the 

upper margin of the propodus by a pubescent 

area that is much wider than the area separating 

it from the lower margin. 

7. The merus of the fifth pereiopod is longer 

and more slender in B. maravigna than in B. 

piperitus. The dactylus of this leg in B. maravigna 

is slightly more slender than in B. piperitus and 

has the lower margin somewhat convex and joining 

the distal tooth under a concave curve. 

8. The coloration of living B. maravigna is quite 

different from that of B. piperitus. The colored 

plate of B. maravigna published by A. Milne Edwards 

and Bouvier (1899, pi. 2: fig. 19) shows a 

uniformly orange red crab. Costa's figure (in 

Costa and Costa, 1838-1871, pi. 7) of the species 

has the carapace irregularly mottled with reddish 

brown and yellowish, the pereiopods 1 to 4 are 

uniformly reddish brown, only the dactyli of the 

walking legs and a few spines being yellowish; the 

fifth pereiopods are shown entirely yellowish. A 

male specimen of B. maravigna (cl 61 mm, cb 73 

mm without lateral spines, 96 mm with lateral 

spines) from east of Madeira (11-12 Mar 1976, 

L) was examined rather soon after capture, when 

the colors were still visible and a color photograph 

of the living specimen was made. This specimen 

showed the following color pattern: The animal 

is dark brownish orange dorsally. The margins of 

the frontal teeth are white. A very short median

NUMBER 306 81 

line extends from between the bases of the median 

frontal teeth backward. The tips of the three short 

anterolateral spines of the carapace (not the outer 

orbital angle) have the tips white. One to three 

small white dots are seen on the middle of the 

large lateral spine. A few very small white spots 

are placed in the cervical groove. A large lateral 

white spot stands at each side of the carapace 

about halfway between the lateral spine and the 

posterior margin of the carapace. A narrow white 

line extends along the posterior carina and a 

small median spot is placed somewhat before this 

line. The chelipeds have dorsally the same color 

as the carapace, with a small white spot near the 

tips of many of the spines. The tips of the fingers 

and the teeth on their cutting edges are black. 

Also the tips of various of the spines are black. 

The lower surface of the chelipeds is lighter than 

the upper. The following legs are of the same 

brownish orange color as the carapace. The second 

and third legs have the basal part slightly 

lighter than the rest. There may be some small 

white spots at the articulations and at the tips of 

the second to fourth pereiopods, but otherwise 

the legs are of a uniform brownish orange color. 

The dactyli of the fifth leg show two very small 

spots: one just before the tip, the other near the 

middle of the lower margin, otherwise the leg is 

of a uniform dark brownish orange color. At first 

view B. maravigna seems to be uniformly brownish 

orange, and the white spots are rather small and 

inconspicuous. They evidently have been overlooked 

by the artist of A. Milne Edwards and 

Bouvier's colored plate. Costa's specimen may 

have been made after a specimen that was preserved 

or dead for a considerable time. The most 

important color difference between B. maravigna 

and B. piperitus seems to be the less striking color 

pattern of the carapace and the absence of white 

and orange bands on the pereiopods in the former 

species. 

9. The range of B. maravigna is in the N Atlantic 

Ocean between Iceland and Norway south to the 

Mediterranean (as far as Greece), Madeira, the 

Azores, and NW Morocco. We have examined 

specimens of this species from several localities 

between France and Morocco, as noted above. 

Bathynectes piperitus is a West African species, 

known from the Cape Verde Islands and Senegal 

to Angola. 

The differences between B. piperitus and B. 

longispina are as follows: 

1. In B. longispina the outer frontal teeth are 

more acute and are more triangular in shape; the 

inner teeth are narrowly triangular. 

2. The anterolateral teeth of B. longispina are 

longer, more robust, and relatively narrower than 

in B. piperitus. The length of the second tooth 

after the exorbital angle is greater than half the 

distance between it and the first tooth; in B. 

piperitus it is narrower than half that distance. 

3. The lateral spine of the carapace is straight 

in B. longispina and is longer than in B. piperitus. 

4. The transverse ridge that connects the lateral 

spines of the carapace in B. longispina is 

practically straight and does not show the posterior 

incurvations at the level of the branchiocardiac 

grooves. 

5. The two mesogastric ridges are each divided 

in two smaller ridges like in B. piperitus, but 

they are usually shorter, less distinct (sometimes 

hardly visible), and lie in one line. 

6. The last somite of the male abdomen in B. 

longispina is more triangular than in B. piperitus, 

and has the distal margin less broadly rounded. 

7. The fourth and fifth abdominal somites in 

B. longispina do not show a sharp transverse carina. 

8. The first male gonopod of B. longispina is 

more slender distally and more strongly curved 

than in B. piperitus. 

9. In B. longispina the chelae are relatively 

shorter and higher: the height of the palm (inclusive 

of the dorsal spine) is more than its dorsal 

length. In B. piperitus the dorsal length of the 

palm is definitely greater than the height. 

10. In B. longispina the subdistal tooth on the 

upper margin of the palm is much higher, being 

large, compressed, almost wing-like, reaching 

practically to the base of the dactylus. Before this 

large tooth a small but distinct spine is present 

on the dorsal margin. In B. piperitus the dorsal

82 

tooth is much shorter and less wide and does not 

reach the base of the dactylus; the dorsal margin 

bears no distal spine. 

11. The inner surface of the propodus of the 

second pereiopod in B. longispina shows a single 

deep longitudinal groove in the middle, and the 

two other grooves are inconspicuous. The legs 

have hardly any pubescence. Also the upper 

groove on the inner surface of the carpus is very 

obscure. 

12. The fifth legs in B. longispina have the 

merus more slender and the propodus less slender 

than in B. piperitus. 

13. Also in the color there are marked differences 

between the two species. In B. longispina the 

carapace is reddish because all or practically all 

the tubercles are of that color; the lateral spine is 

dark red in the larger distal part, the anterolateral 

teeth are whitish with a red tip or with a red ring. 

The front is pale and the orbital margin is often 

darker red than the surroundings. A median 

white line extends from the transverse ridge of 

the carapace forwards reaching to the postfrontal 

ridges. The mesogastric ridges, at least the inner 

two, are white and form a more or less distinct 

cross with the median white line. The chelipeds 

are pink; the large teeth on the inner margin of 

the carpus and the upper margin of the palm are 

white at the base and dark red distally. The 

dactylus is red with a white base and a light spot 

in the middle of the dorsal margin; the tip of the 

finger and practically the entire cutting edge (in 

the large chela) or the distal part of it (in the 

smaller) are black. The fixed finger has the distal 

part and the lower surface red, the tip black. The 

legs are pink because of the presence of scattered 

red spots; the articulations are pale or white. Very 

striking is the presence of a solid deep red color 

on the propodus and dactylus of the second to 

fourth leg; this color, which persists long in alcohol-preserved 

specimens, occupies practically the 

entire segments with the exception of a very 

narrow area near the articulation. The carpus 

sometimes shows a narrow red band over its 

middle. In the fifth leg the dactylus, but not the 

propodus, has this same strikingly deep red, al- 


most lacquer-like, color. The presence of these 

deep red bands on the legs distinguishes this 

species immediately from the others belonging to 

this genus. The lower surface of the body is white. 

TYPE-LOCALITY.—Angola, 10°36'S, 13° 12% 

ca 366 m {Undaunted Sta 111). 


D.31505) is a male, carapace width 79 mm, in the 

collection of the Rijksmuseum van Natuurlijke 

Historie, Leiden. The remainder of the specimens 

examined are paratypes, which have been deposited 

in the Rijksmuseum and in the Division of 

Crustacea, National Museum of Natural History, 

Smithsonian Institution. 

ETYMOLOGY.—The Latin name piperitus is given 

in allusion to the bright red and white color, 

resembling that of a peppermint cane. 


depths between 200 and 628 m, and has been 

taken most often between 300 and 450 m. The 

bottom on which it was found was described as 

mud (Guinot and Ribeiro, 1962; Forest, 1963), 

hard, dark gray mud (Voss, 1966), muddy sand 

(Bouvier, 1922), mud and sand, brown mud and 

sand, green mud and sand (Capart, 1951), sand 

and gravel (A. Milne Edwards and Bouvier, 1899, 

1900), corals (Guinot and Ribeiro, 1962), green 

muddy sand (Maurin, 1968b). A. Milne Edwards 

and Bouvier (1900) mentioned the bottom at 

Talisman Sta 113 as "Sable, roche," but in the 

official list of stations of the expedition it is 

indicated as "Sable. Gravier," exactly as in Sta 

110 of the same expedition. 

Ovigerous females of this species have been 

collected in February (Monod, 1956), March, 

April, and May (Capart, 1951), May and June 

(p. 77). Several specimens carry Lepadidae on the 

dorsal surface of the carapace and on the legs. A 

male from Geronimo Sta 198 carries a number of 

hydroid colonies on the merus of the last pereiopod. 

DISTRIBUTION.—Bathynectes piperitus is a West 

African species, known from the Cape Verde 

Islands and the coast of Africa from Senegal to 

Angola. Records of B. superbus (= B. maravigna) 

from Spanish Sahara and Mauritania given by

NUMBER 306 83 

Maurin (1968a,b) and of Bathynectes given by 

Filhol (1885a) may be referable to this species. 

Records in the literature include the following: 

Cape Verde Islands: No specific locality (Filhol, 1885a). 

16°53'N, 27°3O'W of Paris [= 25°10'W of Greenwich] to 

16°54'N, 27°30'W of Paris [= 25°10'W of Greenwich], 410- 

460 m (A. Milne Edwards and Bouvier, 1899, 1900). 16°- 

52'N, 27°30'W of Paris [= 25°10'W of Greenwich] to 16°- 

52'N, 27°32'W of Paris [= 25°12'W of Greenwich], 550-760 

m (A. Milne Edwards and Bouvier, 1900). 15°14'N, 23°- 

03'45"W, 628 m (Bouvier, 1922). 

Senegal: Off Saint-Louis, 300 m (Maurin, 1968b). Fosse 

de Kayar, 85-660 m (Monod, 1956), 300-350 to 600 m 

(Maurin, 1968b). Off Pointe des Almadies, 150-245 m, 250- 

300 m, ±300 m, and 270-500 m (Monod, 1956). 

Guinea-Bissau: 10°05'N, 17°00'W, 320-360 m (Capart, 

1951). 

Guinea: 09°05'N, 15°10'W, 310-380 m (Monod, 1956). 

Liberia: 04°40'N, 09°20'W, 311-366 m (Voss, 1966; 

Lewis and Haefner, 1978). 

Ivory Coast: No specific locality (Intes and Le Loeuff, 

1976). 05°05'N, 04°00'W to 05°04'N, 04°02'W, 403-586 m 

(Voss, 1966). 04°32.5'N, 06°31'W, 300-455 m, and 04°54'N, 

03°23'W, 380-400 m (Forest, 1963). 

Ghana: 04°39'N, 02°46'W, 300-400 m (Forest, 1963). Off 

Accra, 200 m (Longhurst, 1958; Gauld, 1960). 

Gabon: Off Port-Gentil, 00°15'S, 08°47'E, 290-390 m 

(Capart, 1951). 0O°02.2'S, 08°50.2'E, 293 m (cited as Liberia); 

01°26.4'S, 08°24'E, 400 m (cited as 01°6.4'S, 08°- 

44'E, 396 m); 01°28'S, 08°24.5'E, 300 m (as 297 m); 02°01'S, 

08°50.5'E, 200 m; 02°31'S, 08°51'E, 300 m (as 288 m); 

04°38.4'S, ll°01.2'E, 400 m (all Lewis and Haefner, 1978). 

Cabinda: 05°23'S, 11°32'E, 290-350 m, and 05°39'S, 

11°25'E, 470 m (Capart, 1951). 

Angola: Off Ponta da Moita Seca, 06°08'S, 11°24'E, 350- 

380 m, and 06°23'S, 11°29'E, 400-430 m; off Ambrizete, 

07°16'S, 12°02'E, 380-420 m; off Ponta do Morro (as Cap 

Morro), 10°45'S, 13°O7'E, 400-500 m; off Egito [Praia], 

11°53'S, 13°28'E, 300 m, 11°53'S, 13°20'E, 480-510 m, and 

11°53'S, 12°23'E, 400-500 m (all Capart, 1951). Off Benguela, 

405-505 m; off Baia dos Tigres, 320-400 m (Guinot 

and Ribeiro, 1962). 

Genus Uocarcinus Stimpson, 1871 

Liocarcinus Stimpson, 1871a: 145 [type-species: Portunus holsatus 

Fabricius, 1798, by original designation; gender: masculine]. 

DEFINITION.—Carapace broader than long, 

with 5 anterolateral teeth, lacking iridescent 

patches on surface. Front with 3 lobes, 7-10 irreg- 

ular teeth or tubercles, or entire. Dorsal margin 

of orbit with 2 closed incisions, ventral margin 

(Figure 18a) with narrow, V-shaped, incision. 

Basal antennal article touching front, scarcely or 

not at all movable. Chelipeds unequal, generally 

shorter than pereiopods; merus without distal 

ventral tooth; carpus with strong inner tooth, 

generally lacking outer tooth; dactylus (Figure 

18c) usually with 3 dorsal ridges. Propodus of 

second to fourth pereiopods each with 1 ventral 

ridge (Figure 18*,g). Third to fifth abdominal 

somites fused in male. 

REMARKS.—It is with some hesitation that we 

separate Liocarcinus, including most of the European 

species assigned to Macropipus for the last 20 

years or so, from Macropipus; but as Guinot (1961: 

7) pointed out, the species of Macropipus sensu 

stricto, restricted to three species, share certain 

features that serve to distinguish them readily 

from those species here assigned to Liocarcinus. 

Indeed, 50 years ago Palmer (1927, fig. 1) came 

to the conclusion that "Portunus" tuberculatus [the 

type-species of Macropipus] occupied an isolated 

position from the remainder of the British species 

of "Portunus" [here assigned to Liocarcinus]. Liocarcinus 

has not been used by European zoologists 

but was used by Bennett (1964:65) for L. corrugatus 

from New Zealand. 

Liocarcinus differs from Macropipus as follows: 

(1) The ventral margin of the orbit has one 

narrow, V-shaped incision (Figure 18a); in Macropipus 

(Figure 18b) the ventral margin is open and 

deeply U-shaped (see also Guinot and Ribeiro, 

1962, fig. 8). (2) Liocarcinus lacks iridescent patches 

on the carapace and pereiopods. (3) The ornamentation 

of the cheliped is different: Liocarcinus 

lacks a distal ventral spine on the merus, generally 

lacks a well-developed outer tooth on the carpus 

(such as in L. marmoreus and L. corrugatus), and 

usually has three rather than two longitudinal 

ridges on the dactylus (there may be only two in 

the highly ornamented L. corrugatus). (4) The 

propodi of the second to fourth pereiopods are 

ornamented with a single ventral ridge in Liocarcinus 

(Figure I8g), two distinct ridges in Macropipus 

(Figure \8h). (5) The walking legs are distinctly

84 


FIGURE 18.—Comparison of morphological features. Liocarcinus holsatus (Fabricius), male, cb 

36.5 mm, England: a, orbit, ventral view; c, chela; e, propodus of second pereiopod, lateral 

view; g, propodus of second pereiopod, ventral view. Macropipus rugosus (Doflein), male, cb 36.7 

mm, Guinea-Bissau: b, orbit, ventral view; d, chela; /, propodus of second pereiopod, lateral 

view; h, propodus of second pereiopod, ventral view. 

longer in Macropipus (compare figure 26 (Macropipus 

tuberculatus) with figures 20-25 (Liocarcinus 

spp.) in Christiansen, 1969). 

Liocarcinus, Macropipus, and Bathynectes form a 

progression from generally shallow-water, shortlegged 

species to deep-water, long-legged species. 

One species of Liocarcinus, L. bolivari (Zariquiey 

Alvarez, 1968) apparently is endemic in the Mediterranean. 

Two species, L. holsatus (Fabricius, 

1798) and L. marmoreus (Leach, 1814), are not 

known to occur in the Mediterranean; records of 

these species in the latter area probably are referable 

to L. vernalis (Risso, 1816). 

As Christiansen (1969:60) pointed out, there 

has been some confusion regarding identity of 

Liocarcinus spp. off the northwest African coast, 

due to similarities between Xaiva mcleayi and L. 

pusillus. 

Eleven species of Liocarcinus occur in European- 

Mediterranean waters; Zariquiey Alvarez (1968: 

367) provided a key to the species then known. 

Liocarcinus arcuatus (Leach, 1814), 


Portunus arcuatus.— Capart, 1951:115 [Mauritania].—Monod, 

1956:173 [references]. 

Macropipus arcuatus.—Zariquiey Alvarez, 1968:369, figs. 

116d-h, 120a, 122c, 123a [Spain; references].—Christiansen, 

1969:57, fig. 21, map 15 [Scandinavia]. 

SYNONYMS.—Portunus emarginatus Leach, 1814; 

Portunus guttatus Risso, 1816; Portunus rondeleti 

Risso, 1816; Portunus infractus Otto, 1828. 

DISTRIBUTION.—Eastern Atlantic from the 

North Sea and the British Isles to Mauritania, 

Mediterranean, in depths to 108 m, usually between 

10 and 50 m. 

Liocarcinus corrugatus (Pennant, 1777), 


Portunus corrugatus.—Bouvier, 1911:226 [Mauritania].—Capart, 

1951:120 [Spanish Sahara].—Monod, 1956:174, fig. 

202 [Senegal, Guinea, Gabon, Angola; references].— 

Maurin, 1968b:484 [Spanish Sahara]. 

Macropipus corrugatus.—Guinot and Ribeiro, 1962:31 [Cape 

Verde Islands, Angola].—Ribeiro, 1964:5 [Cape Verde 

Islands].—Forest and Guinot, 1966:59 [Senegal].— 

Maurin, 1968b:484 [Spanish Sahara].—Zariquiey Alvarez, 

1968:372, figs. 13e, 118c-e, 120c, 122d, 123e, 124c 

[Spain; references].—Bas, Arias, and Guerra, 1976, table 

3 [Spanish Sahara].—Tiirkay, 1976b:61 [listed], 66 [Madeira].

NUMBER 306 85 

SYNONYM.—Portunus leachi Risso, 1827. 


to Senegal, Angola, including the Azores, 

Canary Islands and Cape Verde Islands, Mediterranean; 

also Indo-West Pacific (Stephenson, 

1972:23); littoral to 60 m. 

Genus Macropipus Prestandrea, 1833 

Macropipus Prestandrea, 1833:5 [type-species: Portunus macropipus 

Prestandrea, 1833, a subjective junior synonym of 

Portunus tuberculatus Roux, 1830, by tautonymy; gender: 

masculine; name 987 on Official List]. 

Elliptodactylus Doflein, 1904:93 [type-species: Elliptodactylus 

rugosus Doflein, 1904, by monotypy; gender: masculine]. 

DEFINITION.—Carapace broader than long, 

with 5 anterolateral teeth, iridescent patches present 

on surface and on pereiopods. Front with 3 

lobes. Dorsal margin of orbit with 2 closed incisions, 

ventral margin (Figure 186) deeply Ushaped. 

Basal antennal article touching front, 

usually freely movable. Chelipeds unequal, 

shorter than pereiopods 2 to 5; merus with distal 

ventral tooth; carpus with strong inner tooth and 

smaller outer tooth; dactylus (Figure ISd) with 2 

dorsal ridges. Propodus of second to fourth pereiopods 

each with 2 ventral ridges (Figure I8f,h). 

Third to fifth abdominal somites fused in male. 

REMARKS.—As noted above (p. 83), we are 

extremely reluctant to separate the genus Liocarcinus, 

from Macropipus sensu stricto. However, as 

Guinot (1961:7) pointed out in her account of the 

three species of Macropipus, "Ce trois especes ont 

en commun [in contrast with other species then 

assigned to Macropipus] certains caracteres, en particulier 

l'orntmentation de la face dorsale de la 

carapace et des appendices, la disposition des 

epines sur le chelipede, la forme du maxillipede 

externe, la presence de zones iridescentes nacrees 

sur la carapace et les appendices." These features, 

and those discussed under Liocarcinus, warrant the 

recognition of two genera: Liocarcinus, including 

11 relatively short-legged, shallow water species, 

and Macropipus, including 3 relatively longlegged, 

deep-water species from the Mediterranean 

and West Africa. 

Macropipus austral is Guinot, 1961 

Elliptodactylus rugosus.—Barnard, 1950:817 [not Elliptodactylus 

rugosus Doflein, 1904 = Macropipus rugosus]. 

Portunus tuberculatus.—Capart, 1951:117, fig. 41 [part, specimens 

from Angola (Sta A.S. 110) and South-West Africa 

(Sta A.S. 108) only].—Barnard, 1954:123 [listed]; 1955:3 

[listed]. [Not Portunus tuberculatus Roux, 1830 = Macropipus 

tuberculatus.] 

Macropipus australis Guinot, 1961:5, figs. 1, 2, 5, pi. 1, pi. 2: 

fig. 1— Crosnier, 1970:1216. 

Macropipus sp.—Guinot and Ribeiro, 1962:44, 45, figs. 13- 

15, 24a,b, pi. 2: fig. 3, pi. 3: fig. 3. 


Undaunted Material: Angola: Sta 94, 90 m, \6 (L). Sta 95, 

126 m, 56, 6? (W). Sta 96, 162 m, 1$ (L). Sta 103, 90 m, 

20(5, 6? (L). Sta 104, 126 m, 16, 1? (L). Sta 105, 155 m, 36 

(L). 

South-West Africa: Sta 106, 225 m, 1$ (L). 

DESCRIPTION.—Capart, 1951:117; Guinot, 

1961:5. 

Figures: Capart, 1951, fig. 41; Guinot, 1961, 

figs. 2, 5, pi. 1, pi. 2: fig. 1; Guinot and Ribeiro, 

1962, figs. 13-15, 24a,b, pi. 2: fig. 3, pi. 3: fig. 3. 

Male Pleopod: Guinot and Ribeiro, 1962, fig. 

24a,b (South-West Africa). 

Color: "Brick-red, under surface greyish; bright 

iridescent patches as follows: on antero-lateral 

margin between each pair of teeth, and a larger 

crescentic patch behind the last lateral tooth, in 

the smooth grooves between the regions of the 

carapace, anterior surface of the palp of mxp. 3, 

on the smooth distal portion of upper surface of 

arm of cheliped, the spine and other smooth areas 

on wrist, the smooth areas on upper surface of 

hand, and in the groove on upper margin of 

finger" (Barnard, 1950:818). 

BIOLOGY.—Macropipus australis is a sublittoral 

species living on the outer shelf and upper slope 

in depths between 90 and 238 m. Of ten depth 

records available, two are from less than 100 m 

(both 90 m), five are from between 100 and 

200 m (110, 126, 126, 155, and 162 m), and three 

are from depths greater than 200 m (220, 225, 

and 238 m). The only information on the nature 

of bottom on which the species has been found is 

given by Capart (1951); both of the lots he re-

86 

ported were taken on muddy sand in 110 and 

220 m. Ovigerous females have not been collected. 

DISTRIBUTION.—Off West Africa, from the 

coasts of Angola and South-West Africa, in 

depths between 90 and 238 m. Records in the 

literature include: 

Angola: 16°27'S, 11°35'E, 90 m (Crosnier, 1970). 18 mi 

[29 km] WSW of Baia dos Tigres, 16°36'S, 11°27'E, 110 m 

(Capart, 1951; Guinot, 1961). 16°37'S, 11°22'E, 126 m; 

16°41'S, 11°21'E, 162 m; 17°06'S, 11°35'E, 90 m; 17°09'S, 

ll°30'E, 126 m; 17°13'S, 11°27'E, 155 m (all Crosnier, 

1970). 

South-West Africa: 17°18'S, 11°24'E, 225 m (Crosnier, 

1970). 52 mi [84 km] SxW Fort Rock Point, 19°52'S, 12°- 

20'E, 220 m (Capart, 1951; Guinot, 1961). 26°36'S, 14°37'E, 

130 fm (238 m) (Barnard, 1950; Guinot, 1961; Guinot and 


* Macropipus rugosus (Doflein, 1904) 

Portunus sp.—Leach, 1818, in 1817-1818:413.—Monod, 

1970:66. 

Elliptodoctylus rugosus Doflein, 1904:94, figs. 7, 8, pi. 30: figs. 

1-3, pi. 32: fig. 7. 

Portunus tuberculatus.—Capart, 1951:117 [part, not fig. 41, not 

specimens from A.S. 108, 110].—Monod, 1956:180, 632, 

figs. 207-209.—Rossignol, 1957:80, 123 [key].—Longhurst, 

1958:87.—Rossignol, 1962:115.—Maurin, 1968b: 

484, fig. 5.—Le Loeuff and Intes, 1968:44, table 1, figs. 

49, 63. [Not Portunus tuberculatus Roux, 1830.] 

Macropipus rugosus.—Guinot, 1961:2, 9, figs. 1, 4, 7, pi. 2: fig. 

3 [synonymy].—Guinot and Ribeiro, 1962:32-34, 35, figs. 

7-9, 17, 19, 20, 22a,b, pi. 2: fig. 1, pi. 3: fig. 1.—Crosnier, 

1964:34.—Forest and Guinot, 1966:60.—Le Loeuff and 

Intes, 1969:63, 65.—Turkay, 1976a:25 [listed], 37. 

Macropipus.—Voss, 1966:27. 

Portunus.—Maurin, 1968b, fig. 4. 

Portunus (Macropipus) tuberculatus.—Maurin, 1968b:486, 489, 

491 [not Portunus tuberculatus Roux, 1830]. 

Macropipus tuberculatus.—Maurin, 1968b:491.—Bas, Arias 

and Guerra, 1976, table 3. [Not Portunus tuberculatus Roux, 

1830.] 


Sta 68, 70 m, broken shell, 3 juv (W). Sta 83, 156-220 m, 2? 

ov (L). 


Foraminifera, 1 juv (W). Sta 64, 68 m, 1 carapace (L). Sta 

65, 46-49 m, 19 ov (dry) (L). 

Ghana: Sta 23, 42 m, foliate brown to orange bryozoans, 

2$ (L). Sta 24, 35-37 m, dark red bryozoans, 1$ (W). 


Nigeria: Sta 241, 59-63 m, mud and shell, 56*, 69, 9 juv 

(W). 

Cameroon: Sta 259, 59 m, mud and broken shell, 96, 9? 

(L). Sta 260, 46 m, 16*, 19 (L). 

Gerommo Material: Gabon: Sta 202, 100 m, 366*, 30? (W). 

Sta 211, 100 m, 36*, 7° (W). Sta 213, 300 m, 16", 2$ (W). Sta 

228, 300 m, 16*, 29 (W). 

Other Material: Guinea-Bissau: 10°19'N, 16°34'W, 174 m, 

mud, sand, 16 May 1956, Calypso Sta 5, 16* (W). 

Liberia: Off Monrovia, 400 m, 27 Apr 1964, Guinean 

Trawling Survey 14/8, 1 specimen (L). 

Gulf of Guinea: Between Ghana and Sierra Leone, Feb 

and Apr 1964, Guinean Trawling Survey, 16* (L). 

DESCRIPTION.—Guinot, 1961, table 1 (comparison 

with M. tuberculatus (Roux, 1830) and M. 

australis Guinot, 1961; Guinot and Ribeiro, 1962: 

37-42 (comparison with M. tuberculatus). 


Male Pleopod: Monod, 1956, fig. 209 (Senegal); 

Guinot and Ribeiro, 1962, fig. 22a,b (Guinea- 

Bissau). 

Color: According to Capart (1951:118), this species 

is "gris-jaune avec taches roses, prenant un 

aspect nacre apres fixation." His color notes may 

have been based on M. australis, for part of his 

material is referable to that species (see p. 85). 


widths of 8 to 39 mm; ovigerous females 

have carapace widths of 32 to 39 mm. 

BIOLOGY.—Macropipus rugosus inhabits moderate 

depths and seems to prefer depths of less than 

100 m; the shallowest record is of 5 m and the 

deepest is 400 m. 78% of the records for which 

depth is given are from depths of less than 100 m, 

and, in those depths, records are equally divided 

between those in 50 m or less and those in 51- 

100 m. All but two of the Pillsbury specimens came 

from depths of less than 100 m and the species 

was found on bottoms with bryozoans, Foraminifera, 

broken shell, and mud and shell or mud 

and broken shell. Guinot (1961:10) noted that 

"Son habitat le plus commun semble etre le sable 

ou la vase." Forest and Guinot (1966) reported it 

from mud, rocks and calcareous algae, mud, mud 

and sand or broken shell, mud and calcareous 

algae and shells, sand, and algae and calcareous 

algae. Maurin (1968b) found it on mud in 40-

NUMBER 306 87 

60 m, on detrital mud in 50-90 and 200 m, on Mauritania: Bane d'Arguin, 40-60 and 90-100 m; off 

fine detrital muddy sand in 200-400 m, and on Tamzak (as Tamxat), 200-400 m (Maurin, 1968b). 

Senegal: Off Saint-Louis, 35-40 m; off Mboro, 35-40 m 

fluid mud with fine sand or mud and very muddy 

(Maurin, 1968b). 13°01'N, 17°24'W, 51-55 m, and 

fine sand in 35-40 m. 

12°55.5'N, 17°33'W, 65-75 m (Guinot and Ribeiro, 1962; 

Le Loeuffand Intes (1968:44) studied this spe- Forest and Guinot, 1966). SE of lie de la Madeleine, 48 m; 

cies (as Portunus tuberculatus) off the Ivory Coast S border, Fosse de Kayar, 300 m; off Camberene, 50 m (all 

and commented: 

Guinot and Ribeiro, 1962). 

Guinea-Bissau: 10°32'N, 16°53.5'W, 174 m, and 10°19'N, 

La zone des 50 m constitue l'habitat privilegie de l'espece 16°34'W, 60-73 m (Guinot and Ribeiro, 1962; Forest and 

qui accompagne cependant les eaux froides jusqu'a 35 m Guinot, 1966). 

quand elles remontent le long du plateau continental en Sierra Leone: No specific locality, 15-236 m (Longhurst, 

Mai, puis Aout-Septembre-Octobre et meme Janvier-Fev- 1958). 

rier. En fait P. tuberculatus vit dans des eaux bien determinees Liberia: 04°59'N, 09°37'W, to O4°57.5'N, 09°33'W, 156- 

(22°C-16°5C de temperature, 35,55%o a 35,75%o de salinite) 220 m (Voss, 1966). 

qui, en Cote d'lvoire correspondent a l'eau subtropicale. Ivory Coast: No specific locality (Le Loeuff and Intes, 

C'est sans doute, parmi tous les organismes benthiques lit- 1969). Off Grand-Lahou, off Grand-Bassam, 35-100 m (Le 

toraux de cette etude, l'espece la plus stenotherme et la plus Loeuff and Intes, 1968). 

infeodee a une masse d'eau. 

Cameroon: No specific locality, in more than 50 m (Crosnier, 

1964). 

Principe: 01°38'25"N, 07°22'05"E, 31 m (Guinot and 

Ribeiro, 1962; Forest and Guinot, 1966). 

Sao Tome: 00°25'40"N, 06°40'10"E, 50 m (Guinot and 

Ribeiro, 1962; Forest and Guinot, 1966). 

Gabon: 00°38'25"S, 08°46'E, 5 m; 00°38'20"S, 08°48'- 

30"E, 35 m (Guinot and Ribeiro, 1962; Forest and Guinot, 

1966). W of Pointe Panga, 70 m (Rossignol, 1962). 

Congo: Off Pointe-Noire (Rossignol, 1957). 

Zaire: 06°16'S, 12°07'E, 50 m; 06°28'S, 12°05'03 w E 

(Guinot, 1961). 

Angola: 10°S, 13°3O'E, 60-70 m (Guinot, 1961). Bai'a 

Farta, Benguela, 90 m; Baia da Caota, Benguela, 13 m (both 


Tvirkay's (1976a) record of this species from off 

Morocco seems questionable to us, not only because 

it is from so far north (33°37.5'N) but also 

because of the relatively great depth, 952- 

1038 m, at which his material is supposed to have 

been collected. That depth is more than twice as 

great as the deepest record otherwise observed for 

this species. 

Ovigerous females have been collected in June 

and September (Monod, 1956; Forest and 

Guinot, 1966; Pillsbury). 

DISTRIBUTION.—Off West Africa, from Mauritania 

to Angola, in moderate depths, from 5 to 

400 m. Monod (1956) identified this species with 

the Mediterranean M. tuberculatus (Roux) and 

reported material from Mauritania, Senegal, 

Sierra Leone, and Guinea. Guinot (1961) differentiated 

earlier records and provided synonymies 

for each of the three nominal species: M. australis, 

M. rugosus, and M. tuberculatus. Subsequent records 

include: 

West Africa: Gulf of Guinea (Leach, 1818, in 1817-1818; 

Monod, 1970). 

Morocco: 33°37.5'N, 09°02.2'W, 952-1038 m (?) (Turkay, 

1976a). 

Spanish Sahara: No specific locality (Guinot and Ribeiro, 

1962). Between Cabo Barbas and Cabo Blanco, 59-90 m; 


1968b). 25°37.5'N, 14°52'W to 25°39.2'N, 14°51.5'W, 72- 

78 m, and 24°06.3'N, 16°20.2'W to 24°06'N, 16°24.8'W, 


Subfamily PORTUNINAE Rafinesque, 1815 

REMARKS.—This subfamily is characterized by 

the presence of a strong tooth on the upper, outer 

surface of the palm of the cheliped near the upper 

articulation of the carpus. 

Genus Callinectes Stimpson, 1860 

Callinectes Stimpson, 1860:220 [type-species: Callinectes sapidus 

Rathbun, 1896, by designation under the plenary powers 

of the International Commission on Zoological Nomenclature, 

Opinion 712 in Bulletin of Zoological Nomenclature, 

21(5):337, 1964; gender: masculine; name 1613 on Official 

List]. 

REMARKS.—The genus Callinectes has been reviewed 

recently in detail by Williams (1974); not 

all of his distribution records are duplicated here.

88 

Key to West African Species of Callinectes* 

(modified from Williams, 1974) 


1. Granules of carapace (Figure 19a) anterior to epibranchial ridges larger 

and placed wider apart than those posterior to ridges. Epibranchial 

ridges without distinct inflection in middle, almost straight. Submedian 

pair of frontal teeth well developed, usually half or more than half as 

long as lateral pair (measuring from base of lateral notch between teeth). 

First abdominal somite laterally ending in triangular, rather blunt point, 

not sharply pointed nor curved upward. Tips of male gonopods reaching 

beyond suture between thoracic sternite vi and mesially expanded 

sternite vn C. amnicola, new combination 

Granulation of carapace not different anterior and posterior to epibranchial 

ridges. Frontal teeth decidedly unequal in size, submedian pair no more 

than half as long as the lateral pair. Tips of male gonopods falling well 

short of suture between thoracic sternite vi and mesially expanded 

sternite VII 2 

2. Carapace (Figure 196) coarsely granulated. Epibranchial ridges with a 

distinct deflection in the middle. First abdominal somite ending laterally 

in a triangular point, which is neither sharply drawn out nor curved up. 

Male gonopods well separated from each other, not touching or 

crossed C. marginatus 

Carapace (Figure 19

NUMBER 306 89 

Other Material: Senegal: No specific data, 1 juv

90 

species, distinguishing it from other African species 

of the genus. 

An excellent account of the species has been 

given by Williams (1974:775). 

Figures: Rathbun, 1921, fig. 4, pi. 15: fig. 2, pi. 

21, pi. 22: fig. 1; Capart, 1951, fig. 47; Monod, 

1956, figs. 240-243; Williams, 1974, figs. 15, 19c, 

20p, 23a, 27. 

Male Pleopod: Rathbun, 1921, fig. 4e (Zaire); 

Williams, 1974, figs. 19c, 20p (Congo). 

Color: Williams (1974:776) summarized some 

earlier observations on color and noted that "a 

mottled olive coloration persists at least as long 

as 20 yr in some preserved specimens." Capart 

(1951:133) noted that in his material the color 

was "uniforme bleuatre (exemplaires fixes, orange 

clair)." Gauld (1960:69) observed that off Accra 

material of this species is "olive-brown with pale 

legs." De Rochebrune's (1883) color descriptions 

of the adult and juvenile are cited on p. 91. 


widths of 15 to 126.5 mm; the ovigerous 

female has a carapace width of 120 mm. 

REMARKS.—In the recent literature this species 

has usually been given the name Callinectes latimanns 

Rathbun, 1897, but there is an older name 

(or actually two older names) available for it. De 

Rochebrune (1883:168-171) described three new 

species of swimming crabs from Senegal: Neptunus 

amnicola (p. 168), N. edwardsi (p. 169), and N. 

pallidus (p. 170). The identity of these species has 

puzzled many authors. Miers (1886:175) listed 

them in the subgenus Amphitrite de Haan, 1833 

[= Monomia Gistel, 1848] of the genus Neptunus de 

Haan, 1833 [= Portunus Weber, 1795], without 

any comment other than that amnicola perhaps 

should be placed in the subgenus Neptunus. Rathbun 

(1900a: 290) placed amnicola with some doubt 

in the synonymy of Callinectes bocourti A. Milne 

Edwards, and treated edwardsi (p. 289) and pallidus 

(p. 290) as good species of Portunus. Later 

Rathbun (1921:398) reassigned the West African 

specimens that she at first had thought to be C. 

bocourti to C. latimanus, but at that occasion she 

did not deal with amnicola. Monod (1956:215) 

listed all three species of De Rochebrune as "spe- 


cies inquirendae" under Callinectes and stated that 

he had not been able to locate De Rochebrune's 

type material in the Paris Museum. So far as we 

know all later authors ignored De Rochebrune's 

species. 

A study of De Rochebrune's descriptions makes 

clear that Monod was right in considering all to 

belong to the genus Callinectes. The species N. 

amnicola was described from a fullgrown specimen 

(cb 141 mm, cl 76 mm), the other two species 

after juveniles, which makes their identification 

especially difficult. Neptunus amnicola was described 

by De Rochebrune (1883:168-169) as 

follows (only the French description is cited and 

the abbreviated Latin diagnosis is omitted). 

Carapace beaucoup plus large que haute, faiblcment 

bombee, fortement granuleuse, a granulations plus grosses 

et plus espacees dans la region anterieure; lignes epigastriques 

et hypogastriques bien marquees, les epibranchiales 

peu indiquees; bords latero-anterieurs plus courts que les 

latero-posterieurs; premiere dent de forme trapezoidale, obtuse 

au sommet; les suivantes epaisses, brusquement terminees 

en pointe courte et aigue; la neuvieme tres longue, 

etroite, aceree, faiblement incurvee en dehors; front decoupe 

en six dents, les deux medianes petites et obtuses, les mitoyennes 

plus longues, subaigues, les externes tres larges courtes 

arrondies au sommet; apophyse epistomienne grosse, atteignant 

le niveau du front; pattes anterieures tres robustes, 

celles de droite plus volumineuses que celles de gauche; bras 

arme a la partie anterieure de trois dents en forme d'epine 

de Rosier, la premiere faible, les deux suivantes tres fortes; 

partie posterieure du bras aplatie, carenee, a carene obtusement 

denticulee et terminee en haut par une epine courte; 

avant-bras tricarene en dessous et en cote, chaque carene 

terminee par une dent obtuse; mains quadrangulaires, a 

angles fortement granuleux, armees en dessous d'une epine 

courte, en cote de deux tubercules et a Particulation de 

l'avant-bras, d'une dent triangulaire robuste et aigue. 

Teinte generale d'un beau bleu d'outre-mer nuance de 

rose, deux taches arrondies, rouges, de chaque cote et a la 

base de la carapace. 

This description agrees quite well with the 

present species, especially the very characteristic 

granulation of the carapace shows that Neptunus 

amnicola is a synonym of Callinectes latimanus. De 

Rochebrune's remark that the chelae are "armees 

en dessous d'une epine courte" evidently is a 

misprint for "armees en dessus. ..." The fact 

that Callinectes latimanus is a common species in

NUMBER 306 91 

fresh and brackish waters of Senegal makes the 

identification even more likely. Rathbun's 

(1900a) tentative identification of De Rochebrune's 

Neptunus amnicola with what she at that 

time considered C. bocourti A. Milne Edwards thus 

proves to have been correct. The epithet amnicola 

De Rochebrune, 1883, being older than latimanus 

Rathbun, 1897, has priority and thus must be 

used for the species. 

De Rochebrune's second species, Neptunus edwardsi, 

is based on a very small specimen (cb 

26 mm, cl 14 mm), much smaller than N. amnicola 

which measured 141 mm in carapace width and 

76 mm in carapace length. De Rochebrune's 

(1883:169-170) description of N. edwardsi runs as 

follows. 

Carapace plus large que haute, aplatie, subhexagonale, 

tres faiblement granuleuse a la partie anterieure; lignes 

epigastriques et hypogastriques a peine indiquees, lignes 

epibranchiales fortement accusees; bords latero-anterieurs 

plus longs que les latero-posterieurs; premiere dent droite 

epaisse, les suivantes tres obtuses a bords libres tres finement 

creneles; la neuvieme aigue un peu incurvee en haut, relativement 

courte; front decoupe en six dents, les deux 

medianes tres petites, les mitoyennes et les externes egales et 

obtuses; apophyse epistomienne courte, ne depassant pas le 

front; pattes anterieures faibles; bras arme sur le bord anterieur 

de trois epines, et de deux autres epines plus faibles 

a l'extremite articulaire; avant-bras portant sur la face externe 

une crete saillante limitee en dehors par une epine 

courte et accompagnee en dedans d'un tubercule obtus; 

mains anguleuses, a cretes longitudinales assez saillantes 

faiblement rugueuses, et armees de deux petites epines en 

dessus de l'articulation du pouce. 

Teinte generate d'un roux canelle marbre de bleuatre et 

de blanc. 

The description of Neptunus edwardsi fits juvenile 

specimens of Callinectes latimanus very well. In 

specimens of the size of that of Rochebrune's the 

carapace is smooth with a few granules in the 

area before the epibranchial ridge. The lateral 

spine is short, being only somewhat longer than 

the last anterolateral tooth, which gives the carapace 

indeed a "subhexagonal" shape. The lateral 

frontal and inner orbital teeth are rounded 

and similar, while the epistomial spine does not 

show in dorsal view. In juvenile Callinectes gladiator 

the lateral spines are well developed and long. It 

seems quite safe to consider Neptunus edwardsi 

De Rochebrune a synonym of N. amnicola De 

Rochebrune, being based only on a juvenile specimen 

of that species. 

De Rochebrune's third species, Neptunus pallidus, 

proves to belong to the species that is generally 

known as Callinectes gladiator, and as De 

Rochebrune's name is older it has to be known as 

Callinectes pallidus (De Rochebrune) (p. 95). 

The material that Irvine (1947:297) referred to 

Callinectes latimanus proves to belong to Portunus 

validus (p. 103); his figure 202, however, represents 

Callinectes pallidus (p. 95). 

The specimens recorded by Herklots (1851), 

De Man (1883), and Biittikofer (1890) are in the 

collections of the Rijksmuseum van Natuurlijke 

Historie in Leiden. Herklots' specimen, from Elmina 

(as St. George del Mina), Gold Coast 

(Ghana), is a male C. amnicola (Crust. D. 376). De 

Man's material included a specimen of C. amnicola 

from the Congo (Crust. D. 1863), and Biittikofer's 

material included a C. amnicola from Grand Cape 

Mount (Crust. D. 1873), as well as from Lake 

Piso (as Fisherman's Lake), Liberia (Crust. D. 

1864). Monod (1956:214) had listed the records 

by Herklots and De Man as unidentifiable. 

There is a syntype of C. latimanus Rathbun 

(USNM 19877), a female 52 X 114 mm, from 

Lagos, in the collection of the National Museum 

of Natural History, Smithsonian Institution. 

Pauly (1957:75) noted that in the Sakumo 

lagoon, Ghana, this species was common enough 

to support a subsistence trap fishery. 

BIOLOGY.—As pointed out by Williams (1974) 

and earlier authors, this species is primarily estuarine, 

although it is not restricted to estuaries. 

De Rochebrune (1883:171) commented on the 

"habitat exclusivement fluviatile" of this species, 

and said that it is common in the Senegal River 

and disappears when the water becomes salt, to 

reappear when it turns again completely fresh. 

Actually the species can stand brackish water and 

Monod (1956:214) classified it as occurring in 

marine, brackish, and freshwater habitats. Generally 

it is considered to be more characteristic of 

estuaries than C. pallidus, but in the Pillsbury

92 

collections from Lagos harbor, a brackish to fluviatile 

habitat, the latter species (55 specimens) 

far outnumbers C. amnicola (2 specimens). 

Le Loeuff and Intes (1968:44) made the following 

observations on this species: 

C'est un element important de la faune des milieux 

lagunaires de l'ouest Africain. Nous l'avons cependant 

recolte a 9 reprises en mer dans la zone littorale au-dessus de 

30 metres. Les individus etaient tous des femelles de grande 

taille, ovigeres a une exception pres. Ces collectes ont eu lieu 

de juillet a aout, puis de decembre a fevrier. Les deux sorties 

de decembre 1967 a Grand-Bassam ont permis encore de 

ramener onze femelles ovigeres. C. latimanus migre done en 

mer pour pondre; et il y aurait deux saisons de ponte, l'une 

en saison froide, apres la periode des fortes precipitations de 

Mai et Juin, l'autre au moment des petits upwellings, apres 

les crues des fleuves de Septembre et Octobre. 

In a subsequent paper these same authors commented 

(1969:63): 

Le Grabe de lagune Callinectes latimanus Rathbun vient 

egalement pondre en mer; des femelles de grande taille, en 

grande majorite ovigeres ont ete recoltees a plusieurs reprises 

sans qu'il soit possible encore d'affirmer que la sortie en mer 

et la ponte ont lieu a certaines periodes definies ou tout au 

long de l'annee. 


February, March, May, June, July, August, 

October, and December (Monod, 1956; Ribeiro, 

1964; Le Loeuff and Intes, 1968; Williams, 1974; 

Pilbbury). 

DISTRIBUTION.—Callinectes amnicola is an inshore, 

estuarine species, occurring off West Africa 

from Baie de Saint-Jean (19°27'N, 16°22'W), 

Mauritania to at least as far south as a lagoon N 

of Luanda (08°47'S, 13°16'E), Angola. Williams 

(1974) summarized earlier records; distributional 

records from the literature not in Williams include 



Cape Verde Islands: Baia [Porto] de Sal Rei, Boavista 

(Guinot and Ribeiro, 1962; Ribeiro, 1964). Porto da Praia, 

SaoTiago (Ribeiro, 1964). 

Senegal: Senegal River, Faleme River [?], Bakoy [?] [= ? 

Bako, 14°00'N, 15° 16^, or Bakor River 12°55'N, 14°- 

M'W], Bating [?] [= Baling ?] "et tous les Marigots tributaires," 

Casamance River (all De Rochebrune, 1883). 

PresquTIe du Cap Vert; Rosso; Dakar (Sourie, 1954a). 

Dakar; Saloum (Sourie, 1957). 


Gambia: Gambia River (13°28'N, 16°34'W) (De Rochebrune, 

1883). 

Sierra Leone: Sierra Leone River (Longhurst, 1957). 


1906). 

Ivory Coast: Off Sassandra, off Grand-Lahou, off Grand- 

Bassam, 15-30 m (Le Loeuff and Intes, 1968). 

Ghana: Off Accra, in shallow water (Gauld, 1960). Sakumo 

lagoon (Pauly, 1975). 

Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). Elechi 

Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell, 

1979). 


* Callinectes marginatus 

(A. Milne Edwards, 1861) 

FIGURES 19b, 20a,b 

Neptunus diacanthus.—Pechiiel-Loesche, 1882:287.—Osorio, 

1892:199. [Not Portunus diacantha Latreille, 1825 = Callinectes 

sapidus Rathbun, 1896]. 

PCallinectes diacanthus.—Gruvel, 1913:78, 106 [this species or 

C. amnicola?]. 

PCallinectes diacanthus var. africanus.—Gruvel, 1913:168 [listed] 

[this species or C. amnicola ?]. 

Callinectes marginatus.—Gruvel, 1913:168 [listed].—Capart, 

1951:134, fig. 48.—Monod, 1956:208, figs. 238, 239.— 

Rossignol, 1957:82.—Buchanan, 1958:20, 23.—Gauld, 

1960:69.—Guinot and Ribeiro, 1962:48.—Rossignol, 

1962:116.—Ribeiro, 1964:6.—Forest and Guinot, 1966: 

65.—Monod, 1967:180, pi. 15: fig. 6 [no material].—Bott, 

1968:169.—Williams, 1974:722 [part, not figs. 3, 18a,b, 

20a, 22b, 27] [review of genus].—Hartmann-Schroder and 

Hartmann, 1974:19. 

Callinectes.—Bayer, 1966:102. 

SYNONYM.—Callinectes diacanthus var. africanus A. 

Milne Edwards, 1879. 

MATERIAL EXAMINED.—Pillsbury Material: Nigeria: 

Sta 1, Lagos harbor, washed ashore, 1 carapace (L). 

Fernando Poo: Sta 257, shore, 26, 1 juv (L). 

Annobon: Sta 281, shore, 26, 2 juv (W). 

Other Material: Cape Verde Islands, Porto Grande, Sao 

Vicente, U.S. Eclipse Expedition, 1 juv (W). Porto da Praia, 

Sao Tiago (as La Praya, Santiago), Jul 1883, Talisman, 16* 

(W). 

Ghana: Mouth of Hwini River, Takoradi, 27 Jul 1961, 

Bane and Richards, Id (W). Virgins Pool, Takoradi, 27 Jul 

1961, Bane and Richards, 16" (W). 

Zaire: Banana, mouth of Congo River, American Museum 

Congo Expedition 1909-1915, Aug 1915, H. Lang, 56", 

49 (W). Data same, Jul-Aug 1915, 1? (W).

NUMBER 306 93 

Angola: Luanda (as St. Paul de Loanda), U.S. Eclipse 

Expedition, 26 (W). 

DESCRIPTION.—Carapace (Figure \9b) somewhat 

elevated, generally flatter than in C. amnicola, 

width slightly more (2.1 to 2.3 times) than 

twice as great as length (lateral spines included). 

Granulation dense, uniform, granules anterior to 

epibranchial ridge as dense and large as those 

posterior to ridge; granules finer than in C. amnicola 

but distinct. Epibranchial ridge between cervical 

groove and lateral spine of carapace not 

straight, with strong inflection near midlength. 

Length of metagastric area equal to or shorter 

than posterior width, latter half anterior width. 

Front with submedian teeth very narrow, short, 

much shorter than half length outer teeth; latter 

bluntly triangular, extending somewhat beyond 

more widely triangular inner orbital angles. Epistomial 

spine distinct in dorsal view, extending 

about as far forward as outer frontal teeth. Anterolateral 

margins of carapace more strongly 

convex than in C. amnicola. Anterior teeth triangular, 

curved somewhat forward, blunt; posterior 

teeth also triangular, more sharply pointed. Lateral 

spine stout, about twice as long as preceding 

tooth. 

Carinae on cheliped of adults distinct, granular. 

Chelae of adults slender, occasionally somewhat 

swollen in basal part of fingers, less strongly 

so than in C. amnicola. Basal tooth of dactylus of 

large chela slightly enlarged. 

First somite of abdomen in both sexes terminating 

laterally in triangular or blunt point, neither 

prolonged nor turned upward. 

First gonopods of male short, failing by far to 

reach end of sternite of third pereiopod. Distal 

part of gonopods somewhat curved externally, 

curve gradual, gonopods generally not touching 

(Figure 20a,b). 

Figures: Rathbun, 1921, fig. 2, pi. 19: fig. 1, pi. 

20: fig. 1; Capart, 1951, fig. 48; Monod, 1956, 

figs. 238, 239. 

Male Pleopod: Rathbun, 1921, fig. 2d (Zaire). 

Color: Capart (1951:135) noted that "coloration 

apres fixation non uniforme, des marbrures plus 

foncees sur la carapace et sur P.I." Rossignol 

(1957:82) remarked that the species can best be 

distinguished from C. amnicola by its color, namely 

"carapace marbree." 


widths of 15 to 97 mm. 

REMARKS.—Since Rathbun (1896:358, 359) referred 

African specimens of this species to the 

East American Callinectes larvatus, the specific 

identity of African and American specimens of 

the two forms have not been questioned. After 

having examined the types of Callinectes marginatus 

(A. Milne Edwards, 1861) from Gabon, Callinectes 

larvatus Ordway, 1863, from Key West, the Tortugas, 

the Bahamas and Haiti, and Callinectes 

diacanthus africanus A. Milne Edwards, 1879, from 

the Cape Verde Islands, Rathbun (1897a: 149) 

synonymized the three species and used the oldest 

available name, marginatus, for it. Rathbun's views 

on the taxonomy and nomenclature of the species 

have since been followed by most authors, including 

Williams (1974) in his fundamental revision 

of the genus Callinectes. 

However, a comparison of our African material 

with American specimens identified with C. marginatus 

showed a number of seemingly constant 

differences, which in our opinion necessitates the 

recognition of two distinct species. The American 

species should be given the name Callinectes larvatus 

Ordway, 1863; the well-known name Callinectes 

marginatus (A. Milne Edwards, 1861) is retained 

for the African species. The differences 

between these species are the following: 

1. In C. marginatus (Figure 196) the granulation 

of the carapace before the epibranchial ridges is 

as dense as that behind these ridges, the granules 

there are of the same size as those in the posterior 

part of the carapace. In C. larvatus the granules 

before the epibranchial ridges are larger and 

placed much wider apart than those behind the 

ridges. 

2. The cervical groove behind the orbits is 

much deeper in C. larvatus than in C. marginatus. 

3. The ventral inner orbital angle in C. marginatus 

is narrower than in C. larvatus. 

4. The male gonopods are short in both species, 

but in C. marginatus (Figure 20a,b) they are more

94 

slender and the distal part that is curved outward 

is relatively longer and less abruptly bent; the 

two gonopods as a rule do not touch. In C. larvatus 

(Figure 20c,d) the distal outward curved part of 

the gonopod is shorter and more abruptly bent; 

the two gonopods touch each other in the midline 

of the body. 

BIOLOGY.—Monod (1956:210) indicated that 

this species is a shallow water form that prefers a 

sandy or sandy mud bottom, and is found in 

marine habitats. The latter in contrast to C. 

amnicola, which is known from fresh, brackish, and 

salt water, and C. pallidus, which Monod reported 

from salt and brackish waters. Buchanan (1958: 

20, 23) indicated C. marginatus as a characteristic 

epifaunistic species of the inshore fine sand community. 

Rossignol (1962:116) described the species 

as a "forme littorale (eaux saumatres, lagunes)," 

but possibly based this on the literature 

in which more than one species is confused under 

the name C. marginatus. Gauld (1960) mentioned 

the species to be caught intertidally. Guinot and 

Ribeiro (1962:48, 49) listed an abundant material, 

mostly from sandy beaches, either taken by 

hand or by beach seines; the species was mentioned 

once by them from the hull of a ship, and 


once from under rocks near the shore; there is no 

indication that any of this material is estuarine. 

Ribeiro (1964:6) reported the species from "zona 

intercotidal." Forest and Guinot (1966:65) mentioned 

the species from a sand beach (0-9 m) and 

from intertidal rocks. West African material of 

Callinectes marginatus was listed by Williams (1974) 

from "rock pool" and "beach." The Pillsbury specimens 

were collected from sandy beaches (Sta 1 

and 281) and from the seashore (Sta 257). All 

these data confirm Monod's statement that the 

species is marine rather than estuarine. The data 

are too few, however, to definitely fix the habitat 

of the species; specimens have been reported from 

the mouth of the Congo River (Rathbun, 1921; 

and p. 92), and from the mouth of the Hwini 

River (p. 92), localities that might be estuarine. 

Off West Africa ovigerous females have been 

collected in February, May, June, September, 

October, and December (Guinot and Ribeiro, 

1962; Ribeiro, 1964; Forest and Guinot, 1966; 

Williams, 1974). 

DISTRIBUTION.—The species is known from the 

Cape Verde Islands and Port Etienne, Mauritania 

to central Angola. Monod (1956) summarized 

earlier records; distribution records from the 

literature not in Monod include the following. 

FIGURE 20.—Position and shape of gonopods: a,b, Callinectes marginatus (A. Milne Edwards), 

male, cb 90 mm, Congo; c,d, Callinectes larvatus Ordway, male, cb 116 mm, Jamaica.

NUMBER 306 95 


Cape Verde Islands: Baia das Gatas, Sao Vicente (Guinot 

and Ribeiro, 1962; Ribeiro, 1964). Porto da Praia, SaoTiago 

(Guinot and Ribeiro, 1962; Ribeiro, 1964; Williams, 1974). 

Senegal: Dakar (Williams, 1974). 

Gambia: Near Gunjur (Williams, 1974). 

Guinea: Fotoba, 6 miles [10 km] W of Conakry (Williams, 

1974). 

Sierra Leone: Murray Town (Williams, 1974). 

Ghana: Off Accra (Buchanan, 1958). Apam (Gauld, 

1960). 

Togo: No specific locality (Gruvel, 1913). 

Dahomey: No specific locality (Gruvel, 1913). 

Nigeria: Lagos harbor (Williams, 1974). 

Fernando Poo: No specific locality (Williams, 1974). 

Principe: No specific locality (Forest and Guinot, 1966). 

Sao Tome: No specific locality (Forest and Guinot, 1966; 

Williams, 1974). Iogoiogo (Osorio, 1892; Forest and Guinot, 

1966). Morro Peixe (Forest and Guinot, 1966; Williams, 

1974). 

Annobon: 01°24'S, O5°37'E, shore (Bayer, 1966). 

Gabon: No specific locality (Williams, 1974; syntypes). 

Congo: No specific locality (Williams, 1974). Loango 

(Pechiiel-Loesche, 1882). Djeno, lagoon (Rossignol, 1957). 

Near Pointe-Noire (Rossignol, 1962; Williams, 1974). 

Zaire: Mouth of Congo River near Banana (Williams, 

1974). 

Angola: No specific locality (Forest and Guinot, 1966). 

Luanda (Forest and Guinot, 1966; Bott, 1968; Williams, 

1974). Baia do Lobito (Guinot and Ribeiro, 1962; Williams, 

1974). Praia da Rocha, Benguela (Guinot and Ribeiro, 

1962). Between Cacuaco and Lobito-Benguela (Hartmann- 

Schroder and Hartmann, 1974). 

* Callinectes pallidus (De Rochebrune, 1883), 


FIGURE 19C 

Neptunus pallidus De Rochebrune, 1883:170.—Miers, 1886: 

175.—Balss, 1921:59 [listed].—Monod, 1956:215. 

Neptunus diacanthus.—De Man, 1883:150 [part].—Biittikofer, 

1890:466, 487 [part]. [Not Portunus diacantha Latreille, 1825 

= Callinectes sapidus Rathbun, 1896]. 

Portunus pallidus.—Rathbun, 1900a:290. 

Callanectes Sp.?.—Irvine, 1932, fig. 13. 

Callinectes Gladiator.—Irvine, 1932:15. 

Callinectes lalimanus.—Irvine, 1947, fig. 202 [not C. latimanus 

Rathbun, 1897 = C. amnicola (De Rochebrune, 1883)]. 

CallinecUs gladiator.—Irvine, 1947:298.—Frade, 1950:11, 

26.—Bruce-Chwatt and Fitz-John, 1951:117.—Capart, 

1951:130, fig. 46.—Monod, 1956:205, figs. 236, 237.— 

Rossignol, 1957:82.—Sourie, 1957:13, 51.—Longhurst, 

1958:87.—Gauld, 1960:69.—Guinot and Ribeiro, 1962: 

48.—Rossignol, 1962:116.—Crosnier, 1964:32.—Forest 

and Guinot, 1966:64.—Monod, 1967:180, pi. 15: fig. 3 [no 

material].—Bayer, 1966:98.—Le Loeuff and Intes, 1968: 

40, p. 2 of table 1, figs. 50, 61; 1969:64, 65.—Monod, 

1970:66.—Uschakov, 1970:439, 455 [listed].—Ejike, 1973: 

253.—Williams, 1974:735, figs. 5, 18c, 20b, 22c, 24 [review 

of genus]. 

Callinectes.—Bayer, 1966:98. 

Lupa Smythiana.—Monod, 1970:66. 

SYNONYM.—Callinectes tumidus var. gladiator Benedict, 

1893. 


Coast: Sta 43, surface, 29 (W). 

Nigeria: Sta 2, Lagos harbor, surface, 41 specimens (L,W). 

Sta 226: Lagos harbor, surface, 16* (W). Sta 228, Lagos 

harbor, surface, 16", 2$ (W). Sta 229, Lagos harbor, surface, 

46\ 6$ (W). Sta 250, 24 m, brackish water, mud, 9

96 

seum Congo Expedition, 1909-1915, Aug 1915, H. Lang, 

1

NUMBER 306 

the carapace very short, but in juveniles of the 

present species that spine is very elongate and 

straight, being much longer than the anterolateral 

teeth. The rest of the description also fits the 

present species, except for the measurements, for 

even C. amnicola specimens with cl 9 mm are much 

broader than 15 mm; one wonders whether 

15 mm for the width is not a typographical error 

for 25 or 19 mm. Because both C. amnicola and C. 

gladiator are estuarine species, and both are common 

in Senegal, it seems most likely that Neptunus 

pallidus belongs to one of these two species; the 

shape of the lateral spines and the granulation of 

the carapace makes clear that it can only be 

identified with C. gladiator Benedict, 1893. As 

Benedict's name gladiator was proposed 10 years 

after De Rochebrune's publication of pallidus the 

latter name has priority and has to be used for 

the species. 

As the type specimen of N. pallidus is no longer 

extant, we propose to fix the identity of Neptunus 

pallidus De Rochebrune, 1883, by selecting as its 

neotype the holotype of Callinectes tumidus gladiator 

Benedict, 1893 (USNM 14879), a male measuring 

29 X 63 mm (one lateral spine broken). In this 

way N. pallidus and C. gladiator become objective 

synonyms. The name pallidus is quite appropriate 

for the species, as preserved material usually is of 

a very pale color. 

The specimens reported as Neptunus diacanthus 

by De Man (1883) and Buttikofer (1890) are in 

the collection of the Rijksmuseum van Natuurlijke 

Historie, Leiden. Both author's accounts 

were based on material of C. amnicola as well as C. 

pallidus. De Man's material included two specimens 

of C. pallidus from the Congo (Crust. D. 374, 

1876), and Biittikofer's collection included one 

specimen of C. pallidus from Liberia (Crust. D. 

1871). 

As pointed out by Capart (1951:130, 132) the 

figures that Irvine (1932, 1947) published with 

his "Callinectes gladiator" and "Callinectes latimanus" 

have been interchanged: the figures indicated by 

Irvine in 1932 (fig. 13) as "Callanectes Sp.(?)" and 

in 1947 (fig. 202) as Callinectes latimanus actually 

represent C. pallidus. Irvine's text of "Callanectes 

Sp.(?)" and C. latimanus refers to Portunus validus 

(p. 103). 

BIOLOGY—Williams (1974:736) has summarized 

most of the available ecological information 

on this species, an inshore form usually found on 

mud, sand, or sand and shell in depths of less 

than about 30 m. It is less often encountered in 

estuaries than C. amnicola, although the majority 

of the Pillsbury specimens were taken in brackish 

water in Lagos harbor during outgoing tides. 

Monod (1956:208) indicated this species as being 

found in the sea and in brackish water. 

Sourie (1954b) found the species on bottoms of 

fine shelly sand with mud and Molgula hannensis 

Peres in 2-7 m in the Baie de Dakar; Uschakov 

(1970) found it in turbid water in depths of less 

than 20 m on unstable mud off Guinea. Gauld 

(1960) noted that off Ghana it was very common 

from low water mark to 18 m, usually on sand. 

Bruce-Chwatt and Fitz-John (1951) noted that in 

Nigeria it lives on muddy bottoms of creeks. 

Le Loeuff and Intes (1968:40) made the following 

observations on this species from their studies 

off the Ivory Coast: 

Extremement abondant et frequent au-dessus de 30 

metres, ce Callinectes est tres littoral car il ne descend pas 

au-dela de 35 m; son aire de repartition est tres stable: C. 

gladiator ne se deplace pas quelle que soit la saison et est done 

extremement tolerant aux variations de temperature, salinite, 

teneur en oxygene des eaux. CAPART (1951) a signale C. 

gladiator comme vivant sur des fonds vaseux. En fait l'espece 

est indifferente a la nature du substrat. Tous les sediments 

lui conviennent, sables ou vases. Sur les petits fonds C. 

gladiator domine nettement en nombre d'individus les autres 

Portunidae coders: Portunus inaequalis, Cronius ruber, Neptunus 

validus et contribue sans doute a repousser le maximum 

d'abondance des deux premiers au-dela de 25 metres et a 

limiter la densite de N. validus. 

Williams (1974) noted that earlier authors had 

commented on the aggressiveness of this species, 

a characteristic of most Portunidae. 

Ovigerous females have been recorded in January, 

February, March, April, May, June, October, 

and December (Capart, 1951; Forest and 

Guinot, 1966; Monod, 1956; Williams, 1974; 

Pillsbury). 

DISTRIBUTION.—West Africa, from numerous 

97

98 

localities between Baie de Saint-Jean (19°27'N, 

16°22 / W), Mauritania, to Bafa do Lobito (12°- 

20'S, 13°34'E), Angola, generally in depths of 

35 m or less. Williams (1974) reported material 

from numerous localities and summarized earlier 

records. Records not in Williams include the 

following. 


Senegal: Lac de Nguer (as N'Guer), marigots de MBao 

[= Grand Mbao], Thionk [Pointe], Leybar (all De Rochebrune, 

1883). Anse de Hann, Baie de Dakar, 2-7 m (Sourie, 

1954b), Saloum (Sourie, 1957). 

Guinea: No specific locality, less than 20 m (Uschakov, 

1970). 


1958). 

Liberia: No specific locality (Frade, 1950). 

Ghana: No specific locality, low water mark to 18 m 

(Gauld, 1960). 


1969). Off Sassandra, off Fresco, off Grand-Lahou, off 

Jacqueville, off Grand-Bassam, 15-35 m (Le Loeuff and 

Intes, 1968). 

Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). Lagos 

harbor, 06°28'N, 03°23'E, surface (Bayer, 1966). 7 miles 

[11 km] off Lagos harbor (Ejike, 1973). 



Coast: Sta 46, 38-42 m, mud with dense Jullienella, 1 juv 

(W). Sta 48, 22 m, 4o\ 39 (L, W). 

Annobon: Sta 275, 9-69 m, rubble of coralline algae, 2 

juv (L). 

Other Material: Ghana: Accra, 1868-1869, M. Sintenis, 1

NUMBER 306 99 

FIGURE 21.—Gonopods of two West African portunids. Cronius 

ruber (Lamarck), male, cb 72 mm, Pillsbury Sta 48: a, 

sternal view; b, apex, enlarged. Portunus validus Herklots, 

male, cb 151 mm, Pillsbury Sta 252: c, sternal view; d, apex, 

enlarged. 

imens in the color pattern and ornamentation of 

the chelae. In adults from West Africa the dark 

color of the movable finger extends proximally 

on the cutting edge, but is not expanded dorsally 

on the outer margin at the base of the finger. In 

specimens from Brazil, the dark color extends 

proximally as in the West African population, 

but at the base of the finger it expands dorsally 

so the finger appears to have a large, dark spot 

basally on the outer margin. This enlarged proximal 

spot is present in juveniles from both areas. 

The enlarged proximal spot is present in some 

eastern Pacific specimens, absent in others. 

The ornamentation of the propodus of the 

claw, which appears to be similar in American 

specimens from both coasts, differs significantly 

in the one large specimen from the Gulf of Guinea 

available to us (male, cb 72 mm, Pillsbury Sta 48). 

In American specimens the prominent longitudinal 

ridges on the propodus of the claw, especially 

the lower one on the outer face and the two 

ventral ones, are composed of short, transverse 

lines of tubercles or ridges which are inflated 

along the main axis of the longitudinal ridges so 

that the latter appear to be broad ridges with 

tuberculate edges. In our large specimen from 

West Africa the transverse lines of tubercules 

forming the main ridges are not inflated in the 

larger chela but are distinct throughout the 

length of the longitudinal ridge, as illustrated by 

Monod (1956, fig. 220), whereas in the smaller 

chela the ventral longitudinal ridges are ornamented 

with tubercles scattered in no apparent 

pattern, and there is no trace of the transverse 

lines of granules present on the major chela. 

In American specimens of C. ruber, the tubercles 

forming the lower, outer ridge on the chela also 

are inflated, so that the ridge appears smooth 

along its midline, with numerous marginal tubercles 

dorsally, fewer ventrally. In our West African 

specimen, that ridge is made up of 2 distinct rows 

of tubercles with a space between them. 

Although the male pleopods of specimens from 

both sides of the Atlantic appear to be similar in 

shape and ornamentation, that of our largest 

specimen from West Africa is more curved laterally 

and has the apex directed anterolaterally 

rather than anteriorly. In available specimens 

from Brazil the pleopod is less strongly curved 

and the apex is directed anteriorly. 

BIOLOGY.—This is a shallow water species occurring 

on a variety of bottom types off West 

Africa; it has been recorded from the intertidal 

zone to a depth of 69 (9-69) m, but the majority 

of depth records are from depths of 30 m or less. 

Sourie (1954a) characterized it as a sand dwelling 

species associated with the understone fauna of 

rocky shores; he also found it in the Dakar region 

(1954b) on coarse shelly sand, bottom with Area 

and Pyura, and noted that juveniles were found 

on fine shelly sand, with mud, bottom with Molgula 

hannensis Peres in 2-7 m. Off Guinea, Uschakov 

(1970) found it on the inferior mesolittoral 

part of rocky shores, associated with red algae 

and Padina. Buchanan (1958) characterized it as 

a member of the active epifauna, inshore fine 

sandy community, in 3-8 fm (5-15 m) off Ghana. 

Longhurst (1958) found it on shelly sand in 14- 

25 m off Sierra Leone. Le Loeuff and Intes (1968) 

reported that off the Ivory Coast it reached its 

maximum abundance at a depth of 35 m, but 

that on one transect in May, 465 individuals were 

taken at a depth of 50 m. They characterized the 

species as eurythermic and euryhaline and noted

100 

that it occurred on all types of bottom. Forest 

and Guinot (1966) reported several lots from the 

offshore islands of the Gulf of Guinea from the 

following habitats and depths: calcareous algae 

in 4-5 to 10-12 m (7 stations), calcareous algae 

and shells in 5-6 m, sand, algae, and calcareous 

algae in 8-30 m, mud, calcareous algae and shells 

in 31 m, and mud and sand in 35 m. The Pillsbury 

specimens were taken on mud bottom with Jullienella 

in 38-42 m and in the coralline algae habitat 

off Annobon in 9-69 m. 


recorded in January, March, and September (Capart, 

1951; Monod, 1956; Guinot and Ribeiro, 

1962; Ribeiro, 1964). Off West Africa, juveniles 

are more often found than adults; of more than 

80 lots recorded by Monod (1956) more than 50 

were made up of juveniles. 

DISTRIBUTION.—Atlantic-East Pacific. It is 

widely distributed in the eastern Pacific (Garth 

and Stephenson, 1966), as well as in the western 

Atlantic (Rathbun, 1930). Off West Africa it has 

been recorded from localities between Mauritania 

and Angola; shallow water to a depth of 69 m, 

usually between 10 and 30 m. Monod (1956) 

summarized earlier West African records and 

reported material from Mauritania (questionable 

data), Senegal, Guinea, Sierra Leone, Ivory 

Coast, Ghana, Gabon, and Angola; since 1956 it 

has been recorded from the following. 


Cape Verde Islands: Porto da Furna, Brava, 6-20 m and 

Baia de Porto Grande, Sao Vicente, 4-6 m (Guinot and 

Ribeiro, 1962; Ribeiro, 1964). 

Guinea: Conakry (Uschakov, 1970). 


1958). 


1969). O5°O5'N, 04°59.5'W, 22 m (Voss, 1966). Off Sassandra, 

off Fresco, off Grand-Lahou, and off Grand-Bassam, 

15-60 m (Le Loeuffand Intes, 1968). 

Ghana: Off Accra, 3-8 fm (5-15 m) (Buchanan, 1958). 

Off Accra, Winneba, and Tenkpobo (as Tenpobo), low water 

mark to 12 m (Gauld, 1960). 

Principe: O1°37'2O W N, 07°21'45"E, 35 m; 01°38'25"N, 

07°22'05"E, 31 m; between Ponta da Mina and Ilheu Santana, 

8-10 and 10-12 m; and in front of Praia Pequena, 5- 



Sao Tome: 00°20'N, 06°46'E, 10 m; O0°25'15"N, 06°. 

43'05"E, 8-30 m; in front of Ponta Oquedelrei, 6 m; off 

Ponta Diogo Nunes, 4-5 m; off Sao Tome, 5 m (Forest and 


Annobon: N of San Antonio, 9 m (Forest and Guinot, 

1966). 

Congo: Pointe-Indienne and Baie de Pointe-Noire (Rossignol, 

1957, 1962). 

Angola: Baia Farta, Benguela, 22-28 m, and Baia do 

Lobito, intertidal (Guinot and Ribeiro, 1962). 

Genus Portunus Weber, 1795 

Portunus Weber, 1795:93 [type-species: Cancer pelagicus Linnaeus, 

1758, by selection by Rathbun, 1926:75; see Opinion 

394, International Commission on Zoological Nomenclature, 

1956; gender: masculine; name 986 on Official 

List}. 

Portunus Fabricius, 1798:325, 363 [type-species: Cancer pelagicus 

Linnaeus, 1758, by selection by Latreille, 1810:94, 422; 

gender: masculine; name 410 on Official Index}. 

Lupa Leach, 1814:390 [type-species: Cancer pelagicus Linnaeus, 

1758, by monotypy; gender: feminine; name 411 

on Official Index}. 

Lima Leach, 1814:429 [possibly an erroneous spelling of 

Lupa; type-species: Cancer pelagicus Linnaeus, 1758, by 

present selection; gender: feminine; name 413 on Official 

Index}. 

Lupania Rafinesque, 1818:272 [substitute name for Lupa 

Leach, 1814; type-species: Cancer pelagicus Linnaeus, 1758; 


Neptunus de Haan, 1833:3, 7 [type-species: Cancer pelagicus 

Linnaeus, 1758, by selection by Miers, 1886:172; gender: 

masculine; name 414 on Official Index}. 

Achelous de Haan, 1833:3, 8 [type-species: Portunus spinimanus 

Latreille, 1819, by monotypy; gender: masculine]. 

Amphitrite de Haan, 1833:3, 8 [invalid junior homonym of 

Amphitrite O. F. Miiller, 1771 (Polychaeta); type-species: 

Portunus gladiator Fabricius, 1798, by selection by Miers, 

1886:172; gender: feminine]. 

Pontus de Haan, 1833:3, 9 [type-species: Portunus (Pontus) 

convexus de Haan, 1833, by monotypy; gender: masculine]. 

Lupea H. Milne Edwards, 1834:445 [erroneous spelling of 

Lupa Leach, 1814]. 

Monomia Gistel, 1848:viii [substitute name for Amphitrite de 

Haan, 1833; type-species: Portunus gladiator Fabricius, 


Posidon Herklots, 1851:3 [invalid junior homonym of Posidon 

Illiger, 1801 (Crustacea); type-species: Portunus (Posidon) 

validus Herklots, 1851, by monotypy; gender: masculine]. 

Xiphontctes A. Milne Edwards, 1873b: 157 [type-species: Amphitrite 

vigilans Dana, 1852, by selection by Rathbun, 1930: 

33; gender: masculine].

NUMBER 306 101 

Hellenus A. Milne Edwards, 1874, in 1873-1881:210, 221 

[type-species: Achelous spinicarpus Stimpson, 1871, by selection 

by Rathbun, 1930:33; gender: masculine]. 

Lupocycloporus Alcock, 1899a:31, 32, 44 [type-species: Achelous 

whitei A. Milne Edwards, 1861, by monotypy; gender: 

masculine]. 

Cycloachelous Ward, 1942:79 [type-species: Lupea granulata H. 

Milne Edwards, 1834, by monotypy; gender: masculine]. 

Portunus hastatus (Linnaeus, 1767) 

Lupa hastata.—Barrois, 1888:14. 

Neptunus (Amphitrite) hastatus.—Lenz and Strunck, 1914:278. 

Neptunus hastatus.—Capart, 1951:125, fig. 44 [part, not specimen 

from Guinea].—Chapman and Santler, 1955:374.— 

Monod, 1956:203, figs. 232-235 [no material]. 

Portunus hastatus.—Figueira, 1960:8.—Zariquiey Alvarez, 

1968:384, figs. 125d,e, 126c, 128a,b [Spain; references].— 

Turkay, 1976b:61 [listed], 64, pi. 1: figs. 1, 2. 

SYNONYMS.—Cancer ponticus Herbst, 1790; Portunus 

Dufourii Latreille, 1819; Eriphia prismaticus 

Risso, 1827; Neptunus hastatus rubromaculatus Steinitz, 

1932. 


Other Material: Madeira: Canical, 30 fm (55 m), seine, 18 

Apr 1916, A. C. de Noronha, 16* (W). Machico Bay, seine, 

16 Apr 1916, A. C. de Noronha, lcJ (W). Same data, 30 Jun 

1916, 2$ (W). Same data, 5 Nov 1921, 126\ 11? (6 ov) (W). 

Angola: 8 mi [13 km] W of Rio Cuanza, 09°20'S, 13°04'E, 

20-22 m, muddy sand, 31 Jan 1949, Expedition Oceanographique 

Beige, Sta A. S. 116, lcJ, 1$ ov (Brussels). 


Figure: Capart, 1951, fig. 44. 

Male Pleopod: Monod, 1956, figs. 232-235 (Lebanon) 

. 



of the ovigerous females from Madeira range 

from 18 to 26.5 mm, that from Angola is 42 mm. 

REMARKS.—In large males, the abdomen is narrower 

than in P. inaequalis; the last somite is 

triangular in P. hastatus, ovate in P. inaequalis. In 

P. hastatus the width of the terminal somite of the 

male abdomen is half or more than half the width 

of the preceding somite; in P. inaequalis it is less 

than half the width of the preceding somite. 

Finally, the ventral incision of the orbit is a closed 

V in P. hastatus, an open V or U in P. inaequalis, as 

noted by Monod (1956:203). 

The two specimens from Angola are those reported 

by Capart (1951), who correctly identified 

them with Portunus hastatus. We have compared 

these specimens with material of this species from 

several Mediterranean localities, as well as with 

the material from Madeira reported above, and 

we could find no constant differences in specimens 

from different areas. The specimens from 

Angola at first sight appear to be different from 

those from the Mediterranean, for, as figured by 

Capart (1951, fig. 44), the dorsal sculpturing of 

the carapace is much less pronounced on them. 

In addition, the lateral spine of the carapace is 

shorter than in most specimens of P. hastatus from 

the Mediterranean we have examined, and the 

chelae appear stouter with a slightly shorter movable 

finger. 

There are relatively few records for this species 

outside of the Mediterranean and we have tried 

to compile all of them here. One record, that of 

Miers (1886:175), as Neptunus (Amphitrite) hastatus, 

from Tenerife or Isla de la Gomera, Canary Islands, 

appears to be based on specimens of Portunus 

sayi. Miers noted that his specimens were 

"very prettily mottled with purple on a yellowish 

ground" (this is the color of P. sayi as described 

by Rathbun, 1930:42) and that the same jar 

contained specimens of Planes minutus. Miers' was 

the only record for this species from the Canary 

Islands that we could find. 

BIOLOGY.—Portunus hastatus is a sublittoral species, 

occurring on sandy bottoms subtidally to a 

depth of at least 55 m. The two specimens from 

Angola were taken on muddy sand in 20-22 m. 


recorded in January (Angola) and November 

(Madeira). 

DISTRIBUTION.—Eastern Atlantic, where it occurs 

primarily in the Mediterranean. Outside of 

that sea it has been recorded from the following: 

Azores: Ponta Delgada, Uha de Sao Miguel (Barrois, 

1888; Lenz and Strunck, 1914). Ilha do Faial (as Fayal) 

(Chapman and Santler, 1955). Ilha do Pico (Figueira, 1960).

102 

Madeira: Funchal, from fish market, Prainha[?], and 

Ponta de Sao Lourenco, 15-20 m (all Tiirkay, 1976b). 

Angola:8mi[13km]WofRioCuanza,09°20'S, 13°O4'E, 

20-22 m (Capart, 1951). 

* Portunus inaequalis (Miers, 1881) 

Neptunus hastatus.—Capart, 1951:125 [part, not fig. 44].— 

Rossignol, 1957:123 [key]. [Not Portunus hastatus (Linnaeus, 

1767).] 

Neptunus inaequalis.—Monod, 1956:198, figs. 225-231.—Buchanan, 


69.—Rossignol, 1962:116.—Le Loeuff and Intes, 1968:40, 

44, table 1, figs. 49, 61; 1969:63, 65.—Tiirkay, 1976b:61 

[listed] 64, pi. 1: figs. 3,4. 

Portunus inaequalis.—Guinot and Ribeiro, 1962:47.—Ribeiro, 

1964:6.—Forest and Guinot, 1966:63. 


Coast: Sta 47, 37 m, bottom with Jullienella, 126*, 7$, 3 juv 

(W). 

Ghana: Sta 24, 35-37 m, dark red bryozoans, 1 juv (L). 

Sta 26, 27 m, shell bottom (scallops), 36*, 19 ov (L). Sta 27, 

33 m, 1$ (L). 

Nigeria: Sta 246, 37 m, lcJ (L). Sta 248, 33 m, Id, 1$ ov 

(L). Sta 250, 24 m, brackish water, mud, 16*, 19 (L). Sta 252, 

30 m, mud, Id, 19 (L). 

Other Material: Ghana: Chorkor, near Accra, Dec 1950, R. 

Bassindale, Id (L). 

DESCRIPTION.—Carapace more than twice as 

broad as long, regions distinct, surface roughened. 

Gastric region with long patch of raised granules, 

with curved branches extending onto protogastric 

region. Metagastric region with 2 raised submedian 

lobes ornamented with enlarged tubercles. 

Cardiac region with 2 raised submedian lobes, 

often united posteriorly in form of V- Epibranchial 

ridges raised, granular, sinuous. Mesobranchial 

region with raised prominence mesially, 

flanked laterally by 1 or more oblique granular 

ridges. Hepatic region with curved, granular 

ridges. Frontal teeth triangular, submedians 

smallest, sharpest, second pair larger and more 

triangular than blunt inner orbital spines. Interantennular 

spine small, visible in dorsal view. 

Anterolateral margin with 9 sharp, separate 

spines, outer orbital (first anterolateral) blunter 

than next spine; lateral spine long, at least as long 

as space occupied by preceding 4 or 5 spines, 

slender, curved upward and forward. Small patch 


of iridescence present between bases of spines. 

Posterolateral margins unarmed. Chelae long, 

slender, merus longer than carapace, subequal. 

Dorsal surface of merus irregular, anterior margin 

with 4 or 5 spines, occasionally 1 smaller spine 

distally, posterior and ventral margin each with 

1 distal spine. Carpus with inner and outer spine. 

Propodus with 1 spine proximally and 2 spines 

distally on upper margin. Fingers slender, shorter 

than palm. Pereiopods 2-5 with iridescent lines. 

Merus of fifth leg unarmed. Sternum and abdomen 

smooth. 



Sierra Leone). 

Color: The following observations on color were 

provided by Monod in his list of material of this 

species: 

Tres marbres; en alcool une tache rouge plus ou moins 

marquee sur le dactyle p5 [1956:200, Senegal] . . . ; rouge 

brique; tache carminee sur la partie posterieure du dactyle 

p5; ligne carmin sur la crete inf. du propode pi, se prolongeant 

sur le doigt fixe; face interne du dactyle egalement 

carmin [1956:202, Guinea] . . . ; traces de tache rouge au 

dactyle p5 [1956:202, Ghana]. 

The dark red color on the dactylus of the last leg 

also is characteristic of P. hastatus (Linnaeus). 


widths of 16 to 53 mm; the measurable 

ovigerous female has a carapace width of 25 mm. 

BIOLOGY.—Portunus inaequalis is a shallow water 

species, living near shore to a depth of 60-73 m, 

generally in depths of less than 40 m. It can 

tolerate waters of reduced salinity. The Pillsbury 

took it in brackish water on mud in 24 m off 

Nigeria, and Longhurst (1958), who reported it 

from Sierra Leone, characterized it as an estuarine-shelf 

species; he found it on stones, sand, and 

shelly sand in 10-25 m. Sourie (1954b) reported 

it from fine shelly sand with mud, bottom with 

Molgula hannensis Peres, in 2-7 m in the Baie de 

Dakar. Buchanan (1958) found it in the sandy 

silt community in 8-20 fm (15-37 m) off Ghana. 

Guinot and Ribeiro (1962) reported it from algae 

and rocks and on sand in 10 m. It was reported 

from a variety of bottom types by Forest and


Guinot (1966): shell in 20-25 and 27 m; mud, 

shells and Cidaris in 60-73 m; sand or muddy 

sand and Foraminifera in 5 and 21-27 m; calcareous 

algae in 5, 6, and 12 m; rocks and coral in 

3-10 m; sand and calcareous algae in 4-5 m; and 

from a sand beach, 0-4 m. 

Le Loeuff and Intes (1968:44) found that off 

the Ivory Coast the species lived on sand or 

muddy sand and was rarely observed on mud. 

There it generally lives at depths between 30 and 

35 m, not going below 40 m, although during 

periods of upwelling or cooling part of the population 

was found at 15-20 m; in warm periods it 

generally was found below 25 m. 

Only two of the Pillsbury samples were taken at 

stations on mud; the species was found on shell 

bottom with scallops, on bottom with dark red 

bryozoans, and bottom with Jullienella. 


March, April, May, June, July, September, and 

November (Monod, 1956; Guinot and Ribeiro, 

1962; Ribeiro, 1964; Forest and Guinot, 1966; 

Pillsbury). 

DISTRIBUTION.—Off West Africa, from Madeira 

and from Senegal to Angola, in shallow 

water, from shore to a depth of 60-73 m, generally 

in less than 40 m. Monod (1956) recorded material 

from the Cape Verde Islands, Senegal, 

Guinea, Sierra Leone, Ghana, Gabon, and Principe. 

Since 1956 it has been recorded from the 


Madeira: No specific locality (Tiirkay, 1976b). 

Cape Verde Islands: Baia de Porto Grande and Baia de 

Calheta, 8-16 m, Sao Vicente; Baia da Faja di Agua (? = 

Porto da Faja) and Porto da Furna, 4 m, Brava; Tarrafal 

and Porto da Praia, Sao Tiago; and Tarrafal, Sao Nicolau 

(all Guinot and Ribeiro, 1962; Ribeiro, 1964). 




1958). 


1969). 05°00'N, 05°28.5'W, 27 m; 05°03'N, 05°25"W, 20-25 

m; and 05°02.5'N, 05°25'W, 21-27 m (all Forest and Guinot, 


Jacqueville, and off Grand-Bassam, 8-40 m (Le Loeuff and 

Intes, 1968). 

Ghana: Off Accra, 8-20 fm (15-37 m) (Buchanan, 1958); 

10-32 m (Gauld, 1960). 

Principe: Between Uheu Santana and Ponta Capitao, 12 

m (Forest and Guinot, 1966). 

Sao Tome: Baia de Ana de Chaves; Praia de Santa 

Catarina, W coast, 3-10 m; in front of Ponta Oquedelrei, 6 

m; Morro Peixe, 0-4 m; Ilheu das Cabras, 4-5 m; off Sao 

Tome, 5 m (all Forest and Guinot, 1966). 

Annobon: 01°26'20"S, 05°36'25"E, 22-25 m (Forest and 


Congo: W of Pointe-Noire (Rossignol, 1962). 

Gabon: 00°38'25"S, 08°46'E, 5 m (Forest and Guinot, 

1966). 

Angola: Baia de Caota, Benguela, 10 m (Guinot and 


*Portunus validus Herklots, 1851 

FIGURES 2lc,d, 22, 23a,c,e,g 

Lupa Cranchiana White, 1847a:27 [nomen nudum; under 

Neptunus sanguinolentus].—Monod, 1970:66, 72. 

Neptunus validus.—Pechiiel-Loesche, 1882:287.—Bvittikofer, 

1890:466, 487, fig. on p. 465.—Johnston, 1906:862.— 

Frade, 1950:11, 26.—Capart, 1951:123, fig. 43.—Monod, 

1956:1%, fig. 224.—Rossignol, 1957:80, 123 [key], pi. 2: 

fig. 6.—Buchanan, 1958:20.—Longhurst, 1958:87.— 

Gauld, 1960:69.—Rossignol, 1962:115.—Crosnier, 1964: 

32, 87, 90, 92, 98, 105, 106, 110, 112, 120, 121.—Crosnier 

and Berritt, 1966:68, 100, 101, 102, 109, 123, 127, 131, 

132, 136.—Le Loeuff and Intes, 1968:40, 46, table 1, figs. 

49,61; 1969:64, 65. 

Callanectes sp.?.—Irvine, 1932:14. 

Callinectes Gladiator.—Irvine, 1932: fig. 9 [not C. gladiator 

Benedict, 1893 = C. pallidus (De Rochebrune, 1883)]. 

Callinectes gladiator.—Irvine, 1947: fig. 203 [not C. gladiator 

Benedict, 1893 = C. pallidus (De Rochebrune, 1883)]. 

Callinectes latimanus.—Irvine, 1947:297 [not Callineclus latimanus 

Rathbun — C. ammcola (De Rochebrune, 1883)]. 

Portunus validus.—Monod, 1967:180, pi. 15: fig. 1 [no locality]; 

1970:66, 72.—Uschakov, 1970:439, 455 [listed].— 

Baron, 1975a:3-13 [physiology]; 1975b: 103, figs. 1-4. 


Sta 241, 59-63 m, mud and shell, 19 (L). Sta 252, 30 m, 26, 

2$(W). 

Other Material: Liberia: Off St. Paul River, Monrovia, 3- 

6 fm (5-11 m), 4 Mar 1953, G. C. Miller, lcj (W). Liberia, 

1881, J. Biittikofer, 2$ (L). 

Ghana: Elmina (as St. George-del-Mina), 1840-1855, 

H. S. Pel, lectotype, l

104 

1964, B. de Wilde-Duyfjes, ld\ 29 (L). Kribi, beach seine, 9 

Mar 1964, B. de Wilde-Duyfjes, 1$ (L). Same, 10 Aug 1964, 

B. de Wilde-Duyfjes, 1$ (L). 

Angola: No specific locality, 1879, P. Kamerman, 2$ (L). 

DESCRIPTION.—A. Milne Edwards, 1861:321; 

Capart, 1951, fig. 124. 

Figures: A. Milne Edwards, 1861, pi. 29: fig. 1; 

Capart, 1951, fig. 43; Monod, 1956, fig. 224. 

Color: The carapace is rather uniformly greenish 

gray with some brown tinges. A large conspicuous, 

white triangular spot is present on each side 

just before the middle of the posterolateral margin. 

This white spot is surrounded by a ring of 

darker gray than is on the rest of the carapace. 

The posterior margin also is white. The uniform 

color of the carapace contrasts strongly with the 

brightly marbled upper surface of the chelipeds 

(Figure 22). The upper half of each cheliped is 

very dark purple with numerous rather large 

white spots, which at some places are confluent. 

The lower half of the cheliped is white. The same 

marbled purple color is present on the upper half 

of the merus and carpus of the next 3 legs, the 

purple color and size of the white spots being 

very variable. The propodus and dactylus are 

greenish gray as is the carapace. The fifth leg has 

the upper surface of all segments except the dactylus 

purplish to greenish with white spots. The 

dactylus is gray dorsally with a white streak along 

FIGURE 22.—Portunus validus Herklots, female, 

Pillsbury St&2U. 


the basal half of the outer margin. The lower 

surface of the body and legs is uniformly pale. 

Irvine (1947:297, under Callinectes latimanus) described 

the color of this species as follows: The 

carapace "is khaki in colour, with a bluish tinge. 

The claws are blue. . . . The legs are also 

blue. ..." 

Capart (1951:124) gave the following color 

account of this species: "Carapace bleu-vert, avec 

deux taches blanches sur les aires branchiales." 

Rossignol (1957:81) gave the following color 

description: 

Le 6 et la 9 sont identiques. Face dorsale: 

a. Carapace: marron ou brun violace plus ou moins irrise, 

avec deux taches arrondies d'un blanc creme sur chaque aire 

branchiale en arriere des dents posterieures. Sou vent, une 

autre petite tache blanche de forme imprecise juste au-dessus 

de la precedente. Extremite de la dent posterieure blanche. 

b. Chelipedes: brun-violace marbre de blanc (merus, 

carpe, propode, dactyle). Face interne de la pince: moitie 

inferieure 4e la paume blanche: pouce bleu turquoise. 

c. Pattes: bleu turquoise avec des taches blanches sur le 

merus (P2, P3). P4 et P5: merus et carpe brun-violace marbre 

de bleu et de blanc. Dactyle de P5 beige legerement rose. 

Face ventrale de la carapace et face externe des pinces: 

blanc creme. 

MEASUREMENTS.—The carapace width of our 

specimens varies between 62 and 180 mm. In the 

literature males with carapace lengths of 46 to 

114 mm, carapace widths of 85 to 205 mm, 

females with carapace lengths of 41.5 to 93 mm, 

widths 82 to 190 mm, and ovigerous females with 

carapace lengths of 83 to 97 mm, widths of 150 

to 170 mm, have been reported. 

REMARKS.—Portunus validus is remarkable in 

having the carapace finely and evenly granular, 

almost evenly convex, lacking sharp ridges or 

grooves. It falls within the genus Portunus sensu 

lato, as currently defined, but a careful revision 

of this heterogeneous genus may necessitate the 

recognition of a new genus for the preoccupied 

genus Posidon Herklots, 1851, originally established 

for Portunus validus. Judging from the accounts 

of Stephenson and Campbell (1959) and 

Stephenson (1972), Portunus validus shows little 

similarity to any of the many Indo-West Pacific 

species of the genus, and it shows little affinity


with any of the American species of the genus 

(Rathbun, 1925; Garth and Stephenson, 1966). 

Indeed, a comparison of P. validus with P. pe- 

lagicus, the type-species of Portunus, reveals some 

interesting differences in addition to the ornamentation 

of the carapace and the structure of 

the male pleopod. In P. validus the interantennular 

spine is very short and does not extend to the 

front; it is not at all visible in dorsal view (Figure 

23a), whereas in P. pelagicus the entire spine, 

including its base, is visible in dorsal view, and 

the apex of the spine extends well beyond the 

level of the frontal spines (Figure 236). In P. 

validus there are two strong spines on the posterior 

margin of the merus of the claw, one subdistal 

(Figure 23c), whereas in P. pelagicus (and in Callinectes 

as well) the subdistal spine is absent (Figure 

23

106 

terminates in a bifid tip. The male abdomen has 

been well figured by A. Milne Edwards (1861). 

This species has some limited commercial importance 

and great commercial potential. Baron 

(1975b) has summarized observations on catch, 

size distribution, sex ratio, and growth of claw in 

specimens taken commercially off Senegal and 

Gambia. 

A male collected off Elmina (St. George-del- 

Mina), Ghana, by H. S. Pel, between 1840 and 

1855, is here selected as the lectotype of the 

species (Crust. D.395). It is in the collection of the 

Rijksmuseum van Natuurlijke Historic 

Irvine (1932) mentioned two species of Callinectes 

from Ghana; one he indicated correctly as 

C. gladiator Benedict (= C. pallidus (De Rochebrune)), 

the other was referred to as "Callanectes 

Sp.(?)" In 1947 Irvine again described and figured 

the two species and identified the latter as 

Callinectes latimanus Rathbun. The true identity of 

the two forms puzzled many subsequent authors, 

because Irvine had interchanged the figures illustrating 

these two species, as was first pointed out 

by Capart (1951:130, 132). The figures said to 

represent Callinectes sp. (Irvine, 1932, fig. 13) and 

Callinectes latimanus Rathbun (Irvine, 1947, fig. 

202) actually are those of C. gladiator (= C. pallidus). 

Irvine's text of "Callanectes Sp.(?)" (1932:14) 

and his figure 9, as well as his later text of C. 

latimanus (1947:297) do not represent Callinectes 

latimanus as was thought by Capart (1951), but 

Portunus validus. The great size of the species, the 

shape of the front, the fact that the merus of the 

chelipeds bears two distal teeth on the posterior 

margin, all this proves the identity of Irvine's 

material with P. validus. Even the peculiar white 

spots on the carapace are correctly indicated in 

the drawing, which, however, shows the carapace 

grooves somewhat overaccentuated. Monod 

(1956:196) already recognized Irvine's figures of 

"Callinectes gladiator" as representing Portunus validus, 

but evidently did not realize that they did 

not belong to the text of C. gladiator, but to that 

of the other species of "Callinectes." Williams 

(1974:737) recognized that something was wrong 

and referred to Capart and Monod, but con- 


cluded that "since the features [of Irvine's figures] 

are sketchy, it is best to accept the author's designation 

with allowance for error." 

BIOLOGY.—This species inhabits shallow water 

in depths mostly between 0 and 40 m. Among the 

almost 50 depth records for this species known to 

us there are only two records of the capture of 

specimens that originate from hauls made entirely 

in water deeper than 40 m, viz. 50 m (Crosnier, 

1964:91, 92) and 50-55 m (Crosnier and Berritt, 

1966:135, 136); more than two-thirds of the records 

are from depths between 10 and 30 m. 

Rossignol (1957:81) therefore is mistaken in giving 

the range as 0-70 m and sublittoral (from 50 

to 99 m). 

The species is frequently taken with beach 

seines. Buchanan (1958) found it in the inshore 

fine sandy community in 5 to 15 m off Accra. 

Crosnier (1964) characterized it as a warm water 

species, living in depths between 0 and 30 m off 

Cameroon. Le Loeuff and Intes (1968:46) made 

the following observations on the species off the 

Ivory Coast: 

II est assez frequent dans les traits mais jamais en grandes 

quantites: 10 individus constituent un maximum. Espece 

cotiere qui ne descend pas au-dessous de 35 m, N. validus 

tolere de gros ecarts des facteurs physico-chimiques des eaux, 

ainsi que tous les types de sediments puisqu'on le peche aussi 

bien sur les sables de Fresco que sur les vases de Grand- 

Lahou. 

Off Guinea, Uschakov (1970) found the species 

in depths of less than 20 m on unstable mud in 

turbid water. It has been reported from bottoms 

consisting of shells and sand, sand and Gorgonaria, 

sand, muddy sand, fine sand and mud, 

sandy mud, mud, mud and Foraminifera, or on 

shells and mud. 


March, September, and December (Capart, 1951; 

Monod, 1956; Baron, 1975a). 

Irvine (1947:297) commented on the "ferocious 

disposition" of this species and the harm it can 

do to fishes caught in fish pots. In Ghana the 

crabs are caught and sold as food. According to 

Irvine (1947:297) "there is reason to believe that 

they migrate to the sea at times and it is very


probable that they do so during the breeding 

season." 

DISTRIBUTION.—Off tropical West Africa, from 

Mauritania to Angola, sublittoral to about 55 m. 

Monod (1956), who summarized earlier records, 

reported material from Mauritania, Senegal, 

Guinea, Sierra Leone, Liberia, Ivory Coast, 

Ghana, and the Congo. In addition, the species 


West Africa: No specific locality (White, 1847a; Balss, 

1921; Monod, 1967, 1970). 

Senegal: 13°25'N, 16°55'W, 15 m; 12°25'N, 17°15'W, 

26 m; 12°20'N, 17°5O'W (all Baron, 1975a). Saint-Louis; off 

the Casamance River (Baron, 1975b). 

Gambia: Off the Gambia River (Baron, 1975b). 

Guinea: No specific locality, less than 20 m (Uschakov, 

1970). 


1958). 


1906). 



Jacqueville, off Grand-Bassam, 8-35 m (Le Loeuff and Intes, 

1968). 

Ghana: Elmina [as S. Jorge da Mina] (Frade, 1950). Off 

Accra, 3-8 fm (5-15 m) (Buchanan, 1958); from shallow 

water to 25 m (Gauld, 1960). 

Togo: 06°15'N, 01°53'E, 15-17 m; O6°07'N, 01°53'E, 50- 

55 m (Crosnier and Berritt, 1966). 

Dahomey: 06°10'40"N, 02°02'E, 35 m; 06°10'30"N, 

02°02'E, 35-40 m; 06°17'N, 02°22'E, 20 m; 06°18'30"N, 

02°24'E, 16 m; 06°21'N, 02°37'E, 12-14 m; 06°16'N, 

02°37'E, 22 m; 06°12'30"N, O2°37'E, 35-36 m (Crosnier 

and Berritt, 1966). 

Cameroon: 03°55'N, 09°00'E, 50 m; 03°47'N, 

09°12'30"E, 20 m; 03°39'30"N, 09°14'E, 30 m; 03°32'N, 

09°34'E, 13-15 m; 03°27'N, 09°38'30"E, 12 m; 03°15'N, 

09°51'30"E, 9 m; 03°09'N, 09 o 44'30 w E, 30 m; 02°41'30"N, 

09°49'30"E, 20 m; 02°28'N, 09°45'30"E, 7-10 m (Crosnier, 

1964). 

Congo: Loango (Pechuel-Loesche, 1882). Off Pointe- 

Noire (Rossignol, 1957, 1962). 

Portunus vocans (A. Milne Edwards, 1878) 

Neptunus means.—Monod, 1956:194, figs, 222, 223 [Cape 

Verde Islands].—Forest, 1959:19 [Annobon, Sao Tome]. 

Portunus vocans [Neptunus vocans].—Guinot-Dumortier and Dumortier, 

1960:120, 144 [table 2] [islands of Gulf of Guinea; 

discussion of stridulation]. 

Portunus [Neptunus] vocans.—Guinot-Dumortier and Dumortier, 

1960:142 [listed]. 

Portunus vocans.—Forest and Guinot, 1966:62, fig. 4 [Annobon, 

Sao Tome].—Tiirkay, 1976b:61 [listed], 66, pi. 2: 

figs. 1, 2 [Madeira]. 

DISTRIBUTION.—An insular species in the Atlantic. 

In the eastern Atlantic it is known from 

Madeira, the Cape Verde Islands, and Annobon 

and Sao Tome Islands in the Gulf of Guinea. It 

also has been recorded from Ascension Island in 

the central Atlantic and from several western 

Atlantic localities; in moderate depths, 7-10 m to 

about 309 m (Rathbun, 1925). 

Genus Thalamita Latreille, 1829 

Thalamita Latreille, 1829:33 [type-species: Cancer admete 

Herbst, 1803, by monotypy; gender: feminine; name 195 


Thalamita poissonii (Audouin, 1826) 

Portunus Poissonii Audouin, 1826:84 [Egypt]. 

Thalamita integra var. africana Miers, 1881a:218. 

Thalamita africana.—Sourie, 1954b: 151.—Monod, 1956:186, 

figs. 213-217.—Rossignol, 1957:124 [key].—Guinot and 

Ribeiro, 1962:46.—Crosnier, 1962:116 [discussion].—Ribeiro, 

1964:5.—Forest and Guinot, 1966:61.—Uschakov, 

1970:455 [listed].—Stephenson, 1972:19 [key], 44 [listed]. 

Thalamita integra africana.—Stephenson and Hudson, 1957: 

319 [key]. 

Thalamita poissonii.—Holthuis and Gottlieb, 1958:89, 118, pi. 

2: fig. 10a,b [Eastern Mediterranean references].—Stephenson, 

1976:23. 

Thalmita africana.—Hartmann-Schroder and Hartmann, 

1974:19 [erroneous spelling]. 


Other Material: Gulf of Guinea, no specific locality, 1956, 

Calypso, 1

108 

life of this species. Monod (1956) gave the following 

notes in his list of material: "rougeatre" (p. 

187); "doigts des chelipedes (en alcool) rouges 

avec les apex noirs, la couleur se poursuivant, en 

lisere, le long des bords internes, dentigeres" (p. 

188). 




REMARKS.—Crosnier (1962:117) noted the similarity 

between the male pleopods of T. poissonii 

and T. africana. Stephenson (1976:23) synonymized 

T. africana with T. poissonii and noted that 

"these [specimens from Angola] are identical in 

all respects with T. poissonii." We have compared 

our material from Angola with specimens from 

other areas and can find no differences. 

This is one of a few Indo-West Pacific species 

of brachyurans to occur off West Africa; it has 

also colonized the eastern Mediterranean via the 

Suez Canal (Holthuis and Gottlieb, 1958). 

BIOLOGY.—Thalamita poissonii is a shallow water 

species, occurring off West Africa from the littoral 

zone to a depth of about 30 m. Rathbun (1921: 

404) noted that it was "the most common species 

in the quiet stretches of the bay near town of St. 

Paul de Loanda, and easily caught at low tide on 

the many submerged sand flats. Their behavior 

is much the same as that of Callinectes . . . but 

their habitat there is typically marine." Sourie 

(1954b) found this species on fine shelly sand, 

bottom with Molgula hannensis Peres, in the Baie 

de Dakar. The specimens collected by the Calypso 

were taken on the following types of bottom 

(Forest and Guinot, 1966): mud, 18 m; calcareous 

algae, 11 m; sand, 0-4 m; sand, algae, and calcareous 

algae, 8-30 m; mud and calcareous algae, 

4 and 5 m. 


collected in January, February, March, April, 

June, August, September, and October, suggesting 

that there the species breeds all year (Rathbun, 

1921; Monod, 1956; Forest and Guinot, 

1966). 

DISTRIBUTION.—Eastern Atlantic and Indo- 

West Pacific. In eastern Atlantic, from eastern 


Mediterranean and off West Africa at scattered 

localities between the Canary Islands and Angola, 

in depths between the littoral zone and about 

30 m; Indo-West Pacific from localities between 

Japan and Madagascar and the Red Sea (Crosnier, 

1962). Monod (1956) summarized the West 

African literature and reported material from 

localities off Mauritania, Senegal, Guinea, and 

Angola. Records since then include the following. 

Cape Verde Islands: Bala de Porto Grande, Sao Vicente, 

4-6 and 8 m (Guinot and Ribeiro, 1962; Ribeiro, 1964). 

Senegal: Baie de Dakar (Sourie, 1954b). 

Guinea: No specific locality (Uschakov, 1970). 

Principe: In front of [Cais de] Santana, llm (Forest and 


Sao Tome: 00°25'15"N, 06°43'05"E, 8-30 m; Baia de 

Ana de Chaves, 5 m; in front of Ponta Diogo Nunes, 4 m; 

Morro Peixe, 0-4 m (all Forest and Guinot, 1966). 

Gabon: 00°40'S, 08°46'25"E, 18 m (Forest and Guinot, 

1966). 

Angola: Baia de Luanda (Guinot and Ribeiro, 1962). 

Luanda (as St. Paul de Loanda) (Stephenson, 1976). Between 

Cacuaco and Lobito-Benguela (Hartmann-Schroder 

and Hartmann, 1974). 

Family GERYONIDAE Colosi, 1923 

GERYONIDAE Colosi, 1923:249. 

EASTERN ATLANTIC GENERA.—Due to the uncertainty 

of the taxonomic status of this family 

and of the genera sometimes assigned to it, we 

are unable to give here a reliable list of the eastern 

Atlantic geryonid genera. Guinot (1971:1077) included 

in the Geryonidae the following three 

genera that are represented in the eastern Atlantic: 

Geryon, Paragalene, and Progeryon; of these only 

Geryon is known from tropical West Africa. 

Paragalene Kossmann (1878:253). Type-species: 

Paragalene neapolitana Kossmann, 1878, a junior 

subjective synonym of Eriphia longicrura Nardo, 

1869, by monotypy; gender: feminine; name 341 

on Official List. 

Progeryon Bouvier (1922:71). Type-species: Progeryon 

paucidens Bouvier, 1922, by monotypy; gender: 

masculine. 

We are not convinced that these two genera 

are correctly assigned to the Geryonidae, but as


we have not had the opportunity to study this 

question in detail, we follow Guinot, at least for 

the time being. The genus Platychelonion (p. 155), 

placed by Guinot (1971:1078) with some doubt 

in the Geryonidae, in our opinion finds a better 

place in the Xanthidae. 

EASTERN ATLANTIC SPECIES.—The above-mentioned 

three genera are represented in the eastern 

Atlantic area by 6 species; only two of these have 

been found in tropical West African waters. The 

other four are as follows: 

Geryon tridens Kr0yer, 1837. North Atlantic from 

N Norway and Iceland to the British Isles and 

the North Sea; records from the Bay of Biscay, 

the Mediterranean and Morocco need to be verified 

and at least partially pertain to G. longipes; 

32-690 m, usually deeper than 100 m (Christiansen, 

1969). 

Geryon longipes A. Milne Edwards, 1881. Bay of 

Biscay, Morocco, Mediterranean; 600-1370 m 


Paragalene longicrura (Nardo, 1869). Mediterranean 

(Algeria, Malta, Naples, Adriatic and Aegean 

Seas) and Madeira; 20-ca. 120 m (Tiirkay, 

1976b). 

Progeryon paucidens Bouvier, 1922. Off Morocco, 

2165 m (Bouvier, 1922). 

Monod (1956:337) recognized a single West 

African species: Geryon quinquedens Smith, synonymizing 

G. affinis with it. In the present publication 

two tropical West African species are distinguished: 

G. affinis and G. maritae, new species; G. 

quinquedens is thought to be restricted to East 

American waters. 

Only one species of Geryon, G. maritae, was taken 

by the Pillsbury in West African waters. 

Genus Geryon Kr0yer, 1837 

Geryon Kr0yer, 1837:10, 20, 21 Itype-species: Geryon tridens 

Kr0yer, 1837, by original designation and monotypy; 


Chalaepus Gerstaecker, 1856:118 [type-species: Cancer trispinosus 

Herbst* 1803, by monotypy; gender: masculine]. 

REMARKS.—The place of the genus Geryon in 

the system of the Brachyura has been subject to 

many different interpretations by zoologists. 

Kr0yer (1837:19, 20), when describing his new 

genus remarked that it should be placed in the 

"famille des Cyclometopes" of H. Milne Edwards 

(1834:363) although in some respects it showed 

some resemblance to the Catometopes, and because 

of the shape of the dactyli of the last pair 

of pereiopods ("if0lge de bageste Tarsers Beskaffenhed") 

thought it should be placed in the former 

of the two tribes (Canceriens and Portuniens) 

in which H. Milne Edwards had subdivided his 

Cyclometopes. Miers (1886:223) placed Geryon in 

the subfamily Carcinoplacinae of the family Ocypodidae. 

Ortmann (1894:685; 1899:1176) left it 

in that subfamily, which he made the nominal 

subfamily of a new family Carcinoplacidae; he is 

followed in this by Stebbing (1905:35). A Milne 

Edwards and Bouvier (1894:41) first placed the 

genus in the family Canceriens sensu lato (= 

Cancridae + Xanthidae), but later (1899:34) assigned 

it to the family Galenidae. Alcock (1899c: 

84) accepted this disposition, but made the Galeninae 

a subfamily of the Xanthidae. Doflein 

(1904:105), without comment, ranged Geryon in 

the Potamidae. Colosi (1923:249) partly agreed 

with Doflein and erected a new (monotypic) family 

Geryonidae, which he placed near the Potamidae. 

Several authors, like Balss (1927:1020, 

1933a:298), Bouvier (1940:261), Stephensen 

(1945:222), Monod (1956:337), Guinot and Ribeiro 

(1962:62), Zariquiey Alvarez (1968:388), 

and Crosnier (1970:1216) placed the genus in the 

family Xanthidae; others, e.g., Rathbun (1937: 

265), Barnard (1950:290) and Capart (1951:173) 

kept it in the Goneplacidae. In recent years the 

family Geryonidae Colosi, 1923 (often attributed 

to Beurlen, 1930) is recognized by many authors. 

It then is usually placed between the Xanthidae 

and Goneplacidae (Balss, 1957:1654; Christiansen, 

1969:83; Glaessner, 1969:R524) or closer 

to the Goneplacidae than to the Xanthidae 

(Kaestner, 1970:349). Serological investigations 

by Leone (1949:284, 1951:44-48) suggested that 

the Geryonidae are more closely related to the 

Xanthidae and Portunidae on the one hand, than 

to the Cancridae, Ocypodidae, Grapsidae, and 

Majidae on the other. All in all, Stebbing's (1893:

110 

93) remark, made 80 years ago, still holds good: 

"The genus Geryon, Kr0yer, 1837, may claim a 

passing notice as one of those instances in which 

systematic arrangement finds itself at fault. It is 

sometimes placed among the Cyclometopa and 

sometimes among the Catometopa.. . . That, on 

the theory of the evolution of different groups 

from a common stem, such inosculant forms are 

almost sure to occur, has long been recognized. 

Darwin himself humorously admits that while as 

a theorist he delighted in coming across them, as 

a naturalist engaged in classification he found 

them an unmitigated nuisance." 

In studying the present material, we were 

struck by the great resemblance in general and in 

detail of Geryon with the Portunidae. In practically 

every point our specimens resemble Portunidae, 

except in the shape of the last pereiopods which 

lack the paddle-shape of the distal segments. 

Were the distal segment of these pereiopods 

broadened, no one would have hesitated to assign 

the genus to the Portunidae. The resemblance 

with species of Benthochascon is especially striking. 

The resemblance of the Geryonidae to the Portunidae 

is much closer, we think, than to either 

Xanthidae or Goneplacidae. In our opinion the 

family Geryonidae is close to the Portunidae, 

perhaps between Portunidae and Xanthidae, 

which incidentally would also agree with the 

serological findings by Leone (1949, 1951) and a 

study of the larvae oi Geryon tridens by Ingle (1979: 

230), who noted that "the larvae of G. tridens 

possess many portunid features...." The taxonomy 

of the Brachyrhynchous crabs, especially 

at the family level, is still highly unsatisfactory 

and a thorough revision is badly needed. 

The strong resemblance of the chelae of Geryon 

to those of Portunidae is shown in Figure 24. As 

in Portunidae the basal tooth of the dactylus of 

the large chelipeds in Geryon is blunt, enlarged 

and often directed somewhat backward. The 

other teeth are often flanked by small teeth; in 

Geryon the larger teeth usually have a single 

smaller tooth separating them, while in portunids 

the larger teeth often are flanked each with two 

teeth. 


FIGURE 24.—Chelae: a, Geryon maritae, new species, paratype, 

female, cb 59.5 mm, Pillsbury Sta 51; A, Benthochascon schmitti 

Rathbun, female, cb 50.3 mm, Florida; c, Scylla serrata (Forskal), 

male, cb 55.1 mm, China. 

Guinot (1969c:692) mentioned the fact that 

between the first two abdominal segments of 

Geryon and the coxa of the fifth pereiopod a very 

small section of the 8th thoracic sternite is visible, 

and considered this as a goneplacid character. 

However, in portunid genera like Ovalipes and 

Benthochascon a similar situation exists, and as 

pointed out by Guinot (1971:1066) also in 

Xanthidae, like Panopeus and Rhithropanopeus, part 

of the 8th thoracic sternite is visible, in the last 

mentioned genus even "une assez importante partie 

laterale." In Geryon the third abdominal somite 

of the male covers the basal part of the coxa of 

the fifth pereiopod and the 8th thoracic sternite 

is only visible lateral to the first and second 

abdominal somites. All this demonstrates that the 

taxonomy of these crabs is still far from clear. 

Geryon affinis A. Milne Edwards and Bouvier, 

1894 

Geryon affinis A. Milne Edwards and Bouvier, 1894:41, figs. 

A, C, pi. 1; 1899:35.—Hansen, 1908:18, pi. 1: fig. 1.—


Bouvier, 1922:70.—Rae and Lamont, 1963:24.—Kjennerud, 

1967:193, fig. 1.—Sankarankutty, 1968:50.—Christiansen, 

1969:87, fig. 35, map 29.—Mason and Davidson, 

1969:208.—Tiirkay, 1976b:61 [listed], 70. 

Geryon quinquedens.—Bouvier, 1922:70, pi. 6: fig. 7.—Monod, 

1956:337 [part, not fig. 441]. [Not Geryon quinquedens Smith, 

1879.] 

Geryon tridens.—Saemundsson, 1937:21 [not Geryon tridens 

Kr0yer, 1837]. 


Other Material: Azores: E of Corvo, 39°41'35"N, 31°04'- 

07"W of Paris (= 28°44'07"W of Greenwich), 844 m, sand 

and gravel, 7 Aug 1888, Princesse Alice Sta 222, syntypes, 1(5, 

1$ (MP). 

SE of Madeira: 32°42'N, 16°43'W, 670 m, tent trap, 13 

Mar 1976, Onversaagd Sta 63, 1(5, 19 ov (L). 

Cape Verde Islands: 16°44'N, 24°48'05"W, 692 m, hard 

bottom, 21-22 Jul 1901, Princesse Alice Sta 1138, 16* (MP). 

DESCRIPTION.—A. Milne Edwards and Bouvier, 

1894:41-45; Christiansen, 1969:87. 


figs. A, C, pi. 1 (color); Christiansen, 1969, fig. 

35. 

Male Pleopod: We have found no illustrations of 

the male pleopod of this species. Sankarankutty 

(1968:51) remarked that it was similar to that of 

G. maritae as figured by Doflein (1904). 

Color: A colored figure of one of the type-specimens 

was published by A. Milne Edwards and 

Bouvier (1894, pi. 1). It shows a pale brown crab 

with the cervical groove and posterolateral margins, 

as well as the spine on the carpus, with a 

pink hue; the central part of the carapace is 

somewhat greenish. Kjennerud (1967:194) remarked 

of her specimen: "The colour of the 

specimen before it was preserved in alcohol was 

dull yellow with patches of red and red brown, 

the tips and the margins of the dactyls in the 

walking legs were dark brown. In general it may 

be said that the colour was very like the colour of 

one of the type-specimens illustrated by Milne- 

Edwards and Bouvier." Christiansen (1969:85) 

on the other hand described the color as "red to 

brick-red." A. Milne Edwards and Bouvier (1899: 

35) mentioned a probably abnormal specimen 

"a pattes blanches." 

MEASUREMENTS.—The male and female syntypes 

examined have the carapace lengths 109 

and 112 mm, the carapace widths 134 and 

135 mm, respectively. In the male and female 

from Madeira the carapace widths are 140 and 

145 mm, respectively. In the male from the Cape 

Verde Islands the carapace length is 133 mm, the 

carapace width 158 mm. A. Milne Edwards and 

Bouvier (1894) gave the following measurements 

of some of the types: males, carapace lengths 15, 

97, and 133 mm, carapace widths 17, 112, and 

153 mm, respectively; females, carapace length 

128 mm, carapace width 14-(?) mm. (Due to a 

printer's error the last digit of this number was 

not printed so that the width may be anywhere 

between 140 and 149 mm). One of Saemundsson's 

(1937) specimens was 160 mm long by 

180 mm wide; Kjennerud's (1967) male had a 

length of 150 mm and a width of 180 mm. The 

5 males reported upon by Mason and Davidson 

(1969) had carapace widths ranging from 192 to 

210 mm. In contrast to these large specimens are 

the specimens reported upon by Hansen (1908), 

the male of which was 40 mm long, the ovigerous 

female 42 mm. The only other ovigerous female 

of which the measurements are known is the one 

(cb 145 mm) from Madeira (above). A. Milne 

Edwards and Bouvier, (1894:43) mentioned the 

eggs as very small. Hansen (1908:19) gave the egg 

size in his specimen as 0.5-0.6 mm. 

REMARKS.—The differences between the present 

species and G. maritae are enumerated under 

the latter species. Most records of Geryon qffinis or 

Geryon quinquedens from West Africa pertain to G. 

maritae. It was a surprise to find that the male 

from the Cape Verde Islands listed above, and 

already reported upon by Bouvier (1922), belongs 

to G. affinis rather than to G. maritae; this shows 

that both species do inhabit the tropical West 

African region. The Princesse Alice specimens were 

examined by Holthuis in Paris on 4 October 

1971. 

The specimen from the Azores assigned by 

Bouvier (1922) to Geryon quinquedens is, as shown 

by Bouvier's illustration, a good Geryon qffinis. 

BIOLOGY.—In the eastern Atlantic the species 

is known from depths between 130 and 2047 m; 

almost 50% of the records is from 1165 to 2047

112 

m, the other half is from 130 to 844 m. The 

bottom on which the species was found was described 

as sand and gravel (A. Milne Edwards 

and Bouvier, 1894); muddy sand, black sand, 

sand and rock (A. Milne Edwards and Bouvier, 

1899); hard bottom (Bouvier, 1922). 

Ovigerous females have been collected in August 

(A. Milne Edwards and Bouvier, 1894); our 

ovigerous female from Madeira was taken in 

March. 

DISTRIBUTION.—In the eastern Atlantic the species 

has been reported from the following localities: 

Near Iceland: SW edge of Hvalsbaksbanki (approximately 

64°N, 13°W), 140 m (Saemundsson, 1937). SW of 

Geirfuglasker, Vestmanneyjar (approximately 63°N, 21°W), 

130 m (Saemundsson, 1937). SW of Vestmanneyjar (approximately 

63°N, 21°W), 240 fm (440 m) (Rae and Lamont, 

1963). S of Iceland, 61°30'N, 22°3O'W, 975 fm (1785 m), 

and 61°33'N, 19 O (AV, 1089 fm (2000 m) (Hansen, 1908). 

Off SW Norway: Nyegga Bank (approximately 63°38'N, 

05°52'E), 410 m (Kjennerud, 1967; Sankarankutty, 1968). 

Faroe Islands: ? Off Enniberg (approximately 62°24'N, 

06°33'W) (Mason and Davidson, 1969). 

Atlantic Ocean, NW of Scotland: Lousy Bank (approximately 

60°N, 13°30'W), 440 m (Mason and Davidson, 

1969). Rosemary Bank (approximately 59°15'N, 10°W), 

480 m (Mason and Davidson, 1969). George Bligh Bank 

(approximately 59°N, 13°30'W), 240 fm (440 m) (Rae and 

Lamont, 1963; Mason and Davidson, 1969). 30 miles [48 

km] SW of Rockall, 57°18'N, 14°25'W (Mason and Davidson, 

1969). 

Azores: 39°39'10''N, 31°03'40"W, 1300 m; 39°21'20"N, 

31°06'08"W, 1360 m; 38°27'N, 26°31'W, 1165 m; 38°25'- 

50"N, 28°34'30 w W, 785 m; 38 o 01'N, 29°22'30"W, 1260 m; 

37°43'N, 25°06'W, 1385 m (all A. Milne Edwards and 

Bouvier, 1899). Near Ilha de Sao Miguel, 37°37'N, 25°2O'- 

45"W, 1187 m (Bouvier, 1922). E of Ilha do Corvo, 39°41'- 

35"N, 28°44'07"W, 844 m; SE of Ilha do Corvo, 1386 m; 

between Ilha do Pico and Ilha de Sao Jorge, 38°38'N, 

28°08'15"W, 620 m (A. Milne Edwards and Bouvier, 1894). 

Madeira: No specific locality, ca. 1000 m (Tiirkay, 

1976b). 

Cape Verde Islands: W of the Cape Verde Islands, 16°- 

44'N, 24°48'05"W, 692 m; near Ilha de Maio, 15°17'40"N, 

23°02'45"W, 1300 m (Bouvier, 1922). 

Christiansen (1969, map 29) provided distribution 

charts of the species. Outside the eastern 

Atlantic region Geryon affinis has been reported 

from the east coast of America, from South Africa, 


the Indian Ocean, and Australia. A comparison 

of material from those areas with specimens from 

the eastern Atlantic is highly desirable in order 

to ascertain whether or not a single species is 

involved in all these records. The taxonomic confusion 

within the genus Geryon is notorious. Many 

authors (including Rathbun, 1937:271; Barnard, 

1950:291; Monod, 1956:337) considered Geryon 

affinis and G. quinquedens synonymous, notwithstanding 

the fact that A. Milne Edwards and 

Bouvier (1894:41-45, figs. A-D) and Chace 

(1940:38-40) had convincingly shown the differences 

between the two species. A revision of the 

genus is highly desirable. 

* Geryon maritae, new species 

FIGURES 24a, 25, 26 

Geryon chuni Doflein, 1903:21 [nomen nudum]. 

Geryon affinis.—Doflein, 1903:23; 1904:106, 110, 111, 175, 

177, 180, 186, 190, 204, 206, 208, 210, 211,213, 214, 219, 

220, 251, 253, 255, 257, 258, text figs. 9, 14, 32, 62, 

unnumbered pi.: figs. 1, 2, pis. 3, 33, 34, 38: figs. 1-5, 9, 

pi. 41: figs. 3-7, pi. 43: figs. 2, 8, pi. 49/50: fig. 1, pi. 52: 

fig. 7, pi. 55: fig. 6, pi. 58: figs. 2, 4 [part]. [Not Geryon 

affinis A. Milne Edwards and Bouvier, 1894.] 

Geryon quinquedens.—Capart, 1951:173, fig. 66.—Monod, 

1956:337, fig. 441 [part].—Guinot and Ribeiro, 1962: 

62.—Forest, 1963:628.—Monod, 1967:180, pi. 17: fig. 2 

[no material].—Le Loeuff and Intes, 1969:66.—Crosnier, 

1970:1216.—Le Loeuff, Intes, and Le Guen, 1974:73, figs. 

1-4.—Intes and Le Loeuff, 1976:101. [Not Geryon quinquedens 

Smith, 1879.] 

Geryon.—Voss, 1966:19.—Maurin, 1968b, figs. 1,4. 

Geryon quinquidens.—Maurin, 1968a:50; 1968b:484, 487, 491, 

492, fig. 6 [erroneous spelling.] 

"Third form of Geryon".—Christiansen, 1969:87. 

Geryon sp.—Dias and Seita Machado, 1973:1.—Bas, Arias, 

and Guerra, 1976, table 3. 


Sta 74, 641-733 m, Id (holotype, L), 1$ (W). 

Ivory Coast: Sta 41, 641-842 m, 1(5, 1$, 10 juv (L). Sta 

44, 403-586 m, hard, dark gray mud, 4

NUMBER 306 

FIGURE 25.—Geryon maritae, new species (from Monod, 1956, 

fig. 441). 

times wider than long, greatest width at posterior 

pair of anterolateral teeth. Median pair of frontal 

teeth about as long as but narrower than lateral 

pair, overreaching lateral teeth by half their 

length. Median sinus U-shaped. Orbital margin 

L-shaped, short leg of L forming lateral margin of 

front, longer leg curving distally to join exorbital 

tooth. Orbital margin, apart from 2 fissures, 

smooth, lacking tubercles. Exorbital tooth and 

second and last (= fourth) anterolateral teeth of 

carapace distinct but low and triangular. First 

anterolateral tooth vaguely visible as low angle, 

third tooth absent or visible as low, indistinct 

elevation. Distances between anterolateral teeth 

subequal, only slightly greater than distance between 

exorbital tooth and first anterolateral 

tooth. Grooves of carapace very faint, much less 

distinct than in G. quinquedens. Blunt ridge extending 

anteriorly from near base of fifth leg, running 

more or less parallel to lateral margin of carapace. 

Anterior half of dorsal surface of carapace relatively 

smooth, with some groups of low, eroded 

tubercles on elevated parts. Posterior part of carapace 

with many distinct, coarse tubercles, becoming 

finer laterally. 

Suborbital margin terminating in strong inner 

tooth, directed anteriorly, extending to or just 

beyond second segment of antennular peduncle. 

Lower orbital margin distinctly tuberculate, ending 

before reaching exorbital tooth. 

113 

Abdomen triangular, closely resembling that of 

G. quinquedens, differing in lacking transverse carina 

on fourth somite. Telson almost an equilateral 

triangle. 

Epistome shorter than in G. quinquedens, with 

deep pit in middle of basal part. 

Mouth parts of this species have been well 

figured by Doflein (1904, pi. 38: figs. 1-5, 9). 

Mandible with heavy molar part and rather slender 

3-segmented palp, with last segment somewhat 

sickle-shaped. Maxillule (Figure 26) with 

very slender lower lacinia, upper widening distally, 

twice as wide at distal margin as at base; 

palp 2-segmented, basal segment about as wide 

as long or slightly wider, distal segment about 

half as wide as basal segment, slender in comparison. 

In Doflein's (1904, pi. 38: fig. 5) illustration 

an evidently mutilated maxillule is shown, which 

lacks part of the upper lacinia; therefore a new 

figure of the maxillule is provided here (Figure 

26). Maxilla with 2 inner laciniae deeply incised, 

forming together 4 unequal slender lobes; basal 

part of palp almost circular, distal part very 

narrow, ending in slender sharp point, somewhat 

curved; scaphognathite broad, truncated posteriorly. 

First maxilliped with 2 laciniae, upper 

about twice as high as lower; endopod 2-segmented, 

distal segment inversely triangular with 

FIGURE 26.—Maxillule of Geryon maritae, new species, paratype, 

female, cb 59.5 mm, Pillsbury Sta 51.

114 

distal margin wide; faint lobe visible on inner 

margin; exopod with well-developed flagellum; 

epipod large, ending in elongate narrow distal 

part. Second maxilliped pediform; peduncle of 

exopod reaching beyond endopod; well-developed 

flagellum present. Merus of third maxilliped 

with outer angle rounded, not produced. 

Larger of 2 chelipeds with dactylus 5/4 to 4/3 

as long as palm (measured along upper margin). 

As in Carcinus proximal teeth of fingers wide, 

molar-like, distal fingers flattened, tooth-like. Distal 

teeth alternatingly large and small. Upper 

surface of dactylus minutely granular, with some 

coarse tubercles basally; normal ridges and rows 

of pits present on fingers. Palm lacking anterodorsal 

spine, outer surface with transverse rows of 

tubercles and with broad longitudinal groove on 

upper part and 1 or 2 low, inconspicuous ridges 

over middle. Smaller chela resembling larger, but 

with fingers proportionately slightly longer and 

lacking molar-like teeth. Dorsal surface of carpus 

(of both chelipeds) with very strong, rather sharp 

tubercles, inner margin with large, sharp tooth, 

anterior and outer margins unarmed. Merus of 

usual shape, triangular in transverse section, with 

strong subdistal dorsal spine, distal spine absent. 

Merus less slender than in G. quinquedens. 

Walking legs slender, merus of last leg 4 times 

as long as high. Dorsal part of anterior margins 

of walking legs angular, unarmed. Subdistal tubercle 

present just behind transverse dorsal 

groove on distal part of merus, with smaller tubercles 

present on dorsal surface, restricted to 

distal part, more numerous and more distinct on 

anterior legs than on posterior ones. Distinct tubercles 

present on upper margin of carpus and 

propodus. Propodus of fifth leg about 4 times as 

long as high, 5/4 as long as dactylus (length 

measured on dorsal margin). Dactylus dorsoven- 

trally flattened, resembling that of G. quinquedens. 

Posterior margin of dactylus with distal groove, 

most distinct on second pereiopod (first walking 

leg). Lower posterior margin of dactylus of second 

pereiopod with row of closely placed short hairs 

over proximal two-thirds. Third pereiopod with 


few scattered tufts of hairs on dactylus, tufts 

completely absent on fourth and fifth pereiopods. 

Female, carapace length 89 mm, width 95 mm: 

First and third anterolateral teeth of carapace 

more distinct than in male, but far less conspicuous 

than second and fourth teeth. First tooth a 

low triangle, third a slight elevation. Otherwise 

carapace similar to that of male but slightly 

smoother. 

Young male (Pillsbury Sta 41), carapace length 

27 mm, width 34 mm: Carapace relatively wider 

than in adult male, still widest at level of last pair 

of anterolateral teeth. Median pair of frontal 

teeth short, bluntly rounded, placed close together, 

extending about as far as slightly broader 

lateral frontal teeth. Exorbital tooth and second 

and fourth anterolateral teeth slender, spiniform, 

far more distinct than in adult. First and third 

anterolateral teeth scarcely visible, presence indicated 

by slight blunt hump; anterior end of 

hump of third tooth often abrupt, more clearly 

marked than that of first. Grooves of carapace 

more distinct than in adult, branchiocardiac 

grooves particularly well developed. Entire surface 

of carapace finely tuberculated, but elevated 

portions of anterior half and most of posterior 

part with coarse tubercles. Abdomen similar to 

that of adult, but with faint low transverse carina 

on fourth and less distinct carina on fifth somite. 

Orbit and epistome as in adult. Merus of third 

maxilliped with anterolateral angle slightly more 

auricularly produced. 

Chelipeds slenderer than in adult, with fingers 

somewhat longer. First tooth of dactylus large, 

rounded, directed proximally, exactly as in many 

Portunidae; other teeth also quite portunid-like. 

Tuberculation and pitting of chela as in adult 

male. No trace of spine present at broadly 

rounded distal end of upper margin of palm. 

Carpus as in large male, but spine at inner margin 

longer, more slender; carpus lacking any trace of 

spine on anterior margin. Merus only with subdistal 

dorsal spine, distal dorsal spine absent. 

Pereiopods as slender as in adult. Sharp tubercles 

present on upper margin of merus, carpus, 

and propodus, merus lacking distal dorsal spine.


Dactylus scoop-shaped, as in adults. On second 

pereiopod row of hairs visible along either margin 

of flattened area, rows interrupted on third and 

fourth pereiopods, scarcely discernible on fifth 

pereiopod. 

Very small specimens, carapace length 11- 

17 mm: carapace slightly wider, 4/3 as wide as 

long. Legs slightly slenderer, merus of fifth leg 

about 4.3 times as long as wide. First and third 

anterolateral teeth of carapace visible as small 

lobes, first more distinct than third. 

Male Pleopod: Doflein, 1904, pi. 41: fig. 6 (locality 

not stated). 

Color: Color photographs were taken of the 

large male holotype of Pillsbury Sta 74, immediately 

after it came aboard. The ground color of 

the body is pale cream. The carapace is grayish 

brown in the anterior part, with a pale median 

longitudinal zone. Black spots of irregular size, 

which are irregularly arranged, are found in the 

middle and in the anterolateral parts of the carapace; 

they probably are caused by external agencies 

and do not form part of the actual color 

pattern. The chelipeds are cream with a black 

color along the cutting edges of the fingers. The 

propodi and dactyli of the walking legs are reddish 

brown, contrasting with the cream color of 

the merus; the carpus is somewhat intermediate 

in color. In alcohol the male specimens and the 

juveniles are pale cream colored, the females in 

several instances are more of a reddish brown. 

Doflein (1904, pi. 3) published a color plate of a 

male, showing it entirely reddish brown. Capart 

(1951:174) gave the following color description of 

his material: "Couleur generate bistre; zone rouge 

brique a rose sur l'arriere de la carapace et sur les 

pattes; dents laterales plus foncees. Les grands 

specimens generalement a coloration plus intense." 

MEASUREMENTS.—The males examined have 

the carapaces up to 140 mm long and 160 mm 

wide. The largest female has a carapace length of 

89 mm, a width of 95 mm. The smallest specimen 

has a carapace length of 11 mm, a width of 

15 mm. The records in the literature mostly deal 

with animals smaller than our largest. Doflein 

(1904) indicated that his largest specimen was 

about 90 mm long, about 110 mm wide; as 

Doflein's material included ovigerous females 

they must have been smaller than this. Le Loeuff, 

Intes and Le Guen (1974) gave the carapace 

width of their largest female as 110 mm, and that 

of their largest male as 164 mm. Dias and Seita 

Machado (1973) reported upon a male with carapace 

width of 165 mm. The species can attain a 

weight of up to 1650 g (Le Loeuff, Intes and Le 

Guen, 1974). 

REMARKS.—Geryon maritae closely resembles Ger- 

yon qffinis A. Milne Edwards and Bouvier, but 

differs in the following points. 

1. The front between the orbits is narrower, 

because the inner margins of the orbits are less 

strongly divergent. In G. maritae this margin is 

short, straight, and runs only slightly obliquely 

backwards. It then abruptly turns into the posterior 

margin of the orbit, which from there goes 

in an almost straight line to the exorbital tooth. 

In G. qffinis the inner orbital margin is much 

longer, diverges widely, and has a little hump in 

the middle. It gradually merges with the posterior 

margin of the orbit, which at first is directed 

obliquely laterally and then turns more anteriorly 

to the apex of the exorbital tooth. Capart's (1951, 

fig. 66) drawing very clearly shows the situation 

as it is in G. maritae. 

2. The first anterolateral tooth usually is more 

distinct in G. qffinis than in G. maritae. 

3. The inner infraorbital tooth in G. maritae is 

shorter and reaches hardly past the second segment 

of the antennula; in G. qffinis it is much 

larger and stronger and reaches far beyond that 

segment. 

4. In G. affinis the lower orbital angle is finely 

and closely tuberculate, in G. maritae these tubercles 

as a rule are placed wider apart. 

5. The main character distinguishing the two 

species, however, is shown by the dactyli of the 

walking legs, which in G. qffinis are laterally 

compressed, while in G. maritae they are dorsoventrally 

flattened. 

In the fifth character the new species resembles 

G. quinquedens, a species with which G. qffinis has

116 

often been confused; however, G. quinquedens differs 

from G. maritae in the following points: 

1. The submedian frontal teeth reach with 

their entire length beyond the lateral frontal teeth 

and are placed on a prolongation of the front. 

2. The upper margin of the palm of the chelipeds 

ends in a distinct tooth. 

3. The outer anterior margin of the carpus of 

the cheliped has a distinct sharp tooth. 

4. The dorsal margin of the merus of the cheliped 

has a distinct distal and subdistal tooth. 

5. The pereiopods are longer: the merus of the 

fifth leg is more than five times as long as high. 

6. The meri of the walking legs each end in an 

anterodorsal tooth. 

7. The male abdomen is broader and the lateral 

margin shows incisions at the lines of separation 

of the segments. 

The species was first described and figured by 

Doflein (1904), who had about 40 specimens from 

the Valdivia Bank in the South Atlantic Ocean, 

which he incorrectly assigned to Geryon qffinis. A 

specimen from E Africa also identified by him 

as this species evidently is specifically distinct. 

Doflein's extensive description and illustrations of 

his Atlantic specimens leave little doubt that they 

belong to the present new species. Doflein described 

many of the important characters of the 

species, but attached little importance to the 

systematic value of features as the shape of the 

dactylus of the walking legs; he was inclined to 

unite all the species of Geryon into a single variable 

one. His figures (1904, pi. 41: figs. 5, 7) of the 

dactylus of the pereiopod distinctly show that his 

specimens belong in the present species and not 

to G. affinis. 

A year before the publication of his large paper 

on the Valdivia crabs, Doflein (1903) published a 

short note on the eyes of deep sea crabs, using the 

Valdivia material as a basis. In this preliminary 

paper he remarked (Doflein, 1903:21) "Bei Geryon 

chuni konnte ich in den Eihiillen wohlentwickelte 

Zoeen konstatieren." Two pages beyond this he 

(1903:23) made the observation: "Zudem fand 

ich spater bei Geryon qffinis aus mehr als 1000 m 

Tiefe, dass die noch in den Eischalen eingeschlos- 


senen Larven deutlich pigmentierte Augen besassen." 

As Doflein's (1903:4) observations are evidently 

based on Valdivia material, and as the only 

ovigerous specimens of Geryon collected by the 

Valdivia were those of the present species obtained 

from the Valdivia Bank, the specimens referred 

to by Doflein (1903) under both the names Geryon 

chuni and G. affinis evidently belong to G. maritae. 

This is confirmed by Doflein's (1904:258) remark 

in his Valdivia report: "Dagegen konnte ich z.B. 

bei Geryon affinis M.-E.u.Bouv., dessen Eier klein 

sind, die im Ausschliipfen begriffenen Zoeen auffinden." 

This sentence being practically identical 

with the one from his 1903 (p. 21) paper cited 

above. It seems likely that Doflein at first intended 

to describe the South Atlantic form as a 

new species Geryon chuni, but that he later changed 

his mind and considered it conspecific with G. 

affinis. Everywhere he changed the epithet chuni 

to affinis, but evidently forgot to do so in the entry 

on p. 21 of his 1903 paper; otherwise it is difficult 

to understand why he indicated one species in 

that paper with two different names. The name 

G. chuni was published without any description 

and must be considered a nomen nudum. Its 

identity can only be ascertained by inference. 

Bouvier (1922) reported a small specimen of 

Geryon from the Azores as G. quinquedens; the figure 

given by Bouvier, however, shows that it is G. 

affinis (p. 110). A large male and female from the 

Cape Verde Islands were referred by the same 

author (Bouvier, 1922:70) to G. affinis. Of these 

two specimens the male is in the collection of the 

Paris Museum and could there be examined by 

us. Contrary to our expectations, it proves to be 

a true Geryon affinis. 

Capart (1951), under the name Geryon quinquedens, 

gave a good description and figure of the 

present species. Monod (1956:337, 338) incorrectly 

united Geryon affinis and G. quinquedens to a 

single species. His specimens clearly belong to G. 

maritae as shown by his figure. Crosnier (1970) 

dealt with our above-mentioned specimens from 

Angola, which are now in the collection at Leiden. 

It also seems most likely that the material 

listed by Guinot and Ribeiro (1962), and Forest


(1963) belong to the present species. All these 

authors thought the synonymy of Geryon quinquedens 

and G. affinis very likely, but they had reservations. 

This is the more peculiar as both A. 

Milne Edwards and Bouvier (1894:41-45, figs. 

A-D) and Chace (1940:38-40) had enumerated 

very clearly the differences between these two 

species. The cause of the confusion evidently is 

that the present new species is somewhat intermediate 

between G. affinis and G. quinquedens: it 

shows the dactyli of the walking legs as in G. 

quinquedens, but in most of the other characters it 

more closely resembles G. affinis. 

The anatomy of this species, especially that of 

the eye, the statocyst, etc., has been extensively 

dealt with by Doflein (1904). 

ECONOMIC IMPORTANCE.—An active trawling 

fishery for this species has been developed in 

Angola since about 1970 (Dias and Seita Machado, 

1973; Le Loeuff, Intes and Le Guen, 1974), 

the annual catch amounting to 2000 tons. Spanish 

trawlers, which operate off the West coast of 

Africa to fish for shrimps, often catch Geryon, but 

only sell the chelipeds; in recent years the Japanese 

have also developed an interest in the species 

and have done some exploratory fishing for it off 

South-West Africa (Le Loeuff, Intes and Le Guen, 

1974). Le Loeuff, Intes, and Le Guen (1974) and 

Intes and Le Loeuff (1976) described exploratory 

fishing for this species off the Ivory Coast, in 400 

to 650 m and 300 to 700 m depth. As the rough 

bottom configuration there made trawling impractical, 

lobster pots were used with reasonable 

success (from 1.6 to 53 kg crabs were caught per 

pot; the best catches being in depths between 400 

and 500 m, where 30 to 50 kg crabs were obtained 

per pot). 

TYPE-LOCALITY.—Off Liberia, 04°20'N, 09°. 

26'W to 04°30'N, 09°22'W, 641-733 m. 


D. 21052), a male, carapace length 134 mm, is in 

the Rijksmuseum van Natuurlijke Historie, Leiden. 

The remaining specimens are paratypes; 

they have been divided between the Rijksmuseum 

van Natuurlijke Historie, Leiden, and the 

National Museum of Natural History, Smithsonian 

Institution, Washington, D.C. 

ETYMOLOGY.—Dr. Marit E. Christiansen, who 

saw our Pillsbury material, was the first to publish 

that the present species is distinct from G. affinis 

and pointed this out in her excellent book on the 

Scandinavian Brachyura (1969:87). For this reason 

we take great pleasure in naming this interesting 

species for her. 


depths between 100 and 936 m. Only three records 

are definitely from less than 250 m (150- 

245 m, Monod, 1956; no deeper than 200 m on 

talus, Le Loeuff and Intes, 1969; 200 m, p. 112 

herein), and only four are definitely from more 

than 600 m (936 m, Doflein, 1904; 800 m, Dias 

and Seita Machado, 1973; 650 m, Le Loeuff, 

Intes and Le Guen, 1974; and 641-842 m, p. 112 

herein). The majority, 24 of 31, of the records are 

partly or entirely from between 250 and 600 m. 

The bottom on which the species has been taken 

has been given as mud (Guinot and Ribeiro, 

1962; Forest, 1963); mud, sandy mud, slightly 

muddy sand, greenish muddy sand (Maurin, 

1968b); green mud and sand (three records), 

brown mud and sand, green mud (Capart, 1951); 

hard, dark gray mud (Voss, 1966; p. 112 herein); 

corals (Guinot and Ribeiro, 1962). 

Dias and Seita Machado (1973) found that at 

300 m depth the catch consisted of 96.8% females, 

but that this percentage dropped considerably in 

deeper waters being 38% in 500 m, 2.2% in 

600 m, and 1.6% m in 800 m. Le Loeuff, Intes, 

and Le Guen (1974) found in their catches (400- 

650 m) that the females were very scarce. The 

males were more numerous and probably represented 

two age classes. In 1976 Intes and Le 

Loeuff observed that the percentage of males 

increased with increasing depth. 

DISTRIBUTION.—Geryon maritae is a West African 

species, known from localities between Spanish 

Sahara and Valdivia Bank, South-West Africa. A 

fishing survey reported by Dias and Seita Machado 

(1973) encountered the species in depths 

between 300 and 800 m from 29 transects made 

between just N of Point-Noire, Congo, and off

118 

the Rio Cunene, southern Angola; their individual 

records are not repeated below. Other records 

in the literature include the following. 


Spanish Sahara: Off Villa Cisneros, 300-500 m (Maurin, 

1968a, 1968b). 23°35.5'N, 16°58'W to 23°32'N, 16°57'W, 


Mauritania: Between Cap Timiris and Tamzak (as 

Tamxat), 350-600 m (Maurin, 1968b). 

Senegal: Off Saint-Louis, 300 m (Maurin, 1968b); S of 

Saint-Louis, 100-300 m (Monod, 1956). Fosse de Kayar, 

300-350 to 600 m (Maurin, 1968b). Off Senegal, 14° 46'- 

51"N, 150-245 m (Monod, 1956). 11 mi [18 km] NW of 

Pointe des Almadies, ca. 300 m (Monod, 1956). 

Liberia: No specific locality (Christiansen, 1969). 

Ivory Coast: No specific locality (Christiansen, 1969; Le 

Loeuff and Intes, 1969). Numerous localities at transects 

across the Ivory Coast, 03° to 07.5°W, 300-700 m (Intes and 

Le Loeuff, 1976). Off Grand-Bassam, 03°49'W, 400-650 m 

(Le Loeuff, Intes and Le Guen, 1974). 05°05'N, 04°00'W to 

05°04'N, 04°02'W, 403-586 m (Voss, 1966). 04°32.5'N, 

06°31'W, 300-455 m and 04°54'N, 03°23'W, 380-400 m 

(Forest, 1963). 


Angola: SW of Moita Seca, 06°08'S, 11°24'E, 250- 

380 m; W of Ambrizete, 07°16'S, 12°02'E, 380-420 m; W of 

Ponta do Morro, 10°45.5'S, 13 b 07'E, 400-500 m; NW of 

Egito, 11°53'S, 12°23'E, 400-500 m, and 11°53'S, 13°20'E, 

480-510 m (all Capart, 1951). Baia dos Tigres, 320-400 m, 

453-478 m (Guinot and Ribeiro, 1962). 

South-West Africa: 17°23'S, ll°20'E, 359 m (Crosnier, 

1970). Valdivia Bank, 25°27'S, 06°08.2'E, 936 m (Doflein, 

1904). 

Family PLATYXANTHIDAE Guinot, 1977 

PLATYXANTHIDAE Guinot, 1977:1052. 

This family, comprising three genera, is not 

represented in the eastern Atlantic. 

Family XANTHIDAE MacLeay, 1838 

PILUMNIDAE Samouella, 1819:86 [by direction, under the 

Plenary Powers of the International Commission on Zoological 

Nomenclature, not to be used in preference to 

Xanthidae MacLeay, 1838, but available for use by those 

who consider that Pilumnus Leach and Xantho Leach belong 

to different family-group taxa; Opinion 423 in Opinions 

and Declarations of the International Commission on Zoological 

Nomenclature, volume 14, 1956; name 74 on Official List]. 


XANTHIDAE MacLeay, 1838:59 [name 73 on Official List; 

there dated from Dana, 1851, in error]. 

ERIPHIDAE MacLeay, 1838:59, 60. 

TRICHIDEA de Haan, 1839, pi. H. 

CHLORODINAE Dana, 1851b: 125. 

POLYDECTINAE Dana, 1851b: 127. 

OziNAEDana, 1851b: 127. 

ACTUMNINAE Dana, 1851b: 128. 

Carpilides A. Milne Edwards, 1862a:40. 

Trapezides A. Milne Edwards, 1862a:40. 

Liagorides A. Milne Edwards, 1862a:41. 

Trapeziden Nauck, 1880:64. 

TRAPEZIINAE Miers, 1886:163. 

MENIPPIDAE Ortmann, 1893b:428, 431. 

MYOMENIPPINAE Ortmann, 1893b:429, 432. 

CARPILINAE Ortmann, 1893b:429, 463. 

ETISINAE Ortmann, 1893b:429, 470. 

PANOPAEINAE Ortmann, 1893b:429, 473. 

DOMOECIINAE Ortmann, 1893b:429, 478. 

ZOZYMOIDA Alcock, 1898:77, 94. 

EUXANTHOIDA Alcock, 1898:77, 109. 

HALIMEDOIDA Alcock, 1898:77, 134. 

ACTAEINAE Alcock, 1898:78, 137. 

XANTHODIOIDA Alcock, 1898:78, 156. 

CYMOIDA Alcock, 1898:78, 172. 

PSEUDOZIOIDA Alcock, 1898:176, 180. 

RUPPELLIOIDA Alcock, 1898:176, 186. 

HETEROPANOPIOIDA Alcock, 1898:177, 207. 

MELIOIDA Alcock, 1898:177, 230. 

LYBIOIDA Serene, 1965:9, 12, 26, 37. 

ZALASHNAE Serene, 1968:62. 

LIOMEROIDA T. Sakai, 1976:xvii, 385, 390. 

EASTERN ATLANTIC GENERA.—Thirty-two, of 

which all but five are represented by species 

occurring off tropical West Africa. The extralimital 

genera are the following: 

Atergatis de Haan (1833:4, 17). Type-species: 

Cancer integerrimus Lamarck, 1818, by subsequent 

designation by the International Commission on 

Zoological Nomenclature in Opinion 73 in Smithsonian 

Miscellaneous Collections, 73(1) :26, 1922; gender: 


Neopanope A. Milne Edwards (1880, in 1873- 

1881:329). Type-species: Neopanope pourtalesii A. 

Milne Edwards, 1880, a subjective junior synonym 

of Panopeus packardii Kingsley, 1879, by 

subsequent designation by Fowler (1912:400); 

gender: feminine. 

Pilumnoides H. Milne Edwards and Lucas 

(1843:21). Type-species: Hepatus perlatus Poeppig,



on Official List. 

Rhithropanopeus Rathbun (1898:273). Type-species: 

Pilumnus harrisii Gould, 1841, by original 

designation and monotypy;. gender: masculine; 


Sphaerozius Stimpson (1858a:32). Type-species: 

Sphaerozius nitidus Stimpson, 1858, by subsequent 

designation by the International Commission on 

Zoological Nomenclature in Opinion 85 in Smithsonian 

Miscellaneous Collections, 73(3): 17, 1925; gender: 


EASTERN ATLANTIC SPECIES—Fifty-seven, of 

which 44 occur off tropical West Africa. Monod 

(1956) recorded the following species: 


Menippe nodifrons 

Globopilumnus africanus 

Globopilumnus stridulans 

Epixanthus helleri 

Pseudozius bouvieri 

Pilumnus stebbingi 

Pilumnus perrieri 

Pilumnus inermis 

Pilumnus spinifer 

Parapilumnus pisifer 

Parapilumnus boletifer 

Heteropanope (Heteropanope) 

tuberculidens 

Heteropanope (Pilumnopeus) 

caparti 

Heteropanope (Pilumnopeus) 

africana 

Domecia hispida 

Xantho (Xantho) incisa 

Xantho (Xantho) pilipes 

Xantho (Xantho) sexdentata 

Xantho (Xantho) denticulata 

Xantho (Leptodius) inaequalis 

punctata 

Xantho (Leptodius) inaequalis 

convexa 

Xantho (Leptodius) floridana 

Actaea (Actaea) rufopunctata 

Actaea (Actaea) margaritaria 

Actaea (Glyptoxanthus) 

angolensis 

Menippe nodifrons 

Globopilumnus africanus 

Globopilumnus stridulans* 

Epixanthus hellerii* 

Euryozius bouvieri 

Euryozius pagalu, 

new species* 

Pilumnus stebbingi* 

Pilumnus perrieri* 

Pilumnus inermis 

Pilumnus spinifer 

Leopoldius pisifer* 

Nanopilumnus boletifer* 

Heteropanope tuberculidens 

Pilumnopeus caparti 

Pilumnopeus africanus* 

Domecia acanthophora 

africana* 

Xantho incisus 

Xantho pilipes 

Xantho sexdentatus 

Xanthodius denticulatus* 

Xanthodius inaequalis 

inaequalis* 

Xanthodius inaequalis faba 

Cataleptodius floridanus* 

Paractaea rufopunctata 

africana* 

Paractaea monodi 

Paractaea margaritaria* 

Glyptoxanthus angolensis* 

Actaea (Glyptoxanthus) 

cavemosa 

Actaea (Glyptoxanthus) corrosa 

Platypodia picta 

Cycloxanthops occidentalis 

Paraxanthias eriphioides 

Medaeus africanus 

Medaeus couchi 

Medaeus (?) rouxi 

Micropanope rufopunctata 

Micropanope parvula 

Micropanope melanodactyla 

Panopeus africanus 

Panopeus parvulus 

Panopeus sp. 

Glyptoxanthus cavemosus 

Glyptoxanthus corrosus 

Platypodiella picta 

Cycloxanthops occidentalis 


Pseudomedaeus africanus* 

Monodaeus couchii 

Monodaeus rouxi* 

Microcassiope minor* 

Coralliope parvula 

Nanocassiope melanodactyla* 

Panopeus africanus* 

Eurypanopeus blanchardi* 

Panopeus africanus* 

The extralimital species include the following: 

Atergatis roseus (Riippell, 1830). Eastern Mediterranean, 

Israel; an immigrant from the Red 

Sea; littoral (Lewinsohn and Holthuis, 1964). 

Heteropanope laevis (Dana, 1852). Eastern Mediterranean, 

Egypt, an immigrant from the Red 

Sea; sublittoral (Holthuis and Gottlieb, 1958; 

Ramadan and Dowidar, 1976). 

Monodaeus guinotae Forest, 1976. Mediterranean, 

from the Balearic Isles, the Gulf of Taranto, and 

off Israel, shore (?) to 800 m (Forest, 1976). 

Neopanope sayi (Smith, 1869). An East American 

species introduced into England; littoral (Naylor, 

1960). 

Pilumnoides perlatus (Poeppig, 1836). An eastern 

Pacific species (Rathbun, 1930:537) introduced 

to England, South-West Africa (L), and South 

Africa (Barnard, 1950:257). 

Pilumnopeus vauquelini (Audouin, 1826). Eastern 

Mediterranean, from Egypt and Israel, an immigrant 

from the Red Sea; sublittoral (Holthuis 

and Gottlieb, 1958; Ramadan and Dowidar, 

1976). 

Pilumnus aestuarii Nardo, 1869. Mediterranean, 

from Cadaques, Spain and Venice, Italy; shallow 

water (Zariquiey Alvarez, 1968). 

Pilumnus hirsutus Stimpson, 1858. Eastern Mediterranean, 

Egypt, an immigrant from the Red 

Sea; sublittoral (Holthuis and Gottlieb, 1958; 

Ramadan and Dowidar, 1976). 

Pilumnus villosissimus (Rafinesque, 1814). Med-

120 

iterranean; shallow water (Zariquiey Alvarez, 

1968). 

Rhithropanopeus harrisii (Gould, 1841). Scattered 

localities between the Baltic, the southern North 

Sea, NW and SW France, Black Sea, Sea of Azov, 

and Caspian Sea; introduced from North America; 

estuarine (Christiansen, 1969). 

Sphaerozius nitidus Stimpson, 1858. Eastern 

Mediterranean, Egypt, an immigrant from the 

Red Sea; shallow water (Ramadan and Dowidar, 

1976). 

Xantho granulicarpus Forest, 1953. Mediterranean; 

sublittoral (Zariquiey Alvarez, 1968). 

Xantho poressa (Olivi, 1792). Canary Islands, 

Portugal, and Mediterranean; shallow water 


We believe that two species included in the 

West African fauna by Monod (1956) are based 

either on misidentifications or erroneously labeled 

specimens: 

Chlorodiella longimana (H. Milne Edwards, 

1834). A western Atlantic species reported from 

Sao Tome by Osorio (1887:221) (see Monod, 

1956:303), and not subsequently recollected 

there. 

Pilumnoides hassleri A. Milne Edwards, 1880. A 

western Atlantic species reported by Monod 

(1956:262) on the basis of a specimen supposedly 

from Gabon collected by Heurtel. The origin of 

most or all of the Heurtel collection from Gabon 

is doubtful. 

REMARKS.—The status of subfamilies and genera 

within the family Xanthidae is currently 

being studied by Mme. D. Guinot of the Museum 

national d'Histoire naturelle, Paris. Preliminary 

observations of Mme. Guinot have appeared in a 

series of shorter papers published between 1967 

to 1971 and the first of more comprehensive 

papers, detailing observations on three subfamilies 

of xanthids, Actaeinae, Polydectinae, and 

Trichiinae, was published in 1976. Rather than 

try to accomodate the West African genera within 

the partial framework established by Mme. 

Guinot to date or one of the unsatisfactory older 

classifications, we have arranged accounts of all 

of the West African genera alphabetically. 


Genus Cataleptodius Guinot, 1968 

Cataleptodius Guinot, 1968a:704 [type-species: Chlorodius floridanus 

Gibbes, 1850, by original designation; gender: masculine].—Guinot, 

1971:1068 [list of species]. 

* Cataleptodius floridanus (Gibbes, 1850) 

Xantho (Leptodius) flondana.—Monod, 1956:291, figs. 353- 

355. 

Xantho (Leptodius) floridanus.—Forest and Guinot, 1966:75.— 

Guinot, 1968a:706 [discussion]. 

Cataleptodius a.ft. floridanus.—Guinot, 1968a:706, 708 [discussion], 

fig. 23; 1971:1068 [listed]. 


Sta 275, 9-69 m, rubble of coralline algae, lcJ, 2$ (1 ov) 

(L,W). 

DESCRIPTION.—We have found no full description 

of this species based on West African specimens, 

but we anticipate that Mme. Guinot, in 

future studies, will provide a description. A description 

based upon western Atlantic specimens 

may be found in Rathbun (1930:297). 

Figure: Monod, 1956, figs. 353-356. 

Male Pleopod: Monod, 1956, figs. 354-355 (Annobon), 

fig. 356 (Florida); Guinot, 1968a, fig. 23 

(Annobon). 


widths of 6 to 7 mm; that of the ovigerous 

female is 6.5 mm. Monod (1956) mentioned specimens 

from West Africa 8 to 20 mm wide, and 

Rathbun (1930) reported a carapace width of 

33 mm in an American specimen. 

REMARKS.—In her generic revision, Guinot 

(1968a: 706) designated the West African population 

of this species as Cataleptodius aff. floridanus 

and noted: 

Sous ce nom, nous separons les representants ouest-africains 

rapportes a Xantho (Leptodius) floridanus par Monod 

(1956, p. 291, fig. 353-356) et par nous-meme (Forest et 

Guinot, 1966, p. 75), car nous relevons de petites differences, 

tout au plus d'ordre sous-specifique, entre les echantillons 

americains et ouest-africains: aff. floridanus serait de plus petit 

taille et en particulier presenterait des fosses orbitaires moins 

arrondies et moins profondes, plus inclinees lateralement et 

munies a Tangle externe d'une dent moins saillante; de plus, 

les pattes ambulatoires semblent un peu plus greles. II y aura


lieu de revenir sur cette question a Paide d'un materiel plus 

important que celui dont nous disposons aujourd'hui. 

Unfortunately, the Pillsbury collections are so 

small that we cannot add anything to the solution 

of this problem at this time. The question can 

only be settled when much more material becomes 

available. 

BIOLOGY.—The West African population of 

this species lives sublittorally, in depths between 

4 and 12 or more meters; the Pillsbury specimens 

were taken in the beds of coralline algae off 

Annobon in 9 to 69 m. All of the specimens taken 

by the Calypso also were taken from bottoms with 

coralline algae (Forest and Guinot, 1966). 


collected in May and June (Forest and Guinot, 


DISTRIBUTION.—Atlantic Ocean: western Atlantic 

from Bermuda and Florida to Brazil; eastern 

Atlantic from off Gabon and the offshore 

islands of the Gulf of Guinea: Annobon, Principe 

and Sao Tome; sublittoral in the eastern Atlantic, 

in depths between 4 m and more than 12 m (9- 

69 m). Monod (1956) was the first to report the 

species from the eastern Atlantic, based on material 

from Gabon and Annobon. The only other 

records that we have found in the literature are: 



Sao Tome: 00°20'N, 06°46'E, 10 m; in front of Ponta 

Diogo Nunes, 4 m; off Ponta Diogo Nunes, 4-5 m; in front 

of Ponta Oquedelrei, 6 m; in front of Praia Lagarto, 5-6 m; 

Iogoiogo (all Forest and Guinot, 1966). 

Genus Coralliope Guinot, 1967 

Coralliope Guinot, 1967c:353 [type-species: Actumnus parvulus 

A. Milne Edwards, 1869, by original designation; gender: 

feminine].—Guinot, 1971:1076 [list of species]. 

Coralliope parvula (A. Milne Edwards, 1869) 

Micropanope parvula.—Monod, 1956:317, figs. 393-400.— 

Guinot and Ribeiro, 1962:59.—Ribeiro, 1964:11.—Forest 

and Guinot, 1966:82.—Guinot, 1967c:348 [discussion]. 

Actumnus parvulus.—Guinot, 1967c:355 [listed].—Garth, 

1968:314 [discussion]. 

Coralliope parvula.— Guinot, 1967c, figs. 3, 11; 1971:1076 

[listed]. 

SYNONYM.—Xanthodes talismani A. Milne Edwards 



Other Material: Cape Verde Islands: Sao Vicente, 20 m, 26 

Jul 1883, Talisman, syntypes of Xanthodes talismani A. Milne 

Edwards and Bouvier, 1898, 3c5, 7? (2 ov) (L, Crust. D.1567; 

W, USNM 22957). 


1900:91 (as Xanthodes talismani). 


Male Pleopod: Monod, 1956, figs. 397-400 (Cape 

Verde Islands, Senegal); Guinot, 1967c, fig. 1 la,b 

(Senegal). 


widths of 2.4 to 6.0 mm; the carapace widths 

of the ovigerous females are 6.0 mm. 

BIOLOGY.—Coralliope parvula is a sublittoral species, 

inhabiting relatively shallow water, 4-5 m, 

to a depth of 355 m. More than 90% of the 

records in the recent literature are from depths of 

less than 50 m. The deepest record is that from 

Cabo Bojador from 355 m on shelly sand and 

coral (Monod, 1956). The Calypso collected the 

species on calcareous algae in 4-5 m, calcareous 

algae, sand and coral in 7-10 m, and rocks and 

coral in 25-40 m. 


July, September, November, and December 

(Monod, 1956; Guinot and Ribeiro, 1962; Ribeiro, 

1964; Forest and Guinot, 1966). 

DISTRIBUTION.—West Africa, from the Cape 

Verde Islands and Cabo Bojador, Spanish Sahara 

southward to Angola, including the offshore islands 

Principe, Sao Tome, and Annobon in the 

Gulf of Guinea. On the mainland it is known 

only from off Cabo Bojador, several localities off 

Senegal, and one locality off Angola. Monod 

(1956) summarized the literature and reported 

material from the Cape Verde Islands, Spanish 

Sahara, and Senegal. Subsequent records include 


Cape Verde Islands: Sao Vicente (Guinot, 1967c). Baia 

da Matiota, Sao Vicente; Baia do Porto Grande, Sao Vicente, 

(3)3.5-11 m, 4-6 m, 8-10 m; Baia do Monte Trigo,


Santo Antao; Porto dos Carvoeiros (as Baia do Porto Novo), 

Santo Antao, 12 m; Baia do Tarrafal, Sao Nicolau, 16 m; 

channel between Santa Luzia and Branco, 18 and 30 m; 

Porto Ingles, Maio, 14 m; S of Ilheu Luis Carneiro, 22 m; 

Baixo Joao Leitao, 25 m; Porto da Furna, Brava, 6-20 m 

(all Guinot and Ribeiro, 1962; Ribeiro, 1964). 

Senegal: Dakar (Guinot, 1967c). 

Principe: Tinhosa Grande (as Hermano Grande) island, 

12 mi [19 km] SSW of Principe, 01°20'45"N, O7°17'37"E, 


Sao Tome: Off Ponta Diogo Nunes, 4-5 m (Forest and 


Annobon: 01°24'04"S, 05°36'45"E, 7-10 m (Forest and 


Angola: Praia Azul, Ponta da Caruita (Guinot and Ribeiro, 

1962). 

Genus Cycloxanthops Rathbun, 1897 

Cycloxanthus A. Milne Edwards, 1863:278 [invalid junior 

homonym of Cycloxanthus H. Milne Edwards, 1850 (Crustacea); 

type-species: Xantho sexdecimdentatus H. Milne Edwards 

and Lucas, 1843, by original designation and 


Cycloxanthops Rathbun, 1897b: 164 [substitute name for Cycloxanthus 

A. Milne Edwards, 1863; type-species: Xantho 

sexdecimdentatus H. Milne Edwards and Lucas, 1843; gender: 

feminine].—Guinot, 1968a:699; 1971:1067 [list of 

species, C. occidentalis not included]. 

Cycloxanthops occidentalis (A. Milne Edwards, 

1867) 

Cycloxanthops occidentalis.—Monod, 1956:301, figs. 368-370 

[Senegal; references].—Gauld, 1960:70 [Ghana].—Guinot 

and Ribeiro, 1962:57 [Angola].—Guinot, 1968a:700 [discussion].—Crosnier, 

1969:530 [Congo].—Guinot, 1971: 

1068 [listed]. 


Verde Islands and Senegal to Angola; intertidal 

to 23 m. 

Genus Domccia Eydoux and Souleyet, 1842 

Domecia Eydoux and Souleyet, 1842:234 [type-species: Domecia 

hispida Eydoux and Souleyet, 1842, by monotypy; 


Neleus Desbonne and Schramm, 1867:35 [type-species: Neleus 

acanthophorus Desbonne and Schramm, 1867, by monotypy; 

gender: masculine]. 

* Domecia acanthophora africana Guinot, 1964 

Domecia hispida.—Monod, 1956:273.—Forest, 1959:22.— 

Guinot and Ribeiro, 1962:52. [Not Domecia hispida Eydoux 

and Souleyet, 1842.] 

Domecia sp.—Ribeiro, 1964:8. 

Domecia acanthophora.—Forest and Guinot, 1966:73 [not Domecia 

acanthophora Desbonne and Schramm, 1867]. 

Domecia acanthophora forma africana Guinot, 1964:272, figs. 6, 

10-12, 16, 17.—Forest and Guinot, 1966:73 [discussion]. 


Sta 273, shore, Id (L). Sta 282, 18-37 m, nodular coralline 

algae, 1$ (W). 

Other Material: Annobon: S coast, 01°28.5'S, 05°37.5'E, 

35-55 m, 16 Jun 1967, F. Poinsard, 19 (W). 

DESCRIPTION.—Guinot, 1964:273-278 (comparison 

with the Indo-West Pacific D. hispida 

Eydoux and Souleyet, 1842 and D. glabra Alcock, 

1899). 

Figures: Guinot, 1964, figs. 6, 10-12, 16. 

Male Pleopod: Guinot, 1964, figs. 11, 12 (Cape 

Verde Islands; Annobon). 


widths of 4 mm. 

REMARKS—In 1964 Guinot clarified the status 

of the species of Domecia and recognized three 

species, two of which, D. hispida Eydoux and 

Souleyet, 1842 and D. glabra Alcock, 1899, are 

restricted to Indo-West Pacific localities. She 

pointed out that the oldest available name for the 

Atlantic species, previously thought to be D. hispida, 

actually was D. acanthophora (Desbonne and 

Schramm, 1867), a species originally described 

from Guadeloupe in the Antilles. She further 

recognized Domecia acanthophora forma africana, an 

infrasubspecific name without standing in nomenclature, 

for the smaller West African population 

of D. acanthophora, and noted (Guinot, 1964: 

281) that "les petites differences (pilosite, spinulation 

[and size]) separant nos exemplaires ouestafricains 

des specimens est-americains sont sans 

doute liees a des conditions ecologiques differentes." 

Our material is too limited to add any new 

information but we believe that east and west 

Atlantic subspecies should be recognized in this 

case.


BIOLOGY.—This species usually is associated 

with coral; it appears to be quite abundant in 

rocks and coral as well as in the beds of coralline 

algae off Sao Tome and Annobon islands in the 

Gulf of Guinea (Forest and Guinot, 1966), based 

on collections made by the Calypso. Off West 

Africa it occurs from the intertidal zone to a 

depth of 35-55 m; of 14 stations at which this 

species was taken by the Calypso, 10 were in depths 

of 10 m or less. 


collected in June and August (Guinot and Ribeiro, 

1962; Ribeiro, 1964; Guinot, 1964; Forest 


DISTRIBUTION.—Atlantic; an insular form off 

West Africa, occurring in the Cape Verde Islands 

and the offshore islands of the Gulf of Guinea, 

Annobon, Principe, and Sao Tome, in depths 

from shore to about 35 m (15-40 m and 35- 

55 m). Records in the literature include the following: 

Cape Verde Islands: No specific locality (Monod, 1956; 

Guinot, 1964). Channel between Santa Luzia and Ilheu 

Branco, 18 m (Guinot and Ribeiro, 1962; Ribeiro, 1964). 

Baia de Santa Clara, Sao Tiago, 15-35 m; Porto da Praia, 

Sao Tiago, 10 m; Ponta da Areia, Fogo, 2.5 m; Ponta 

Garbeiro [?], Brava; Ilheu de Sal Rei, Boavista, 7 m (all 


Gulf of Guinea: No specific locality (Forest, 1959). 

Principe: Ilheu Caroco, 2-8 m; 01°38'25"N, 07°21'35"E, 


Sao Tome: Between Ponta Oquedelrei and Ponta de Sao 

Sebastiao, 5 m (Forest and Guinot, 1966). Ponta Diogo Vaz, 

W coast, 0-6 and 30 m; Praia de Santa Catarina, W coast, 

3-10 m; Baia de Sao Miguel, 6-10 m; Ilheu Macaco [as ilot 

dos Cocos], 3-8 m; Morro Peixe, 2-6 m; Ilheu das Cabras, 

0-2 m; and Ponta Furada, 3-8 m (all Guinot, 1964; Forest 


Annobon: Off Annobon, 12 m (Monod, 1956; Guinot, 

1964). Santa Cruz bay, 8-12 m (Guinot, 1964). 01°24'04"S, 

05°36'45"E, 7-10 m; Isla Tortuga, NW side, 15-40 m; 

01°25'10"S, 05°36'10"E, 20-25 m (Guinot, 1964; Forest and 


Genus Epixanthus Heller, 1861 

Epixanthus Heller, 1861:323 [type-species: Epixanthus kolschii 

Heller, 1861, a subjective junior synonym of Ozius frontalis 

H. Milne Edwards, 1834, by monotypy; gender: masculine; 


* Epixanthus hellerii A. Milne Edwards, 1867 

Epixanthus hellen.—Monod, 1956:236, figs. 270-272.—Longhurst, 

1958:88.—Gauld, 1960:70.—Guinot and Ribeiro, 

1962:51.—Ribeiro, 1964:8.—Forest and Guinot, 1966: 

68.—Garth, 1968:314 [discussion].—Uschakov, 1970:445, 

447, 455 [listed]. 

SYNONYM.—Ozius corrugatus Osorio, 1887. 

MATERIAL EXAMINED.—Pillsbury Material: Fernando 

Poo: Sta 258, shore, 36*, 4$, 4 juv (L,W). 

Other Material: Sierra Leone: Aberdeen, J. Cadenat, Mar 

1948, 19 (W). 


1900:83. 


Male Pleopod: Monod, 1956, figs. 271, 272 

(Sierra Leone). 



BIOLOGY.—Epixanthus hellerii is a primarily intertidal 

species which may also occur subtidally 

in shallow water; Forest and Guinot (1966) reported 

one specimen from a depth of 3-10 m off 

Principe. Sourie (1954a) noted that this species 

was a characteristic component of the understone 

fauna of rocky shores of Senegal, and Uschakov 

(1970) found it in the inferior mesolittoral zone 

of rocky shores off Guinea, a zone with red algae 

and Padina. 

Ovigerous females have been recorded in 

March and November (Monod, 1956). 


Verde. Islands and Senegal to Angola, including 

Fernando Poo, Principe, Sao Tome, and Annobon. 

Monod (1956), who reported material from 

the Cape Verde Islands, Senegal, Guinea, Sierra 

Leone, and Ghana, summarized earlier records. 

Records in the literature since 1956 include the 

following: 

Cape Verde Islands: Baia das Gatas, Sao Vicente (Guinot 

and Ribeiro, 1962; Ribeiro, 1964). 

Guinea: lie Tamara, lies de Los; Conakry (Uschakov, 

1970). 

Sierra Leone: No specific locality (Longhurst, 1958). 

Ghana: Tenkpobo (as Tenpobo), Dixcove (Gauld, 1960). 

Principe: Ponta da Mina, beach; Ilheus dos Mosteiros, 3- 

10 m (Forest and Guinot, 1966).

124 

Sao Tome: Off Sao Tome; Iogoiogo; W of Ponta Diogo 

Nunes; Ilheu das Cabras, 0-2 m; and Santana (Forest and 


Annobon: No specific locality (Forest and Guinot, 1966). 

Genus Eriphia Latreille, 1817 

Eriphia Latreille, 1817a:404 [type-species: Cancer spinifrons 

Herbst, 1785, a subjective junior synonym of Cancer verrucosus 

Forskal, 1775, by selection by H. Milne Edwards, 

1842, in 1836-1844, pi. 14: fig. 1; gender: feminine; name 

1621 on Official List}. 

Eriphia verrucosa (Forskal, 1775) 

Cancer verrucosa Forskal, 1775:93. 

Eriphia spinifrons.—Stimpson, 1907:72 [Madeira].—Bouvier, 

1911:226 [Mauritania].—Chapman and Santler, 1955:374 

[Azores]. 

Eriphia verrucosa.—Figueira, 1960:9 [Azores].—Forest and 

Games, 1960:352 [Morocco].—Zariquiey Alvarez, 1968: 

393, figs, li, 135b [Spain; references].—Tiirkay, 1976b:61 

[listed], 67 [Madeira, Ilhas Desertas]. 

SYNONYMS.—Cancer spinifrons Herbst, 1782; Eriphia 

spinifrons angusta Czerniavsky, 1884; Eriphia 

spinifrons mediterranea Czerniavsky, 1884; Eriphia 

spinifrons orientalis Czerniavsky, 1884. 

DISTRIBUTION.—Eastern Atlantic from the Bay 

of Biscay to Mauritania, including Madeira, the 

Azores, and the Mediterranean; intertidal. 

Genus Euryozius Miers, 1886 

Euryozius Miers, 1886:142 [type-species: Xantho bouvieri A. 

Milne Edwards, 1869, by monotypy; gender: masculine].—Guinot, 

1968c:320 [listed], 325 [discussion][genus 

restricted to type-species]; 1971:1077 [included species]. 

Gardineria Rathbun, 1911:236 [type-species: Gardinena canora 

Rathbun, 1911, by monotypy; gender: feminine].— 

Guinot, 1967a:264; 1968b: 156; 1968c:325; 1971:1077. 

DISCUSSION.—Guinot (1967a: 264, footnote) 

noted that "Gardineria canora appartient au meme 

genre que l'espece ouest-africaine Pseudozius bouvieri 

(A. MILNE EDWARDS). Ces deux especes appartiennent, 

selon nous, au genre Euryozius etabli 

par MIERS en 1886 et dont le plus proche parent 

serait Carpilius." In that same paper Gardineria was 

assigned to the Xanthinae whereas Pseudozius 


sensu stricto, was assigned to the Menippinae. In 

another paper, part of an overall review of some 

of the more difficult xanthid genera, Guinot 

(1968b: 156) resurrected Euryozius from the synonymy 

of Pseudozius and placed both Euryozius 

and Gardineria in the Menippinae. In a subsequent 

paper in that same series, Guinot (1968c:320, 325) 

recognized the subfamily Carpilinae Ortmann, 

1893, for the Recent genera Carpilius, Euryozius, 

and Gardineria, and for two other genera based on 

fossils. 

In her account of Gardineria and Euryozius, 

Guinot (1968c:325-330) pointed out the numerous 

similarities and some differences between the 

species of these two monotypic genera, concluding 

(p. 328): "la plupart des ces differences ne semblent 

que d'ordre specifique." Earlier (p. 326) she 

noted, 

En fait, canora est tellement proche de bouvieri que notre 

premier mouvement fut d'unir les deux genres Gardineria et 

Euryozius. Pratiquement, tous les caracteres concordent. Nous 

avons maintenu separe Gardineria surtout pour la raison que 

le male de l'espece de RATHBUN n'est pas connu. 

Finally, Guinot (1971:1077), in her summary 

of her xanthid revision, noted that Gardineria is 

"tres proche, sinon synonyme, d'Euryozius." 

We have examined the unique female holotype 

of Gardineria canora (USNM 41535) from Providence 

Island, western Indian Ocean, and conclude 

that it should be known as Euryozius canorus 

(Rathbun, 1911). As Guinot pointed out, E. canorus 

shares all essential features with E. bouvieri, 

including the stridulating ridge under the carapace, 

as well as the short branchial ridges. Miers 

(1886:142) and Rathbun (1911:236) both commented 

on the well-developed but incomplete 

branchial ridges, a unique feature of this genus. 

The Indian Ocean species differs from its Atlantic 

counterparts in having two faint notches in the 

orbit; at most one such notch is visible in our 

material of the Atlantic species. 

Three species of Euryozius can be recognized in 

the Atlantic, E. sanguineus (Linnaeus, 1767) from 

Ascension and Saint Helena islands in the central 

Atlantic; E. bouvieri (A. Milne Edwards, 1869) 

from Madeira, the Azores, and the Cape Verde

NUMBER 306 

Islands; and a new species, E. pagalu, from the have carapace lengths of 7.4 and 8.0 mm. The 

Gulf of Guinea. Euryozius canorus, an insular spe- male from Madeira has a carapace length of 

cies like those in the Atlantic, is the only repre- 28.8 mm, carapace width of 46.5 mm. Barrois' 

sentative of the genus from the Indo-West Pacific (1888) specimen was 25.8 mm long, 38.5 mm 

region. 

wide. Guinot (1968c) figured a female 30 mm 

long, 47 mm wide. This species, like E. sanguineus, 

Euryozius bouvieri (A. Milne Edwards, 1869) is a large species. 

FIGURE 27a 

REMARKS.—See comments under account of E. 

pagalu, new species. 

Xantho Bouvieri A. Milne Edwards, 1869:377.—Stebbing, BIOLOGY.—Almost nothing is known of the 


ecology of this species. Barrois's (1888) specimen 

Pseudozius bouvieri.—Miers, 1886:142 [listed].—Monod, 1956: 

239, figs. 275-278 [part].—Figueira, 1960:9.—Guinot-Du- 

from the Azores was taken on a jetty among 

mortier and Dumortier, 1960:121, 144 [table 2], fig. 4 rocks. The specimen reported by Figueira (1960) 

[stridulation].—Guinot, 1967a:264 [footnote], 272 [discus- from the Azores was found dead on a beach. One 

sion]; 1968b: 156 [discussion]; 1968c:325 [discussion]. of the specimens collected by the Talisman in the 

Ozius Edwardsi Barrois, 1888:10, pi. 1 [fig. 1 in color].— Cape Verde Islands was taken at a depth of 

Stebbing, 1893:63 [discussion]. 

Pseudorius Bouvieri.—A. Milne Edwards and Bouvier, 1898: 

20 m, and one of the specimens identified with 

185 [discussion; error for Pseudozius]. 

this species by Tiirkay (1976b:67) was taken on 

Pseudozius Bouvieri.—A. Milne Edwards and Bouvier, 1900: an "aus 5-6 m Tiefe gegreiferten Steinblock." 

82, pi. 15: figs. 11-14.—Bohn, 1901:297, 298, 302, 308, DISTRIBUTION.—Euryozius bouvieri is an insular 

309, 315, 316, 318 [morphology].—Monod, 1933b:529 

[part of synonymy only; listed]. 

species occurring in the central eastern Atlantic, 

Euryozius bouvieri.—Guinot, 1968b: 156 [discussion], fig. 8 [nofrom 

the littoral zone to a depth of about 20 m. 

locality]; 1968c:325, 326, 327, 329, 330 [all discussion], fig. Records in the literature include the following: 

8 [no locality], pi. 2: fig. 2; 1971:1077 [listed].—Tiirkay, 

1976b:61 [listed], 67. 

Azores: No specific locality (A. Milne Edwards and Bouvier, 

1900). Ponta Delgada, Ilha de Sao Miguel, jetty 

MATERIAL EXAMINED.—Pillsbury Material: None. (Barrois, 1888). Volcano Capelhinos [= Rochas dos Capel- 

Other Material: Madeira: No specific locality, 1(5 (BM). hinos], on beach (Figueira, 1960). 

Cape Verde Islands: No specific locality, syntypes, 2 dry Madeira: No specific locality (Monod, 1956). Funchal, 

specimens (MP). 

fish market (from stomachs of littoral fishes and Phycis phycis 

DESCRIPTION.—A. Milne Edwards, 1869:377; 

Barrois, 1888:10 [as Ozius Edwardsi]. 

(Linnaeus)); between Ponta da Oliveira and Ponta da Garajau, 

5-6 m (Tiirkay, 1976b). 

Cape Verde Islands: No specific locality (A. Milne Ed- 

Figures: Barrois, 1888, pi. 1; Monod, 1956, figs. wards, 1869; A. Milne Edwards and Bouvier, 1900; Bohn, 

275-278; Guinot, 1968c, pi. 2: fig. 2. 

1901; Monod, 1956; Guinot, 1968c). Baia das Gatas (as Baie 

Male Pleopod: Monod, 1956, figs. 276-278 

(Azores). 

Color: Barrois (1888:12, pi. 1: fig. 1) described 

the color in the type of Ozius edwardsi as follows: 

de la Gatta), Sao Vicente [indicated as type-locality] 

(Monod, 1956). Between Ilheu Branco and Ilheu Raso (as 

ilot Razzo), Sao Vicente (A. Milne Edwards and Bouvier, 

1900; Monod, 1956). 

Un beau rouge orange, legerement teinte de jaune en 

arriere; les pattes ravisseuses sont de meme teinte que la 

carapace, sauf les doigts qui sont d'une noir intense, tandis 

According to Barrois (1888:12), A. Milne Edwards 

informed him that Barrois's specimen was 

conspecific with a species collected by the Talis- 

que les pattes ambulatoires sont d'un nuance plus claire et 

plus rosee. 

man in the Canary Islands in 1883. Monod (1956: 

240, footnote) correctly pointed out that the Ca- 

The pereiopods are shown to be of uniform nary Islands were not among the localities listed 

color in his figure. 

for the species by A. Milne Edwards and Bouvier 

MEASUREMENTS.—The two syntypes, juveniles, (1900) in their report on the Talisman collections. 

125

126 

h 


FIGURE 27.—Atlantic species of Euryozius. Euryozius bouvieri (A. Milne Edwards), dry syntype, cl 

7.4 mm: a, carapace. Euryozius pagalu, new species, paratype, male, cl 9.0 mm, Pillsbury Sta 275: 

b, carapace; c, second pereiopod; d, fifth pereiopod; e, abdomen. Euryozius sanguineus (Linnaeus), 

male, cl 11.7 mm, Ascension Island:/, carapace; g, second pereiopod; h, fifth pereiopod; i, 

abdomen. 

* Euryozius pagalu, new species 

FIGURES 27b-e, 28 

Pseudozius Bouvieri.—Balss, 1921:63 [not Xantho bouvieri A. 

Milne Edwards, 1869]. 

Pseudozius caystrus.—Capart, 1951:19 [error for bouvieri] [not 

Panopeus caystrus Adams and White, 1848]. 

Pseudozius bouvieri.—Capart, 1951:164, fig. 62.—Monod, 

1956:239 [part, not figs. 275-278].—Rossignol, 1962: 

116.—Forest and Guinot, 1966:68. 

young crab.—Cousteau, 1958:393 [color photograph]. 

Euryozius bouvieri.—Guinot, 1968c, figs. 15, 16, pi. 1: fig. 3. 


Sta 275, 9-69 m, rubble of coralline algae, 66, 4$, 16 juv (L, 

W) (largest male is holotype). Sta 282, 18-37 m, nodular 

coralline algae, 36, 10$ (1 ov) (L, W). Sta 283, 51-55 m, 

nodular coralline algae, 36, 19, 9 juv (L). 

Other Material: Annobon: 01°28'S, 05°38'30"E, 80 m, 11 

Dec 1965, A. Crosnier, Ombango, \6 (W). 01°28'40"S, 05° 

35'50"E, 11 Dec 1965, A. Crosnier, 39 (W). 

DESCRIPTION.—Size small, maximum carapace 

width about 18 mm. Carapace (Figure 21b, 28a) 

oval, length 0.6 to 0.7 times width. Front broad, 

fronto-orbital margin 0.54 to 0.7 times carapace 

width. Front composed of 2 sinuate lobes, almost 

straight in dorsal view, sinuate in oblique anterior 

view in adults (convex mesially, concave laterally), 

less sinuate in smaller specimens; lobes 

separated by a distinct median notch. Front 

flanked dorsally by shallow groove and low, irregular 

ridge above groove. Orbits smooth, with 

indications of 1 incision in dorsal margin of 

smaller specimens, upraised margins continuing 

laterally and ventrally into suborbital margin,


FIGURE 28.—Euryoziuspagalu, new species, paratypes. Female, 

cb 13.5 mm, Pillsbury Sta 282: a, dorsal view. Male, cb 14.4 

mm, Pillsbury Sta 275: b, gonopod; c, apex of gonopod. 

lateral part of ridge appearing as tubercle in 

dorsal view. Anterolateral margin cristate, unarmed 

anterior to lateralmost point of carapace, 

there produced into 2 low anterolateral teeth, 

anteriormost, anterior to midlength of carapace, 

occasionally not developed, posteriormost distinct 

behind midlength, each with short ridge on carapace, 

posteriormost the longer. Low irregular 

swelling extending laterally from orbit above lateral 

margin. Carapace regularly convex from 

front to back, regions indistinct, dorsal surface 

punctate, anterior surface, in arc behind front, 

rougher than posterior surface. Stridulating ridge 

under anterolateral border of carapace arising 

under first anterolateral tooth, extending about 

to midlength of lower margin of orbit, composed 

of about 60 striae. 

Chelipeds stout, unequal. Carpus with dorsal 

and outer surface irregular, deeply punctate; inner 

margin produced into rectangular lobe (the 

stridulating lobe) terminating anteriorly in blunt 

inner tooth. Surface of palm deeply punctate and 

longitudinally sulcate dorsally, outer surface 

smooth. Fingers dark, evenly toothed, with bilobed 

enlarged basal tooth. 

Walking legs slender, third (Figure 27c) much 

the longest, overreaching second by about 1/2 to 

1/3 of dactylus. First walking leg with at most a 

low tubercle distally on inner margin of carpus. 

Dactylus of each walking leg longer than respective 

propodus, grooved laterally and dorsally, 

ornamented with a few slender, simple setae. 

Propodi of third and fourth walking legs grooved 

laterally, length of propodus of third about 3 

times depth, length of that of fourth 2 times depth 

(measured dorsally). 

Abdomen (Figure 27*) of 7 distinct somites in 

both sexes, terminal somite of male abdomen 

longer than penultimate. Male pleopods as figured 

by Guinot (1968c, figs. 15, 16). 

Color: Like the other Atlantic species of Euryozius, 

E. pagalu is reddish. Capart (1951:165) noted 

that in his material the general color was "rose 

orange, les doigts bruns." Cousteau (1958) illustrated 

a juvenile from Sao Tome; it is red with 

the pereiopods banded red and white, the latter 

at the articulations. The color pattern of our 

material has completely faded. 


widths ranging from 8.0 to 16.2 mm; the 

carapace width of the ovigerous female is 11.2 

mm. The largest specimens examined by Capart 

(1951) were a male, 10.5 X 17 mm, and a female, 

11.5 X 18 mm; the latter is the largest known 

representative of the species. Guinot (1968c) figured 

a male, 9.7 X 14.5 mm, from the Calypso 

collection. 

REMARKS.—Specimens from different geographic 

areas identified with Euryozius bouvieri in 

the past have exhibited an order of magnitude 

difference in size. For example, a syntype of Xantho 

bouvieri from the Cape Verde Islands measured 

8 X 12 mm (A. Milne Edwards, 1869), whereas 

another specimen from the same area measured 

30 X 47 mm (Guinot, 1968c). The type of Ozius 

edwardsi, from the Azores, measured 25.8 X 38.5

128 

mm (Barrois, 1888), and syntypes of Pseudozius 

mellissi, from Ascension and Saint Helena Islands 

in the central Atlantic, measured 25 X 38 mm 

and 35.5 X 59 mm (Miers, 1881b). Our specimens 

from Annobon, more than 33 in all, comprising 

an ovigerous female as well as adult males, do not 

exceed 10 X 16 mm, and are obviously adult at 

a size of 7 X 11 mm. 

This size disparity, considered together with 

two features reported in the literature, pereiopod 

length and color, suggested to us that the species 

now known as E. bouvieri actually might consist of 

a complex of species. Euryozius bouvieri is considered 

to be a widely distributed insular species in 

the central and eastern Atlantic (Monod, 1956: 

240), and, as noted by Monod "Cette distribution 

presque uniquement insulaire est singuliere." 

Such an insular distribution pattern for a single 

species is difficult to understand or explain. We 

decided to investigate the possibility that different 

species occurred in each of the three geographic 

areas from which E. bouvieri was reported: 

the mid-Atlantic, on Ascension and Saint Helena 

islands; the central eastern Atlantic, from Madeira, 

the Azores, and the Cape Verde Islands; and 

the Gulf of Guinea. 

The problem is further complicated by the fact 

that large specimens from both the mid-Atlantic 

{Pseudozius mellissi), as well as the central eastern 

Atlantic (Ozius edwardsi and a female from the 

Cape Verde Islands taken by the Talisman, see 

Guinot, 1968c, pi. 2: fig. 2), have a very narrow 

front, which in the specimens from the mid-Atlantic, 

at least, is distinctly four-lobed. In our 

specimens from Annobon, as well as in the syntypes 

of Xantho bouvieri, the front is comparatively 

quite broad and is produced into two transverse 

or sinuous lobes. 

In his original account of Xantho bouvieri from 

the Cape Verde Islands, A. Milne Edwards (1869: 

378) remarked that in his specimens the fourth 

pereiopods were quite long, exceeding all of the 

other legs by the length of the dactylus. This is 

not the case in our series from Annobon or in the 

young male (6.6 X 9.8 mm) from Ascension that 

we have examined; in our specimens the fourth 


legs exceed the third by no more than half the 

dactylus. The small specimens from the Cape 

Verde Islands reported by Bouvier appear to 

represent a species distinct from that occurring 

on Annobon or that occurring in the mid-Atlantic. 

Those small specimens from the Cape Verde 

Islands represent either a distinct small species or 

the juveniles of the large species also known from 

the Cape Verde Islands. We believe that it is 

unlikely that two species of Euryozius occur in the 

Cape Verde Islands and that the types of Xantho 

bouvieri are based on the young of a species, which 

in that area attains a size of at least 30 X 47 mm. 

The second feature mentioned in the literature 

that attracted our attention is color. Euryozius 

bouvieri, sensu lato, is known to be a bright red 

crab. Melliss (1875:275) referred to an orange-red 

crab. A. Milne Edwards (1869), Miers (1886), 

Barrois (1888), and Capart (1951) all referred to 

the red color of this species. Barrois (1888, pi. 1: 

fig. 1) illustrated his Ozius edwardsi in color and 

showed the pereiopods to be uniform in color, not 

banded. Cousteau (1958: 393 photograph) illustrated 

a juvenile specimen from Sao Tome in 

color, showing walking legs distinctly banded red 

and white. The walking legs in all of our specimens 

from Ascension are distinctly banded red 

and white. 

The available material suggests to us that there 

are three species of Euryozius in the Atlantic: a 

large species with banded legs which lives on 

Ascension and Saint Helena; a large species with 

uniform coloration which is found at Madeira, 

the Azores, and the Cape Verde Islands; and a 

small species with banded legs which lives in the 

Gulf of Guinea. 

Four names are available for two of these taxa. 

The large central Atlantic species from Ascension 

and Saint Helena was named Pseudozius mellissi 

by Miers (1881b:432); until now it was considered 

a synonym of Xantho bouvieri A. Milne Edwards 

from the Cape Verde Islands. However, Miers' 

name is not the oldest available name for this 

species. Linnaeus (1767:2970) described an 

oceanic crab, Cancer sanguineus, a species with an 

obsoletely bidentate carapace and a four-lobed


front that, judging from the name, is red. Herbst 

(1785, in 1782-1804) noted that this species occurred 

at Ascension. The central atlantic form is 

identifiable as Euryozius sanguineus, which, as noted 

by Linnaeus, has two low anterolateral teeth on 

the carapace and a distinctly four lobed front. 

Two names also are available for the species 

occurring in the central part of the eastern Atlantic, 

Xantho bouvieri A. Milne Edwards, 1869 and 

Ozius edwardsi Barrois, 1888, the former, based on 

juveniles, having priority. 

The third species, from the Gulf of Guinea, is 

named herein. 

The possibility exists that Barrois was incorrect 

in showing his species to have a uniform color 

pattern or that A. Milne Edwards was incorrect 

in his account of the walking legs of his species. 

Moreover, all of the Atlantic species of Euryozius 

might prove to be conspecific in which case the 

name E. sanguineus would have to be used. Alternatively, 

the same large species might occur in 

the central as well as in the eastern Atlantic and 

E. bouvieri also might occur in the Cape Verde 

Islands and the Gulf of Guinea. Additional fresh 

material will have to be studied before this problem 

can be settled. 

A detailed account of E. sanguineus is in preparation 

by Manning in collaboration with Fenner 

A. Chace, Jr., in a review of the marine decapods 

of Ascension Island. 

We have been able to compare our material of 

E. pagalu with three male specimens of E. sanguineus 

(Figure 21f-i) from Ascension Island (6.6 X 

9.8 mm, 17.6 X 27.8 mm, and 28.6 X 45.8 mm) 

and one female of E. sanguineus from Saint Helena 

(42.5 X 71.1 mm) in the collections of the Smithsonian 

Institution. The specimens from Ascension, 

newly collected, all have the walking legs 

banded red and white. The fronto-orbital width 

of the three larger specimens of E. sanguineus 

ranges from 0.45 times the carapace width in the 

smaller male (17.6 X 27.8 mm) to 0.37 times the 

carapace width in the large female. In the smallest 

male from Ascension, of comparable size with our 

specimens of E. pagalu from Annobon, it is 0.67 

times the carapace width; it ranges from 0.54 to 

0.7 times the carapace width in E. pagalu. 

A comparison of our material with the smallest 

specimen of E. sanguineus from Ascension shows 

the following differences: the frontal lobes are less 

sinuous; the carapace and dorsal surfaces of the 

carpus and propodus of the claw are rougher; the 

third pereiopod overreaches the second by less of 

its length, 1/3 to 1/2 its length in E. pagalu, 2/3 

of its length in E. sanguineus; the pereiopods appear 

to be slenderer in E. pagalu; there is a distinct 

tubercle (almost a spine in larger specimens) on 

the inner, distal surface of the carpus of the 

second and third periopods in E. sanguineus; this 

tubercle is scarcely or not at all detectable in even 

the largest specimens of E. pagalu. 

Guinot (1968c:330) commented on differences 

in pilosity, possibly related to sex or size, in 

material available to her. Our material exhibits 

no obvious differences. 

TYPE-LOCALITY.—Off Annobon Island, Gulf of 

Guinea, 01°24'S, 05°37'E to 01°24'S, 05°38'E, in 

9 to 69 m. 

DISPOSITION OF TYPES.—The holotype (Crust 

D. 31539), the largest male from Pillsbury Sta 275, 

measuring 10.3 X 16.2 mm, is in the Rijksmuseum 

van Natuurlijke Historie, Leiden. Paratypes are 

in the collections of that museum and in the 


ETYMOLOGY.—The specific name is derived 

from the modern name for Annobon Island, Pagalu. 

BIOLOGY.—Euryozius pagalu is generally a sublittoral 

species, living from shore to a depth of 

80 m. Of 16 depth records available, only one is 

from shore (shore to 2 m); that and two others, 

2-6 m and 7-10 m, are in 10 m or less. Five 

records (9-69 m, 51-55 m, 60 m, 60 m, and 

80 m) are from depths generally greater than 50 

m. Fifty percent of the records are from depths 

between 15 and 40 m, suggesting a preference for 

that depth range. 

Only two records are from localities other than 

Sao Tome and Annobon islands: one specimen 

was found dead on a beach off the Congo (Rossi 

gnol, 1962), and Capart (1951) reported 4 males

130 

and 3 females from sandy mud in 20-22 m off 

Angola. 

Apparently the preferred habitat for this species 

is the nodular coralline algae found off Annobon 

and Sao Tome; all of the Pillsbury specimens 

and eight of the ten lots (25 specimens) 

collected by the Calypso were taken in coralline 

algae. Other kinds of bottom noted for the Calypso 

collections (Forest and Guinot, 1966) include: 

rocks and coral; one specimen from a sponge on 

rocky bottom with calcareous algae; sand, with 

calcareous algae; sand, algae, rocks, or coral. 

The only ovigerous female known was taken 

by the Pillsbury in May. 

DISTRIBUTION.—Euryozius pagalu is primarily an 

insular species occurring in the Gulf of Guinea; it 

lives from the shore to a depth of 80 m. Records 

in the literature include the following: 

Sao Tome: No specific locality (Cousteau, 1958). Ilheu 

Gago Coutinho (as Ilha das Rolas) (Balss, 1921). 00°25- 

15"N, 06°43'E, 8-30 m; Morro Peixe, 2-6 m (Forest and 

Guinot, 1966). W of Ponta Diogo Nunes, shore to 2 m (Forest 

and Guinot, 1966; Guinot, 1968c). 

Annobon: OfT Annobon, 60 m (Monod, 1956). 01°24'- 

04"S, 05°36'45"E, 7-10 m; 01°25'10"S, 05°36'10"E, 20- 

25 m; 01°25'12"S, 05°36'05"E, 20 m; 01°26'15"S, 

05°35'40"E, 60 m; O1°27.5'S, 05°36.5'E, 35 m; N of San 

Antonio, 23 m; Isla Tortuga, NW coast, 15-40 m (all Forest 


Congo: Pointe Indienne, beach (Rossignol, 1962). 

Angola: 8 mi [12.8 km] W of Rio Cuanza, 09°20'S, 

13°04'E, 20-22 m (Capart, 1951). 

Genus Eurypanopeus A. Milne Edwards, 1878 

Eurypanopeus A. Milne Edwards, 1878, in 1873-1881:318 

[type-species: Panopeus crenatus H. Milne Edwards and 

Lucas, 1843, by subsequent designation by Fowler, 1912: 


* Eurypanopeus blanchardi 


FIGURE 296 

Xantho parvulus.—Dana, 1852b: 170.—Miers, 1881a:214 [discussion]. 

[Not Cancer parvulus Fabricius, 1793 = Xanthodius 

parvulus; not Xantho parvulus sensu H. Milne Edwards, 1834 

= Eurypanopeus abbreviate (Stimpson, 1860); not Actumnus 

parvulus A. Milne Edwards, 1869 = Coralliope parvula.] 


Xantho parvula.—Stimpson, 1858a:31; 1907:47 [= Xantho parvulus 

sensu Dana, 1852, not parvulus auctorum]. 

Panopeus Blanchardi A.. Milne Edwards, 1880, in 1873-1881: 

308, footnote [nomen nudum]; 1881, in 1873-1881:323. 

Panopeus blanchardi.—Miers, 1886:129 [discussion]. 

Eurypanopeus parvulus.—A. Milne Edwards and Bouvier, 1900: 

99, pi. 17: fig. 7 [not Cancer parvulus Fabricius, as in first 

entry above]. 

Eurypanopeus blanchardiQ).—Rathbun, 1921:440, fig. 20, pi. 

36: figs. 2, 3. 

Panopeus parvulus.—Monod, 1956:329, figs. 416-434.—Gauld 

and Buchanan, 1959:127.—Gauld, 1960:70.—Guinot and 

Ribeiro, 1962:62—Rossignol, 1962:118.—Monod, 1963, 

fig. 35 [no material].—Ribeiro, 1964:13.—Forest and 

Guinot, 1966:85.—Uschakov, 1970:447, 455 [listed]. 


Poo: Sta 257, shore, 5(5 (W). Sta 258, shore, 5


Elle [E. abbreviatus] ressemble beaucoup a un Panope de 

la cote occidentale d'Afrique que j'ai designe dans les cata- 

logues du Museum sous le nom de Panopeus Blanchardi, dont 

les pinces sont depourvues de dent basilaire et dont les bords 

latero-anterieurs sont un peu plus courts. 

Rathbun (1921:440, 441) who used the name 

Eurypanopeus blanchardi (?) for the species and who 

gave the only description of the species, commented 

in her account based on a specimen from 

Angola: 

I think, however, that this is with little doubt the species 

named blanchardi. Bouvier makes it a synonym of the Amer- 

ican parvulus or abbreviatus [in A. Milne Edwards and Bouvier, 

1900:99]. Our African specimen, however, differs from abbreviatus 

of equal size as follows. The carapace is slightly 

narrower in proportion to its length, but is wider in its 

posterior half, the posterolateral margins less convergent, so 

that the carapace appears less regularly oval than in abbreviatus. 

The anterolateral teeth are more pronounced and more 

elevated; the chelipeds more rugose, fingers more deeply 

grooved. A small piece of the sternum shows at each end of 

the second abdominal segment, between it and the coxa of 

the last leg; this is not the case in abbreviatus. 

Rathbun (1930) did not synonymize E. blanchardi 

with E. abbreviatus in her account of the 

latter species. 

With one exception, the characters pointed out 

by Rathbun to differentiate E. blanchardi from E. 

abbreviatus are well marked in the materal available 

to us. In addition to the difference in proportion 

of the body, in our specimens of E. blanchardi 

on the carapace the regions are more distinct, 

the surface is much more uneven, and the 

lines of granules are much better developed, as 

shown by Monod (1956, fig. 417). A character 

that appears to have some value is the width of 

the third anterolateral tooth. In Eurypanopeus abbreviatus 

this tooth is broader than the fourth 

tooth, whereas in E. blanchardi it is subequal in 

length to the fourth tooth. The male pleopods of 

the two species, as figured by Monod (1956, figs. 

422-434) for E. blanchardi and by Williams (1965, 

fig. 183K) for E. abbreviatus are essentially similar, 

except that in E. blanchardi the slender, subterminal, 

triangular lobe overreaches the irregularly 

shaped median lobe, whereas in E. abbreviatus it 

does not. 

The character mentioned by Rathbun, which 

is of relatively little value, is the exposure of the 

small piece of sternum between the second abdominal 

somite and the coxa of the last leg. In E. 

blanchardi the part of the sternum is clearly visible, 

but in material of E. abbreviatus examined for 

comparison the sternum is usually concealed in 

specimens from the Antilles (USNM 15658) 

whereas it usually is visible in specimens from 

Brazil (USNM 59844). 

That A. Milne Edwards (1881, in 1873-1881) 

described this species as lacking an enlarged basal 

tooth on the dactylus of the chela is puzzling, for 

it is clearly present in our material. Monod (1956: 

334), who examined what might be the types of 

E. blanchardi from Gabon, did not mention 

whether or not he observed the tooth. 

Although Rathbun (in Stimpson, 1907:47, 

footnote) identified Dana's Xantho parvulus and 

Stimpson's Xantho parvula, both from the Cape 

Verde Islands, with Xanthias melanodactylus [= Nanocassiope 

melanodactylus, below], we believe that 

Monod (1956:329) was correct in identifying 

those records with E. blanchardi (as Panopeus parvulus). 

Dana's description fits E. blanchardi quite 

well. 

The carapace of E. blanchardi (Figure 296) does 

appear to be less oval in shape than that of E. 

abbreviatus (Figure 29a) as noted by Rathbun, but 

the length/width ratio of representatives of the 

two species overlaps broadly. Dana (1852b) noted 

that in his specimens from the Cape Verde Islands 

the length/width ratio was 1 to 1.53; his reported 

measurements were 5 X 7% lines (10.6 X 16.2 

mm). Monod (1956) included measurements of 

numerous specimens in his account; the length/ 

width ratio for all material reported by Monod 

ranged from 1:1.17 to 1:1.64. The ratio for 22 

larger specimens, carapace length 8.5 mm or 

more, ranged from 1:1.33 to 1:1.64, with a mean 

of 1.44. The ratio for 13 specimens from the 

western Atlantic from Brazil and Trinidad ranges 

from 1:1.47 to 1:1.56; the mean is 1:1.53. The 

difference in shape of the carapace of E. blanchardi 

and E. abbreviatus is shown in Figure 29. 

The specific epithet parvulus has been applied

132 

a 

FIGURE 29—Carapaces: a, Eurypanopeus abbreviatus (Stimpson), 

male, cb 20.0 mm, Brazil; b, Eurypanopeus blanchardi (A. Milne 

Edwards), male, cb 19.1 mm, Pillsbury Sta 257. 

to three distinct species of xanthid crabs from the 

Atlantic by three different authors. Fabricius 

(1793:451) named Cancer parvulus which Rathbun 

(1930:305), who had examined the type, placed 

in Leptodius. H. Milne Edwards (1834:395) identified 

a second species as Xantho parvulus (Fabricius); 

Rathbun (1930:404) placed this species in 

Eurypanopeus. Actumnus parvulus was described by 

A. Milne Edwards (1869), was placed in Micropanope 

by Monod (1933b, 1956), and finally was 

assigned to Coralliope by Guinot (1967c). 

At least three authors, Dana (1852b: 170), A. 

Milne Edwards (1881, in 1873-1881:322), and 

Benedict and Rathbun (1891:369), considered 

Cancer parvulus Fabricius and Xantho parvulus sensu 

H. Milne Edwards to be synonymous. Rathbun 

(1930), without discussing the problem, correctly 


replaced parvulus sensu H. Milne Edwards with 

the next oldest available name, Panopeus abbreviatus 

Stimpson, 1860, which she, as A. Milne Edwards 

(1881, in 1873-1881:320) had before her, considered 

to be a Eurypanopeus. 

The problem of the two identical specific 

names has extended even further. Monod (1956: 

285), in discussing the synonymy and affinities of 

Xanthodius inaequalis [as Xantho (Leptodius)], compared 

it with an American species which Rathbun 

(1930:305) called Leptodius parvulus (Fabricius, 

1793), and concluded that the eastern and 

western Atlantic populations represented distinct 

species. Because of the confusion over the use of 

parvulus, Monod suggested that the American 

population be called Xantho (Leptodius) americana 

(Saussure, 1858): "Comme par ailleurs le binome 

"Xantho parvulus" ne peut que provoquer de nouvelles 

confusions, je serais d'avis d'utiliser Xantho 

(Leptadius) [sic] americana (SAUSSURE)." Guinot 

(1968a: 712; 1971:1068) followed Monod in substituting 

americanus for parvulus; she assigned De 

Saussure's species to Xanthodius. That species 

should be known as Xanthodius parvulus (Fabricius, 

1793). 

BIOLOGY.—Eurypanopeus blanchardi is an intertidal 

species, which may occur subtidally to a depth 

of 10-30 m; the majority of records in the literature 

are based on specimens taken intertidally on 

rocky shores. Gauld (1960) noted that it is common 

on rocky shores off Ghana, and Gauld and 

Buchanan (1959) characterize it as a member of 

the understone fauna. Uschakov (1970) found it 

in the mesolittoral zone of rocky shores of Guinea, 

in habitats with red algae and Padina. 

It has been taken together with Panopeus ajricanus, 

but in general that species prefers lagoons 

and estuaries rather than rocky shores. 


February, March, May, June, and November 

(Monod, 1956; Forest and Guinot, 1966; present 

paper). The species probably spawns all year. 

DISTRIBUTION.—West Africa, where it occurs 

from the Cape Verde Islands and Mauritania 

southward to Angola, including the offshore islands 

in the Gulf of Guinea, Principe, Sao Tome,


and Fernando Poo (new record); usually intertidal, 

rarely to a depth of 10-30 m. Records in the 

literature include the following. 

West Africa: No specific locality (A. Milne Edwards, 

1880, 1881, in 1873-1881; Monod, 1963). 

Cape Verde Islands: No specific locality (Dana 1852b). 

Sao Tiago (as St. Jago) (Stimpson, 1858a). Porto da Praia, 

Sao Tiago, 12 fm (22 m) (Stimpson, 1907); beach (Guinot 

and Ribeiro, 1962; Ribeiro, 1964); same, (as La Praya), 10- 

30 m (A. Milne Edwards and Bouvier, 1900). Sao Vicente 

(Monod, 1956). 

Mauritania: No specific locality; Bilaouakh [?] (Monod, 

1956). 

Senegal: Goree (?); Ngazobil, shore (Monod, 1956). 

Guinea: Conakry, shore (Monod, 1956). lie Tamara and 

ile Roume, lies de Los, shore (Uschakov, 1970). 

Sierra Leone: Aberdeen, near Freetown, shore (Monod, 

1956). 

Ghana: Accra, shore (Monod, 1956). Christiansborg, Accra 

(Monod, 1956). Tenkpobo (as Tenpobo), shore or reef 

(Monod, 1956; Gauld and Buchanan, 1959). Apam, Prampram; 

Dixcove (Monod, 1956). Tenkpobo to Dixcove, shore 

(Gauld, 1960). 

Principe: Praia Ponta da Mina, shore (Forest and Guinot, 

1966). 

Sao Tome: Harbormaster's dock, shore (Forest and 


Gabon: No specific locality (Monod, 1956). 

Congo: Baie de Pointe-Noire (Rossignol, 1962). 

Angola: Santo Antonio do Zaire (as San Antonio), less 

than 20 ft [6.1 m] (Rathbun, 1921). Lucira, beach (Guinot 


Genus Globopilumnus Balss, 1933 

Globopilumnus Balss, 1933b:6 [type-species: Pilumnus globosus 

Dana, 1852, by original designation; gender: masculine]. 

Globopilumnus afiricanus 


MATERIAL EXAMINED—Pillsbury Material: None. 

Other Material: Senegal: No other data, 2? (W). 


Figures: Capart, 1951, fig. 51, pi. 1: fig. 15, pi. 

3: figs. 7, 8; Monod, 1956, figs. 249-257. 

Male Pleopod: Capart, 1951, pi. 3: figs. 7, 8 

(Angola); Monod, 1956, figs. 254-257 (Senegal); 

Turkay, 1976b, fig. 4 (Madeira). 

Color: "Rougeatre plus ou moins fonce, a maculatures 

brun chocolat ou violettes; epines antero-laterales 

et doigts de chelipedes noirs" 

(Monod, 1956:235). The black color of the fixed 

finger of the chela extends well onto the palm 

(Monod, 1956, fig. 251). 

MEASUREMENTS—Our specimens have carapace 

widths of 20 and 22 mm. 

BIOLOGY.—Globopilumnus africanus is a shallow 

water species, generally found intertidally or subtidally; 

of 22 records for which depth is given in 

the recent literature, seven are from shore, 11 are 

from depths between shore and 12 m, and four 

are from depths greater than 12 m (15, 20-22, 30, 

and 35 m). Intertidally it appears to be part of 

the rocky shore fauna, where it lives among rocks 

or in algae; some specimens have been taken with 

bryozoans. Subtidally it has been taken on sandy 

mud, rocks, rocks and coral, rocks and sand, or 

on sand with calcareous algae, rocks and coral. 

Ovigerous females have been collected in January 

and September (Capart, 1951; Guinot and 


DISTRIBUTION.—Off West Africa, from Madeira, 

the Cape Verde Islands, Senegal, offshore 

islands of the Gulf of Guinea, and Angola, intertidal 

to 35 m, generally in shallower water. 

Monod (1956:230) commented on its apparent 

absence on the mainland between Senegal and 

Pilumnus qfricanus.—Capart, 1951:142, fig. 51, pi. 1: fig. 15, 

pi. 3: figs. 7, 8.—Garth, 1968:314 [discussion]. 

Angola; he summarized earlier records and re- 

Globopilumnus qfricanus.—Monod, 1956:227, figs. 249-257.— ported material from the Cape Verde Islands, 

Guinot-Dumortier, 1959:98, 99, 116, 118, fig. 14 [discus- Senegal, Sao Tome, and Angola. Subsequent recsion; 

review of Indo-West Pacific Globopilumnus].—Guinot- ords in the literature include the following: 

Dumortier and Dumortier, 1960:138, fig. 21 [stridulation].—Guinot 

and Ribeiro, 1962:51.—Ribeiro, 1964:7.— West Africa: No specific locality (Guinot-Dumortier and 

Forest and Guinot, 1966:66.—Garth, 1968:312, 313 [dis- Dumortier, 1960). 

cussion].— Turkay, 1976b:61 [listed], 67, fig. 4, pi. 2: figs. Madeira: Funchal, fish market (from stomach of Phycis 

3,4. 

phycis (Linnaeus)) (Turkay, 1976b).

134 

DESCRIPTION.—Monod, 1956:231. 



Guinea, Annobon). 

Color: "Blanc sale" (Monod, 1956:235). Our 

specimens have a reddish body, with red and 

white banded legs. 



REMARKS.—The larger specimens taken by the 

Pillsbury off Annobon differ from Monod's account 

in having the anterolateral spines of the 

carapace and those on the outer face of the wrist 

and palm of the chela dark; in none of the 

specimens does the dark color of the fixed finger 


Cape Verde Islands: Matiota, Sao Vicente (Guinot and of the chela extend onto the palm. The large male 


Principe: Ilheu Caroco (as ilot Carioco), 2-8 m; 01°38'- 

35"N, O7°21'35"E, 35 m; and S of Ilheu Santana (as tie Sta. 

Ana), 12 m (all Forest and Guinot, 1966). 

Sao Tome: 00°25'N, 06°37'E, 5-6 m; Ponta Diogo Vaz, 

0-6 m; Praia de Santa Catarina, 3-10 m; Baia de Sao 

from Pillsbury Sta 282 (carapace width 17 mm) 

differs from the specimen from Annobon illustrated 

by Monod (1956, fig. 261) in having the 

terminal segment of the abdomen semicircular 

rather than subtriangular. 

Miguel, 6-10 m; Ilheu Macaco (as Hot dos Cocos), 3-8 m; E 

of the beach, Morro Peixe, 2-6 m; Ilheu das Cabras, 0-2 m; 

Santana, shore; Ponta Furada, 3-8 m (all Forest and Guinot, 

1966). 

Annobon: Bay of Santa Cruz, 8-12 m (Forest and Guinot, 

1966). 

BIOLOGY.—In contrast with G. africanus, this 

species prefers sublittoral habitats, as noted by 

Forest and Guinot (1966:67, 68). Of 16 depth 

records in the literature, 6 are from depths ranging 

from 0-6 m to 10 m whereas 10 are from 

Angola: Baia da Caota, Benguela, shore; Baia das Vacas, depths ranging from 15-40 m to 73 m; none was 

shore; Lucira; Praia das Conchas, Mocamedes, shore (all 


from the intertidal. The Calypso took both species 

together at three stations, on sand with calcareous 

algae, rocks, and coral in 35 m and on rocks and 

* Globopilumnus stridulans Monod, 1956 coral in 0-6 m and 30 m. This species, like G. 

Globopilumnus stridulans Monod, 1956:230, figs. 258-269.— africanus, apparently prefers rough bottom with 

Guinot-Dumortier, 1959:98, 99, 116, 118, fig. 14 [discus- rocks and coral or coralline algae. Crosnier (1964: 

sion; review of Indo-West Pacific Globopilumnus].—Guinot- 31) found it off Cameroon in sponges, coral, and 

Dumortier and Dumortier, 1960:138, 139, figs. 20, 21 

[stridulation].—Rossignol, 1962:116.—Crosnier, 1964: 

31.—Forest and Guinot, 1966:67.—Garth, 1968:312, 313, 

314, 315, 316 [discussion]. 

marl on rocky bottom with gorgonians. Part of 

Monod's material was taken from a wash of 

sponges. 

No ovigerous females have been recorded. 


Sta 275, 9-69 m, rubble of coralline algae, 4(3, 12$ (L). Sta 

282, 18-37 m, nodular coralline algae, 16\ 2$ (W). 

Other Material: Principe: 12 mi [19 km] SSW of Tinhosa 

Grande (as Hermano Grande) Island, 01°20'45"N, 07° 17'- 

37"E, 25-40 m, rocks and coral, 7 Jul 1956, Calypso Sta 110, 

DISTRIBUTION.—Off West Africa, from a few 

mainland localities between Senegal and Gabon 

and from the offshore islands of Principe, Sao 

Tome, and Annobon; intertidal to a depth of 

73 m. Monod (1956) described the species on 

13, 1$ (W). 

Annobon: 01°28.5'S, 05°37.5'E, 35-55 m, F. Poinsard, 

2(5, 89, 8juv(W). 

material from Senegal, Guinea (the type-locality), 

and Annobon; the holotype from Guinea was 

taken in 40 m. Subsequent records in the literature 

include the following: 

West Africa: No specific locality (Guinot-Dumortier and 

Dumortier, 1960). 


Principe: 01°36'50"N, O7°22'1O"E, 19 m; S part, Baia 

das Agulhas, 4-8 m; 01°38'35"N, 07°21'35"E, 35 m; 01°- 

43'10"N, 07°28'20"E, 73 m; 01°43'N, O7°28'55"E, 37 m; 

and Tinhosa Grande (as Hermano Grande) Island, 12 mi 

[19 km] SSW of Principe, O1°2O'45"N, O7°17'37"E, 25- 

40 m (all Forest and Guinot, 1966). 

Sao Tome: 00°20'N, 06°46'E, 10 m; Ponta Diogo Vaz, 

W coast, 0-6 m and 30 m; in front of Ponta Oquedelrei, 6 

m; and off Ponta Diogo Nunes, 6 m (all Forest and Guinot, 

1966). 

Annobon: 01°24'04"S, 05°36'45"E, 7-10 m; 01°25'- 

12"S, 05°36'05"E, 20 m; and NW coast, Isla Tortuga, 15- 

40 m (all Forest and Guinot, 1966).


Gabon: W of Cap Santa Clara, Libreville, 45-47 m 

(Rossignol, 1962). 

Genus Glyptoxanthus A. Milne Edwards, 1879 

Glyptoxanthus A. Milne Edwards, 1879, in 1873-1881:253 

[type-species: Actaea erosa Stimpson, 1859, by selection by 

Rathbun, 1930:263; gender: masculine].—Guinot, 1971: 

1072 [list of species]. 

* Glyptoxanthus angolensis 

(De Brito Capello, 1866) 

Actaea angolensis.—Capart, 1951:157, fig. 59.—Longhurst, 

1958:88.—Gauld, 1960:70.—Rossignol, 1962:117. 

Actaea (Glyptoxanthus) angolensis.—Monod, 1956:296, fig. 

361.—Serene, 1961:198 [listed]. 

Glyptoxanthus angolensis.—Forest and Guinot, 1966:77, fig. 6a, 

b.—Guinot, 1971:1073 [listed]. 

Glyptoxanthus anglolensis.—Guinot, 1967d:556 [listed; erroneous 

spelling]. 


Sta 282, 18-37 m, nodular coralline algae, let, 19 (L, W). 

Other Material: Annobon: S coast, 01°28.5'S, O5°37.5'E, 

35-55 m, 16 Jun 1967, F. Poinsard, \6 (W). 



361; Forest and Guinot, 1966, fig. 6a,b. 

Male Pleopod: Forest and Guinot, 1966, fig. 6a,b 

(Annobon). 

Color: "Rouge orange avec des taches plus 

claires" (Capart, 1951:158). 


widths of 12, 17 and 25 mm. 

BIOLOGY.—Glyptoxanthus angolensis is a shallow 

shelf species, occurring sublittorally in depths 

between 4-5 m and 40 m, with one record of 35- 

55 m. Like the West African species of Paractaea, 

habitats on rough bottom are preferred. The 

Pillsbury specimens were collected in nodular calcareous 

algae off Annobon, and the material 

collected by the Calypso (Forest and Guinot, 1966) 

was found on the following types of bottom: 

calcareous algae, 4-5 m; calcareous algae, sand 

and coral, 7-10 m; rocks and calcareous algae, 

20 m; and rocks and coral, 15-40 m. Longhurst 

(1958) found the species on shelly sand in 25 m 

off Sierra Leone. 

Ovigerous females have been recorded in January 

and June (Monod, 1956; Forest and Guinot, 

1966). 

DISTRIBUTION.—West African coast, from scattered 

localities between Senegal and Angola, including 

Annobon and Sao Tome islands in the 

Gulf of Guinea; sublittoral, from 4-5 m to 40 m 

(also 35-55m). Monod (1956) summarized earlier 

records and reported material from Senegal, 

Sierra Leone, Congo (Congo Francais), and Annobon; 

records in the literature since 1956 include 



1958). 

Ghana: Off Accra, sublittorally; Tenkpobo (asTenpobo), 

intertidal (Gauld, 1960). 

Sao Tome: Off Ponta Diogo Nunes, 4-5 m (Forest and 


Annobon: 02°24'04"S, 05°36'45"E, 7-10 m; 01°25'12"S, 

05°36'05"E, 20 m; and Isla Tortuga, NW coast, 15-40 m 

(Forest and Guinot, 1966). 

Congo: Pointe Indienne (Rossignol, 1962). 

Angola: No specific locality (Forest and Guinot, 1966). 

Glyptoxanthus cavernosus 


Xantho vermiculatus.—Osorio, 1906:149 [Cape Verde Islands; 

see Forest and Guinot, 1966:79] [not Cancer vermiculatus 

Lamarck, 1818]. 

Actaea (Glyptoxanthus) cavemosa.—Monod, 1956:298, fig. 362 

[Cape Verde Islands; references].—Serene, 1961:198 

[listed]. 

Glyptoxanthus cavemosa.—Forest and Guinot, 1966:79 [Cape 


Glyptoxanthus cavemosus.—Guinot, 1967d:551, 553 [discussion], 

556 [listed], fig. 31 [Cape Verde Islands]; 1971:1073 

[listed]. 

DISTRIBUTION.—Known only from the Cape 

Verde Islands. 

Glyptoxanthus corrosus 


Actaea (Glyptoxanthus) corrosa.—Monod, 1956:298 [Cape 

Verde Islands; references]. 

Glyptoxanthus corrosus.—Guinot, 1967d:556 [listed], fig. 21 

[Cape Verde Islands]; 1971:1073 [listed]. 

DISTRIBUTION.—Cape Verde Islands.


Genus Heteropanope Stimpson, 1858 

Heteropanope Stimpson, 1858a:33 [type-species: Heteropanope 

glabra Stimpson, 1858, by subsequent selection by Balss, 

1933b:32; gender: feminine; name 1627 on Official List]. 

Heteropanope acanthocarpus Crosnier, 1967 

Heteropanope acanthocarpus Crosnier, 1967:325, figs. 3, 4, 7-10, 

28 [Dahomey]; 1971:570 [Congo]. 

DISTRIBUTION.—West Africa, from Dahomey 

(06°10'N, O2°27'E), 55 m, and off Pointe-Noire, 

Congo, 20 m. 

Heteropanope tuberculidens Monod, 1956 

Heteropanope tuberculidens Monod, 1956:265, figs. 309-313.— 

Gauld, 1960:70.—Crosnier, 1967:325, figs. 1, 2, 6, 27.— 

Uschakov, 1970:444, 455 [listed]. 

Heteropanope tubulicidens.—Longhurst, 1958:88 [erroneous 

spelling]. 

MATERIAL EXAMINED.—PMsbury Material: None. 

Other Material: Togo: 06° 1 l'N, 01 °28'30"E, 14-17 m, sand 

with gorgonians, mud, 17 Oct 1963, A. Crosnier, 16\ 2$ (W). 

Congo: Pointe-Noire, 5-10 m, lobster net, 27 Jan 1967, J. 

Marteau, Id, 2$, 1 juv (W). 


Figures: Monod, 1956, figs. 309-313; Crosnier, 

1967, figs. 1, 2, 6, 27. 


(Ghana); Crosnier, 1967, fig. 27 (Guinea-Bissau). 

Color: "Plus ou moins brun rouge, traces 

d'annelures aux pattes" (Monod, 1956:268). 


widths of 2.6 to 5.2 mm. 

REMARKS.—Our specimens agree well with accounts 

of this species, except that in our smaller 

specimens from the Congo the outer face of the 

chela is relatively smooth, and in one of the 

females from the same locality the tubercles on 

the palm are aligned in rows. 

BIOLOGY.—Heteropanope tuberculidens is a sublittoral 

species living in relatively shallow water. 

Depth records for the species range from 0-2.5 m 

off Guinea (Uschakov, 1970) to 72 m off Sierra 

Leone (Longhurst, 1958); the latter is the only 

record for the species in depths below 25-30 m, 

and the majority of depth records are from less 

than 20 m; records include: 0-2.5, 8-16, 10-12, 

14-17, 15 (2X), 25-30, and 72 m. The species 

apparently prefers rough bottom; it has been 

recorded from bottom with coral; gravel, sand 

and shell; muddy sand with gorgonians; sand, 

rock or mud, and gorgonians; and shelly mud. 


March, April, and June (Monod, 1956; Crosnier, 

1967). 

DISTRIBUTION.—Off West Africa, from a few 

localities between Guinea-Bissau and the Congo, 

in depths to 72 m, generally in less than 20 m. 


Guinea-Bissau: Channel between Ilha de Rubane (as 

Rouban) and Ilha de Bubaque, Bissagos Islands, 25-30 m, 

and channel near Ilha de Soga, Arquipelago dos Bijagos (as 

Bissagos Islands), 8-16 m (Monod, 1956; Crosnier, 1967). 

Guinea: Conakry, 0-2.5 m (Uschakov, 1970). Between lie 

Tamara and lie de Corail, 10-12 m (Monod, 1956). 


1958). 

Ghana: Off Accra, 15 m (Gauld, 1960). 2 mi [3 km] off 

Densu, near Accra, 15 m (type-locality) (Monod, 1956). 

Togo: 06°ll'N, 01°28'30"E, 14-17 m (Crosnier, 1967). 

Congo: Off Pointe-Noire (Crosnier, 1967). 

Genus Leopoldius Serene, 1971 

Leopoldius Serene, 1971:908 [an invalid junior homonym of 

Leopoldius Rondani, 1843 (Insecta); type-species: Parapilumnus 

leopoldi Gordon, 1934, by original designation; gender: 

masculine]. 

REMARKS.—R. Serene has informed us (in litt.) 

that he will propose a replacement name for this 

genus. 

•Leopoldius pisifer (MacLeay, 1838), 


Parapilwnnus pisifer.—Capart, 1951:146, fig. 53, pi. 3: fig. 

18.—Sourie, 1954b: 150.—Monod, 1956:254, figs. 298- 

301.—Longhurst, 1958:88.—Gauld, 1960:70.—Rossignol, 

1962:117.—Guinot and Ribeiro, 1962:52.—Forest and 

Guinot, 1966:71.—Crosnier, 1969:535.—Uschakov, 1970: 

439, 455 [listed].—Takeda, 1974:216 [discussion]. 

Synonym.—Pilumnus verrucosipes Stimpson, 

1858.



Coast: Sta 46, 38-42 m, mud with dense Julltenella, 66, 5$ 

(4 ov) (W). Sta 47, 37 m, bottom with Jullienella, 26, 2$ 

(1 ov) (L). 

Nigeria: Sta 248, 33 m, 16, 1$ (L, W). Sta 253, 33-40 m, 

mud, 1(5 (L). 

Other Material: Senegal: Les Almadies, Dakar, 0-0.5 m, 

under stones, 9 Jun 1964, F. M. Bayer, R. B. Manning, and 

L. B. Holthuis, Ic5, 1$ (L). Goree, lobster net, 15 Sep 1950, 

M. Paraiso, 216, 26$ (12 ov) (W). Same locality, 6 Oct 1950, 

M. Keita, 26, 19 ov (W). 

Congo: Pointe-Noire, rocks with gorgonians, 25 Jan 1967, 

A. Crosnier, 46 (W). 


Figures: Capart, 1951, fig. 53, pi. 3: fig. 18; 

Monod, 1956, figs. 298-301. 

Male Pleopod: Capart, 1951, pi. 3: fig. 18 (Senegal); 

Monod, 1956, figs. 300, 301 (Senegal). 




REMARKS.—As suggested by Takeda (1974: 

216), this species appears to belong to Leopoldius 

Serene, 1971, rather than to Parapilumnus Kossmann, 

1877. In L. pisifer the carapace is covered 

with a thick coat, arranged more or less symmetrically, 

the lobes of the front are sinuous rather 

than truncated, the three posterior anterolateral 

teeth of the carapace are broad, not spiniform, 

and their borders are granulated, the greatest 

width of the carapace is at the level of the posteriormost 

tooth, the pereiopods are short and irregularly 

carinate, and the lateral palatal ridges 

are completely absent. 

This is the only species of Leopoldius to occur in 

the Atlantic; the remainder of the species are 

from the Indo-West Pacific region. 

BIOLOGY.—Like Nanopilumnus boletifer (p. 145), 

Leopoldius pisifer is a shallow shelf species, living 

from the intertidal zone to a depth of 50 m; the 

majority of records in the literature are from 

depths of 30 m or less. It can utilize a wide range 

of substrates. Sourie (1954b) found it on shelly 

sand in the Baie de Dakar and Longhurst (1958) 

reported in from muddy sand in 22 m off Dakar. 

Uschakov (1970) reported it from hard sand bottom 

in depths greater than 20 m in clear water. 

It was reported from rocks ornamented with gorgonians 

in 10 m off Pointe-Noire, and was taken 

by the Calypso on mud and shells in 18 m, on mud 

in 18-30 m and 50 m, on muddy sand with 

Foraminifera in 21-27 m, and on mud with Area 

in 32 m (Forest and Guinot, 1966). The Pillsbury 

specimens were taken on mud or mud with Jullienella. 

Ovigerous females have been recorded from all 

months (Monod, 1956; Forest and Guinot, 1966). 

DISTRIBUTION.—West Africa, from Mauritania 

southward to Gabon, including Ilha do Principe 

in the Gulf of Guinea; and southern Africa, from 

South Africa and Mozambique (Barnard, 1954); 

intertidal zone to a depth of 50 m. Monod (1956) 

summarized earlier records and reported material 

from Mauritania, Senegal, Gambia, Guinea, 

Guinea-Bissau, Sierra Leone, Ghana, Gabon, and 

Principe. In addition the species has been reported 

from the following localities: 

Senegal: Baie de Dakar (Sourie, 1954b). Dakar, 22 m 


Guinea: No specific locality, in depths greater than 20 m 

(Uschakov, 1970). 09°40'N, 14°05'W, 18 m, and 09°36'N, 


Ivory Coast: 05°02.5'N, 05°25'W, 21-27 m (Forest and 


Ghana: Tenkpobo (as Tenpobo), Christiansborg, Apam, 

Winneba and Axim, rarely sublittoral to 37 m (Gauld, 1960). 

04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m (Forest and 




Gabon: W of Libreville, 20-40 m (Rossignol, 1962). 

Congo: Baie de Pointe-Noire, 6-8 m (Rossignol, 1962). 

Pointe-Noire, 10 m (Crosnier, 1969). 

Angola: Baia da Caota, Benguela, 13 m and 30 m (Guinot 


Genus Menippe de Haan, 1833 

Menippe de Haan, 1833:21 [type-species: Cancer rurnphii Fabricius, 

1798, by subsequent designation by Glaessner, 

1929:253; gender: feminine].—Guinot, 1971:1076 [list of 

species]. 

Menippe nodifrons Stimpson, 1859 

Menippe nodifrons.—Frade, 1950:11, 26 [Congo].—Capart, 

1951:138, fig. 49 [Guinea].—Monod, 1956:222, figs. 244-

138 

248 [Senegal, Guinea, Sierra Leone, Ghana, Rio Muni; 

references].—Longhurst, 1957:374 [Sierra Leone].—Rossignol, 

1957:82 [Congo].—Buchanan, 1958:20 [Ghana]. 

—Longhurst, 1958:88 [Sierra Leone].—Gauld and Buchanan, 

1959:127 [Ghana].—Gauld, 1960:70 [Ghana].— 

Rossignol, 1962:116 [Congo].—Guinot and Ribeiro, 1962: 

50 [Angola].—Monod, 1967:180, pi. 17: fig. 3 [no material].—Le 

Loeuff and Intes, 1968, table 2 [Ivory 

Coast].—Guinot, 1968b: 156 [discussion].—Uschakov, 

1970:445, 446, 447, 455 [listed; Guinea].—Guinot, 1971: 


Menippe nanus.—Capart, 1951:140 [discussion]. 

Menippe.— Gauld and Buchanan, 1959:128 [Ghana]. 

SYNONYMS.—Menippe rudis A. Milne Edwards, 

1879; Menippe nanus A. Milne Edwards and Bouvier, 

1898. 

DISTRIBUTION.—Eastern Atlantic from the 

Cape Verde Islands and Senegal to Angola; western 

Atlantic from Florida to Brazil (Rathbun, 

1930); littoral and sublittoral, from shore to about 

20 m. 

Genus Microcassiope Guinot, 1967 

Microcossiope Guinot, 1967c:358 [type-species: Xanthodes rufopunctatus 

A. Milne Edwards, 1869, by original designation; 

gender: feminine]; 1971:1076 [list of species]. 

* Microcassiope minor (Dana, 1852) 

FIGURE 30 

Xmtho minor Dana, 1852b: 169; atlas, 1855, pi. 8: fig. 7.— 

Miers, 1881a:214; 1886:124 [listed]. 

Pilumnus granulimanus Stimpson, 1871a: 143 [Cuba]. 

Muropanopt granulimanus.—Rathbun, 1930:439, pi. 180: figs. 

1, 2 [western Atlantic records]. 

Micropanope granosa.—Chapman and Santler, 1955:374. 

Micropanope rufopunctata.—Monod, 1956:313, figs. 386-392.— 

Gauld, 1960:70.—Guinot and Ribeiro, 1962:59.—Monod, 

1963, fig.34 [no material].—Ribeiro, 1964:10—Forest and 

Guinot, 1966:81.—Chace, 1966:639, fig. 8.—Guinot, 

1967c:348 [discussion], 358 [listed].—Le Loeuff and Intes, 

1968, table 1—Tiirkay, 1976b:61 [listed], 69. 

Microcassiope rufopunctata.—Guinot, 1967c, figs. 10, 15; 1971: 


Xanthodes rufopunctata.—Guinot, 1967c:359 [listed]. 

Xanthodes rufopunctatus.—Garth, 1968:314 [discussion]. 

Microcassiope granulimanus.— Guinot, 1971:1076 [listed; considered 

to be distinct from M. rufopunctata]. 


SYNONYMS.—Xanthodes rufopunctatus A. Milne 

Edwards, 1869; Xanthodes granosus A. Milne Edwards 



Sta 282,18-37 m, nodular coralline algae, 3d, 59, 2 juv (L, 

W). 

Other Material: Madeira: Ponta de Sao Lourenco, 32°44'N, 

16°44'W, littoral, rocky shore with tide pools, 29 Feb 1976, 

Onversaagd Sta 16, 19 (L). SE coast, near Canical, 32°44'N, 

16°44'W, 0-22 m, shore collecting, snorkeling, diving, 11 

Mar 1976, Onversaagd Sta 48, Ic5 (L). 

Cape Verde Islands: Porto da Praia (as La Praya), Sao 

Tiago, 10-30 m,Jun-Jul 1883, Talisman, syntype of Xanthodes 

granosus A. Milne Edwards and Bouvier, 1898, Id, 19 (L, W). 

Annobon: S coast, 01°28.5'S, 05°37.5'E, 35-55 m, 16 Jun 

1967, F. Poinsard, 19 (W). 


1900:87. 



(Azores, Cape Verde Islands); Chace, 1966, fig. 8 

(St. Helena, Curacao, Cape Verde Islands); 

Guinot, 1967c, fig. 15 (Cape Verde Islands). 


widths of 2.5 to 9.1 m. 

REMARKS.—In his account of the Crustacea of 

the U.S. Exploring Expedition, Dana (1852b: 169) 

described a small xanthid crab that had been 

collected at Madeira or the Cape Verde Islands, 

Xantho minor. His account is based on an ovigerous 

female 2.1 lines long, 3.1 lines wide (approximately 

4.5 X 6.6 mm): 

Near X. parvulus [= Eurypanopeus blanchardi\. Carapax anteriorly 

areolate, areolets slightly raised, 2M, 3M, 5L, 6L 

circumscribed behind; antero-lateral margin thin, fourtoothed, 

three posterior teeth subtriangular. Anterior feet of 

moderate size, subequal, carpus and hand above slightly 

granulous, hand exteriorly faint granulato-costate, and 

above sulcate, moveable finger not armed with a large basal 

tooth. Eight posterior feet sparsely pubescent. 

Two small species of xanthids, both formerly 

placed in Micropanope, are now known to live in 

the Cape Verde Islands and one or more of the 

more northern islands or island groups, Madeira, 

the Azores, and the Canary Islands: Microcassiope 

rufopunctata (A. Milne Edwards, 1869) and Nanocassiope 

melanodactyla A. Milne Edwards, 1867). 

Earlier authors have identified Xantho minor with


FIGURE 30.—Microcassiope minor (Dana) (from Dana, 1852b, 

pi. 8: fig.7). 

one or the other of these species without adopting 

the specific epithet minor. A third small species of 

xanthid, also at one time placed in Micropanope, 

CoralHope parvula (A. Milne Edwards, 1869), also 

occurs in the Cape Verde Islands; that species, as 

noted above, has but three anterolateral teeth 

whereas X. minor was described as having four, 

and has an additional inner spine on the carpus 

of the chela; the spine is absent in X. minor. 

Miers (1886:124) listed Xantho minor with other 

species now assigned to Xantho sensu stricto, and 

retained both melanodactyla and rufopunctata in Xanthodes, 

the genus to which they were assigned in 

Milne Edwards' original accounts. 

A. Milne Edwards and Bouvier (1898:190), in 

their description of Xanthodes talismani [= Coralliope 

parvula], pointed out similarities between that species 

and Xantho minor: 

Cette espece se rapproche beaucoup du Xantho minor 

Dana, de Madere et des lies du Cap Vert; mais cette derniere 

appartient a un genre different; sa dent anterieure est encore 

nette, sa carapace est depourvue de touffes de poils, ses 

pinces sont un peu costulees et il n'y a pas d'epines sur le 

bord interne du carpe. 

Later A. Milne Edwards and Bouvier (1900: 

93) repeated this observation in their account of 

Xanthodes talismani, in spite of the fact that on p. 

85 of the same work they identified Xantho minor 

Dana, 1852 with Xanthodes melanodactylus A. Milne 

Edwards, 1867 (and used the junior synonym!) 

and on page 87 of the same work reported Xan- 

thodes melanodactylus var. rufopunctata and noted: "il 

y a probablement lieu d'y rattacher [that mate- 

rial] plus specialement le Xantho minor de Dana." 

They also synonymized Dana's Xantho parvulus, 

which we believe should be identified with Eurypanopeus 

blanchardi, with Xanthodes melanodactylus. 

Monod (1956:320, in synonymy) identified 

Xanthodes melanodactylus var. rufopunctatus of A. 

Milne Edwards and Bouvier, 1900, with Micropanope 

melanodactyla. Three females from one of 

the lots (in 75 m) reported by A. Milne Edwards 

and Bouvier and labeled Xanthodes melanodactylus 

var. rufopunctata are in the collection of the Smithsonian 

Institution; they are Nanocassiope melanodactyla. 

Monod (1933b:519) pointed out the multiple 

use of the epithet minor by A. Milne Edwards and 

Bouvier and further noted: "En fait la figure de 

Xantho minor donnee par DANA semblerait plutot 

representer Xanthias granosus [ = Microcassiope rufopunctata] 

ou X. tuberculatus [— Monodaeus couchii] 

que X. melanodactylus. L'examen du type est indispensable." 

We believe that Monod (1956:314, synonymy) 

was correct in identifying Xantho minor Dana with 

Xanthodes rufopunctatus A. Milne Edwards. Dana's 

account mentioned the small size, the wellmarked 

regions, the triangular anterolateral teeth 

of the carapace, the granular chela lacking a large 

basal tooth, and, in addition, clearly showed in 

his plate 8: figure 7 (see Figure 30) several transverse 

lines of granules on the carapace. All of 

these features are characteristic of the species now 

known as Microcassiope rufopunctata. We believe 

that the older name, Microcassiope minor (Dana, 

1852), must be used for the species. 

Chace (1966:639), in recording this species 

from Saint Helena in the south central Atlantic, 

showed that Pilumnus granulimanus Stimpson, 1871, 

from the western Atlantic, placed in Micropanope 

by Rathbun (1930), is conspecific with M. minor 

(as M. rufopunctata); the latter is now known to 

have a relatively wide range on both sides of the 

Atlantic. Guinot (1971:1076), in her summary of 

nomenclatural changes resulting from her studies 

of the xanthids, removed M. granulimanus from the 

synonymy of M. minor without stating her reasons. 

She noted: "Cette espece a ete mise en synonymie

140 


avec Micropanope rufopunctata par CHACE (1966, p. insohn and Holthuis, 1964:59, for records), the 

639, fig. 8), mais pour l'instant nous la mainten- Azores, the Canary Islands, Madeira, the Cape 

ons separee de l'espece ouest-africaine." 

Verde Islands, the African mainland from Span- 

In discussing the distribution patterns of some ish Sahara, Ivory Coast, and Ghana, and the 

Atlantic-East Pacific xanthids, Garth (1968:314) offshore islands of the Gulf of Guinea, Principe, 

was in error in stating that Microcassiope did not Sao Tome, and Annobon; central Atlantic from 

occur in the western Atlantic. It is represented Saint Helena (Chace, 1966); western Atlantic 

there by M. minor (as Micropanope granulimanus). from the Bahamas, Cuba, Curacao, and Islas Los 

Garth apparently was misled by the omission of Roques (Chace, 1966); intertidal to a depth of ca 

this species in Guinot's (1967c:358) original ac- 220 m, commoner in shallower water. Monod 

count of Microcassiope. 

(1956) summarized earlier West African records 

A male syntype of Xanthodes granosus (see "Syn- and reported material from the Azores, the Caonyms") 

is in the collection of the National Munary Islands, the Cape Verde Islands, and Ghana. 

seum of Natural History, Smithsonian Institu- Other West African records in the literature intion, 

under catalog number USNM 22956. A clude the following: 

female syntype is in the collection of the Rijksmuseum 

van Natuurlijke Historie, Leiden, under 

Crust. D.1564. 


Azores: Pasteleiro, Feteira, and Horta, Ilha do Faial, 

intertidal (Chapman and Santler, 1955). 

BIOLOGY.—Off West Africa, Microcassiope minor Madeira: Porto do Funchal, ca 220 m, on telegraph cable; 

lives from the intertidal zone to a depth of ca 

Ilheu do Gorgulho, littoral; between Ponta da Garajau and 

Ponta da Oliveira, 5-6 m (all Tiirkay, 1976b). 

220 m. Of the recent records for which depth is 

Cape Verde Islands: No specific locality (Guinot, 1967c). 

available, 19 out of 26 (or 73%) are from depths Baixo Joao Leitao, 25 m (Guinot and Ribeiro, 1962; Ribeiro, 

of 10 m or less, and seven (or 27%) are from 1964). Porto da Praia (as La Praya), Sao Tiago (Chace, 

depths ranging from between 15 and 45 m to ca 1966). 

220 m (15-40 m, 43-45 m, and ca 220 m). Eight 

of the 19 shallower collections are from shore. 

Forest and Guinot (1966:84) pointed out that 

Nanocassiope melanodactyla (as Micropanope melanodactyla) 

and Microcassiope minor both are common 

in the Gulf of Guinea, where they may occur 

Spanish Sahara: 21°05'N, 17°14'W, 43-45 m (Forest and 


Ivory Coast: Lagoon of Abidjan, 05°16'N, 04 

together, but that the former lives in deeper water 

generally on less stable substrates whereas "la 

seconde, plus littorale, a souvent ete capturee 

dans la zone intercotidale ou jusqu'a une dizaine 

de metres, principalement sur des fonds durs, 

roches et coraux," a statement borne out by the 

capture of these two species off the offshore islands 

of the Gulf of Guinea. The deepest record is that 

of Tiirkay (1976b) from Madeira, in ca 220 m on 

a telegraph cable; he also recorded specimens 

from the littoral zone, under stones. 


collected in June (Forest and Guinot, 1966). 

Chace (1966) recorded ovigerous females from 

Saint Helena in January and April. 

DISTRIBUTION.—Atlantic Ocean. Eastern Atlantic 

from the eastern Mediterranean (see Lew- 

o 01'20 w W, 

(Forest and Guinot, 1966). Off Jacqueville, 40 m (Le Loeuff 


Ghana: Off Tenkpobo (as Tenpobo), littoral; off Accra, 

37 m (Gauld, 1960). 

Principle: Ilheu Caroco, 2-8 m; Ponta da Mina, beach 



W coast, 0-6 m; Praia de Santa Catarina, W coast, 3-10 m; 

off Sao Tome, 8 m; off Ponta Diogo Nunes, shore and 

4-5 m; in front of Ponta Oquedelrei, 6 m; Morro Peixe, 2- 

6 m; Ilheu das Cabras, shore; in front of the harbormaster's 

office, shore; in front of Praia Lagarto, 5-6 m (all Forest and 


Annobon: No specific locality (Guinot, 1967c). 01°24' 

04"S, 05°36'45"E, 7-10 m; O1°25'12"S, 05°36'05"E, 20 m; 

Isla Tortuga, NW coast, 15-40 m (all Forest and Guinot, 

1966). 

Saint Helena: Rupert's Bay, 0-75 m; James Bay (Chace, 

1966). 

Genus Monodaeus Guinot, 1967 

Monodaeus Guinot, 1967c:369 [type-species: Xantho couchii 

Couch, 1851, by original designation; gender: masculine]; 

1971:1074 [ list of species].


Monodaeus couchii (Couch, 1851) 

Xantho couchii Couch, 1851:13. 

Medaeus couchi.—Monod, 1956:310, figs. 383, 877, 878.— 

Peres, 1964:20.—Forest and Guinot, 1966:80.—Crosnier, 

1967:331, figs. 13, 14.—Zariquiey Alvarez, 1968:400, figs. 

9, 15e, 133, 134a [Spain; references].—Crosnier, 1970:1215 

[listed], 1216. 

Micropanope (?) couchi.—Forest and Gantes, 1960:352. 

Xantho couchi.—Peres, 1964:27, 28, 29.—Guinot, 1967c:348, 

349, 371 [all discussion], 373 [listed].—Maurin, 1968a: 19, 

116 [Spain, Mediterranean].—Forest, 1976:66 [discussion]. 

Monodaeus couchi.—Guinot, 1967c:372 [discussion], figs. 23, 

32 [Spain]; 1971:1074 [listed].—Turkay, 1976a:25 [listed], 

38, fig. 24 [Portugal, in part].—Forest, 1976:68 [discus- 

SYNONYM.—Xantho tuberculatus Bell, 1852. 


Undaunted Material: South-West Africa: Sta 106, 225 m, 1$ 

(L). 


08°45'W, Agassiz trawl, 28 Mar 1976, Onversaagd Sta 154, 1

142 


Sta 68, 70 m, broken shell, 16* (W). 

Ivory Coast: Sta 45, 73-97 m, 16* (L). Sta 49, 73-77 m, 

16*, fragments (L). Sta 50, 128-192 m, 1(5, 2$ ov (W). Sta 

62, 46 m, brown, branched and foliate Foraminifera, 16*, 1$ 

(L). 

Nigeria: Sta 237, 101 m, coral ground, rough, 16* (L). Sta 

239, 73 m, 19 (L). 

Other Material: Ivory Coast: Off Sassandra, 11 m, 3 Apr 

1964, Guinean Trawling Survey, Tr 22, Sta 1, 16* (L). 

04°33'N, 06°36'W, 100-109 m, sand, mud, shells, rocks, 21 

May 1956, Calypso Sta 16, 26*, 39 (1 ov) (W). 

Congo: Off Pointe-Noire, 04°56'S, 11°31'E, 95-97 m, 

trawl, 21-22 Sep 1965, Id (W). 



Monod, 1956, figs. 384, 385; Guinot and Ribeiro, 

1962, fig. 26; Crosnier, 1967, figs. 25, 26. 

Male Pleopod: Capart, 1951, pi. 3: fig. 17 

(Congo); Monod, 1956, fig. 385 (Congo); Guinot 

and Ribeiro, 1962, fig. 26 (Ivory Coast). 



of ovigerous females is 8 mm. Crosnier (1967:336) 

pointed out that this is a relatively small species; 

he examined one ovigerous female 6.6 mm wide, 

and the ovigerous female reported by Capart 

(1951) was only 6 mm wide. 

BIOLOGY.—Monodaeus rouxi is an offshore species 


Ivory Coast: SW of Grand-Bassam, 100-400 m (Crosnier, which lives on the continental shelf and upper 

1967). 

slope. It usually occurs in depths between 46 m 

Congo: 51.5 mi [83 km] WNW of Banana [Zaire], 

05°50'S, 11°32'E, 215-220 m (Capart, 1951). Off Pointe- 

Noire, 80-100 m (type-locality) (Crosnier, 1967); in 255 m 

(Crosnier, 1969). 04°53'S, 11°38'E, 75 m; 05°00'S, 11°26'E, 

115 m (Crosnier, 1969). 

and 215-220 m; the extremes of depth recorded 

so far, 11 m herein and ?500 m (Guinot and 

Ribeiro, 1962), require verification. Nine of the 

14 available depth records are for depths below 

100 m, and seven of those are for depths between 

64 and 97 m. The five deeper records are 100- 

* Monodaeus rouxi (Capart, 1951) 

109 m, 101 m, 128-192 m, 215-220 m, and 

Micropcmope rouxi Capart, 1951:153, fig. 57, pi. 3: fig. 17.— ?500 m. The species has been collected on brown 

Forest, 1965a:380.—Forest and Guinot, 1966:81.— mud in 215-220 m and on muddy sand and rocks 

Guinot, 1967c:348, 349 [discussion], 371 [listed].—Forest, in 80 m (Capart, 1951); mud, sand and com- 


pacted sand (sable construit) in 65-75 m, sand, 

Medaeus (?) rouxi.—Monod, 1956:312, figs. 384, 385.—Guinot 

mud, and shells in 64 m, and on sand, mud, 

and Ribeiro, 1962:58, fig. 26.—Crosnier, 1967: 335, figs. 

25, 26. [Not Medaeus rouxi Balss, 1935.] 

shells, and rocks in 100-109 m by the Calypso 

Medaeus rouxi.—Forest, 1959:15 [not Medaeus rouxi Balss, (Forest and Guinot, 1966). In the latter depth a 

1935]. 

large series of 56 specimens was collected. The 

Monodaeus rouxi.— Guinot, 1967c:371, 372, 373 [all discus- specimens taken by the Pillsbury were taken on 

sion], fig. 24; 1971:1074 [listed].—Forest, 1976:68 [discusion]. 

brown, branching, and foliate Foraminifera in 46 

m, on broken shell in 70 m, and on coral on rough 

bottom in 101 m. 

Ovigerous females have been collected in May 

and December (Capart, 1951; Forest and Guinot, 


DISTRIBUTION.—West Africa, from scattered localities 

between Senegal and Angola, in depths 

between 46 m and 215-220 m (11 m to ?500 m). 

No material was available to Monod (1956). Records 

in the literature include the following: 

West Africa: No specific locality (Forest, 1976). 

Senegal: No specific locality (Forest, 1965a); in 65-75 m 

(Forest, 1959). 12°55.5'N, 17°33'W, 65-75 m (Forest and 


Liberia: 04°34.5'N, 08°31'W, 64 m (Forest and Guinot, 

1966). 

Ivory Coast: No specific locality (Guinot and Ribeiro, 

1962; Forest, 1965; Crosnier, 1967). 04°33'N,06°36'W, 100- 


Congo: 51.5 mi [83 km] WNW of Banana [Zaire], 

05°50'S, 11°32'E, 215-220 m (Capart, 1951). Off Pointe- 

Noire (Crosnier, 1967). 

Angola: 11 mi [18 km] W of Cap Ledo, 09°40'S, 13°02'E, 

80 m (Capart, 1951). Benguela, ?500 m (Guinot and Ribeiro, 

1962). 

Genus Nanocassiope Guinot, 1967 

Nanocossiope Guinot, 1967c:355 [type-species: Xanthodes melonodactylus 

A. Milne Edwards, 1867, by original designation;

NUMBER 306 

gender: feminine]; 1971:1075 [list of species].—Takeda, 

1976:85 [definition]. 

* Nanocassiope melanodactyla 


Micropanope polita Rathbun, 1893b:238; 1930:440, fig. 70, pi. 

180: figs. 3, 4.—Garth, 1946:459, pi. 77: fig. 4 [eastern 

Pacific; considered distinct by Guinot, 1971]. 

Panopeus tanneri Faxon, 1893:154 [eastern Pacific]. 

143 

Cape Verde Islands: Sao Vicente, 75 m, 29 Jul 1883, 

Talisman, 39 (W) [labelled Xanthodes melanodactylus var rufopunctatus]. 

Annobon: 01°24'S, 05°37.5'E, 11 Dec 1965, A. Crosnier, 

26, 39 (W). O1°27'S, 05°35'50"E, 50-60 m, 11 Dec 1965, A. 

Crosnier, 26*, 39 (W). O1°27'S, 05°35'48 w E, 50-60 m, 11 Dec 

1965, Ombango, A. Crosnier, 16*, 89 (2 ov) (W). 



Xanthodes melanodactylus var. rufopunctatus.—A. Milne Edwards Monod, 1956, figs. 401-405. 

and Bouvier, 1900:87, pi. 16: figs. 4, 5 [part?] [not Xan- Male Pleopod: Capart, 1951, pi. 3: fig. 16 (Anthodes 

rufopunctatus A. Milne Edwards, 1869 = Xantho minor 

gola); Monod, 1956, figs. 403-405 (Senegal); 

Dana, 1852]. 

Micropanope melanodactylus.—Capart, 1951:151, fig. 56, pi. Chace, 3; 1966, fig. 7 (Cape Verde Islands, Baja 

figs. 15, 16.—Chace, 1966:637, fig. 7 [Saint Helena]. California, Cocos Island); Guinot, 1967c, fig. 13 

Micropanope melanodactyla.—Monod, 1956:320, figs. 401- (Senegal). 

405.—Gauld, 1960:70.—Guinot and Ribeiro, 1962:60.— Color: "Coloration dans l'alcool, gris tachete de 

Ribeiro, 1964:12.—Forest and Guinot, 1966:83.—Guinot, 

rose; les doigts des chelipedes parfois bruns, par- 

1967c:348 [discussion], 355 [listed].—Le Loeuff and Intes, 

1968, table 1. 

fois noirs" (Capart, 1951:152). 

Xanthodes melanodactylus.—Guinot, 1967c:358 [listed]. MEASUREMENTS.—Our specimens have cara- 

Nanocassiope melanodactyla.—Guinot, 1967c, figs. 8, 13; 1971: pace widths of 2.5 to 13 mm; the carapace widths 

1075 [listed].—Tiirkay, 1976b:61 [listed], 68. 

of ovigerous females ranges from 4 to 9 mm. 

Nanocassiope polita.—Guinot, 1971:1075 [listed]. 

REMARKS.—Chace (1966:637-638) reported 


Sta 70, 33 m, branched Foraminifera, 36*, 1$ ov (L). 

Micropanope melanodactylus from Saint Helena in 

the south central Atlantic and noted: "I agree 

Ivory Coast: Sta 42, 62-75 m, mud with brown, branched with Monod (1956, p. 324) that M. melanodactylus 

Foraminifera, 16* (W). Sta 46, 38-42 m, mud with dense 

Jullienella, 16*, 29 (1 ov) (L). Sta 47, 37 m, bottom with 

Jullienella, 66, 9$ (3 ov) (L). 

Ghana: Sta 22, 51 m, rough bottom, 16*, 1? (L). Sta 23, 

42 m, foliate brown to orange bryozoans, 136*, 79 (1 ov) (L). 

Sta 24, 35-37 m, dark red bryozoans, 26*, 179 (6 ov), 1 juv 

is probably not a synonym of Xantho minor Dana, 

1852b. Unfortunately, the type-specimen of 

Dana's species is probably no longer extant, and 

his name is therefore likely to remain a nomen 

dubium indefinitely." We agree that Xantho minor 

(L,W). 

cannot be identified with Xanthodes melanodactylus., 

Nigeria: Sta 250, 24 m, brackish water, mud, 19 (W). 

Annobon: Sta 271, shore, sand beach, 16* (W). Sta 275, 

9-69 m, rubble of coralline algae, 86, 89 (1 ov), 2 juv (L). 

Sta 282, 18-37 m, nodular coralline algae, 26, 19 (L, W). 

Sta 283, 51-55 m, nodular coralline algae, 426\ 369 (7 ov) 

we have identified it with Microcassiope rufopunctata, 

above. 

We believe that Chace (1966:637) correctly 

synonymized Micropanope polita Rathbun, 1893, 

(W). Sta 284, 73 m, black basaltic rocks, 29 ov (L). from the eastern Pacific, with M. melanodactyla. 

Other Material: Madeira: Porto da Abra, SE coast, Guinot (1971:1075) did not accept this action, 

32°45'N, 16°41'W, to 12 m, diving, 13 Mar 1976, Onversaagd apparently preferring, without stating her rea- 

Sta 68, 16 (L). Near Canical, SE coast, 32°44' N, 16°44' W, 

sons, to keep the two taxa separate. As Garth 

0-22 m, shore collecting, snorkeling, diving, 10 Mar 1976, 

Onversaagd Sta 39, 1 juv

144 

this species. The front is transversely grooved in 

all specimens examined by us. 

BIOLOGY.—Nanocassiope melanodactyla is a sublittoral, 

usually shallow shelf species that apparently 

prefers a bottom with coralline algae or Foraminifera 

on mud or muddy sand. Of 70 recent depth 

records for the species off West Africa, 56 or 80% 

are from depths of 50 m or less; two of those are 

from shore, the remainder in depths from 3-11 m 

to 40-54 m or 9-69 m. Of the 14 records from 

deeper water, two, 200 m and 225 m, are from 

depths greater than 100 m and the remainder 

range from 60 to 85 m. The deepest records in 

the literature are in A. Milne Edwards and Bouvier 

(1900), who provided 11 depth records. Only 

three of these are from depths of less than 50 m, 

three are from depths generally more than 50 m 

but less than 100 m (75, 80, and 80-115 m), and 

the remainder are 100-150 m, 110-180 m, 225 m, 

355 m, and 627 m. The reliability of these deeper 

records is questionable, but the species may stray 

into water that deep. Subsequently, Tiirkay 

(1976b) recorded it from 80 m on a telephone 

cable in the harbor of Funchal; in other samples 

he recorded it from blocks of stone. It generally 

occurs in depths between 20 and 50 m, as pointed 

out by Forest and Guinot (1966:84). They compared 

the apparent habitat preferences of N. 

melanodactyla and Microcassiope minor and noted: 

"La premiere a ete recueillie a des profondeurs 

comprises entre 9 et 65-75 m, mais surtout entre 

20 et 50 m, sur des fonds meubles, le plus souvent 

parmis les algues calcaires." 

Apparently this species spawns all year off 

West Africa. Ovigerous females have been collected 

in all months but April, August, and December 

(Capart, 1951; Monod, 1956; Guinot and 

Ribeiro, 1962; Ribeiro, 1964; Forest and Guinot, 


DISTRIBUTION.—Eastern Pacific, from Baja California, 

Mexico, Cocos Island, and the Galapagos 

Islands; central Atlantic, from Saint Helena and 

Ascension Island; and eastern Atlantic, from the 

Azores, Madeira, Ilhas Desertas, the Canary Islands, 

the Cape Verde Islands and Senegal south- 


ward to Angola, including the offshore islands of 

the Gulf of Guinea, Principe, Sao Tome, and 

Annobon; sublittoral, subtidal to a depth of more 

than 600 m. Monod (1956) summarized the literature 

and reported material from the Azores, 

Madeira, the Cape Verde Islands, the mainland 

from localities between Senegal and Gabon, and 

from the offshore islands of Principe and Annobon; 

other records, including those published 

since 1956, include the following. 

Madeira: No specific locality (Dana, 1852b; Miers, 

1881a). Funchal harbor, ca. 80 m; near Ponta da Garajau; 

between Ponta da Garajau and Ponta da Oliveira; and 

Ponta de Sao Lourenô (all Tiirkay, 1976b). 

Cape Verde Islands: No specific locality (Dana, 1852b). 

Porto Grande, Sao Vicente (Chace, 1966). Baia de Fateixa, 

Sao Vicente, shore; Baia de Porto Grande, Sao Vicente, 

4-6 m, 8 m, 8-10 m, 3-11 m (3.5-11 m); Baia do Tarrafal, 

Sao Tiago, 14-23 m and 9-17 m; Porto de Sao Francisco, 

Sao Tiago, 9 m; Porto da Furna, Brava, 6-20 m (all Guinot 

and Ribeiro, 1962; Ribeiro, 1964). Baia de Porto Novo, 

Santo Antao, 12 m; Porto Ingles, Maio (Ribeiro, 1964). 

Senegal: No specific locality (Guinot, 1967c). 12°55.5'N, 




Ivory Coast: Off Sassandra, 100 m (Le Loeuff and Intes, 

1968). 

Ghana: Off Accra, 10 m (Gauld, 1960). 04°40'N, 

02°08'W, 48 m; 04°40'N, 02°08'W to 04°39'N, 02°05'W) 

50 m; and 04°36.5'N, 01°31'W, 50 m (Forest and Guinot, 

1966). 

Principe: No specific locality (Guinot, 1967c). 01°35'N, 

07°28'E, 45 m; 01°38'25"N, 07°22'05"E, 31 m; 01°38'35"N, 

07°21'35"E, 35 m; 01°43'10"N, 07°28'20"E, 73 m; 01°43'N, 

07°28'55"E, 37 m; off Tinhosa Grande (as Hermano 

Grande) Island, 12 mi [19 km] SSW of Principe, 01°20'45''N, 

O7°17'37"E, 25-40 m; in front of [Cais de] Santana, 11 m 

(all Forest and Guinot, 1966). 

Sao Tome: 00°20'N, 06°47'E, 40-54 m; 00°20'N, 

06°46'E, 10 m; 00°25'40"N, 06°40'10"E, 50 m; 00°25'15"N, 

06°43'05"E, 8-30 m; in front of Ponta de Sao Sebastiao, 

11 m (all Forest and Guinot, 1966). 

Annobon: 01°27.5'S, 05°36.5'E, 35 m; 01°26'15"S, 

05°35'40"E, 60 m; 01°25'10"S, 05°36'10"E, 20-25 m; N of 

San Antonio, 9 and 23 m (all Forest and Guinot, 1966). 

Angola: Baia Farta, Benguela, 22-28 m (Guinot and 


Saint Helena: Off Rupert's Bay, 0-75 m, 0-2 m (Chace, 

1966).


Genus Nanopilumnus Takeda, 1974 

Nanopilumnus Takeda, 1974:215 [type-species: Medaeus rouxi 

Balss, 1935; gender: masculine]. 

* Nanopilumnus boletifer (Monod, 1956) 

Pampilumnus boletifer Monod, 1956:260, fig. 302; 1963, fig. 36 

[no material].—Forest and Guinot, 1966:72, fig. 5a,b— 

Takeda, 1974:216 [listed; transferred to Nanopilumnus]. 

MATERIAL EXAMINED.—Pillsbury Material: Annobon, 

Sta 275, 9-69 m, rubble of coralline algae, 1$ ov (L). 


Figure: Monod, 1956, fig. 302. 

Male Pleopod: Forest and Guinot, 1966, fig. 5a,b 

(Sao Tome). 

Color: Cream (Monod, 1956:261). 

MEASUREMENTS.—Our ovigerous female has a 

carapace width of 5 mm. 

BIOLOGY.—Nanopilumnus boletifer is a sublittoral 

species, with a recorded depth range extending 

from shore to 9-69 m. All depth records other 

than that of the Pillsbury are from depths of less 

than 12 m. The Pillsbury specimen was collected 

in the rubble of coralline algae. The following 

bottom types were noted for the Calypso collections 

(Forest and Guinot, 1966): rocks and coral; 

rocks and sand; mud and calcareous algae; and 

calcareous algae. 


June, and July (Forest and Guinot, 1966; 

Pillsbury). 

DISTRIBUTION.—West Africa, from the offshore 

islands of the Gulf of Guinea, Annobon (the typelocality) 

, Principe, and Sao Tome, from shore to 

a depth of 9-69 m, usually in 12 m or less. 


Annobon: No specific locality, in 12 m (Monod, 1956). 

Principe: Ilheu Caroc.o, 2-8 m; Ilheus dos Mosteiros, 3- 


Sao Tome: Off Sao Tome, 8 m; Baia de Ana de Chaves, 

5 m; W of Ponta Diogo Nunes, shore; off Ponta Diogo 

Nunes, 4-5 m; in front of Ponta Oquedelrei, 6 m; Ilheu das 

Cabras, 0-2 m; and in front of Praia Lagarto, 5-6 m (all 

Forest and Guinot, 1966). 

Genus Panopeus H. Milne Edwards, 1834 

Panopeus H. Milne Edwards, 1834:403. 

REMARKS.—When H. Milne Edwards (1834: 

403) erected the genus Panopeus, he assigned two 

species to it, viz., Panopeus herbstii (a new species) 

and Panopeus limosus (a new combination based on 

Cancer limosa Say, 1817, a species now known as 

Eurytium limosum (Say)). Two other species were 

doubtfully assigned by H. Milne Edwards to the 

new genus: Cancer trispinosus Herbst, 1803, and 

Cancer ochtodes Herbst, 1783. In the synonymy of 

his Panopeus herbstii H. Milne Edwards cited Cancer 

panope Herbst (1801:40, pi. 54: fig. 5) and Say's 

(1817:58, pi. 4: fig. 3) use of Herbst's name Cancer 

panope for an East American species. H. Milne 

Edwards indicated that his material of Panopeus 

herbstii came from "les cotes de l'Amerique septentrionale." 

H. Milne Edwards did not indicate a typespecies 

for his genus Panopeus, and the first valid 

type selection that we know of is the one by E. 

Desmarest (1852:17), who stated: "Panopeus: 

genre americain, ayant pour type le Cancer panope, 

Herbst, que M. Milne Edwards nomme Panope 

d'Herbst." The type-species of Panopeus thus is 

Cancer panope Herbst, 1801, as H. Milne Edwards, 

1834, cited that species by name in the synonymy 

of Panopeus herbstii. That this type selection was 

most unfortunate is shown below. 

As was first pointed out by S. I. Smith (1869a: 

278), Cancer panope Herbst, 1801, is an Indo-West 

Pacific species, different from the East American 

form which was figured by Say (1817) and described 

by H. Milne Edwards (1834). Von Martens 

(1872:87) examined Herbst's type material 

and came to the conclusion that Herbst's Cancer 

panope belongs to the genus Menippe de Haan, 

1833; he also showed that the type locality of 

Cancer panope is Tranquebar, India, where the 

material was collected by Ingobert Karl Daldorff, 

an officer of the Danish garrison at Tranquebar 

(1790-1793) and a pupil of J. C. Fabricius for 

whom he collected (cf., Zimsen, 1964:12). Balss 

(1932:513) removed Herbst's species from Menippe

146 

and placed it in the genus Sphaerozius; he pointed 

out that Sphaerozius panope is a rare species; apart 

from the type only three specimens were known 

to him. 

Unpublished notes in the Division of Crustacea, 


Institution, Washington, D.C., made by 

the late Mary J. Rathbun, show that Cancer panope 

Herbst, 1801, is a junior subjective synonym of 

Cancer scaber Fabricius (1798:336). Miss Rathbun 

came to this conclusion after having examined 

the type-specimens of both species. Like Herbst's 

material of C. panope, the type-specimens of Cancer 

scaber were collected in India by Daldorff, and it 

thus is possible that the types of both species 

originally formed part of a single lot. The correct 

name of the species thus should be Sphaerozius 

scaber (Fabricus, 1798). 

The acceptance of Desmarest's (1852) selection 

of Cancer panope Herbst as the type-species of 

Panopeus H. Milne Edwards, would cause an enormous 

confusion. Not only would the generic name 

Sphaerozius Stimpson, 1858 (type-species, designated 

in Opinion 85, Smithsonian Miscellaneous Collections, 

73(3): 17, 1925, Sphaerozius nitidus Stimpson, 

1858) have to be replaced by the generic 

name Panopeus H. Milne Edwards, the well known 

and widely distributed genus of mud crabs now 

known as Panopeus would have to be given a 

different generic name. This genus is known from 

the eastern Atlantic (south coast of Portugal to 

Angola), both coasts of America (Massachusetts 

to Brazil and Lower California to Chile), and 

Hawaii; it consists of about 12 species, most of 

which live in the littoral zone and several are very 

common. The name Panopeus has been very consistently 

used for it. The only other name available 

for it is Eupanopeus Rathbun (1898:273; typespecies, 

by original designation, Panopeus herbstii 

H. Milne Edwards, 1834), a name which has not 

been used since 1908. 

In order to prevent the considerable confusion 

that a strict application of the Code would cause, 

Holthuis (1979b) has applied to the International 

Commission on Zoological Nomenclature to use 

their plenary power to set aside all type-selections 


for the genus Panopeus. Having done so, he also 

asked the Commission to select as the type-species 

of that genus the species Panopeus herbstii H. Milne 

Edwards, 1834. In the meantime, as prescribed 

by the Code, the name Panopeus will be used by 

us in the sense in which it is currently employed 

by zoologists. 

As pointed out above, Panopeus herbstii is a 

composite species, its syntypes consisting of (1) 

the North American material that H. Milne Edwards 

had before him when he described the 

species, (2) the North American material described 

and figured by Say (1817) under the 

name Cancer panope Herbst, and (3) the type-specimen 

of Cancer panope figured by Herbst (1801, pi. 

54: fig. 5). The material under (1) presumably is 

the species now generally known as Panopeus herbstii, 

that under (2) is a mixture of Panopeus herbstii 

and Neopanope texana sayi (Smith, 1869) (cf. Rathbun, 

1930:335, 369), and that under (3) is Sphaerozius 

scaber (Fabricius, 1798). In order to legalize 

the current use of the name Panopeus herbstii for 

the common large East American mud crab, 

Holthuis (1979b: 159) selected as the lectotype of 

Panopeus herbstii the specimen from oyster beds of 

the east coast of the United States, that Say 

(1817, pi. 4: fig. 3) figured. 

By these actions it will remain possible to use 

both the generic name Panopeus H. Milne Edwards, 

1834, and the specific epithet herbstii H. 

Milne Edwards, 1834, in the accustomed sense. 

* Panopeus africanus A. Milne Edwards, 1867 

Panopeus africanus.—Frade, 1950:11, 26.—Capart, 1951:148, 

fig. 54, pi. 3: fig. 3.—Monod, 1956:325, figs. 406-415.— 

Rossignol, 1957:82, 83.—Longhurst, 1957:374, 375, 380, 

382; 1958:88.—Gauld, 1960:70.—Forest and Gantes, 

1960:352.—Rossignol, 1962:118.—Guinot and Ribeiro, 

1962:61.—Forest and Guinot, 1966:84.—Zariquiey Alvarez, 

1968:404, fig. 134b [Spain; references].—Uschakov, 

1970:443, 444, 447, 455 [listed].—Hartmann-Schroder 

and Hartmann, 1974:15.—Powell, 1979:127. 

Panopeus sp.—Monod, 1956:335, figs. 435-438. 

Eupanopeus africanus.—Bott, 1964:30. 


Sta 1, Lagos harbor, shore, 4


Other Material: Liberia: Rock Spring, Monrovia, O. F. 

Cook, G. N. Collins, 1$ (W). Free Port area, Monrovia, 

oyster cultch, 22 Apr 1953, G. C. Miller, 16* (W). Locality 

same, 24 Apr 1953, 1$ ov (W). 

Dahomey: Lagoon of Lac Nokoue near Zogbo, W of 

Cotonou, 29 Mar 1964, H. Hoestlandt, 1? (L). Point XI 

Lagoon near Contonou, 29 Mar 1964, H. Hoestlandt, 16* 

(L). Zogbo Lagoon near Contonou, 10 Apr 1964, H. 

Hoestlandt, 1$ (L). 

Nigeria: Harbor of Lagos, 13 Jun 1963, A. R. Longhurst, 

16* (L). S bank of mouth of Escravos River near Ajudaibo, 

Niger delta, 05°34.5'N, O5°11.75'E, 30 Jul 1975, C. B. 

Powell, 1 large 6* (L). W of Forcados town, near confluence 

of Odimodi Creek and Forcados River, 05°22'N, 05°26'E, 

28 Feb 1976, C. B. Powell, 14 specimens (L). Niger delta, 

between Brass and Port Harcourt, May-Aug 1960, H. J. G. 

Beets, 16* (L). 

Gabon: No specific locality, Duparquet, syntype of Panopeus 

afncanus, 1 dry c* (W, USNM 20263). 

Congo: Pointe-Noire, intertidal, Mar 1965, A. Crosnier, 

26*, 4$ (1 ov) (W). 

Zaire: Banana, mouth of Congo River, American Museum 

Congo Expedition 1909-1915, Jul-Aug 1915, H. Lang, 

46*, 5$ (1 ov) (W). Data same, Jul 1915, 56*, 39 (1 ov) (W). 

Angola: Luanda, W coast; American Museum Congo 

Expedition 1909-1915, 21 Sep 1915, H. Lang, 1$, 2juv (W). 

Between Luanda and Cuanza, 23 km from Luanda, 20 Jun 

1967, G. Hartmann, 1$ (L). Lobito, 1899, P. Kamerman, 16* 

(L). 



Monod, 1956, figs. 406-415. 

MalePleopod:Capart, 1951, pi. 3: fig. 3 (Congo); 

Monod, 1956, figs. 410-414 (Senegal, Sierra 

Leone, Ivory Coast). 

Color: "Coloration gris fonce a brun; extremite 

des pinces noir et blanc" (Capart, 1951:148). 

Rossignol (1957:83) gave a more detailed color 

description: "marron plus ou moins foncee. 

Chelipedes: face externe et bord superieur de la 

main, de raeme que le pouce et le doigt, tete de 

negre (bouts de doigts jaunatres). Face inferieure 

et bord inferieure de la main jaunatres." 


widths ranging from 6 to 43 mm; the carapace 

widths of ovigerous females range from 17 

to 30.5 mm. 

REMARKS.—We suspect that Monod's Panopeus 

sp., a damaged male, 12 X 16 mm, from the 

Ivindo River, near Ogooue, a locality in a river 

some distance from the sea in Gabon, actually 

can be identified with this species. One of the 

characters used by Monod to distinguish this 

specimen is the laterally directed fifth anterolateral 

tooth on the carapace, which is also somewhat 

upturned; it is curved upward in some of 

our specimens. Rathbun (1921:439) noted that 

"there is considerable variation in the shape of 

the lateral teeth of the carapace ... ," and she 

figured one specimen (1921, fig. 19a) in which 

the posterior teeth are directed almost laterally. 

The front of Monod's specimen is damaged. The 

deep sinus between the first and second tooth 

shown by Monod is also figured by Rathbun for 

three different specimens. 

The male pleopod of Monod's specimen, which 

seems to terminate in a club-shaped tip rather 

than in the trilobed apex typical of Panopeus, may 

be abnormal or damaged; the specimen is quite 

small, so it may just be undeveloped. A survey of 

the pleopods of species of Panopeus in the Smithsonian 

Institution, in which males of all species 

but P. convexus A. Milne Edwards, 1880, are represented, 

reveals that all of the other species in 

the genus have the typically trilobed male 

pleopod. 

Monod (1956:329) commented on the absence 

of the red spot on the inner surface of the ischium 

of the third maxilliped in the specimen from 

Ivindo. That spot, characteristic of P. ajricanus, is 

not always present in species in which it is known 

to occur (A. B. Williams, pers. comm.). 

The occurrence of Monod's specimen from Gabon 

in a river is consistent with the known habitat 

preference of P. africanus for lagoons and estuaries. 

Monod (1956:329) noted the occurrence of a 

red spot on the inner surface of the ischium of the 

third maxilliped in this species, as well as in 

Eurypanopeus blanchardi (p. 130). That spot also is 

characteristic of the western Atlantic P. herbstii H. 

Milne Edwards, 1834. In that species, Williams 

(1965:197) noted that in a sample of almost 600 

specimens from Beaufort, North Carolina, that 

spot was present in 100% of the males but only in 

55% of the females. The significance of the spot 

is unknown.

148 

BIOLOGY.—Panopeus africanus is a characteristic 

inhabitant of estuaries and lagoons along the 

West African coast; it may also occur littorally 

and sublittorally. Longhurst (1958) found it in 

Sierra Leone in estuaries as well as offshore to a 

depth of 140 m. It is rarely found far from the 

intertidal zone. 

Longhurst (1957) found this species in the 

stomach contents of the following fishes in the 

Sierra Leone River: Arius latiscutatus Giinther, 

Pomadasys jubelini (Cuvier), Diagramma macrolepis 

(Boulenger), Galeoides decadactylus (Bloch), and Cynoglossus 

senegalensis (Kaup). 

Apparently this species spawns all year. Ovigerous 

females have been collected in January, 

March, April, July, and November (Capart, 

1951; Monod, 1956; Rossignol, 1957; p. 147 

herein). 

DISTRIBUTION.—Eastern Atlantic, from S Portugal 

and SW Spain southward to Angola, including 

the offshore islands of Fernando Poo, 

Principe, and Sao Tome; possibly introduced to 

Durban Bay, South Africa (Barnard, 1954, 1955). 

Shallow water, usually in estuaries and lagoons. 


reported many specimens from localities between 

Mauritania and Angola. Since 1956 it has been 

reported from the following localities. 

Morocco: Oued Bou Regreg (Forest and Gantes, 1960). 

Guinea: Conakry, 0-2.5 m; lie Roume, lies de Los, shore 

(Uschakov, 1970). 

Sierra Leone: Sierra Leone River (Longhurst, 1957). Off 

Sierra Leone, 0-140 m (Longhurst, 1958). 

Ghana: Ada; off Accra, 8 m (Gauld, 1960). 

Nigeria: Elechi Creek, Port Harcourt, 04°47'15"N, 

06°48'45"E (Powell, 1979). 

Principe: No specific locality (Frade, 1950; Forest and 

Guinot, 1966). Rio Papagaio, shore (Forest and Guinot, 

1966). 

Sao Tome: No specific locality (Frade, 1950). 

Congo: Lagoon of Loango (Rossignol, 1957). Wof Pointe- 

Noirc (Rossignol, 1962). 

Angola: Mangais, Lobito (Bott, 1964). Baia de Luanda, 

shore; Baia do Lobito, shore; Praia da Rocha, near Benguela, 

shore (Guinot and Ribeiro, 1962). Between Cacuaco and 

Lobito-Benguela (Hartmann-Schroder and Hartmann, 

1974). 


Genus Paractaea Guinot, 1969 

ParactaeaGuinot, 1969d:241 [type-species: Xantho rufopunctatus 

H. Milne Edwards, 1834, by original designation; gender: 

feminine]; 1971:1071 [list of species: Actaea margaritaria A. 

Milne Edwards, 1867, not included]; 1976:249 (revision). 

REMARKS.—The eastern Atlantic Actaea margaritaria 

A. Milne Edwards, 1867, was listed under 

neither Actaea (p. 1070) nor Paractaea (p. 1071) by 

Guinot (1971), who noted (1969d:244): 

Enfin, il convient de dire un mot de quelques autres 

especes susceptibles d'entrer, apres revision, dans le genre 

Paractaea gen. nov. En effet, un certain nombre d'especes 

jusqu'a present rattachees a Actaea et qui ne peuvent etre 

conservees dans ce genre tel que nous l'avons delimite, offrent 

les memes caracteres essentiels que les Paractaea. II s'agit de 

Actaea margaritaria A. Milne Edwards, 1867, espece est-atlantique. 

In 1976 Guinot assigned margaritaria to Paractaea. 

* Paractaea margaritaria 


Actaea margaritaria.—Capart, 1951:159.—Gauld, 1960:70.— 

Serene, 1961:197 [listed].—Rossignol, 1962:117.—Chace, 

1966:637.—Forest and Guinot, 1966:77.—Guinot, 1969d: 

224, 251 [discussion]. 

Actaea (Actaea) margaritaria.—Monod, 1956:294, figs. 357- 

360.—Guinot and Ribeiro, 1962:56.—Ribeiro, 1964:9. 

Paractaea margaritaria.—Guinot, 1976:251. 


Sta 70, 33 m, branched Foraminifera, 1$ (L). 

Annobon: Sta 275, 9-69 m, rubble of coralline algae, 

26* (W). Sta 282, 18-37 m, nodular coralline algae, 1$, 3 juv 

(W). Sta 283, 51-55 m, nodular coralline algae, 3(5, 2? (L). 


Figures: Monod, 1956, figs. 357-360; Guinot, 

1976, pi. 16: fig. 6. 


(Ghana). 

Color: "Rouge intense, une tache jaune en arriere 

des yeux" (Capart, 1951:159). 


widths of 4 to 9 mm.


BIOLOGY.—Paractaea margaritaria is a sublittoral 

species, occurring on rough bottom in depths 

between 4-5 and 91 m. It has habitat preferences 

similar to those of P. rufopunctata africana; the two 

species were taken together at three stations by 

the Calypso (Forest and Guinot, 1966) and at two 

stations by the Pillsbury. Our material was collected 

on bottom with branched Foraminifera or 

the characteristic coralline algae found off Annobon. 

The Calypso specimens were taken in calcareous 

algae at four stations and on rocks and 

coral at two stations. Guinot (1976) reported one 

specimen from sand and gorgonians off Togo. 


collected in June and November (Guinot and 

Ribeiro, 1962; Ribeiro, 1964; Forest and Guinot, 

1966). Chace (1966) recorded an ovigerous female 

from Saint Helena in April. 

DISTRIBUTION.—West coast of Africa and Saint 

Helena (Chace, 1966). Off West Africa it is 

known from the Cape Verde Islands, Ghana, 

Gabon, and the offshore islands of the Gulf of 

Guinea: Annobon, Sao Tome, and Principe; it 

has not previously been recorded from the African 

mainland as far north as Liberia. It is a sublittoral 

species, occurring in depths between 4-5 and 

91 m. Monod (1956) reported material from 

Ghana and Annobon; since 1956 it has been 


Cape Verde Islands: No specific locality (Guinot, 1976). 

Porto da Praia, Sao Tiago (as La Praya) (Guinot, 1976). Sao 

Vicente (as Cap St. Vincent) (Guinot, 1976). Baia do Porto 

Grande, Sao Vicente, 3.5-11 m (Guinot and Ribeiro, 1962; 



Togo: No specific locality (Guinot, 1976). 

Principe: 01°43'N, 07°28'55"E, 37 m (Forest and Guinot, 

1966). 


W coast, 30 m; off Diogo Nunes, 4-5 m (all Forest and 


Annobon: No specific locality, 12 m (Guinot, 1976). 

01°27.5'S, O5°36.5'E, 35 m, and Isla Tortuga, NW coast, 


Gabon: No specific locality, in 50 m (Guinot, 1976). Off 

Libreville, 45-57 m (Rossignol, 1962). 

Congo: Baie de Pointe-Noire, 13 m (Guinot, 1976). 

Paractaea monodi Guinot, 1969 

Actaca (Actaea) rufopunctota.—Monod, 1956:293 [part] [not 

Actaea rufopunctata H. Milne Edwards, 1834]. 

Actaea rufopunctata.—Guinot, 1969d:250 [discussion] [not Actaea 

rufopunctata H. Milne Edwards, 1834]. 

Paractaea monodi Guinot, 1969d:259, fig. 33; 1971:1072 

[listed]; 1976:250 [listed], fig. 7.—Turkay, 1976b:61 

[listed], 67. 


Other Material: Azores: No other data, paratypes, Id, 19 

(W). 

Madeira: S coast near Ponta da Oliveira, 32°39'N, 16°- 

49'W, 0-20 m, diving, 15 Mar 1976, Onversaagd Sta 82, 1

150 

Azores, Madeira, Ilhas Desertas, the Canary Islands, 

the Cape Verde Islands, and the Mediterranean; 

for comments on material from Mediterranean 

localities see Guinot (1969d:260, 261). 

Guinot (1969d) summarized the literature. We 

have found only the following records published 

since then: 

Madeira: Funchal harbor; between Ponta da Garajau 

and Ponta da Oliveira, 5-6 m (Tiirkay, 1976b). 

Canary Islands: Arrecife, E of Lanzarote (Guinot, 1976). 

Egypt: W of Abou Kir; Abou Kir bay region: Rosetta to 

Port Said (Ramadan and Dowidar, 1976). 

*Paractaea rufopunctata africana Guinot, 1976 

Actaea rufopwclata.—Bouvier, 1906:4%.—Balss, 1914:102.— 

Crosnier, 1964:31.—Forest and Guinot, 1966:76. 

Actaea (Actaea) rufopunctata.—Monod, 1956:293 [part]. 

Paractaea rufopunctata forme africana Guinot, 1969d:251, fig. 

26; 1971:1071 [listed]. 

Paractaea rufopunctata africana Guinot, 1976:250, pi. 16: fig. 5. 


Sta 275, 9-69 m, rubble of coralline algae, 36, 1$ (L, W). 

Sta 282, 18-37 m, nodular coralline algae, Ic3, 1? (L). 

DESCRIPTION.—Guinot, 1969d:246 (P. rufopunctata 

rufopunctata), 251 (P. rufopunctata forme africana). 

Figures: Guinot, 1969d, fig. 26; 1976, pi. 16: 

fig. 5. 



REMARKS.—In her revision of Actaea and Paractaea, 

Guinot (1969d) recognized Paractaea rufopunctata 

rufopunctata as an Indo-West Pacific taxon 

and introduced the infrasubspecific name, africana, 

with no standing in nomenclature, for the 

tropical eastern Atlantic population; she later 

validated the name in 1976. In her discussion of 

the problem she noted (1969d:251) that "il y aura 

lieu de revenir sur le statut des representants 

ouest-africains de rufopunctata, en d'autres termes 

de decider si ceux-ci meritent bien de constituer 

une forme distincte de la rufopunctata typique." 

Unfortunately, the Pillsbury material is not adequate 

for us to elucidate the problem. 


BIOLOGY.—Off West Africa this species occurs 

from the intertidal zone to a depth of at least 

45 m; there is one Pillsbury record of 9-69 m. It 

was taken at 10 stations by the Calypso (Forest 

and Guinot, 1966): three were on or near (0-6 m) 

shore, two were from depths of 6-8 m, and five 

were from depths ranging from 15 (15-40 m) to 

45 m. The species apparently prefers rough bottom; 

the coralline algae habitat common around 

the offshore islands of the Gulf of Guinea has 

yielded several specimens from both the Calypso 

and the Pillsbury collections. This species also was 

found in the following habitats by the Calypso 

(Forest and Guinot, 1966): sand and rocks; rocks 

and coral; rocks, calcareous algae, and coral. 

Crosnier (1964) found the species in sponges, coral 

and marl on rocky bottom with gorgonians off 

Cameroon. 


recorded in June (Forest and Guinot, 1966). 

DISTRIBUTION.—West Africa. Guinot (1969d) 

recognized a forme africana for the population 

occurring off tropical West Africa, from the offshore 

islands of the Gulf of Guinea: Sao Tome, 

Annobon, and Principe, which she elevated to 

subspecific status in 1976. She erred, however, in 

considering the West African population to be 

exclusively insular (1969d:251); Crosnier (1964) 

has reported the species from the continental shelf 

off Cameroon. Guinot (1969d:259, fig. 33) assigned 

material from the Cape Verde Islands to 

Paractaea monodi Guinot (p. 149). West African 

records for P. rufopunctata africana are as follows: 


Principe: 01°35'N, 07°28'E, 45 m; 01°43'N, O7°28'55"E, 

37 m; and Ponta da Mina, beach (Forest and Guinot, 1966). 

Sao Tome: No specific locality (Bouvier, 1906; Guinot, 

1969d). Ponta Diogo Vaz, W coast, 0-6 and 30 m; in front 

of Sao Tome, 8 m; W of Ponta Diogo Nunes, shore; in front 

of Ponta Oquedelrei, 6 m (all Forest and Guinot, 1966). 

Annobon: No specific locality (Balss, 1914). 01°25'12"S, 

05°36'05"E, 20 m, and NW side, Isla Tortuga, 15-40 m 

(Forest and Guinot, 1966; Guinot, 1976). 

The identity of the material from Ascension 

Island assigned to this species by Benedict (1893)


and Rathbun (1930) remains to be determined. 

That material will be studied in a review of the 

decapods of Ascension now underway by Manning 

in collaboration with Fenner A. Chace, Jr. 

Genus Paraxanthias Odhner, 1925 

Paraxanthias Odhner, 1925:85 [type-species: Xanthodes notatus 

Dana, 1852, by original designation; gender: masculine].—Guinot, 




Paraxanthias eriphioides.—Monod, 1956:304, figs. 371-375 

[Cape Verde Islands; references].—Guinot and Ribeiro, 

1962:57 [Cape Verde Islands].—Ribeiro, 1964:10 [Cape 

Verde Islands].—Guinot, 1968a:718 [discussion]; 1971: 


Paraxanthias Periphioides.—Guinot, 1968a: 720, figs. 48, 53 

[Cape Verde Islands]. 


Azores and the Cape Verde Islands; sublittoral, 

10-85 m. 

Genus Pilumnopeus A. Milne Edwards, 1863 

Pilumnopeus A. Milne Edwards, 1863:289 [a genus without 

included nominal species; type-species: Pilumnopeus crassimanus 

A. Milne Edwards, 1867, a subjective junior synonym 

of Ozius serratifrons Kinahan, 1858, by subsequent 

designation by Balss, 1933b: 33, 34; gender: masculine; 

name 1643 on Official List, there dated 1867 in error].— 

Takeda and Miyake, 1969:119 [definition; list of Indo- 

West Pacific species]. 

* Pilumnopeus africanus (De Man, 1902) 

Heteropanope (Pilumnopeus) ajricana.—Monod, 1956:270, figs. 

319-325. 

Pilumnopeus africanus.—Uschakov, 1970:455 [listed]. 


Sta 224, Lagos, shore, sand beach, 1$ (W). 

Other Material: Liberia: Free Port area, Monrovia, oyster 

cultch, 13 Mar 1953, G. C. Miller, 1

152 

MATERIAL EXAMINED.—Ptllsbury Material: None. 

Other Material: Nigeria, between Brass and Port Harcourt, 

Niger delta, May-Aug 1960, H. J. G. Beets, 16*, 19 (L). 



Monod, 1956, figs. 314-318. 

Male Pleopod: Capart, 1951, pi. 3: fig. 1 (Congo); 

Monod, 1956, figs. 316-318 (Ghana). 

Color: According to Capart (1951:150) this species 

is "brun-rouge, les pinces plus claires, les 

doigts noirs a leur extremite." 



BIOLOGY.—Pilumnopeus caparti is an inshore, estuarine 

species; most of the material reported in 

the literature was collected at or near the mouths 

of rivers. Capart's (1951) material was taken in 

6-8 m on brown, black mud, and one of Monod's 

specimens was taken from a beacon at a river 

mouth. Longhurst (1957) found this species in the 

stomach contents of Arius latiscutatus Giinther in 

the Sierra Leone River and in 1958 characterized 

P. caparti as an estuarine, shallow shelf species; it 

was found on rocks, muddy sand, shelly mud, 

and sand. It also was reported by Longhurst in 

the latter paper from one shelf station in 72 m; 

that record appears to be questionable. 

Ovigerous females have been collected in August 

(Capart, 1951). 


between Sierra Leone and the Congo, 

usually in estuaries. Records in the literature 


Sierra Leone: No specific locality, in estuaries (Longhurst, 

1958). Freetown (Monod, 1956). Sierra Leone River 


Ghana: U.A.C. beach, Ada, River Volta (type-locality) 

(Monod, 1956; Gauld, 1960). 

Cameroon: Mouth of Wouri (River) (Monod, 1956). 

Congo: Mouth of Songololo River, Pointe-Noire (Rossignol, 

1962). 

Zaire: Crique de Banana, 06°01'S, 12°23'3O"E, 6-8 m 

(Capart, 1951). 


on Official List].—Takeda and Miyake, 1968:2 [list of 

Indo-West Pacific species]. 

Pilumnus hirtellus (Linnaeus, 1761) 

Cancer hirtellus Linnaeus, 1761:493. 

Pilumnus hirtellus.—Forest and Gantes, 1960:352 [Morocco].—Maurin, 

1968a: 107 [Mediterranean].—Zariquiey 

Alvarez, 1968:392, figs. 2g, 128c,d, 129f [Spain; 

references].—Christiansen, 1969:75, fig. 30, map 24 [Scandinavia].—Turkay, 

1976b:61 [listed], 69 [Madeira]. 

SYNONYM.—Pilumnus hirtellus ponticus Czerniavsky, 

1868. 


and British Isles southward to NW Morocco, 

Madeira, the Cape Verde Islands, Mediterranean; 

sublittoral, to a depth of 15-20 m. 

Pilumnus inermis A. Milne Edwards and 

Bouvier, 1894 

Pilumnus inermis.—Monod, 1956:247, figs. 291-295.—Longhurst, 

1958:88.—Gauld, 1960:70.—Rossignol, 1962:117. 

—Crosnier, 1964:31.—Forest and Guinot, 1966:70.—Zariquiey 

Alvarez, 1968:391 [Portugal; references].—Uschakov, 

1970:455 [listed].—Turkay, 1976a:25 [listed], 38, 

fig. 26 [Portugal in part]; 1976b:61 [listed], 69. 


Other Material: Azores: Ilha do Faial, Talisman, syntype of 

Pilumnus hirtellus var. inermis, 1 damaged $ (W, USNM 

22951). 

Madeira: S of Madeira, 32°39'N, 16°49'W, 125-150 m, 

shells and shell agglomerates, triangular dredge, 16 Mar 

1976, Onversaagd Sta 94, 16*, 2$, 1 juv (L). 

Senegal: Around Dakar, 28 Jan 1941, Th. Monod, 16* 

(W). 


1894:40. 


Male Pleopod: Monod, 1956, figs. 293-295 (Senegal); 

Turkay, 1976a, fig. 26 (Morocco). 


widths of 4.5 to 13 mm. The damaged 

female syntype has a carapace width of 11.6 mm. 

Genus Pilumnus Leach, 1815 

BIOLOGY.—Pilumnus inermis is generally a sub- 

Pilumnus Leach, 1815a:321 [type-species: Cancer hirtellus Linlittoral species; Monod (1956:249) recorded one 

naeus, 1761, by monotypy; gender: masculine; name 348 specimen taken at a beach, but the majority of


the records in the literature are from depths below 

4-5 m. Of the eight West African records in 

Monod (1956) for which depth is given, seven are 

from depths between 15 and 30 m. Indeed, literature 

records for this species suggest that it lives 

in shallower water in the Gulf of Guinea than it 

does in the northern part of its range. Nunes- 

Ruivo (1961) found the species in 350 m off 

Portugal, and Tiirkay (1976a) reported it in 

depths between 120 and 375 m off Morocco and 

(1976b) recorded it from ca 150 m, 220 m, and 

110-440 m off Madeira. Forest and Guinot (1966: 

71) pointed out that of 14 stations at which this 

species was taken by the Calypso in the Gulf of 

Guinea, 13 were in depths between 4-5 and 

21 m. 

This species, like most Pilumnus, prefers a rough 

bottom. Longhurst (1958) reported it from shelly 

sand off Sierra Leone, and Crosnier (1964) took 

it in sponges, coral and marl on rocky bottom 

with gorgonians off Cameroon. Rossignol (1962) 

reported it from muddy sand with Antedon off 

Pointe-Noire. The Calypso collected it from a variety 

of bottoms off Sao Tome and Principe islands 

in the Gulf of Guinea, usually on coralline 

algae or a mixture of coralline algae and other 

substrates (Forest and Guinot, 1966). 


collected in February, March, April, June, and 

July (Monod, 1956; Forest and Guinot, 1966). 

Nunes-Ruivo (1961) reported ovigerous females 

taken off Portugal in August. 

DISTRIBUTION.—Eastern Atlantic, from Portugal, 

the Azores, Madeira, the Cape Verde Islands, 

and the African coast from Morocco and Cabo 

Bojador, Spanish Sahara, southward to the 

Congo and possibly Gabon (that record based on 

material collected by Heurtel, whose localities are 

unreliable, as pointed out by Monod, 1956), including 

the offshore islands of the Gulf of Guinea, 

Principe and Sao Tome; sublittoral, in 4-5 m to 

400 m. Monod (1956) reported material from the 

Azores, Madeira, Spanish Sahara, Senegal, 

Guinea, Guinea-Bissau, Sierra Leone, Ghana, 

Principe, Sao Tome, and Gabon; since then the 

species has been recorded from the following: 

Madeira: No specific locality; Funchal harbor, ca. 150 m, 

220 m, and 110-440 m (Tiirkay, 1976b). 

Morocco: 33°19'N, 09°00'W, 120-180 m; 33°05.5'N, 

09°18'W, 160-250 m; 31°35'N, 10°05'W, 150-160 m; 31°- 

01'N, 10°16'W, 360-375 m (all Tiirkay, 1976a). 



1958). 

Ghana: Off Accra, 15 m; Prampram; Dixcove (all Gauld, 

1960). 


Principe: 01°42'30"N, 07°28'E, 21 m; 01°36'50"N, 07°- 

22'10"E, 19 m; 01°43'10"N, 07°28'20"E, 73 m; between 

Ponta da Mina and Ilheu Santana, 8-10 m and 10-12 m; in 

front of [Cais de] Santana, 11 m; in front of Praia Pequena, 

5-6 m (all Forest and Guinot, 1966). 

Sao Tome: Off Sao Tome, 5 m; 00°20'N, 06°46'E, 10 m; 

00°25'15"N, 06°43'05"E, 8-30 m; Baia de Ana de Chaves, 

5 m; in front of Ponta Oquedelrei, 6 m; in front of Praia 

Lagarto, 5-6 m; off Ponta Diogo Nunes, 4-5 m (all Forest 


Congo: Pointe-Noire, 8-9 m (Rossignol, 1962). 

* Pilumnus perrieri A. Milne Edwards and 

Bouvier, 1898 

Pilumnus perrieri.—Capart, 1951:143 [discussion].—Monod, 

1956:244, figs. 288-290.—Rossignol, 1962:117.—Forest 

and Guinot, 1966:70.—Uschakov, 1970:455 [listed]. 


Sta 23, 42 m, foliate brown to orange bryozoans, 1? (L). Sta 

27, 33 m, 1(3, 1$ ov (W). 

Nigeria: Sta 248, 33 m, 1

154 

taken on mud; mud with Area; mud, calcareous 

algae, and shell; rocks; and on calcareous algae 

(five stations). Rossignol (1962:117) noted that 

the species is "assez commun dans le coralligene 

et les fonds rocheux." 


March and May (Monod, 1956; Pillsbury). 


between the Cape Verde Islands and Senegal 

to Gabon; sublittoral, in depths between 20 

and 91 m. Monod (1956) summarized the earlier 

literature and reported material from Senegal 

and Guinea; since 1956 the species has been 

reported from the following: 


Nigeria: Off the mouths of the Niger River, O4°O3'N, 


Principe: Between Ilheu Caroco and Ponta do Pico Negro, 

40 m (Rossignol, 1962). 01°38'25"N, 07°22'05"E, 31 m; 

01°43'10"N, 07°28'20"E, 73 m; 01°43'N, 07°28'55"E, 37 m; 

in front of Baia de Santo Antonio, 50 m (Forest and Guinot, 

1966). 

Annobon: O1°27.5'S, 05°36.5'E, 35 m; N of San Antonio, 


Rio Muni: W Corisco Bay, 40 m (Rossignol, 1962). 

Gabon: W of Libreville, 45-57 m; W of Nyanga, 03°S, 

65-70 m (Rossignol, 1962). 

Pilumnus spinifer H. Milne Edwards, 1834 

?Pilumnus hirtellus.—Capart, 1951:140, fig. 50 [Spanish Sahara] 

[not Pilumnus hirtellus Linnaeus, 1761)]. 

Pilumnus spinifer.—Monod, 1956:251, figs. 296, 297 [Azores, 

Mauritania; references].—Zariquiey Alvarez, 1968:391, 

fig. 129a-e [Spain; references].—Christiansen, 1969:77, 

fig. 31, map 25 [Scandinavia].—Tiirkay, 1976b:61 [listed], 

69 [Madeira]. 

REMARKS.—There is a dry syntype of this species 

(USNM 20262), a male, carapace width 

30.8 mm, from the Mediterranean, in the collections 

of the Smithsonian Institution. 

DISTRIBUTION.—Eastern Atlantic, from Sweden, 

Portugal, Azores, Mediterranean, NW coast 

of Africa to Mauritania; sublittoral, to about 

100 m. 

* Pilumnus stebbingi Capart, 1951 

Pilumnus stebbingi Capart, 1951:144, fig. 52, pi. 3: fig. 6.— 

Monod, 1956:241, 632, figs. 279-287.—Longhurst, 1958: 


88.—Rossignol, 1962:116.—Guinot and Ribeiro, 1962: 



Sta 24, 35-37 m, dark red bryozoans, \S (L). 

Annobon: Sta 283, 51-55 m, nodular coralline algae, 26cJ, 

399 (7 ov), 15 juv (L, W). Sta 284, 73 m, black basaltic rocks, 

2$, 2juv(W). 



Monod, 1956, figs. 279-287. 

Male Pleopod: Capart, 1951, pi. 3: fig. 6 (Angola); 

Monod, 1956, figs. 284-287 (Senegal, 

Guinea). 

Color: Monod (1956:241) made the following 

observations on specimens from Guinea: "Epines 

rouges sur les pattes, qui sont annelees de rouge." 

Rossignol (1962:116) added: "Caracteristique par 

son tomentum epais, feutre, de couleur grise a 

gris-noir avec, sur les pinces, des tubercules epineux 

rouges." 



of ovigerous females is 10 to 12 mm. 

REMARKS.—The ornamentation of the chelae 

in our specimens is very variable: some have only 

pink tubercles, others pink and white tubercles; 

in some specimens the entire outer surface of the 

chela is covered with tubercles, in others the 

tubercles are restricted to the upper half of the 

surface. 

BIOLOGY.—Pilumnus stebbingi is a shelf rather 

than shore species, inhabiting moderate depths 

on relatively rough bottom. All but three of the 

20 depth records in the literature are from depths 

between 30 and 73 m; the exceptions are 25- 

30 m (Capart, 1951), 10-25 m (Longhurst, 1958), 

and 23 m (Forest and Guinot, 1966). Capart's 

(1951) material was collected on rocks; rocks, 

gravel, and coral; and sandy, rocky green mud. 

In addition to several records of the species from 

the coralline algae off Principe and Annobon, the 

same habitat in which the Pillsbury collected a 

relatively large series off the latter island, Forest 

and Guinot (1966) reported the species from mud, 

shells, gorgonians and ascidians; mud and com-


pacted sand (sable construit); mud, shells, and 

Cidaris; and, either mud and shell or sand, rocks 

and coral and calcareous algae. Longhurst (1958) 

found the species on muddy shell or shelly sand. 


June, and September (Monod, 1956; Forest and 

Guinot, 1966; Pillsbury). 

DISTRIBUTION.—Off West Africa, from scattered 

localities between Spanish Sahara and Angola, 

including the offshore islands of Principe 

and Annobon; sublittoral, in depths between 10- 

25 m and 73 m. Capart (1951) reported material 

from Gabon and Angola, and Monod (1956) 

added records from Senegal, Guinea, Sierra 

Leone, Annobon, and Principe. Records in the 

literature since 1956 include the following: 








1958). 

Principe: 01°35'N, 07°28'E, 45 m; 01°38'25"N, 07°22'- 

05"E, 31 m; 01°38'35"N, 07°21'35"E, 35 m; 01°43'10"N, 

07°28'20"E, 73 m; 01°43'N, 07°28'25"E, 37 m (all Forest 


Annobon: 01°27.5'S, 05°36.5'E, 35 m; N of San Antonio, 

23 m (both Forest and Guinot, 1966). 

Gabon: W of Nyanga, 65-70 m (Rossignol, 1962). 

Cabinda: W of Cabinda, 55 m (Rossignol, 1962). 

Angola: Luanda (Guinot and Ribeiro, 1966). 

Genus Platychelonion Crosnier and Guinot, 

1969 

Platychelonion Crosnier and Guinot, 1969:725 [type-species: 

Platychelonion planissimum Crosnier and Guinot, 1969, by 

monotypy; gender: neuter.].—Guinot, 1971:1078 [transferred 

with a question mark to Geryonidae]. 

Platychelonion planissimum Crosnier and 

Guinot, 1969 

Platychelonion planissimum Crosnier and Guinot, 1969:725, figs. 

1-9 [Congo]. 

DISTRIBUTION.—Known only from the typelocality, 

Pointe-Noire, Congo, in 10-20 m. 

Genus Platypodiella Guinot, 1967 

Platypodiella Guinot, 1967d:562 [type-species: Cancerspectabilis 

Herbst, 1794, by original designation; gender: feminine]; 


Platypodiella picta (A. Milne Edwards, 1869) 

Cancer geographicus.—Monod, 1933b:548 [footnote, nomen 

nudum]. 

Platypodia picta.—Monod, 1956:299, figs. 363-367 [Senegal, 

Sierra Leone, Ghana, Congo; references].—Rossignol, 

1957:83, fig. 3 [Congo].—Longhurst, 1958:88 [Sierra 

Leone].—Gauld and Buchanan, 1959:128 [Ghana].— 

Gauld, 1960:70 [Ghana].—Guinot and Ribeiro, 1962:57 

[Cape Verde Islands].—Rossignol, 1962:117 [Congo].— 

Ribeiro, 1964:10 [Cape Verde Islands].—Forest and 

Guinot, 1966:79 [Sao Tome].—Guinot, 1967d:562 [listed], 

563 [discussion; transferred to Platypodiella]. 

Platypodiella picta.—Guinot, 1971:1074 [listed]. 

DISTRIBUTION.—West Africa, from the Canary 

Islands, the Cape Verde Islands, several mainland 

localities between Senegal and the Congo, and 

the offshore islands of Annobon and Sao Tome 

in the Gulf of Guinea; shallow water, intertidal 

and sublittoral. 

Genus Pseudomedaeus Guinot, 1968 

Pseudomedaeus Guinot, 1968a:7l8 [discussion], 726 [type-species: 

Medaeus africanus Monod, 1956, by original designation; 

gender: masculine]; 1971:1069 [list of species]. 

* Pseudomedaeus africanus (Monod, 1956) 

?Paraxanthios eriphioides.—Capart, 1951:161, fig. 61 [fide 

Guinot and Ribeiro, 1962:58] [not Paraxanthias eriphioides 

(A. Milne Edwards, 1867)]. 

Xanthias tuberculidens.—Capart, 1951, pi. 3: fig. 13 [fide 

Guinot and Ribeiro, 1962:58] [not Xanthias tuberculidens 

Rathbun, 1911]. 

Medaeus africanus Monod, 1956:306, fig. 380.—Longhurst, 

1958:88.—Gauld,1960:70.—Rossignol,1962:118.—Guinot 

and Ribeiro, 1962:58, fig. 25.—Crosnier, 1964:34.—Forest 

and Guinot, 1966:80.—Crosnier, 1967:328, figs. 5, 11, 12, 

15.—Guinot, 1968a:718 [discussion], 726, 727 [listed]. 

Pseudomedaeus africanus.—Guinot, 1968a:726 [discussion], 727 

[listed], fig. 57; 1971:1069 [listed].—Williams, 1978:553, 

fig. 4a.

156 


Coast: Sta 64, 68 m, 2c5 (L, W). 

Gerommo Material: Gabon: Sta 211, 100 m, Id (W). 

Other Material: Dahomey: OfT Dahomey, 55 m, Oct 1963, 

A. Crosnier, 2$ (W). 



Monod, 1956, fig. 380; Guinot and Ribeiro, 1962, 

fig. 25; Crosnier, 1967, figs. 5, 11, 12, 15. 


Guinot and Ribeiro, 1962, fig. 25 (Spanish 

Sahara); Williams, 1978, fig. 4a (Gabon). 


widths of 5 to 21 mm; the largest was taken 

by the Geronimo. 

BIOLOGY.—Pseudomedaeus africanus is an offshore 

species, living on the shelf and upper slope in 

depths between 34 and 200 m. Crosnier (1964) 

characterized it as a cold water species, living in 

depths greater than 50 m off Cameroon. Longhurst 

(1958) found it on shelly mud in 34 to 

100 m off Sierra Leone, and Gauld (1960) reported 

it from colonies of Dendrophyllia in 100 m 

off Ghana. A large series of more than 70 specimens 

was collected by the Calypso on mud shells, 

gorgonians and ascidians in 43-45 m off Spanish 

Sahara (Forest and Guinot, 1966). 


March and May (Monod, 1956; Forest and 




depths between 34 and 200 m. Records since 1956 




Senegal: Dakar (Guinot, 1968a). 


1958). 

Ghana: OfT Accra, 100 m (Gauld, 1960). 

Dahomey: No specific locality, 55 m (Williams, 1978). 

Cameroon: No specific locality, deeper than 50 m (Crosnier, 

1964). 

Gabon: No specific locality (Crosnier, 1967). W of Libreville, 

45-57 m, and W of Nyanga, Pointe Panga, 03°10'S 

(Rossignol, 1962). 02°32'S, 09°05'E, 101 m (Williams, 1978). 

Congo: No specific locality (Crosnier, 1967). WSW of 

Pointe-Noire, 120 m (Rossignol, 1962). 


Angola: Ilha de Luanda, 105 m (Guinot and Ribeiro, 

1962). 

Genus Xantho Leach, 1814 

Xantho Leach, 1814:430 [type-species: Cancer incisus Leach, 

1814, by monotypy; gender: masculine; name 1016 on 

Official List].—Guinot, 1971:1067 [list of species]. 

Salax Gistel, 1848:xi [substitute name for Xantho Leach, 1814; 

type-species: Cancer incisus Leach, 1814; gender; masculine]. 

Xantho incisus (Leach, 1814) 

Xanlho floridus.—Chapman and Santler, 1955:374 [Azores] 

[not Cancer Jloridus Linnaeus, 1767]. 

Xantho (Xantho) incisa.—Monod, 1956:274 [Cape Verde Islands, 

Sao Tome(?); references]. 

Xantho incisus.—Figueira, 1960:9 [Azores].—Forest and 

Games, 1960:352 [Morocco]. 

Xantho incisus incisus.—Zariquiey Alvarez, 1968:398 [Spain; 

references].—Guinot, 1968a: 703, figs. 17, 26 [discussion]; 

1971:1067 [listed].—Tiirkay, 1976b:61 [listed], 68 [Madeira, 

Ilhas Desertas]. 

REMARKS.—There is material of this species 

from the Azores and the Canary Islands, but not 

from tropical West Africa, in the collection of the 


In the Mediterranean this species is replaced 

by Xantho granulicarpus Forest, 1953 (Zariquiey 

Alvarez, 1968:398). 


North Sea southward to Morocco, including the 

Azores, Madeira, Ilhas Desertas, Canary Islands, 

and Cape Verde Islands; possibly also from Sao 

Tome and Principe islands in the Gulf of Guinea 

(see Monod, 1956:274). It usually lives in shallow 

water, littorally to 30-40 m, but also has been 

recorded from as deep as about 100 m. 

Xantho pilipes A. Milne Edwards, 1867 

Xantho (Xantho) pilipes.—Monod, 1956:275, figs. 326-329 

[Mauritania, Senegal; references].—Guinot and Ribeiro, 

1962:53 [Angola]. 

Xantho pilipes.—Zariquiey Alvarez, 1968:395, fig. 130b 

[Spain; references].—Guinot, 1968a: 704 [discussion].— 

Christiansen, 1969:79, fig. 32, map 26 [Scandinavia].—


Guinot, 1971:1068 [listed]. —Tiirkay, 1975a:71 [listed], 74, 

figs. 4, 5 [Spanish Sahara]. 

DISTRIBUTION.—W coast of Norway, Shetland 

Isles, and England southward to Angola, Mediterranean; 

shallow water, intertidal to about 

40 m. 

Xantho sexdentatus (Miers, 1881) 

Xantho sexdentatus.—Capart, 1951:157.—Guinot, 1968a:704 

[discussion]; 1971:1068 [listed]. 

Xantho (Xantho) sexdentata.—Monod, 1956:277, figs. 330-334. 


Other Material: Senegal: Les Almadies, Dakar, 0-0.5 m, 

under stones, 9 Jun 1964, F. M. Bayer, R. B. Manning, L. B. 

Holthuis, 16, 1 juv (L). 

DESCRIPTION.—Miers, 1881a:211. 




widths of 6 and 8 mm. 

BIOLOGY.—Xantho sexdentatus is a shallow water 

species, occurring from the intertidal zone to a 

depth of 23 m. Apparently it is a characteristic 

inhabitant of rocky shores of Senegal (Sourie, 

1954a). 


February, July, and October (Monod, 

1956). 

DISTRIBUTION.—West Africa, where it is known 

only from the coasts of Spanish Sahara, Mauritania, 

and Senegal, from the intertidal zone to a 

depth of 23 m. Monod (1956) summarized earlier 

records. We have found no references to this 

species published since 1956, other than the remarks 

of Guinot (1968a) as part of her revision of 

the xanthids. 

Genus Xanthodius Stimpson, 1859 

Xanthodius Stimpson, 1859:52 [type-species: Xanthodius stemberghii 

Stimpson, 1859, by monotypy; gender: masculine; 

name 379 on Official List].—Guinot, 1971:1068 [list of 

species; Xantho denticulata White not included]. 

* Xanthodius denticulatus (White, 1848) 

Xantho (Xantho) denticulata.—Monod, 1956:280, figs. 335- 

339. 

Xantho denticulata.— Gauld, 1960:70. 

Xantho (Xantho) denticulatus.—Forest and Guinot, 1966:74. 

Xantho denticulatus.—Guinot, 1968a:700 [discussion]. 

Xanthodius denticulatus.—Guinot, 1968a:712 [discussion]. 

[Xantho] denticulatus.—Guinot, 1971:1068 [listed; "Parfois 

aussi, range dans Xanthodius"]. 


Poo: Sta 258, shore, 1$ (L). 

Annobon: Sta 271, shore, sand beach: 1$ (L). Sta 275, 9- 

69 m, rubble of coralline algae, 19, 1 juv (W). 




(Ghana). 



REMARKS.—We assign this species, which was 

not assigned to any genus in Guinot's preliminary 

revision (1968a, 1971), to Xanthodius. Guinot 

(1971:1068) noted that it and three other xanthids, 

including the West African Cycloxanthops 

occidentalis, were "au voisinage de Macromedaeus 

[Ward, 1942], et presentant egalement des affinites 

avec Leptodius et peut-etre aussi avec Cycloxanthops, 

trois-quatre especes apparentees." Apparently 

this problem is under study by Guinot. 

BIOLOGY.—Xanthodius denticulatus is a shallow 

water species, living from the intertidal zone to a 

depth of 21 m or more; it was taken in depths of 

11 m or less in 11 of 12 of the stations occupied 

by the Calypso (Forest and Guinot, 1966). One of 

the collections taken by the Pillsbury was in 9- 

69 m, so it may also occur somewhat deeper than 

21 m. It appears to be fairly abundant sublittorally 

off Sao Tome, where it was taken at 9 

stations by the Calypso (Forest and Guinot, 1966). 

The Pillsbury specimens were taken from shore 

stations and in the offshore bed of coralline algae 

off Annobon. The following bottom types were 

noted for the Calypso collections: calcareous algae 

(5 stations); rocks or sand and rocks (4 stations); 

and algae and calcareous algae (1 station). 

Off West Africa ovigerous females have been

158 

collected in February and June (Monod, 1956; 


DISTRIBUTION.—Atlantic. Western Atlantic 

from Bermuda and the Bahamas southward to 

Brazil. Eastern Atlantic, from Ghana on the 

mainland and the offshore islands of the Gulf of 

Guinea: Fernando Poo, Principe, Sao Tome, and 

Annobon; shallow water, intertidal to at least 

21 m, possibly as deep as 69 m. Monod (1956) 

was the first to record the species from West 

Africa; he reported material from Ghana, Sao 

Tome, and Annobon. Records in the literature 

since 1956 include the following: 

Ghana: Prampram and Winneba, intertidal (Gauld, 

1960). 

Principe: 01°42'30"N, 07°28'E, 21 m; Ponta da Mina, 

beach (Forest and Guinot, 1966). 

Sao Tome: No specific locality, shore; 00°20'N, 06°46'E, 

10 m; off Sao Tome 8 m; in front of Ponta Oquedelrei, 6 m; 

Ilheu das Cabras, 0-2 m; in front of the harbormaster's 

office, shore; in front of Praia Lagarto, 5-6 m; off Ponta 

Diogo Nunes, 4-5 m; in front of Ponta Sao Sebastiao, 11 m 



1966). 

Xanthodius inaequalis faba (Dana, 1852) 

Aciaeodes faba Dana, 1852b: 195; atlas, 1855 pi. 11: fig. 1. 

Chlorodius (Leptodius) convexus A. Milne Edwards, 1869:410. 

Xantho (Leptodius) inaequalis convexa.—Monod, 1956:284 [part 

of synonymy], 290, figs. 341a, 349, 350.—Guinot and 

Ribeiro, 1962:55.—Ribeiro, 1964:8. 

Xanthodius (Leptodius) inaequalis convexus.—Guinot, 1968a: 712 

[listed]. 

Xanthodius inaequalis convexus.—Guinot, 1968a: 714 [discussion], 

fig. 31; 1971:1068 [listed]. 


Other Material: Cape Verde Islands, Sao Vicente, 26 Jul 

1883, Talisman, \6, 1? (L). 


1900:97. 


pi. 17: figs. 1-6. 

Male Pleopod: Monod, 1956, figs. 349, 350 (Cape 

Verde Islands). 

Color: "Coloration vert olivatre, yeux grenat, 

pattes ambulatoires lavees de violet" (Monod, 

1956:290). 


REMARKS.—At least two authors, Odhner 

(1925:37, footnote, and 80) and Monod (1933b: 

511-513), have pointed out that Chlorodius convexus 

A. Milne Edwards, 1869, from the Cape Verde 

Islands, is conspecific with Actaeodes faba Dana, 

1852, from the same locality; Dana's name has 

priority and must be used for the taxon from the 

Cape Verde Islands. 

Monod (1956:284, synonymy) noted that A. 

Milne Edwards (1869) included material from 

Gabon as well as from the Cape Verde Islands in 

his original account of Chlorodius convexus; we can 

find only localities from the Cape Verde Islands 

in his account (Sao Vicente, Santa Luzia, Maio, 

Santiago). 

BIOLOGY.—Like the nominate subspecies, X. 

inaequalis faba is a littoral species inhabiting rocky 

shores. Monod (1956) recorded one specimen 

taken in pools with coral, and Guinot and Ribeiro 

(1962) and Ribeiro (1964) found it in rocks, on 

calcareous algae, and in worm tubes on rocks. 


August, September, and October (Monod, 1956; 

Guinot and Ribeiro, 1962; Ribeiro, 1964). 

DISTRIBUTION.—Known only from the Cape 

Verde Islands, where it occurs in the littoral zone. 

The nominate subspecies, Xanthodius inaequalis inaequalis 

occurs on the African mainland and the 

offshore islands of the Gulf of Guinea. Monod 

(1956) summarized the literature. Records since 

his paper were published include the following: 

Cape Verde Islands: No specific locality (Guinot, 1968a). 

Baia das Gatas, Praia da Matiota, and Baia da Calheta, Sao 

Vicente; Baia de Sal Rei, Boavista; Pedra Lume, Sal; and 

Baia do Tarrafal and Porto da Praia, Sao Tiago (all Guinot 


* Xanthodius inaequalis inaequalis 

(Olivier, 1791) 

Xantho faba.—Capart, 1951:155, fig. 58, pi. 3: fig. 14 [not 

Actaeodes faba Dana, 1852]. 

Xantho (Leptodius) inaequalis punctata.—Monod, 1956:284 [part 

of synonymy], 286, figs. 340, 341b, 342-348. 

Xantho inaequalis.—Gauld and Buchanan, 1959:127.—Gauld, 

1960:70.—Bassindale, 1961:491, fig. 5. 

Xantho (Leptodius) inaequalis inaequalis.—Guinot and Ribeiro, 

1962:54.—Forest and Guinot, 1966:75.—Guinot, 1968a: 

712 [listed].


Leptodius angolensis Bott, 1964:30, fig. 3, pi. 1: figs. 4, 5.— 

Guinot, 1968a:712 [discussion]. 

Xanthodius inaequalis inaequalis.—Guinot, 1968a:7l4 [discussion]; 

1971:1068 [listed]. 

SYNONYM.—Leptodius punctatus Miers, 1881. 


Poo: Sta 257, shore, 26, 1$ (L). Sta 258, shore, 15

160 

EASTERN ATLANTIC SPECIES.—Seven, six of 

which occur off tropical West Africa. Several of 

the names used by Monod (1956) have been 

changed, as follows: 


Pilumnoplax atlantica 

Pilumnoplax oxyacantha 

Carcinoplax bamardi 

Goneplax angulata 

Typh locarcinodes 

Machaerus atlanticus* 

Machaerus oxyacantha* 

Carcinoplax bamardi* 

Goneplax rhomboides 

Typhlocarcinodes inUgrifrons* 

integrifrons 

Acidops cessaci Acidops cessacii 

Hexapus (Thaumastoplax) Thaumastoplax anomalipes 

anomalipes (Hexapodidae) 

(Goneplacidae) 

Hexapus (Pseudohexapus) Pseudohexapus platydactylus 

platydactylus (Hexapodidae) 


Hexapus (Parahexapus) Parahexapus africanus 

africanus (Hexapodidae) 


Hexapus {Hexapus) Hexapinus buchanani 

buchanant (Hexapodidae) 


The four species assigned by Monod to Hexapus 

(Goneplacidae: Hexapodinae) are herein assigned 

to the Hexapodidae (pp. 166-181). 

The extralimital species is Eucrate crenata de 

Haan, 1835. An Indo-West Pacific immigrant 

into the eastern Mediterranean (Holthuis and 

Gottlieb, 1958; Ramadan and Dowidar, 1976). 

Subfamily CARCINOPLACINAE H. Milne 

Edwards, 1852 

Genus Carcinoplax H. Milne Edwards, 1852 

Curtonotus de Haan, 1833:4 [a subgenus established without 

included nominal species; an invalid junior homonym of 

Curtonotus Stephens, 1827 (Coleoptera)]. 

Carcinoplax H. Milne Edwards, 1852:164 [substitute name 

for Curtonotus de Haan, 1833; type-species: Cancer (Curtonotus) 

longimanus de Haan, 1833, by subsequent designation 

by Glaessner, 1929:111; gender: feminine].—Guinot, 

1969b:524 [revision]; 1971:1081 [list of species]. 

• Carcinoplax bamardi Capart, 1951 

Carcinoplax bamardi Capart, 1951:170, fig. 65, p!. 3: figs. 5, 

12.—Monod, 1956:351, figs. 456-461.—Forest, 1963:627, 


628.—Maurin, 1968b:484.—Guinot, 1969b:526 [discussion]; 

1971:1081 [listed]. 


Coast: Sta 44, 403-586 m, hard dark gray mud, 16* (L). Sta 

51, 329-494 m, 26, 19, 1 juv (L). 

Nigeria: Sta 255, 264-269 m, 16\ 19 ov (L). Sta 256, 409- 

485 m, 1$ (L). 

Gerommo Material: Gabon: Sta 179, 293 m, 2? (W). Sta 

198, 300 m, 2c5, 2$ (W). Sta 203, 200 m, 19 (W). 


Figures: Capart, 1951, fig. 65, pi. 3: figs. 5, 12; 

Monod, 1956, figs. 456-461. 


(Angola); Monod, 1956, figs. 460, 461 (Senegal). 

Color: Capart (1951:172) gave the following 

notes on color: "Coloration rose bistre, des taches 

noires a l'extremite des doigts des pinces." 


widths of 15 to 28 mm; the carapace width 

of the single ovigerous female is 23 mm. 

BIOLOGY.—Carcinoplax bamardi is a deep water 

species, living in depths between 200 m and at 

least 500 m; there is one record for the species in 

depths ranging from 403 to 586 m. Of 16 available 

depth records for the species, there are only 

five from depths of less than 300 m, none shallower 

than 200 m. It has been collected on mud, 

sandy mud, or hard dark gray mud. 


March and May (Capart, 1951; Pillsbury). 



depths between 200 m and at least 500 m (403- 

586 m). The species has not previously been 

recorded from off Nigeria or Gabon, although 

these records are well within its known range. 

Monod (1956), who was the second to record the 

species, reported material from Senegal; records 

in the literature since 1956 include the following: 

Spanish Sahara: Off Villa Cisneros, 300-500 m (Maurin, 

1968b). 

Ivory Coast: 04°54'N, 03°23'W, 380-400 m (Forest, 

1963). 

Ghana: 04°16'S, 02°09'30"W, 380-400 m (Forest, 1963).


Subfamily EURYPLACINAE Stimpson, 1871 

Genus Machaerus Leach, 1818 

Machaerus Leach, 1818, in 1817-1818:413 [a genus established 

without included nominal species; type-species: Pilumnoplax 

oxyacantha Monod, 1956, by present designation; 


DEFINITION.—Carapace smooth, convex anteroposteriorly, 

broader than long, with 3 or 4 

pairs of anterolateral teeth, outer orbital tooth 

distinct from first anterolateral tpoth. Frontal 

margin straight, short, less than x k carapace 

width, transversely grooved, with median notch. 

Supraorbital margin with 2 closed incisions. Basal 

antennal segment not fused to front, lacking lateral 

projection into orbit, flagellum in orbit. Male 

abdomen with 7 somites, third covering width of 

sternum, terminal somite elongate triangular. 

Chelipeds stout, slightly unequal. Walking legs 

slender, compressed, elongate, distal 3 segments 

setose dorsally and ventrally. Male first pleopod 

stout, curved, tapering distally, spinulose subapically. 

REMARKS.—Leach (1818, in 1817-1818:413) 

introduced the generic name Machaerus with the 

following remarks: il Machaerus; a new genus allied 

to Gonoplax, but differing in having short peduncles 

to its eyes, which are inserted into the same 

part of the shell as in that genus." The name 

apparently has been overlooked by most carcinologists; 

it was mentioned by Monod (1956:351, 

footnote). We believe that two West African goneplacid 

crabs placed in Pilumnoplax by Monod 

(1956), P. atlantica Miers, 1881, and P. oxyacantha 

Monod, 1956, but subsequently excluded from 

Pilumnoplax and the related Neopilumnoplax, should 

be assigned to Machaerus. 

The genus Pilumnoplax Stimpson, 1858, in 

which the two species here assigned to Machaerus 

were placed by Monod (1956), has as its typespecies 

Pilumnoplax sulcatifrons Stimpson, 1858 

(designated by Rathbun, 1918:21). Tesch (1918: 

154, 155) assigned P. sulcatifrons to Eucrate de 

Haan, 1835, making Pilumnoplax Stimpson a subjective 

junior synonym of Eucrate de Haan. As 

noted by Guinot (1969c:688) carcinologists, including 

Balss (1957:1656) and Glaessner (1969: 

R527) have continued to employ Pilumnoplax 

sensu Miers, 1886. 

Serene recognized that a new genus was required 

for Pilumnoplax sensu Miers, and in his 

checklist of the Indo-West Pacific Brachyura, 

Serene (1968:90) first introduced Neopilumnoplax 

as a manuscript name; it is a nomen nudum. 

Serene considered that five Indo-West Pacific 

species belonged to Neopilumnoplax: Pilumnus heterochir 

Studer, 1882, Pilumnoplax sinclairi Alcock, 

1899, Pilumnoplax abyssicola Miers, 1886, Pilumnoplax 

cooki Rathbun, 1911, and, with a questionmark, 

Pilumnoplax acanthomerus Rathbun, 1911. 

Neopilumnoplax Serene, 1969, was named in 

Guinot (1969c:689, footnote); its type-species is 

Pilumnus heterochir Studer, 1882. Neopilumnoplax 

was "established to include the species [of Pilumnoplax] 

described after Stimpson (1858)" (Serene, 

in Guinot, 1969c:689, footnote). Guinot noted 

that Serene included several species in this genus. 

She, however, restricted it to Pilumnus heterochir, 

Pilumnoplax americana Rathbun, 1898, and, possibly, 

to Pilumnoplax sinclairi Alcock, 1899 (erroneously 

spelled sainclairi throughout Guinot). Of 

the five species listed in Neopilumnoplax by Serene 

in 1968, two, P. heterochir and P. sinclairi, were 

retained in Neopilumnoplax by Guinot (1969c:689, 

footnote), two, Pilumnoplax abyssicola Miers, 1886, 

and P. cooki Rathbun, 1911, were transferred to 

Carcinoplax, and one, Pilumnoplax acanthomerus 

Rathbun, 1911, was not assigned to any genus. 

Guinot (1969c:689) made the following comment 

about that species: "C'est un Crabe enigmatique 

dont nous nous demandons s'il s'agit d'une 

Goneplacidae." 

Guinot (1969b:517) studied the two West African 

species here assigned to Machaerus and noted 

that "les deux especes ouest-africaines decrites 

dans le genre Pilumnoplax Stimpson a savoir P. 

atlantica Miers, 1881 (p. 259; cf. MONOD, 1956, p. 

341, fig. 442-449) et P. oxyacantha Monod, 1956 

(p. 346, fig. 450-455), offrent tous les caracteres 

communs a Euryplax-Eucrate-Frevillea et doivent 

etre rattaches aux Euryplacinae." She went on to

162 

discuss differences between these two species and 

those genera and concluded that "pour le moment, 

nous laisserons atlantica et oxyacantha sans 

attribution generique: en effet, ni Eucrate, ni Hettroplax, 

ni Euryplax, ni Frevillea ne peuvent vraiment 

convenir." 

Machaerus appears to show closest affinities with 

Neopilumnoplax Serene, differing primarily in having 

the outer orbital tooth distinct from the first 

anterolateral tooth and a much slenderer male 

abdomen, the terminal somite of which resembles 

a slender isosceles triangle, much longer than 

broad. 

•Machaerus atlanticus (Miers, 1881), 


Pilumnoplox atlantica.—Monod, 1956:341, figs. 442-449.—Buchanan, 


70.—Forest and Guinot, 1966:85.—Crosnier, 1969:531.— 

Guinot, 1969b:517, 518 [discussion]; 1969c:688 [discussion].—Uschakov, 

1970:455 [listed]. 

"Pilumnoplax" atlantica.—Guinot, 1969b:507, 508 [discussion], 

figs. 36, 53 [atlantica Stimpson in legends]. 

[Pilumnoplax] atlantica.—Guinot, 1971:1081 [listed]. 


Coast: Sta 46, 38-42 m, mud with dense Jullienella, 1


Sierra Leone, and Ghana. Since 1956 the species MEASUREMENTS.—Our specimens have cara- 

has been recorded from the following: 

pace widths of 7 to 27 mm; the single ovigerous 

Senegal: Joal, 10-11 m (Guinot, 1969b). 

Guinea: No specific locality (Uschakov, 1970). 09°40'N, 

14°05'W, 18 m (Forest and Guinot, 1966). 

female has a carapace width of 26 mm. 

BIOLOGY.—Machaerus oxyacantha, like M. atlanticus, 

is a sublittoral, shallow shelf species that 

Sierra Leone: OfT Sierra Leone, 6-30 m (Longhurst, prefers various types of mud bottoms including 

1958). 

sandy mud and mud with Foraminifera or shells, 

Ghana: OfT Accra, 8-20 fm (15-37 m) (Buchanan, 1958); 

10 and 36 m (Gauld, 1960). 04°37'N, 00°50'W, 90-100 m 




Congo: Pointe-Noire, 5-10 m (Crosnier, 1969). 

in depths between 7-8 m and 73 m. Of 50 depth 

records available, only one (7-8 m) is shallower 

than 10 m and only five are deeper than 50 m; 

88% of the depth records fall between 13 m (13- 

26 m) and 50 m. The species may occur in 

brackish water but it apparently prefers normal 

*Machaerus oxyacantha (Monod, 1956), 


sea water: It was collected by the Pillsbury in 

brackish water off Nigeria, and Monod (1956) 

reported one specimen from Ghana taken in a 

Pilumnoplax atlantica.—Capart, 1951:166, fig. 63 [not Pilum- river seine. 

noplax atlantica Miers, 1881]. 

Ovigerous females have been recorded in Jan- 

Pilumnoplax oxyacantha Monod, 1956:346, figs. 450-455.— uary, April, May, and December, suggesting that 

Gauld, 1960:70.—Rossignol, 1962:118.—Guinot and Ri- the species may spawn all year (Monod, 1956). 

beiro, 1962:63.—Crosnier, 1964:38, fig. on pi. A.—Forest 

DISTRIBUTION.—West Africa, from Mauritania 

and Guinot, 1966:85.—LeLoeuff and Intes, 1968:31, table 

1.—Guinot, 1969b:517, 518 [discussion]; 1969c:688 [dis- 

to Angola, possibly including Principe Island in 

cussion]. 

the Gulf of Guinea; sublittoral, from 7-8 m to 

"Pilumnoplax" oxyacantha.—Guinot, 1969b:507, 508 [discus- 73 m. Monod (1956) recorded material from Sension], 

figs. 38, 42, 54, 55. 

egal and Ghana. Records in the literature since 

[Pilumnoplax} oxyacantha.—Guinot, 1971:1081 [listed]. 1956 include the following: 


Coast: Sta 46, 38-4^ m, mud with denseJullienella, 46 (W). 

Sta 47, 37 m, bottom with Jullienella, \6, 1 juv (L). Sta 48, 

22 m, Id (L). 

Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 

several specimens (L). Sta 28, 49-53 m, 1$, 1 carapace (W). 

Nigeria: Sta 241, 59-63 m, mud and shell, 3


rhomboides Linnaeus, 1758, by monotypy; gender: feminine; 

name 313 on Official List].—Guinot, 1969b:520 

[comments on genus]; 1971:1081 [list of species]. 

Goneplat Leach, 1814:393, 430 [invalid original spelling for 

Goneplax Leach, 1814; name 482 on Official Index]. 

Gonoplax Leach, 1816:409, 413 [invalid spelling of Goneplax 

Leach, 1814; name 483 on Official Index]. 

Frevillea A. Milne Edwards, 1880:15 [type-species: Frevillea 

barbata A. Milne Edwards, 1880, by subsequent designation 

by Rathbun, 1918:25; gender: feminine]. 

Goneplax rhomboides (Linnaeus, 1758) 

Cancer rhomboides Linnaeus, 1758:626. 

Goneplax angulata.—Capart, 1951:168, fig. 64, pi. 3: figs. 4, 9 

[England], 10.—Barnard, 1954:126 [discussion].—Monod, 

1956:354, figs. 462-465.—Peres, 1964:27, 28, 29.— 

Maurin, 1968a: 19 [Spain]; 1968b:482, 489. 

Goneplax rhomboides.—Capart, 1951, pi. 3: fig. 11 [Nice].— 

Forest and Games, 1960:353.—Guinot and Ribeiro, 1962: 

63.—Forest and Guinot, 1966:86.—Zariquiey Alvarez, 

1968:414, figs, le, 138a,b [Spain; references].—Crosnier, 

1970:1215 [listed], 1216.—Tiirkay, 1976a:25 [listed], 38, 

fig. 28 [Portugal, in part]; 1976b:61 [listed], 71. 

Goneplax.—Maurin, 1968a: 14 [Spain]. 

Goneplax rhomboides.—Le Loeuff and Intes, 1968, table 1. 

SYNONYMS.—Cancer angulatus Pennant, 1777; 

Ocypode bispinosa Lamarck, 1801; Ocypode longimana 

Latreille, 1803. 


Undaunted Material: Angola: Sta 94, 90 m, 1


33°19.3'N, 08°39'W, 85 m; 33°17.2'N, 08°34.5'W, 65 m; 

31°01'N, 10° 16'W, 360-375 m; and 3O°2O'N, 10°03'W, 255- 

355 m (Turkay, 1976a). Off Agadir, 60-130 m; ofTEssaouira, 

32°08'N, 09°02'W, 33 m; and 31°37'N, 09°54'W, 70 m 

(Forest and Games, 1960). 

Spanish Sahara: Off Cabo Garnet, 400-700 m (Maurin, 

1968b). 

Mauritania: Off Tamzak (as Tamxat), 200-400 m 


Ivory Coast: Off Grand-Bassam, 200 m (Le Loeuff and 

Intes, 1968). 

Ghana: O4°37'N, 00°50'W, 90-100 m (Forest and Guinot, 

1966). 

Angola: Baia dos Tigres, 110 m (Guinot and Ribeiro, 

1962). 16°27'S, 11°35'E, 90 m; and 16°37'S, 11°22'E, 126 

m (Crosnier, 1970). 

Subfamily RHIZOPINAE Stimpson, 1858 

Genus Acidops Stimpson, 1871 

Acidops Stimpson, 1871b: 110 [p. 137 on separate] [type-species: 

Acidops fimbriatus Stimpson, 1871, by monotypy; gender: 

masculine]. 

Epimelus A. Milne Edwards, 1878:227 [p. 8 on separate] 

[type-species: Epimelus cessacii A. Milne Edwards, 1878, by 

monotypy; gender: masculine; name 147 on Official List]. 

Acidops cessacii (A. Milne Edwards, 1878) 

Acidops cessaci.—Monod, 1956:359, figs. 469, 470 [Senegal].—Guinot 

and Ribeiro, 1962:64, figs. 27, 28 [Cape 

Verde Islands].—Ribeiro, 1964:13 [Cape Verde Islands].—Forest 

and Guinot, 1966:88 [Sao Tome, Annobon]. 

Epimelus cessaci.—Garth, 1968:314 [discussion]. 


Verde Islands, Senegal, and Sao Tome and Annobon 

islands in the Gulf of Guinea; intertidal to 

10-30 m. 

Subfamily TYPHLOCARCINOPINAE Rathbun, 

1909 

Genus Typhlocarcinodes Alcock, 1900 

Typhlocarcinodes Alcock, 1900:326 [a genus established without 

included nominal species; type-species: Typhlocarcinus 

integrifrons Miers, 1881, by subsequent designation by 

Tesch, 1918:227; gender: masculine]. 

REMARKS.—Alcock (1900) established this genus 

for an Indian species that he did not name 

and that he compared with the West African 

Typhlocarcinus integrifrons Miers, 1881. The genus 

was established without any included nominal 

species. Tesch (1918:226, 227) considered that 

Typhlocarcinodes included four species: T. integrifrons 

(Miers, 1881), Caecopilumnus hirsutus Borradaile, 

1903, Typhlocarcinops piroculata Rathbun, 

1911, and Typhlocarcinodes crassipes Tesch, 1918; 

he designated T. integrifrons as the type-species. 

According to Article 69(a)(ii) of the International 

Code of Zoological Nomenclature, "if no nominal species 

were included at the time the genus was 

established, the nominal species-group taxa that 

were first subsequently and expressly referred to 

it are to be treated as the only original included 

species." Thus Tesch's designation of Typhlocarcinus 

integrifrons Miers as the type-species of Typhlocarcinodes 

is valid. 

Monod (1956:365) questioned whether the 

Indo-West Pacific species were congeneric with 

T. integrifrons. Tesch (1918:227) and Serene 

(1964b:237) pointed out differences between T. 

integrifrons and Indo-West Pacific species, and 

Takeda (1973:52) accepted the possibility that 

two genera might be involved. 

Those authors who consider the three Indo- 

West Pacific species assigned to this genus, Typhlocarcinodes 

hirsutus (Borradaile, 1902), T. piroculatus 

(Rathbun, 1911), and T. crassipes Tesch, 

1918, to be generically distinct from the eastern 

Atlantic T. integrifrons (Miers, 1881) should refer 

those species to Caecopilumnus Borradaile (1902b: 

267; type-species: Caecopilumnus hirsutus Borradaile, 

1902, by monotypy; gender: masculine). 

* Typhlocarcinodes integrifrons (Miers, 1881) 

Typhlocarcinodes integrifrons.—Monod, 1956:356, 632, figs. 

466-468—Longhurst, 1958:88.—Forest and Guinot, 

1966:87, fig. 8.—Crosnier, 1967:334. 

MATERIAL EXAMINED.—PUlsbury Material. Ghana: 

Sta 24, 35-37 m, dark red bryozoans, 16", 19 (L). 

DESCRIPTION.—Miers, 1881a:260.

166 

Figures: Monod, 1956, figs. 466-469; Forest and 

Guinot, 1966, fig. 8. 

Male Pleopod: Forest and Guinot, 1966, fig. 8 

(Principe). 



BIOLOGY.— Typhlocarcinodes integrifrons is a sublittoral 

species, known to occur in depths between 

12 and 90-100 m. It may prefer shallower water, 

as suggested by the only depth records available, 

viz. 16-44, 18, 30, 30-34, 35-37, 48, 51, and 90- 

100 m. 

Bottom types on which it is known to occur 

include: dark red bryozoans, 35-37 m (Pillsbury); 

shelly mud (2 specimens) and shelly sand (4 

specimens) in 16-44 m (Longhurst, 1958); sand 

and mud in 12 m, mud and shells in 30 m, and 

rocks and shells in 90-100 m (Forest and Guinot, 

1966); and coarse sand in 30 m and green mud in 

48 m (Crosnier, 1967). 

Ovigerous females have been collected in May 


DISTRIBUTION.—West Africa, from localities between 

Senegal and Dahomey as well as Principe 

Island in the Gulf of Guinea; sublittoral, from 

12 m to 90-100 m. Records in the literature 


Senegal: Goree (Miers, 1881a; Monod, 1956). 

Guinea: 1 mi [ 1.6 km] SE of lie Blanche, off Conakry, 

18 m (Monod, 1956). 

Sierra Leone: No specific locality, in 44 m (Monod, 1956); 

in 16-44 m (Longhurst, 1958). 

Ivory Coast: 04°36'N, 06°50^, 48 m (Crosnier, 1967). 

Ghana: 04°37'N, OOÔ'W, 90-100 m (Forest and Guinot, 

1966). 

Dahomey: 06°04'N, 02°26'E, 30 m (Crosnier, 1967). 

Principe: 01°39'35"'N, 07°26'53"E, 12 m, and 01°37'N, 

O7°22'E, 30 m (Forest and Guinot, 1966). 

Family HEXAPODIDAE Miers, 1886 

HEXAPODINAE Miers, 1886:275. 

EASTERN ATLANTIC GENERA.—Four, Hexapinus, 

new genus, Parahexapus, Pseudohexapus, and Thaumastoplax, 

each of which is represented by a tropical 

West African species. 

EASTERN ATLANTIC SPECIES.—Four, all occur- 


ring off West Africa. This family was not represented 

in the Pillsbury collections. The following 

name changes have occurred since Monod (1956): 


Hexapus (Thaumastoplax) Thaumastoplax anomalipes 

anomalipes 

Hexapus (Pseudohexapus) Pseudohexapus platydactylus 

platydactylus 

Hexapus (Parahexapus) Parahexapus afneanus 

africanus 

Hexapus (Hexapus) Hexapinus buchanani 

buchanani 

REMARKS.—The subfamily Hexapodinae was 

established within the Family Pinnotheridae by 

Miers (1886), who included four genera: Hexapus 

de Haan, 1833; Amorphopus Bell, 1859; Thaumastoplax 

Miers, 1881; and Asthenognathus Stimpson, 

1858. The latter genus is now considered to be a 

pinnotherid (Schmitt, McCain and Davidson, 

1973:128). Ortmann (1894:690) retained the 

Hexapodinae within the Pinnotheridae. Alcock 

(1900:287) recognized the affinities of the Hexapodinae 

with the Goneplacidae and included it 

as one of five subfamilies of the latter. He was 

followed in this by Borradaile (1907) and subsequent 

authors. 

We believe that the complete suppression of 

the last pair of pereiopods in members of this 

subfamily is a fundamental character which warrants 

the recognition of the Hexapodidae as a 

family distinct form but related to the Goneplacidae. 

We treat it as a family here. 

The following genera and species have been 

named in the family: Hexapus was erected by de 

Haan (1833:5; 1835:63) for Cancer sexpes Fabricius, 

1798; it is apparent that several different 

species have erroneously been identified with H. 

sexpes by subsequent authors. Other species described 

in Hexapus include: H. williamsi Glassell, 

1938, eastern Pacific; H. stebbingi Barnard, 1947, 

South Africa; H. buchanani Monod, 1956, West 

Africa; H. granuliferus Campbell and Stephenson, 

1970, Australia; and H. estuarinus Sankarankutty, 

1975, India. Each of these latter species are assigned 

to different genera below.


Amorphopus (Bell, 1859:27) was established for 

a single species from an unknown locality, A. 

cylindraceus Bell. According to Bell, it differs from 

Hexapus in having a rudimentary pair of fifth legs, 

a feature shared with no other member of the 

family. Rathbun (1897b: 163) pointed out that 

this name was preoccupied twice [according to 

Neave, Nomenclator Zoologicus, the oldest name is 

Amorphopus Audinet-Serville, 1838 (Insecta)] and 

substituted Paeduma for it. Rathbun's name apparently 

has been overlooked by nearly all zoologists 

dealing with this group; Gordon (1971:108) 

pointed out that Amorphopus was preoccupied. 

Amorphopus Bell has been recognized by Alcock 

(1900:293), Stebbing (1910:315) (who suggested 

that "there seems little justification for the separation 

either of Amorphopus or Thaumastoplax from 

Hexapus."), Tesch (1918:238), Balss (1957:1658), 

and Serene (1968:93). We recognize Paeduma 

Rathbun (p. 173). 

Thaumastoplax Miers (1881a:261) was established 

for a single species from Senegal, 71 anomalipes 

Miers; it was distinguished from Hexapus in 

having a slender third maxilliped with the propodus 

dilated distally. Rathbun (1910:113) described 

two additional species from Thailand, T. 

orientalis and T. chuenensis. A fourth species, T. 

spiralis Barnard (1950:301), was described from 

South Africa. Although Monod (1956:362, 363) 

treated Thaumastoplax as a subgenus of Hexapus, 

Tesch (1918:238), Balss (1957:1658), Glaessner 

(1969:R527) and Sakai (1976:555) all considered 

it to be a distinct genus. We also consider Thaumastoplax 

to be a valid genus; species described in 

it subsequent to 1881 are assigned to other genera 

below. 

Lambdophallus Alcock (1900:329) was based on 

an Indian species, L. sexpes Alcock, in which the 

apices of the male pleopods extended laterally 

well beyond the abdomen, lying in deep lateral, 

transverse grooves in the sternum. A second species, 

Lambdophallus anfractus, was described on material 

from Thailand by Rathbun (1909:113). 

Stephensen (1945:182) considered Lambdophallus 

to be a synonym of Hexapus and synonymized 

both L. sexpes and L. anfractus with H. sexpes. 

Monod (1956:362) and Sakai (1976:554) both 

considered Lambdophallus to be a subgenus of Hexapus; 

Tesch (1918:239) and Balss (1957:1658) both 

considered it to be a distinct genus, as we do. 

Hexaplax Doflein (1904:122) was established for 

a deep water species from India, H. megalops 

Doflein. It is characterized by long slender walking 

legs and very large, hammer-shaped eyes. It 

has been considered a distinct genus by all subsequent 

authors (see Sakai, 1976:556, for synonymy). 

Parahexapus was erected by Balss (1922:77) for 

P. africanus from Cabinda; one of its distinguishing 

features is its slender third maxilliped. Balss 

(1957:1658) retained Parahexapus as a distinct genus, 

but Monod (1956:362) considered it to be a 

subgenus of Hexapus. We believe that it is generically 

distinct. 

Pseudohexapus was recognized by Monod (1956: 

362, 365) as a subgenus of Hexapus. It included 

but one species from West Africa, Hexapus {Pseudohexapus) 

platydactylus, characterized by broadened, 

flattened dactyli on the walking legs. We 

recognize Pseudohexapus here. 

Monod (1956), who studied four species representing 

four different subgenera, reviewed the 

literature and pointed out numerous discrepancies 

in accounts in the literature purporting to 

deal with the same species. He pointed out that 

Stephensen's (1945:182) H. sexpes probably was 

not the same species reported by de Haan and 

provided diagnoses of five subgenera within Hexapus: 

Hexapus, Thaumastoplax, Parahexapus, Pseudohexapus, 

and Lambdophallus. 

West African material available to us includes 

representatives of only two species, Thaumastoplax 

anomalipes and Parahexapus africanus. The collections 

of the Smithsonian Institution, however, 

include specimens identified with several additional 

species from Indo-West Pacific localities: 

Hexapus sexpes (Fabricius) from Japan and the 

Philippine Islands, Thaumastoplax orientalis (Rathbun) 

from Japan, paratypes of Lambdophallus anfractus 

Rathbun from Thailand, and Hexapus williamsi 

Glassell from Mexico. Examination of this 

relatively rich material, which proved to be nee-

Igg SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

essary in order to determine the status of the slender or distally dilated, and the presence or 

species known from West Africa, has suggested to absence of a flagellum on the exopod; eye shape; 

us that all of the subgenera recognized by Monod relative length of walking legs; shape and configshould 

be recognized as distinct genera, that sev- uration of the male abdomen; the shape and 

eral new genera should be recognized, and that extent of sternal grooves in the male, and strucseveral 

name changes are required for known tureof the male pleopod. These characters appear 

species. All existing genera and the new genera to us to afford reliable features for differentiation 

that we believe should be recognized are charac- of genera. 

terized below. Gordon (1971:108) was correct in stating that 

Monod's (1956:361) excellent review of the "the taxonomy of the Hexapodinae poses some 

subgenera of Hexapus stressed the importance of problems which can only be solved by a careful 

several characters that appear to have been over- re-examination of all of the available material." 

looked in the past; several of these have not been We believe that all records in the literature redescribed 

for all species. They include: the shape quire verification. Descriptions of several new 

of the third maxillipeds, whether slender or species and a redescription of the type of Cancer 

broad, and especially the shape of the propodus, sexpes Fabricius are in preparation by Manning. 

Key to Genera of Hexapodidae 

1. Eye hammer-shaped, cornea very large [carpus, propodus, and dactylus 

of third maxilliped slender; walking legs long, slender, merus of longest 

pair longer than carapace] Hexaplax 

Eye small, reduced, cornea scarcely or not at all broader than stalk ... 2 

2. Eyes fixed. Walking legs (Figure 34a) long, slender, merus of longest pair 

longer than carapace. Male with long, narrow transverse grooves in 

sternum extending laterally from abdominal fossa beyond bases of third 

maxillipeds (Figure 34b) (carpus, propodus, and dactylus of third 

maxilliped slender) ". . . Lambdophallus 

Eyes movable. Walking legs short, merus of longest pair shorter than 

carapace. Male lacking long, narrow transverse grooves in sternum 

extending laterally from abdominal fossa beyond bases of third 

maxillipeds 3 

3. Propodus of third maxilliped strongly dilated distally (Figure 31b, 39a) . 4 

Propodus of third maxilliped not strongly dilated distally, slender, like 

carpus and dactylus (Figures 31a, 33a) 6 

4. Male pleopod stout, strongly twisted into a tight spiral (Figure 

37*) Spiroplax, new genus 

Male pleopod variously shaped, not twisted into a tight spiral 5 

5. Third to fifth abdominal somites fused in male. Male pleopod (Figure 

39c) slender, sinuous, apex directed anteriorly Thaumastoplax 

Third and fourth and fourth and fifth abdominal somites fused in male. 

Male pleopod (Figure 35b) slender, strongly recurved posteriorly, apex 

directed posteriorly or recurved anteriorly, shaped like a figure 

° Paeduma 

6. Male abdomen with second through sixth somites fused 

Stevea, new genus


Male abdomen with third to fifth somites fused 7 

7. Dactylus of each walking leg enlarged, triangular, strongly flattened 

dorsoventrally (male pleopod stout, with proximally directed subapical 

spines) Pseudohexapus 

Dactyli of walking legs not enlarged or strongly flattened dorsoventrally 

8 

8. Terminal somite of male abdomen (Figure 40a,d) trilobed or trefoil in 

shape, with distinct lateral angular projections (sixth abdominal somite 

divided longitudinally in type-species; male pleopod stout, bent laterally, 

apex directed anteriorly) Tritoplax, new genus 

Terminal somite of male abdomen (Figures 3\b, 32c) broadly rounded, 

not trilobed 9 

9. Third maxilliped (Figure 36a) slender, almost pediform, ischium width 

about % merus length. Male pleopod (Figure 36c) slender, sinuous, 

tapering abruptly to almost filiform apex, with strong, recurved spines 

near midlength (sixth and seventh somites of male abdomen (Figure 

36b) very slender, width half that of first fused somite) . Parahexapus 

Third maxilliped broad, ischium width usually subequal to or greater 

than merus length 10 

10. Sternum in male (Figure 336) with broad, deep grooves extending anterolaterally 

from end of abdominal fossa to bases of third maxillipeds. 

Male pleopods bent laterally, apices lying in sternal grooves . Hexapus 

Sternum in male (Figure 32c) lacking broad, deep grooves extending 

anterolaterally from end of abdominal fossa to bases of third maxillipeds, 

with at most triangular anterolateral projections on abdominal 

fossa, not extending laterally beyond border of first fused somite. Male 

pleopods not bent, apices concealed under abdomen or visible on either 

side of terminal somite Hexapinus, new genus 

Genus Hexapinus, new genus 

FIGURES 31, 32c,d 

TYPE-SPECIES.—Hexapus latipes de Haan, 1835, 

pi. D. 

ETYMOLOGY.—The masculine name Hexapinus 

is derived from the name Hexapus and the Latin 

suffix-j>iMJ, pertaining to. 

DIAGNOSIS.—Eyes very small, cornea subglobular. 

Third maxilliped (Figure 31a) with broad 

ischium and merus, inner margin of ischium concave 

proximally, straight or convex distally, carpus, 

propodus, and dactylus slender, cylindrical, 

dactylus longer than propodus; exopod with flagellum. 

Walking legs short, stout, merus of longest 

pair shorter than carapace. Male pleopods (Figure 

3\c,d) stout, sinous but not strongly angled 

distally, apices concealed under abdomen or 

scarcely visible next to seventh somite. Male abdomen 

(Figure 3\b) with third to fifth somites 

fused, terminal somite rounded apically, smooth 

laterally. Female abdomen of 7 somites. 

INCLUDED SPECIES.—Hexapus latipes de Haan, 

1835, from Japan; Hexapus buchanani Monod, 

1956, from West Africa; Hexapus granuliferus 

Campbell and Stephenson, 1970, from Australia; 

and one or more species identified with Hexapus 

sexpes by various authors: A. Milne Edwards, 

1873b:253, pi. 12: fig. 1 (New Caledonia); De 

Man, 1888:322, pi. 13: fig. 3 (Amboina); Zehntner, 

1894:159 (Amboina); Nobili, 1906a: 146 (Persian 

Gulf); Tesch, 1918:240, pi. 17: fig. 1 (Indonesia); 

Balss, 1938:74 (Marshall Islands); Stephensen, 

1945:182, fig. 53 (Persian Gulf); Guinot-

a 

170 

FIGURE 31.—Hexapinus buchanam (Monod): a, third maxilliped; 

b, male abdomen; c, male pleopod; d, apex of male 

pleopod (all from Monod, 1956, figs. 498-500). 

Dumortier and Dumortier, 1960:130, fig. 9 (no 

locality); Griffin, 1972:85 (Australia). 

Hexapus sexpes sensu A. Milne Edwards, 1873(b) 

(Figure 32a,6) from New Caledonia differs from 

our material of Hexapinus latipes from Japan in 

having a narrow cornea and the surface of the 

carapace covered with setae. It may well represent 

a new genus. Unfortunately, the female holotype 

could not be located at the Museum national 

d'Histoire naturelle, Paris, in October 1977 (J. 

Forest, pers. comm.). 

We suspect that two distinct Japanese species 

have been referred to H. sexpes by various authors. 

De Haan (1835) reported that his material measured 

3X5 lines (1 line = 2.12 mm) or 6 X 11 

mm; thus it was a relatively small species. Sakai 

(1976) noted that some specimens from Japan 

measured as large as 18 X 29.2 mm. Available 

material of both forms in the collection of the 

Smithsonian Institution exhibit specific differences. 

An examination of the Japanese specimen that 

de Haan (1835:35, 63, pi. 11: fig. 5) described 

and figured as Hexapus sexpes (Fabricius, 1798) 

and which still is preserved dry in the collection 

of the Rijksmuseum van Natuurlijke Historie, 

Leiden (a male, cl 6.7 mm, cb 10.0 mm, Crust. 

D. 31783), shows that it is different from Fabricius' 

type of Cancer sexpes and should be referred 

to the genus Hexapinus. For it the specific epithet 

latipes de Haan is available. On plate D of Fauna 

Japonica, namely, de Haan (1835) used the name 


Hexapus latipes for a figure of the mouthparts of 

his Japanese Hexapus, which he named H. sexpes 

in his text and on pi. 11: fig. 5. That the word 

latipes is not a typographical error is shown by the 

fact that the piece of cardboard on which de 

Haan glued the mouthparts of this specimen, and 

which is also present in the collection of the 

Leiden Museum, bears the following inscription 

in de Haan's handwriting: "Hexapus latipes 

Faun. Jap." Evidently de Haan at first considered 

the species new and gave it the name Hexapus 

latipes, and when he later changed his mind and 

identified the species with Cancer sexpes, the name 

latipes had already been engraved on plate D and 

could not be changed. It is peculiar that de Haan 

under Hexapus sexpes nowhere referred to plate D 

and did not correct the name latipes in the 

erratum. 

The specific epithet latipes, published in the 

combination Hexapus latipes with a figure, is an 

available name and must be used for the present 

Japanese Hexapinus. 

Hexapinus latipes has been discussed under the 

name Hexapus sexpes by Tesch (1918), Sakai (1934, 

1936, 1939, 1976), and Odawara (1965). 

Although de Haan identified his species with 

Cancer sexpes, he noted that in his specimens the 

FIGURE 32.—Hexapus sexpes sensu A. Milne Edwards: a, dorsal 

view; b, front (from A. Milne Edwards, 1873b, pi. 12: fig. 1, 

la). Hexapinus granuliferus (Campbell and Stephenson): c, 

abdomen of male; d, gonopod (apex broken?) (from Campbell 

and Stephenson, 1970, fig. 49g,h).


front was truncate and deflexed and further noted 

(1835:64) that "Fabricius dicit frontem emarginatam." 

Hexapinus differs from the other hexapodid genera 

with short legs, broad maxillipeds, and the 

third to fifth abdominal somites fused in males, 

Hexapus and Tritoplax, as follows: In Hexapus sternal 

grooves extend anterolaterally from the abdominal 

fossa almost to the level of the bases of 

the third maxillipeds, and the male pleopods are 

bent laterally and are situated in these grooves; 

in Hexapinus these grooves are restricted to the 

end of the abdominal fossa and the male pleopods 

are more or less straight and are concealed under 

the abdomen (Figure "52c,d)- In Tritoplax the terminal 

somite of the male abdomen is trilobed or 

trefoil in shape, whereas in Hexapinus it is evenly 

rounded. 

We are not certain that Hexapus granuliferus 

Campbell and Stephenson, 1970, should be assigned 

here, although superficially it resembles 

Hexapinus latipes. Their figure (4%) of the male 

pleopod (Figure 32

172 

tylus longer than propodus; exopod with flagejlum. 

Walking legs short, merus of longest leg 

(third) shorter than carapace. Male pleopods 

(Figure 33c) not concealed under abdomen, lying 

in deep, oblique grooves (Figure 33b) on anterior 

part of sternum, apices setose, directed anterolaterally. 

Male abdomen (Figure 336) with third to 

fifth somites fused, terminal somites rounded, not 

trilobed. Female abdomen with 7 free somites. 

INCLUDED SPECIES.—Cancer sexpes Fabricius, 

1798, from southern India; Lambdophallus anfractus 

Rathbun, 1909, from Thailand and Japan; and 

Hexapus estuarinus Sankarankutty, 1975, from 

India. 

REMARKS.—De Haan (1835:35, 63, pi. 11: fig. 

5, pi. D) included in the genus Hexapus (which he 

had established in 1833 without nominal species) 

two nominal species: Cancer sexpes Fabricius, 1798, 

and his new Hexapus latipes. On plate D, de Haan 

used the name Hexapus latipes for the specimen, 

which in his text (pages 35 and 63, and pi. 11: 

fig. 5), he had indicated with the name Hexapus 

sexpes (Fabricius). Hexapus latipes is an available 

name, and the epithet latipes has to be used for 

the species of Hexapinus to which de Haan's specimen 

belongs. This specimen, namely, is specifically 

(and generically) distinct from the typespecimen 

of Cancer sexpes Fabricius, 1798. As Cancer 

sexpes is the type-species of the genus Hexapus 

it should be known as Hexapus sexpes (Fabricius, 

FIGURE 33 — Hexapus anfractus (Rathbun): a, third maxilliped; 

b, sternum of male; c, male pleopod. (All from Rathbun, 

1910, fig. 36.) 


1798); for de Haan's Japanese species the name 

Hexapinus latipes (de Haan, 1835) is correct. 

De Haan (1835:64) did note differences between 

his Japanese specimen, of which he described 

the front as deflexed and truncate, and 

the description of Cancer sexpes in which "Fabricius 

dicit frontem emarginatam." 

Fabricius (1798:334) gave the following description 

of Cancer sexpes: 

C. thorace laevi integerrimo, fronte emarginata, pedibus 

senis. 

Habitat in India orientali Dom. Daldorff. 

Parvus thorax laevis in medio lineolis duabus impressis. 

Frons deflexa, emarginata. Pedes omnino sex inermes. Chelae 

breves, laeves altera maiore. 

The type of Cancer sexpes Fabricius, in the Universitetets 

Zoologiske Museum, Copenhagen (see 

Zimsen, 1964:650), was examined by Manning in 

Copenhagen in 1977. It is a male, cl 6.2 mm, cb 

9.5 mm, preserved dry, and in very good condition. 

As mentioned by Fabricius, the front is 

faintly emarginate. It was compared with the 

holotype of Lambdophallus anfractus Rathbun, and 

differs from the latter in having the carapace less 

granular, the fingers of the chela longer than the 

palm, and the walking legs are slenderer and less 

setose. There is no doubt, however, that the two 

species are congeneric. Both share the deep, 

oblique sternal grooves, figured well by Rathbun 

(1910, fig. 36a; see Figure 33b herein), with similar 

male pleopods. The abdomen of the male in both 

species has the third to the fifth somites fused. A 

redescription of the type-specimen of Hexapus 

sexpes is in preparation. 

Hexapus estuarinus Sankarankutty, 1975, from 

the Cochin region, SW India, may well prove to 

be synonymous with Hexapus sexpes. In both species 

the chelae are unequal and the second leg is 

the longest of the walking legs. 

This genus has deep sternal grooves like Lambdophallus 

sensu stricto, but differs in having very 

short pereiopods, as in most other genera in the 

family. The sternal grooves are deep and well 

defined but extend anterolaterally almost to the 

base of the maxillipeds rather than extending 

laterally as in Lambdophallus (Figure 34b). In H.


anfractus the third legs are the longest, the male 

abdomen extends almost to the level of the insertion 

of the third maxillipeds, and the carapace is 

relatively broad (width =1.5 times length). 

We do not agree with Campbell and Stephenson 

(1970:291) that Hexapus anfractus (Rathbun) 

can be identified with Hexapinus latipes from Japan. 

The sternal grooves in our material of H. 

anfractus are much better developed than they are 

in our Japanese material of H. latipes, in which 

the grooves are represented only by an emargination 

in the lateral margins of the sternal abdominal 

trench. We refer Japanese specimens previously 

assigned to H. sexpes to a new genus, Hexapinus. 

In L. anfractus the grooves extend anterolaterally 

from the terminal segment of the abdomen 

to the bases of the articulations of the third 

maxillipeds. The size and extent of development 

of these grooves are completely different in the 

two forms, and different from that found in Lambdophallus, 

sensu stricto, in which they extend laterally 

from the last abdominal somite. 

Genus Lambdophallus Alcock, 1900 

FIGURE 34 

Lambdophallus Alcock, 1900:329 [type-species: Lambdophallus 

sexpes Alcock, 1900, by monotypy; gender: masculine]. 

DIAGNOSIS.—Eyes small, immovable, cornea 

slightly broader than stalk. Third rnaxilliped 

(Figure 34A) broad, ischium slightly convex distally, 

carpus, propodus, and dactylus slender, subcylindrical, 

dactylus as long as propodus; flagellum 

of exopod not described. Pterygostomian 

region with row of oblique striae. Walking legs 

(Figure 34a) elongate, merus of longest pair longer 

than carapace. Transverse grooves on sternum 

(Figure 346) for reception of male pleopods present. 

Male pleopods (Figure 34b,c) with apices not 

concealed under abdomen, each bent at right 

angle, distal part of each lying in deep transverse 

groove on sternum, apices naked, with subterminal 

setae, directed laterally. Male abdomen 

(Figure 34b,c) with 5 segments, third to fifth fused, 

terminal segment rounded apically, smooth laterally. 

REMARKS.—This genus contains only the typespecies, 

L. sexpes Alcock. The fixed eyes, long 

pereiopods, and long, L-shaped male pleopods 

extending laterally into transverse grooves in the 

anterior part of the sternum are diagnostic for the 

genus. 

In L. sexpes the third and fourth legs are subequal 

in length, the male abdomen falls short of 

the level of insertion of the third maxillipeds, and 

the carapace is relatively broad (width = 1.6 

times length). 

Another species subsequently described in this 

genus, L. anfractus Rathbun, 1909, from Thailand, 

is transferred to Hexapus (p. 171). 

Genus Paeduma Rathbun, 1897 

FIGURE 35 

Amorphopus Bell, 1859:27 [invalid junior homonym of Amorphopus 

Audinet-Serville, 1838 (Insecta); type-species: Amorphopus 

cylindraceus Bell, 1859, by monotypy; gender: masculine]. 

Paeduma Rathbun, 1897b: 163 [substitute name for Amorphopus 

Bell, 1859; type-species: Amorphopus cylindraceus Bell, 

1859; gender: neuter]. 

DIAGNOSIS.—Eyes small, movable, cornea 

about as broad as stalk. Third maxillipeds slender, 

ischium convex mesially, propodus dilated 

distally, as broad as merus, longer than carpus or 

dactylus; exopod with flagellum. Pterygostomian 

region lacking oblique striae. Walking legs short, 

merus of longest pair (second) shorter than carapace. 

Sternal grooves absent. Apices of male pleopods 

(Figure 3bb) concealed under abdomen, 

pleopods slender, recurved posteriorly or doubly 

recurved into a figure 8, apices apparently naked 

or nearly so. Male abdomen (Figure 35a) with 

third and fourth and fifth and sixth somites fused, 

terminal somite short, subtriangular, rounded apically, 

not trilobed. 

INCLUDED SPECIES.—Amorphopus cylindraceus 

Bell, 1859, locality unknown, and Thaumastoplax 

orientalis Rathbun, 1909 and T. chuenensis Rathbun, 

1909, both from Thailand.

174 


FIGURE 34.—Lambdophallus sexpes Alcock: a, dorsal view; b, ventral view; c, male abdomen and 

apices of pleopods. (From "Illustrations of the Zoology of H. M. Indian Marine Surveying 

Steamer Investigator" Crustacea, 10 (1902), pi. 62: fig. 1-lb.) 

REMARKS.—The genus Amorphopus was recognized 

by Bell (1859:27) for a single species, A. 

cylindraceus, from an unknown locality. Bell's 

name Amorphopus is preoccupied, as pointed out 

by Rathbun (1897b: 163), who substituted Paeduma 

for it. As we noted above, with the exception 

of Gordon (1971:108) and Sankarankutty (1975: 

4), subsequent authors apparently have overlooked 

Paeduma. 

Paeduma has remained unidentifiable since its 

introduction by Bell; no subsequent workers on 

the group have been able to identify any species 

with it. As Gordon (1971:108) pointed out, one of 

the characters mentioned by Bell, the fusion of 

segments of the abdomen, is unusual (i.e., the 

third and fourth segments and the fifth and sixth 

segments are fused, rather than the third, fourth, 

and fifth, as we have found in all other genera of 

the family, save two: in Stevea, new genus, the 

second through the sixth segments are fused and 

in Hexaplax some segments are indistinctly fused). 

FIGURE 35.—Paeduma orientate 

(Rathbun), male: a, abdomen; 

b, pleopod. (Both from Rathbun, 

1910, fig. 33.) 

However, in her expanded account of Tkaumastoplaxorientalis 

Rathbun, 1909, Rathbun (1910:347) 

reported an identical configuration of the abdomen 

in that species, and Sakai (1939:579) mentioned 

it in material from Japan identified with 

T. orientalis. Further, Bell (1859:28) also had 

noted: "the abdomen very long and narrow, the 

first and second joint transversely linear, the third 

and fourth united and forming a triangle truncated 

anteriorly at the articulation of... the fifth 

and sixth joints united, and which with the seventh 

form a very narrow and linear piece extending 

forwards to the posterior margin of the oral 

aperture." This is precisely the condition figured 

by Rathbun (1910, fig. 33) (Figure 35a herein) for 

T. orientalis. In Rathbun's species, the fused third 

and fourth somites taper abruptly, forming a 

truncated triangle anteriorly, and the slender terminal 

sections of the abdomen extend forward to 

a point between the maxillipeds. Inasmuch as all 

other hexapodid genera, with the exception of 

Stevea and possibly Hexaplax, have the third to 

fifth somites of the male abdomen fused, we 

believe that Rathbun's species, T. orientalis, must 

be assigned to Paeduma. This genus also now 

includes P. chuenense (Rathbun, 1909), based on a 

juvenile male, and a species from Japan, identified 

by earlier workers with T. orientalis, which we 

believe represents a new species, the description 

of which is in preparation. Paeduma cyclindraceum 

(Bell), with its unequal chelipeds, remains un-


identifiable; the holotype, long thought to be lost, 

has been rediscovered in the Bell collection at the 

University Museum, Oxford, England, by Angelo 

A. DiMauro, Jr., of the University of Connecticut. 

His study of the Bell collection should allow us to 

be able to identify Bell's species. 

Paeduma resembles Thaumastoplax in having the 

propodus of the third maxillipeds dilated distally, 

in having the second walking legs strongest, and, 

in P. orientale at least, in lacking the oblique striae 

on the pterygostomian region. Paeduma differs 

from other hexapodid genera in having the third 

and fourth and the fifth and sixth somites of the 

abdomen fused in males. 

Genus Parahexapus Balss, 1922 

Parahexapus Balss, 1922:77 [type-species: Parahexapus africanus, 

Balss, 1922, by monotypy; gender: masculine]. 

Hexapus {Parahexapus).—Monod, 1956:362, 370. 


slightly broader than stalk. Third maxillipeds 

(Figure 36a) slender, pediform, ischium convex 

mesially, carpus, propodus and dactylus slender, 

cylindrical, dactylus longer than propodus; exopod 

with flagellum. Pterygostomian region with 

transverse granulate ridge. Walking legs short, 

merus of longest pair shorter than carapace. Sternal 

grooves absent. Apices of male pleopods concealed 

by abdomen, pleopods (Figure 36V) sinuous, 

slender, abruptly tapering distally to slender 

apex, directed anteriorly, with large spines 

near midlength directed proximally. Male abdomen 

(Figure 36b) with third to fifth somites fused, 

slender, subterminal and terminal somites long, 

latter rounded distally, not trilobed. 

INCLUDED SPECIES.—Parahexapus africanus Balss, 

1922. 

REMARKS.—Parahexapus shares slender third 

maxillipeds with Thaumastoplax, Spiroplax, and 

Paeduma; but in Parahexapus the propodus of the 

third maxilliped is very slender, not dilated distally. 

The first male pleopod (Figure 36V) of 

Parahexapus africanus differs from those of known 

species in the other three genera in having strong 

spines near midlength and in abruptly tapering 

distally. 

In P. africanus the interantennular septum is 

scarcely discernible in anterior view, the male 

abdomen extends anteriorly between the bases of 

the third maxillipeds, and the carapace is rather 

broad (width = 1.5-1.9 times length). Although 

the sutures in the abdomen of the single male 

available to us are very indistinct, it appears that 

the third to fifth somites are fused and the sixth 

somite is elongated, as figured by Crosnier (1967, 

fig. 33) (Figure 36A herein), as in Thaumastoplax 

anomalipes. 

Parahexapus africanus Balss, 1922 

FIGURE 36a-c 

Hexapus (Parahexapus) africanus.—Monod, 1956:370, figs. 494- 

496.—Rossignol, 1962:118.—Crosnier, 1967:337, figs. 30, 

33. 

Hexapus africanus.—Longhurst, 1958:88.—Gauld, 1960:71. 


Other Material: Congo: Baie de Pointe-Noire, A. Crosnier, 

26, 1$ (W). 


Figures: Monod, 1956, figs. 494-496; Crosnier, 

1967, figs. 30, 33. 

Male pleopod: Crosnier, 1967, fig. 30 (Congo). 

MEASUREMENTS.—The carapaces of our males 

measure 3.4 X 5.6 and 2.8 X 4.9 mm; that of the 

female measures 3.0 X 4.6 mm. Monod (1956) 

reported ovigerous females measuring 4X7 mm 

and 5 X 7.5 mm, and Crosnier (1967) reported 

males measuring 3.2 X 5 mm and 3.4 X 5.6 mm 

and a female as large as 3.4 X 5.6 mm. Balss 

(1922) reported that the female holotype measured 

4 X 7.5 mm. 

BIOLOGY.—Parahexapus africanus is a sublittoral 

species, which has been collected in depths between 

5 m and about 25 m (16-27 m; 22-25 m). 

Longhurst (1958) found the species off Sierra 

Leone in 16-27 m on shelly sand and sandy mud. 

Crosnier (1967) found it on sandy mud in 22- 

25 m off the mouth of the Congo River.


FIGURE 36.—Parahexapus africanus Balss: a, third maxilliped 

(from Monod, 1956, fig. 4%); b, abdomen of male; c, male 

pleopod (b,c, from Crosnier, 1967, figs. 30, 33). Pseudohexapus 

platydactylus Monod: d, abdomen of male; e, third maxilliped; 

/, male pleopod (d-f, from Monod, 1956, figs. 481, 484, 488). 

Ovigerous females have been collected in February 

(Monod, 1956). 


recorded from localities on the mainland between 

Sierra Leone and Cabinda, in depths between 

5 m and about 25 m (16-27 m is the deepest 

record). Records in the literature include the 

following: 


1958). 

Ghana: Off Accra, 5 m (Monod, 1956; Gauld, 1960). 

Congo: Baie de Pointe-Noire (Rossignol, 1962; Crosnier, 

1967). Off the mouth of the Congo River, 05°56'S, 12°O7'E, 

22-25 m (Crosnier, 1967). 

Cabinda: Landana, 6 m (Balss, 1922). 

Genus Pseudohexapus Monod, 1956 

Hexapus {Pseudohexapus) Monod, 1956:362, 365 [type-species: 

Hexapus (Pseudohexapus) platydactylus Monod, 1956, by 


DIAGNOSIS.—Eyes small, movable, cornea as 

broad as stalk. Third maxilliped (Figure 36e) 

broad, ischium short, mesial margin concave, projecting 

distally, carpus, propodus and dactylus 

slender, cylindrical, propodus longer than dactylus; 

exopod without flagellum. Pterygostomian 

region lacking oblique striae, with hairy ridge. 

Walking legs short, merus of longest pair (fourth) 

shorter than carapace. Dactyli of walking legs 

flattened dorsoventrally. Sternal grooves absent 

(?). Apices of male pleopods completely concealed 

by abdomen; pleopods (Figure 36/) stout, apices 

curved mesially, with terminal spinules and few 

subterminal setae. Male abdomen (Figure 36d) 

with 5 free somites, third to fifth fused, terminal 

somite elongate, rounded apically, not trilobed. 

INCLUDED SPECIES.—Pseudohexapus platydactylus 

Monod, 1956. 

REMARKS.—In Pseudohexapus the carapace is 

subquadrate (width = 1.2-1.3 times length), the 

pterygostomian region lacks a row of oblique 

striae, the propodus of the third maxilliped is 

slender and the exopod lacks a flagellum, the 

sternal grooves are absent, and the male pleopod 

is stout with its apex turned mesially. The flattened 

dactyli of the walking legs apparently are 

unique within the family. 

Pseudohexapus platydactylus Monod, 1956 

FIGURE 36d-f 

Hexapus (Pseudohexapus) platydactylus Monod, 1956:365, figs. 

478-493 [Senegal, Gambia, Ghana].—Gordon, 1971:107, 

fig. 2 [Ghana]. 

Hexapus platydactylus.—Gauld, 1960:71 [Ghana].—Longhurst, 

1958:88 [Sierra Leone]. 

DISTRIBUTION.—West Africa, from Senegal, 

Gambia, Sierra Leone, and Ghana, sublittoral, in 

5-15 m. 

Genus Spiroplax, new genus 

FIGURE 37 

TYPE-SPECIES.—Thaumastoplax spiralis Barnard, 

1950:301, fig. 56h-l, South Africa.

NUMBER 306 

FIGURE 37.—Spiroplax spiralis (Barnard): a, carapace; b, third 

maxilliped; c, sternum; d, abdomen of male; e, male pleopod. 

(All from Barnard, 1950, fig. 56h-l.) 

ETYMOLOGY.—From the Greek, speira (coil, 

twist), in combination with the noun plax (a 

plate), alluding to the form of the male pleopod; 

the gender of the name is feminine. 

DIAGNOSIS.—Eyes movable, cornea slightly 

broader than stalk. Third maxilliped (Figure 37£) 

slender, ischium with obtuse mesial projection, 

propodus shorter than dactylus, with mesial expansion 

distally; exopod flagellum not described. 

Pterygostomian region not described. Walking 

legs not described. Sternal grooves not described. 

Male pleopods (Figure 37*) apparently concealed 

under abdomen, strongly calcified, stout, spiral, 

apices setose. Male abdomen (Figure 37a 1 ) with 

third to fifth somites fused, very broad, terminal 

somite triangular, not trilobed. 

REMARKS.—The dilated propodus of the third 

maxillipeds suggests affinities with both Thaumastoplax 

and Paeduma, but in Spiroplax the carapace 

is much more oval (Figure 37a) (width = 1.3 

times length), the male abdomen is much 

broader, and the stout, spiral male pleopods are 

very different from the slender, almost filamentous 

male pleopods of representatives of the other 

two genera. Spiroplax includes only the typespecies. 

Genus Stevea, new genus 

FIGURE 38 

177 

TYPE-SPECIES.—Hexapus williamsi Glassell, 

1938:445, pi. 35, Guatemala (type locality) and 

Gulf of Tehuantepec, W coast of Mexico ($, 

9.4 X 14.4 mm, W). 

ETYMOLOGY.—The genus is named for Steve A. 

Glassell, the first carcinologist to record a hexapodid 

crab from the Americas; the gender of the 

generic name is feminine. 

DIAGNOSIS.—Carapace suboval (Figure 38a). 

Eyes small, movable, cornea subglobular, narrower 

than stalk. Third maxilliped (Figure 38c) 

broad, ischium flattened or slightly concave distally, 

carpus, propodus and dactylus slender, subcylindrical, 

dactylus longer than propodus; exopod 

with flagellum. Pterygostomian region with 

row of oblique striae. Walking legs short, merus 

of longest pair (third) shorter than carapace. Sternal 

grooves absent. Apices of male pleopods completely 

concealed by abdomen; structure of male 

pleopod not described. Male abdomen (Figure 

3&/) with three free somites, second through sixth 

fused, terminal somite triangular, not trilobed. 

REMARKS.—In the only species assigned to this 

genus, Stevea williamsi (Glasssell, 1938) (incorrectly 

spelled Hexapus williamsoni, in Stephensen, 

1945:182), there is a distinct median lobe on the 

front, visible in anterior view, the interantennular 

septum is poorly developed, the male abdomen 

extends beyond the bases of the third maxillipeds, 

and the carapace is relatively broad (width =1.5 

times length). 

The presence of only three free somites in the 

male abdomen will distinguish this genus from 

all others in the family. 

Genus Thaumastoplax Miers, 1881 

Thaumastoplax Miers, 188la:261 [type-species: Thaumastoplax 

anomalipes Miers, 1881, by monotypy; gender: feminine]. 


rounded, only slightly broader than stalk. Third 

maxilliped (Figure 39a) slender, ischium convex

178 

a 


FIGURE 38.—Stevea williamsi (Glassell): a, dorsal view; b, front; c, third maxilliped; d, ventral 

view. (All from Glassell, 1938, pi. 35.) 

mesially, propodus strongly dilated distally, almost 

as broad as merus, longer than carpus or 

dactylus; exopod with flagellum. Pterygostomian 

region lacking row of oblique striae. Walking legs 

short, merus of longest pair (second) shorter than 

carapace. Sternal grooves absent. Male pleopods 

(Figure 39c) slender, sinuous, apices directed anteriorly, 

completely concealed under abdomen, 

armed with minute spinules. Male abdomen (Figure 

396) with third to fifth somites fused, terminal 

somite elongate, sinuous laterally, rounded apically, 

not trilobed. 

INCLUDED SPECIES.—Thaumastoplax anomalipes 

Miers, 1881, from West Africa. Two other species 

assigned to this genus, T. orientalis Rathbun, 1909, 

and T. chuenesis Rathbun, 1909, are here transferred 

to Paeduma (p. 173). The fourth species 

assigned to this genus, T. spiralis Barnard, 1950, 

is transferred to Spiroplax (p. 176). 

REMARKS.—Several features appear to be diagnostic 

for this genus, now known to include 

only a single species from tropical West Africa. 

The third maxilliped is slender and the propodus 

is strongly dilated distally, being almost as broad 

as the merus; the pterygostomian region is not 

ornamented with oblique striae; the second pair 

of walking legs is much the strongest of the three; 

the interantennular septum is very low, scarcely 

visible in frontal view; and the male pleopod is 

slender, sinuous, and directed anteriorly. 

Two other genera, Paeduma and Spiroplax, share 

the maxilliped with the distally dilated propodus, 

but both can be differentiated readily from Thaumastoplax. 

In Paeduma the third and fourth and 

fifth and sixth, rather than the third to fifth, 

abdominal somites in the male are fused, and the 

male pleopods are strongly recurved posteriorly


or into a figure 8. In Spiroplax the male pleopods 

are much stouter and are twisted into a spiral. 

Thaumastoplax anomalipes Miers, 1881 

FIGURE 39 

Thaumastoplax anomalipes Miers, 188la:261, pi. 14: fig. 2. 

Hexapus (Thaumastoplax) anomalipes.—Monod, 1956:363, figs. 

471_477._Rossignol, 1962:118.—Gordon, 1971:108 [discussion]. 

Hexapus anomalipes.—Longhurst, 1958:88.—Gauld, 1960:71. 


Other Material: Congo: Baie de Pointe-Noire, 7-10 m, A. 

Crosnier, 1

180 

posteriorly on carpus and propodus; inner surface 

of carpus and propodus also with some strong 

tubercles; dactylus slender, curved ventrajly, 

shorter than propodus; latter broad, length only 

1.6 times greatest depth; propodus shorter than 

carpus, merus about 0.8 times as long as carapace, 

0.75 times as long as carpus and propodus combined. 

Third walking leg slightly stouter than 

first; dactylus straight or slightly curved ventrally, 

more than half as long as propodus; latter 1.4 

times as long as broad, shorter than carpus; merus 

slender, 2.7 times as long as broad, 0.6 times as 

long as carapace, as long as carpus and propodus 

combined. 

Male abdomen narrow, composed of 5 free 

somites, third to fifth fused, about as long as sixth 

somite. Latter appearing slender, about 1.3 times 

as long as broad. Terminal somite as long as 

broad, 0.6 times as long as fused somites, lateral 

margins sinuous, apex rounded. Abdominal 

trench in male extending anteriorly to base of 

third maxillipeds. Abdomen of female with 7 free 

somites. 

Male pleopod as figured by Monod (1956, fig. 

476), slender, sinuous, elongate, almost naked, 

completely concealed under abdomen. 


Male Pleopod: Monod, 1956, fig 476 (Senegal). 

MEASUREMENTS.—Our male specimen measures 

3.9 X 5.7 mm, the female 4.8 X 8.0 mm. 

The largest specimen studied by Monod was an 

ovigerous female 11X19 mm. 

BIOLOGY.—Thaumastoplax anomalipes is a sublittoral 

species, living in depths between about 3 m 

and 34 m. The specimens recorded by Monod 

(1956) were taken on mud-sand in 19 m, on shelly 

sand in 8 m, and on muddy sand with gorgonians 

in 10 m. Longhurst (1958) found the species in 

estuaries as well as offshore in depths between 6 

and 34 m; in the estuary the species occurred on 

muddy sand and shelly mud, whereas offshore it 

was taken on shelly sand and shelly mud. 

DISTRIBUTION.— West coast of Africa, from a 

few localities between Senegal and Rio Muni; 

sublittoral, in depths between ca 3 m and 34 m. 

Monod (1956) recorded material from Senegal 


and Guinea; since 1956 the species has been 



1958). 

Ghana: Off Accra, ca. 3 m (Gauld, 1960). 


Genus Tritoplax, new genus 

FIGURE 40 

TYPE-SPECIES.—Hexapus stebbingi Barnard, 

1947. 

ETYMOLOGY.—The feminine name is derived 

from the Greek, tritos (a third) and plax (a plate), 

alluding to the shape of the terminal somite of 

the male abdomen. 

DIAGNOSIS.—Eyes very small, cornea subglobular. 

Third maxilliped with broad ischium and 

merus, ischium concave proximally, straight distally, 

carpus, propodus, and dactylus slender, 

subcylindrical; exopod of flagellum not described. 

Pterygostomian region apparently lacking 

oblique striae. Walking legs short, merus of longest 

pair shorter than carapace. Sternal grooves 

restricted to trilobed anterior terminus of abdominal 

fossa, grooves not extending anterolaterally 

to bases of third maxillipeds. Male pleopods (Figure 

406) slender, sinuous, bent laterally, naked 

apex recurved anteriorly. Male abdomen (Figure 

40a) with third to fifth somites fused, sixth longitudinally 

divided in type-species, terminal 

somite distinctly trilobed, with angled lateral 

projections. 

INCLUDED SPECIES.—Hexapus stebbingi Barnard, 

1947 (see Barnard, 1950:299), from South Africa. 

An unnamed species identified by Stephensen 

(1945, fig. 53B; Figure AQd herein) as Hexapus 

sexpes, which also has a distinctly trilobed abdomen, 

although the sixth somite is not longitudinally 

divided, may also belong here. 

REMARKS.—The trilobed male abdomen and 

the longitudinally divided sixth abdominal somite 

in the male are both quite characteristic and 

will serve to distinguish this genus from others in 

the family. The trilobed shape of the anterior end


of the abdominal fossa is similar to that found in 

species of Hexapinus, but the terminal somite of 

the male abdomen and the shape of the male 

pleopod are different in the two genera. 

FIGURE 40.—Tritoplax stebbingi (Barnard): a, abdomen of 

male; b, male pleopod; c, apex; d, abdomen and apex of 

pleopod of male (a-c, from Barnard, 1950, fig. 56*^/"; d, from 

Stephensen, 1945, fig. 53b). 

Family PINNOTHERIDAE de Haan, 1833 

PINNOTHERIDEA de Haan, 1833:2, 5 [corrected to Pinnotheridae 

by Bell, 1845:119]. 

ASTHENOGNATHIDAE Stimpson, 1858b: 107. 

XENOPHTHALMIDAE Stimpson, 1858b: 107. 

DISSODACTYLIDAE Smith, 1870:172. 

PINNOTHERELINAE Alcock, 1900:294, 335. 

ANOMALOFRONTINAE Rathbun, 1931:84. 

EASTERN ATLANTIC GENERA.—Two, Asthenognathus 

and Pinnotheres, both represented by species 

living off tropical West Africa. 

EASTERN ATLANTIC SPECIES.—Fourteen, ten of 

vvhich are named; some are of doubtful taxonomic 

standing. 

Schmitt, McCain, and Davidson (1973) listed 

2 genera and 10 species of Pinnotheridae from 

West Africa: Asthenognathus atlanticus Monod, Pinnotheres 

pinnotheres (Linnaeus), Pinnotheres pisum 

(Linnaeus), Pinnotheres leloeuffi Crosnier, Pinnotheres 

mccainae Schmitt, Pinnotheres sp. A, B, C, D, of 

Monod (1956) and Pinnotheres sp. of Guinot and 

Ribeiro (1962). Pinnotheres sp. C of Monod is 

described herein as a new species, Pinnotheres conicola. 

Pinnotheres tellinae, which also is described 

here as new, includes in part Monod's species A. 

The only West African records of Pinnotheres 

pinnotheres are those by Balss (1922). Unfortunately, 

Balss did not give any details of his material 

that would confirm the correctness of his 

identification, and it seems likely that his specimens, 

which were collected in Cameroon and 

Gabon, actually belong to one or more of the true 

West African species that were described since 

1922. Until Balss' material can be reexamined, 

Pinnotheres pinnotheres has to be included in the list 

of West African species, be it with great reserve. 

Also the otherwise European species, P. pisum 

(Linnaeus), has been recorded from Morocco and 

Mauritania. At present, therefore, six named species 

of Pinnotheres are known from West Africa (P. 

leloeuffi, P. mccainae, P. conicola, P. tellinae, P. pisum, 

P. pinnotheres; the last doubtfully), and four unnamed 

species (species A (part), B, and D of 

Monod, 1956, and Pinnotheres sp. of Guinot and 


The pinnotherids were not represented in the 

Pillsbury collections. The following species of pinnotherids 

are extralimital: 

Pinnotheres ascidicola Hesse, 1872. Atlantic coast 

of France; in ascidians. 

Pinnotheres marioni Gourret, 1887. Mediterranean 

(Bay of Marseilles); in ascidians. 

Pinnotheres pectunculi Hesse, 1872. Atlantic coast 

of Brittany (France); in Glycimeris (Lamellibranchia). 

Subfamily ASTHENOGNATHINAE Stimpson, 

1858 

Genus Asthenognathus Stimpson, 1858 

Asthenognathus Stimpson, 1858b: 107 [p. 53 in separate; typespecies: 

Asthenognathus inaequipes Stimpson, 1858, by 

monotypy; gender: masculine; name 287 on Official List]. 

Asthenognathus atlanticus Monod, 1933 

Asthenognathus atlanticus.—Monod, 1956:383, figs. 541-545 

[Mauritania, Senegal, Ghana].—Gauld, 1960:71


[Ghana].—Guinot and Ribeiro, 1962:65 [Angola].—Rossignol, 

1962:119 [Congo].—Monod, 1963: fig. 39 [no material].—Zariquiey 

Alvarez, 1968:410, fig. 137 [Spain; 

references].—Schmitt, McCain, and Davidson, 1973:128 

[synonymy].—Noel and Amouroux, 1977:135 [Mediterranean]. 

DISTRIBUTION.—Eastern Atlantic, from Normandy, 

NW France to Angola and Mediterranean; 

on echinoids and commensal with polychaetes; 

sublittoral, in depths between 8 and 70 

m. 

Subfamily PINNOTHERINAE de Haan, 1833 

Genus Pinnotheres Bosc, 1802 

Pinnotheres Bosc, 1802:59, 239 [type-species: Cancerpisum Linnaeus, 

1767, by selection by Latreille, 1810:422; gender: 


Holothunophilus Nauck, 1880:66 [type-species: Holothuriophilus 

trapeziformis Nauck, 1880, by original designation; gender; 


Arcotheres Burger, 1895:361 [type-species: Pinnotherespalaensis 

Burger, 1895, by selection by Rathbun, 1918:62; gender: 

masculine]. 

Zaops Rathbun, 1900b:588, 590 [type-species: Pinnotheres depressum 

Say, 1817, a subjective junior synonym of Pinnotheres 

ostreum Say, 1817, by original designation and monotypy; 


Pinnozoea Aikawa, 1933:130, 246 [type-species: Cancer pisum 

Linnaeus, 1767, by selection by Schmitt, McCain, and 

Davidson, 1973:37; gender: feminine]. 

Pinnotheres conicola, new species 


Pinnoteres sp. C—Monod, 1956:380, figs. 508, 509, 526-538 

[not figs. 524, 525 = Pinnotheres sp. B].—Longhurst, 1958: 

88. 

Pinnotheres sp. C.—Silas and Alagarswami, 1967:1214 [listed].—Schmitt, 

McCain, and Davidson, 1973:92 [synonymy]. 


Other Material: Cameroon: Kribi, in large Conus obtained 

by native fishermen who fished with a canoe ("pirogue") 

near the shore, 10 Mar 1964, B. de Wilde-Duyfjes, holotype, 

Id (L). 

DESCRIPTION.—Carapace (Figure 41) firm, almost 

circular in outline, about 1.1 to 1.3 times 

wider than long. Surface of carapace naked, 

smooth, except for coarsely pitted branchial regions, 

slightly convex dorsally, falling off more 

steeply laterally; anterolateral margin broadly 

and evenly rounded. Lateral surface with long, 

soft hairs, which, with similar hairs on pereiopods, 

enclose naked upper surface in hirsute ring. Front 

almost square, slightly produced in middle, in 

dorsal view anterior margin scarcely produced 

beyond circular outline of carapace. Eyes small, 

short, with reduced cornea; orbit subcircular. 

Third maxilliped (Figure 42a) placed 

obliquely, almost transversely. Merus operculiform, 

very wide, 2/3 as wide as long, greatest 

width in anterior half, anterointernal angle 

evenly rounded. Carpus short. Propodus oval. 

Dactylus very short, narrow, inserted on posterior 

half of lower margin of propodus, falling short of 

apex of propodus. Carpus and propodus together 

less than half as long as merus. 

Chelipeds (Figure 42b-d) well developed, fingers 

about 6/7 as long as palm. Dactylus with 

single large tooth on basal half of cutting edge, 

distal half with some small granular denticles. 

Cutting edge of fixed finger also with large tooth, 

placed before that of dactylus, continued posteriorly 

as crenulated ridge. Base of fingers, upper 

surface of dactylus, lower surface of fixed finger, 

and cutting edges densely covered with soft, slender, 

plumose hairs. Palm with upper half of outer 

surface pilose, lower half naked; inner surface 

completely covered with long, smooth hairs except 

for naked spot anteriorly near base of fingers. 

Carpus short, rounded, upper surface naked, remainder 

covered by long hairs. Merus with upper 

and outer part naked, remainder clothed with 

long hairs. Ischium with distinct lobe on anterointernal 

angle of upper margin. 

Walking legs (Figure 42e-h) subsimilar, all 

rather robust, none longer or slenderer than others. 

Dactylus of each walking leg similar to that 

of other legs, about 2/3 as long as propodus 

(measured dorsally), short, simple, terminating in

NUMBER 306 

narrow, slightly curved tip. Propodus, carpus, 

and merus each high, with thick coat of hairs 

dorsally and ventrally but with lateral surfaces 

bare. Propodus about twice as long as high, merus 

somewhat more slender. 

Male abdomen (Figure 42*) elongate, with 

fifth and sixth somites fused, line of separation 

scarcely distinct but present; Monod (1956:378, 

fig. 508) showed 7 free somites on male abdomen. 

Telson triangularly rounded. 

Male Pleopod: Monod, 1956:382, figs. 537, 538. 

MEASUREMENTS.—The holotype has a carapace 

length of 10 mm and a carapace width of 11.5 

mm. The two male specimens reported upon by 

Monod had cl 9 and 9.5 mm, and cb 11 and 10 

mm, respectively; his ovigerous female had cl 

15 mm, cb 18 mm, and a juvenile female cl 8 and 

cb 9 mm. 

REMARKS.—In Monod's (1956) fundamental 

work on the crabs of West Africa, his figures 508 

and 509 are labeled Pinnotheres sp. B, while figures 

524 and 525 are labeled Pinnotheres sp. C. However, 

a comparison of these figures with the measurements 

of these specimens given in the text 

and with the details shown in his figures 510 to 

523 and 526 to 538, indicate that the explanation 

said to be of his figures 524 and 525 actually is 

that of his figures 508 and 509 and vice-versa. 

Judging by the enlargement indicated, the male 

and female of Monod's figures 508 and 509 have 

FIGURE 41.—Pinnotheres conicola, new species: a, male; b, female. 

(From Monod, 1956, figs. 524, 525.) 

183 

cb 11 and 14 and cl 9 and 11 mm, respectively, 

which agrees with the measurements given for 

Pinnotheres sp. C from Conakry; the male and 

female of Monod's figures 524 and 525 have, 

respectively, cb 20 and 20 mm, and cl 16 and 

18 mm, which agree well with the specimen of 

Pinnotheres sp. B from Goree. There can, therefore, 

be little doubt that Monod's figures 508, 509, 

526-538 show Pinnotheres sp. C, and figures 510- 

525 show Pinnotheres sp. B. Monod's (1956:382, 

figs. 539, 540) Pinnotheres sp. D shows a great 

resemblance to the present species, and might 

well be specifically the same. As Monod did not 

provide a description of his species D, this problem 

can only be solved by an examination of the 

material in question. 

The short stubby legs fringed with heavy rows 

of setae distinguish this species immediately from 

most other named species of this genus known 

from the eastern Atlantic: Pinnotheres pinnotheres 

(Linnaeus), P. pisum (Linnaeus), and P. mccainae 

Schmitt. From P. leloeuffi Crosnier it can be distinguished 

by the less wide carapace, by the 

dorsal and ventral fringes of hair on all the segments 

of the legs except the dactylus, the wider 

merus of the third maxilliped and the shape of 

the male abdomen. Actually it seems closest to 

Monod's (1956) Pinnotheres sp. D with which it 

may prove to be identical. In its pubescence the 

new species shows some resemblance to Pinnotheres

184 


FIGURE 42.—Pinnotheres conicola, new species: a, third maxilliped; b-d, chelipeds; e-h, second to 

fifth pereiopods; i, abdomen of holotype, male, cl. 10.0 mm, Kribi (a-h from Monod, 1956, figs. 

526-532). 

barbatus Desbonne from the West Indies, and also 

resembles that species in the robust walking legs, 

which are of similar size and shape and which 

have all dactyli short and falcate. It differs 

strongly, however, in the shape of the third maxilliped 

and the male abdomen. In P. barbatus the 

dactylus of the third maxilliped is implanted at 

the tip of the propodus, and the male abdomen 

has all somites free. Furthermore the carapace of 

the American species is softer and shows a large 

pit on either side of the middle (see Rathbun, 

1918:88, pi. 19: figs. 8-11). 

TYPE-LOCALITY.—Kribi, Cameroon. 

DISPOSITION OF TYPE.—The holotype (Crust. 

D. 28792) is in the collection of the Rijksmuseum 

van Natuurlijke Historie, Leiden. 

The specimens reported by Monod (1956) from 

Conakry are probably in the collection of the 

Institut Fondamental d'Afrique Noire, Dakar, 

Senegal. The specimen reported by Longhurst 

(1958) may be in the collection of the British 

Museum (Natural History), London. 

ETYMOLOGY.—The specific epithet is derived 

from the Latin generic name Conus, referring to 

the association of the species with a species of 

that genus. 

BIOLOGY.—All specimens of this species known 

so far were found to live commensally with gastropod 

mollusks of the genus Conns; only the host 

of the Sierra Leone specimens is identified to 

species, viz. Conus papillionaceus Hwass. The Cameroon 

specimen was said to be from a large 

Conus. The depths at which the specimens were 

found is not accurately known: Longhurst (1958) 

gave the depth at which his specimens were taken 

as "shallow" and as "0-40 m." The Cameroon 

specimen evidently also was taken in shallow 

water as it was obtained from native fishermen 

fishing near the coast in a canoe. 

One of the specimens reported from Sierra 

Leone was an ovigerous female; the date of collection 

was not recorded. 


recorded from Guinea, Sierra Leone, and now


Cameroon; our specimen represents a considerable 

range extension to the south. Records in the 

literature include the following: 

Guinea: Conakry (Monod, 1956). 

Sierra Leone: No specific locality (Monod, 1956; 

Longhurst, 1958). 

Pinnotheres leloeuffl Crosnier, 1969 

Pinnotheres leloeuffi Crosnier, 1969:531, figs. 1-10, 17 [Ivory 

Coast].—Schmitt, McCain, and Davidson, 1973:52 [synonymy]. 

DISTRIBUTION.—West Africa, from Vridi, Ivory 

Coast, in 20 m. Host unknown. 

Pinnotheres mccainae Schmitt, 1973 

FIGURE 43 

Pinnotheres rouxi Rossignol, 1957:84, fig. 4.—Silas and Alagarswami, 

1967:1208 [listed].—Schmitt, McCain, and 

Davidson, 1973:2, 57 [synonymy]. [Preoccupied by Pinnotheres 

rouxi H. Milne Edwards, 1837.] 

Pinnoteres rouxi.—Rossignol, 1962:118. 

Pinnotheres mccainae Schmitt, in Schmitt, McCain, and Davidson, 

1973:2,5, 11,57. 


Other Material: Congo: Pointe-Noire, from mantle cavity 

of Donax rugosus Linnaeus, 25 Apr 1955, A. G. Humes, 10$ 

ov (L, W). 

DESCRIPTION.—Carapace quadrangular anteriorly, 

semicircular posteriorly. Anterolateral angles 

rounded, about as wide as front, each occupying 

about 1/3 of length of anterior margin of 

carapace. Lateral margins of carapace diverging 

posteriorly toward posterolateral angles. Carapace 

with surface naked, somewhat convex, integument 

thin. Front slightly convex in dorsal 

view, extending anteriorly to level of anterolateral 

angles; front bent strongly downward, coarsely 

pitted, at each anterolateral angle. 

Third maxilliped (Figure 43a,b) placed very 

obliquely, almost transversely. Dactylus narrow, 

elongated, articulated just before middle of lower 

margin of propodus, extending almost to end of 

propodus. Merus about twice as long as palp and 

about twice as long as broad. Exopod almost 

entirely covered by merus, basal segment rather 

broad, more than twice as long as wide. Distal 2 

segments of exopod narrow, together somewhat 

more than half as long as first segment. 

Cheliped of female (Figure 43c) elongate, chela 

more than 3 times as long as high. Fingers about 

3/4 as long as palm, tips sharp, curved. Basal 

part of cutting edges of both fingers with distinct 

tooth, that on fixed finger proximal to that on 

dactylus. Palm twice as long as high. Chela with 

fringe of long hairs on lower part of inner surface 

extending over all of fixed finger and distal part 

of palm. Dactylus with hairs scattered over surface. 

Row of hairs present on upper inner surface 

of carpus, merus with scattered hairs on inner 

surface. 

Pereiopods (Figure 43d-g) similar, of same 

length, on both sides of body. Third leg longest, 

about twice as long as fifth, \ x h times as long as 

second or fourth. Dactyli of second and fifth legs 

shortest, that of fourth slightly longer, that of 

third about \ l h. times length of shortest dactyli. 

In all legs, dactylus terminating in hook-shaped 

claw. Dactylus of second leg slightly more than 

1/3 as long as propodus, half as long as carpus; 

merus twice as long as carpus. Dactylus of third 

leg 2/5 as long as propodus, half as long as carpus; 

merus less than twice as long as carpus. Dactylus 

of fourth leg 2/5 as long as propodus, 2/3 as long 

as carpus, 1/3 as long as merus. Fifth pereiopod 

shortest, stoutest; dactylus 2/5 as long as propodus, 

2/3 as long as carpus, latter 3/4 as long as 

merus. Pereiopods not markedly hairy, only some 

scattered hairs present, densest proximally. 

Abdomen of ovigerous females distinctly wider 

than carapace. 

Eggs numerous, small, spherical, diameter 

about 0.2 mm. 

Figures: Rossignol (1957) provided figures of 

the general shape of an ovigerous female, a rather 

poor figure of the third maxilliped, and one of 

the chela. Some additional sketches are provided 

here (Figure 43). 

Male Pleopod: The shape of the male pleopod is 

unknown. The single type-specimen is a female, 

and our material consists entirely of females. Rossignol 

(1962) evidently had males at his disposal, 

as he reported that many of the Donax specimens

186 

a 


FIGURE 43.—Pinnotheres mccainae Schmitt, ovigerous female, cl 5.3 mm, Pointe-Noire: a,b, third 

maxilliped; c, chela; d, second pereiopod; e, third pereipod; /, fourth pereiopod; g, fifth 

pereiopod. 

collected by him were "habitee par un couple de 

cette espece"; however, he gave no details of the 

male specimens. 

MEASUREMENTS.—Our ovigerous females have 

carapace lengths varying from 4 to 6.5 mm, and 

carapace widths from 5.5 to 7 mm. The holotype, 

also an ovigerous female had the carapace 6 mm 

long and 7 mm wide. The diameter of the eggs is 

about 0.2 mm. 

REMARKS.—The species was first described under 

the name Pinnotheres rouxi by Rossignol (1957) 

based on a single specimen. Silas and Alagarswami 

(1967) correctly pointed out that the name 

Pinnotheres rouxi had already been used by H. 

Milne Edwards (1837) for a different (Indo-West 

Pacific) species and that therefore the West African 

species needed a new name. It was Schmitt 

(in Schmitt, McCain, and Davidson, 1973), who 

provided this new name, Pinnotheres mccainae. 

BIOLOGY.—All the specimens so far mentioned 

in the literature (Rossignol, 1957, 1962) were 

found to be commensals of Donax sp. The present 

specimens were found in the mantle cavity of 

Donax rugosus Linnaeus. Rossignol (1962) indicated 

that a large proportion of the Donax found 

at Pointe-Noire was infested with this commensal;


the bivalves were taken sublittorally in the sand 

at the lower limit of the lowest low tides. 

Rossignol's (1957) ovigerous female was collected 

in November, ours in April. 

DISTRIBUTION.—So far the species is only known 

from the area of Pointe-Noire, Congo. The holotype 

was reported from the beach at Pointe-Noire 

(Rossignol, 1957); additional material was mentioned 

by Rossignol (1962) from the Baie de 

Pointe-Noire. Also the present material is from 

that locality. 

Pinnotheres pinnotheres (Linnaeus, 1758) 

Pinnoteres pinnoteres.—Balss, 1922:79 [Cameroon, Gabon]. 

Pinnoteres pinnotheres.—Monod, 1956:376 [references]. 

Pinnotheres pinotheres.—Zariquiey Alvarez, 1968:409, fig. 

136b,f [Spain; references]. 

Pinnotheres pinnotheres.—Schmitt, McCain, and Davidson, 

1973:68 [synonymy]. 

SYNONYMS.—Pinnotheres veterum Bosc, 1802; Pinnotheres 

pinnae Leach, 1814; Pinnotheres montagui 

Leach, 1815; Pinnotheres pinnophylax H. Milne Edwards, 

1853. 

DISTRIBUTION.—North Sea to Mediterranean; 

possibly Gulf of Guinea; commensal in bivalves 

and ascidians. 

Pinnotheres pisum (Linnaeus, 1767) 

Cancer Pisum Linnaeus, 1767:1039. 

Pinnotheres pisum.—Capart, 1951:175 [Mauritania].—Forest 

and Games, 1960:353 [Morocco].—Zariquiey Alvarez, 

1968:408, figs. 7b, 14f, 135c,d, 136a,c-e [Spain; references].—Christiansen, 

1969:88, figs. 36, 37, map 30 [Scandinavia].—Schmitt, 

McCain, and Davidson, 1973:72 [synonymy]. 

Pinnoteres pisum.—Monod, 1956:375 [references]. 

SYNONYMS.—Cancer nutrix Scopoli, 1763; Cancer 

mytilorum albus Herbst, 1783; Cancer mytilorum juscus 

Herbst, 1783; Cancer varians Olivier, 1791; Pinnotheres 

mytili Leach, 1814; Pinnotheres modioli Leach, 

1814; Pinnotheres cranchii Leach, 1814; Pinnotheres 

latreillii Leach, 1817; Cancer eubolinus Nardo, 1847. 


southward to Port Etienne, Mauritania; Mediter- 

ranean; commensal in bivalves and possibly ascidians; 

intertidal to about 45 m. 

Pinnotheres tellinae, new species 


Pinnoteres sp. A.—Monod, 1956:376 [part; not figs. 502- 

507].—Longhurst, 1958:88. 

Pinnotheres sp. A.—Silas and Alagarswami, 1967:1213 

[part].—Schmitt, McCain, and Davidson, 1973:91 [synonymy; 

part]. 


Other Material: Sierra Leone: Kissy, near Freetown, from 

mantle cavity of Tellina nymphalis Lamarck, 29 Nov 1954, A. 

G. Humes, 16, 109 (holotype; 2 ov) (L, W). Bullom Shore, 

opposite Freetown, from mantle cavity of Tellina nymphalis 

Lamarck, 12 Nov 1954, A. G. Humes, 11

188 


FIGURE 44.—Pinnotheres tellinae, new species, paratypes, Kissy: a, adult female, cl 4.7 mm; 

b, adult male, cl 3.0 mm. 

tooth of fixed finger placed before that of dactylus. 

Remainder of cutting edges of chela with 

widely spaced tubercles; subdistally, cutting 

edges with short ridge, upper margin minutely 

tuberculate. Upper margin of chela with fringe 

of long hairs, latter also present on lower part of 

inner surface of palm and on fingers; shorter hairs 

present on remainder of chela. No tubercles present 

on either surface of palm. Carpus short, conical, 

slightly shorter than merus. Carpus and 

merus both with long hairs on dorsal and lower 

inner surface. 

Pereiopods of right side of equal size and shape 

as those on left side. First 3 walking legs (pereiopods 

2-4) (Figures 45


a 

FIGURE 45.—Pinnotheres tellinae, new species, paratypes, Kissy. Female, cl 5.9 mm: a, front; b, 

third maxilliped; c, cheliped; d, second pereiopod; e, third pereiopod; /, fourth pereiopod; g, 

fifth pereiopod. Male, cl 2.5 mm: h, chela; i, second pereiopod; j, third pereiopod; k, fifth 

pereiopod; /, abdomen; m, gonopod.

190 

his single ovigerous female as 5.5 mm. The diameter 

of the eggs is 0.2 to 0.3 mm. 

REMARKS.—The species is characterized by the 

almost equal walking legs that have the dactylus 

of equal length, by the third maxilliped that has 

the dactylus implanted on the lower margin of 

the propodus, reaching slightly beyond the top of 

the propodus, by the hairiness of the legs, and by 

the naked carapace. It resembles Pinnotheres pholadis 

de Haan from Japan, which, however, has 

more slender and hairless legs. The hairy legs of 

the female P. tellinae distinguish it from the West 

African species P. mccainae and P. pinnotheres; in P. 

leloeuffi the carapace is much wider and the shape 

of the third maxilliped and that of the male 

abdomen are quite different. 

Monod (1956:376) referred an ovigerous female 

of a Pinnotheres found in Tellina sp. in Sierra Leone, 

to his Pinnotheres sp. A. The other specimens referred 

by Monod to his species A were commensals 

of Ostrea tulipa Lamarck (= Ostrea gasar) from 

Senegal. Monod's (1956, figs. 502-507) illustrations 

are made after the Ostrea commensals, and 

show a species that is clearly different from P. 

tellinae. In the Ostrea commensals the carapace is 

wider in the females (length 9 mm, width 10 mm) 

and narrower in the males (length and width 

2 mm), the pereiopods are more slender and 

without long hairs, the shapes of the dactylus and 

propodus of the third maxilliped are different, 

and the size of the females is much larger. Although 

it is highly likely that Monod's specimen 

from Sierra Leone belongs to P. tellinae, as it was 

taken at the type-locality and from the type-host 

of that species, it is certain that the specimens 

from the host Ostrea tulipa belong to a different 

species. Monod (1956:376) himself remarked that 

"la conspecificite de cet exemplaire [i.e., the female 

from Sierra Leone] avec les precedents [i.e., 

those from Senegal] n'est pas certaine." 

TYPE-LOCALITY.—Kissy, near Freetown, Sierra 

Leone. 

DISPOSITION OF TYPES.—The holotype (USNM 

169498) is one of the females from Kissy; all of 

the other specimens are paratypes. The holotype 

is deposited in the National Museum of Natural 



D.C. The paratypes are in the same museum and 

duplicates are placed in the Rijksmuseum van 

Natuurlijke Histoire, Leiden. The specimen mentioned 

by Monod (1956:376) and Longhurst 

(1958:88), also a paratype, most likely is in the 

British Museum (Natural History). 

ETYMOLOGY.—The specific epithet is from the 

generic name Tellina, referring to the association 

of this species with a species of that genus. 

BIOLOGY.—Monod (1956) reported the species 

from Tellina sp. His specimen was collected by A. 

Longhurst in Sierra Leone. Longhurst (1958:88) 

himself gave more detailed information of what 

evidently is the same specimen, indicating that 

his material was found in the mantle cavity of 

Tellina nymphalis Lamarck, taken in an estuarine 

habitat. The specimens dealt with in the present 

paper were obtained by Dr. A. G. Humes from 

the mantle cavity of the same host. 

The date of collecting of Monod's and Longhurst's 

ovigerous female is not known. Our ovigerous 

females were collected in November. 

DISTRIBUTION.—So far the species is only known 

from Sierra Leone (Monod, 1956; Longhurst, 

1958). 

Pinnotheres sp. A 

Pinnoteres sp. A.—Monod, 1956:376, figs. 502-507 [part; 

Senegal]. 

Pinnotheres sp. A.—Silas and Alagarswami, 1967:1213 

[listed].—Schmitt, McCain, and Davidson, 1973:91 

[synonymy]. 

DISTRIBUTION.—Senegal. Commensal of Ostraea 

gasar (Adanson) (= Ostrea tulipa Lamarck) 

(Monod, 1956). 

Pinnotheres sp. B 

Pinnoteres sp. B.—Monod, 1956:378, figs. 510-525 [Senegal]. 

Pinnotheres sp. B.—Silas and Alagarswami, 1967:1213 

[listed].—Schmitt, McCain and Davidson, 1973:91 [synonymy]. 

DISTRIBUTION.—Senegal, in Panopea aldrovandi 

Menard; sublittoral, to at least 5 m.


Pinnotheres sp. D 

Pinnoteres sp. D — Monod, 1956:382, figs. 539, 540 

[Congo].—Rossignol, 1962:118 [Congo]. 

Pinnotheres sp. D.—Silas and Alagarswami, 1967:1214 

[listed].—Schmitt, McCain and Davidson, 1973:92 [synonymy]. 

DISTRIBUTION.—Pointe-Noire, Congo, host and 

depth range unknown. 

Pinnotheres sp. 

Pinnotheres sp.—Guinot and Ribeiro, 1962:64, figs. 30-33 

[Angola].—Schmitt, McCain and Davidson, 1973:94 [synonymy]. 

DISTRIBUTION.—Baia de Santa Marta, Angola, 

among mussels on rocks. 

Family RETROPLUMIDAE Gill, 1894 

RETROPLUMIDAE Gill, 1894:1043. 

PTENOPLACIDAE Alcock, 1899c: 78. 

REMARKS.—This family, which contains a single 

genus, Retropluma Gill, 1894, is not represented 

in the eastern Atlantic. 

Family MICTYRIDAE Dana, 1851 

MYCTIRIDAE Dana, 1851e:247 [corrected to Mictyridae by 

Alcock, 1900:383]. 

REMARKS.—This family, which contains a single 

genus, Mictyris Latreille, 1806, is not represented 

in the eastern Atlantic. 

Family PALICIDAE Rathbun, 1898 

CYMOPOLIDAE Dana, 1854:9 [corrected to Cymopoliidae by 

Faxon, 1895:38; name 406 on Official Index, under Cymopoliidae 

Faxon, 1895]. 

PALICIDAE Rathbun, 1898:280 [name 376 on Official List]. 

PALICAE Bouvier, 1898:56, 58. 

EASTERN ATLANTIC GENERA.—One, Palicus, occurring 

off West Africa and in the Mediterranean 

Sea. 

EASTERN ATLANTIC SPECIES.—One, Palicus caronii 

(Roux, 1830), represented by one lot in the 

Pillsbury collections. It was recorded by Monod 

(1956) as Cymopolia caroni. 

Genus Palicus Philippi, 1838 

Cymopolia P. Roux, 1830, pi. 21 [invalid junior homonym of 

Cymopolia Lamouroux, 1816 (alga); type-species: Cymopolia 

caronii Roux, 1830, by monotypy; gender: feminine; name 

1718 on Official Index]. 

Palicus Philippi, 1838:11 [type-species: Palicus granulatus Philippi, 

1838, a subjective junior synonym ofCymopolia caronii 

Roux, 1830, by monotypy; gender: masculine; name 1640 


REMARKS.—Holthuis and Gottlieb (1958:104) 

explained why Palicus must be used in place of 

the older Cymopolia. 

* Palicus caronii (P. Roux, 1830) 

Cymopolia caroni.—Monod, 1956:387, figs. 546-551. 

Palicus caroni.—Forest and Guinot, 1966:88. 

Palicus caronii.—Zariquiey Alvarez, 1968:411, fig. 135e 

[Spain; references].—Tiirkay, 1976b:61 [listed], 71. 

SYNONYM.—Palicus granulatus Philippi, 1838. 

MATERIAL EXAMINED.—Pillsbury material: Ghana: 

Sta. 24, 35-37 m, dark red bryozoans, 26, 19 ov (L, W). 

Other Material: Canary Islands: No specific locality, 30 m 

(?), Talisman, 16 (W). 

Cape Verde Islands: Ilheu Branco, Talisman, 26, 1? ov 

(W). 

Senegal: Near Bane du Seminole, Baie de Goree, 38 m, I. 

Marche-Marchad, 18 Feb 1954, 26 (W). 

DESCRIPTION.—Bouvier, 1940:5; A. Milne Edwards 

and Bouvier, 1902:46-47 (comparison off. 

affinis A. Milne Edwards and Bouvier and P. 

caronii). 



MEASUREMENTS.—Our males have carapace 

widths of 7.6 to 10 mm; the ovigerous females 

have carapace widths of 9.3 and 11 mm. 

BIOLOGY.—Palicus caronii is a sublittoral species, 

occurring off West Africa in depths between 23 

(8-30) and 100 m. Of 16 recent depth records, 12 

(75%) are from depths between 23 and 50 m and 

4 (25%) are from depths greater than 50 m but 

less than 100 m, namely 73 m, 75 m, 80 m, and

192 

97-98 m. The Pillsbury specimens were taken on 

bottom with dark red bryozoans. The specimens 

collected by the Calypso were taken on bottoms 

with calcareous algae (three stations), algae and 

calcareous algae, and sand, algae, and calcareous 

algae (Forest and Guinot, 1966). 


collected in March, May, June, November, and 

December (Monod, 1956; Forest and Guinot, 



Mediterranean and adjacent Atlantic, including 

the Azores, Ilhas Desertas, Madeira, the Canary 

Islands, the Cape Verde Islands, off the mainland 

from Senegal and Ghana, and from the offshore 

islands of the Gulf of Guinea, Principe, Sao Tome, 

and Annobon, in depths between 23 m and 

100 m. Monod (1956) reported material from 

Senegal and Principe; records since 1956 include 


Madeira: No specific locality; Ponta de Sao Lourengo, 80 

m (Tiirkay, 1976b). 

Principe: 01°43'10"N, 07°28'2O"E, 73 m; and 01°43'N, 


Sao Tome: 00°25'40"N, 06°40'10"E, 50 m; and OO°25'- 

15"N, 06°43'05"E, 8-30 m (Forest and Guinot, 1966). 


1966). 

Family OCYPODIDAE Rafinesque, 1815 

OCYPODIA Rafinesque, 1815:% [corrected to Ocypodidae by 

MacLeay, 1838:63, 64; name 375 on Official List]. 

DOTINAE Dana, 1851c:286 [unavailable, based on a homonym]. 

MACROPHTHALMINAE Dana, 1851c:286. 

UCAINAE Dana, 1851c:289. 

HELOECIACAEA H. Milne Edwards, 1852:153. 

DOTILLIDAE Stimpson, 1858b:98 [p. 44 on separate]. 

CAMPTANDRIIDAE Stimpson, 1858b: 106 [p. 52 on separate]. 

Gelasimiden Nauck, 1880:8, 17, 23, 64, 66. 

GELASIMIDAE Miers, 1886:viii. 

SCOPIMERIOAE Alcock, 1900:290, 295, 363. 

CAMPTANDRIINAE Serene and Kumar, 1971:75. 

CAMPTANDRINI Pretzmann, 1974:443. 

CLEISTOTOMATINI Pretzmann, 1977:66. 

EASTERN ATLANTIC GENERA.—Five, Calabarium, 

new genus, Ecphantor, new genus, Lillyanella, new 


genus (see addendum), Ocypode, Telmatothrix, new 

genus, and Uca, each represented by species off 

tropical West Africa. 

EASTERN ATLANTIC SPECIES.—Six, all occurring 

in tropical West Africa. Two species were taken 

by the Pillsbury: Ocypode cursor and Uca tangeri. 

Three species were recorded by Monod (1956) 

under the same names used here, although he 

used Ocypoda for Ocypode. 

Three species of this family reported from West 

Africa appear to be either erroneously labeled or 

misidentified: 

Ocypode ceratophthalmus (Pallas, 1772). An Indo- 

West Pacific species reported from Fernando Poo 

by Pesta (1911:54) (see Monod, 1956:530). 

Ocypode quadrata (Fabricius, 1787). A western 

Atlantic species reported under the name 0. albicans 

Bosc from "Westafrika" by Balss (1922:79) 

(see Monod, 1956:399). 

Uca burgersi Holthuis, 1967. A western Atlantic 

species reported from Liberia under the name 

Uca mordax (Smith, 1870) by Rathbun (1900a: 

276; 1918:393; see also Monod, 1956:404) and 

Crane (1975:172, 326, 327, 604) and from Cameroon 

by Crane (1975:327, 604). 

It seems highly unlikely that these records are 

correct. We believe that these species should not 

be included in the list of West African Ocypodidae. 

Most modern authors (Balss, 1957:1663-1665; 

Guinot, 1967a:280-283; Serene, 1968:97-101) 

recognized three subfamilies within the family 

Ocypodidae: Ocypodinae, Scopimerinae, and 

Macrophthalminae. As the family group name 

Dotillidae Stimpson, 1858, is older than Scopimerinae 

Alcock, 1900, it has priority and should 

be used for the subfamily which contains both 

the genera Dotilla Stimpson, 1858, and Scopimera 

de Haan, 1833. We agree with Serene (1974:59- 

68) that a fourth subfamily, Camptandriinae 

Stimpson, 1858, should be recognized; it contains 

those genera formerly placed in the Macrophthalminae 

in which the male gonopods are strongly 

recurved. The four subfamilies of Ocypodidae 

that we now recognize are, therefore: Ocypodinae 

Rafinesque, 1815; Dotillinae Stimpson, 1958; 

Macrophthalminae Dana, 1851; and Camptan-

NUMBER 306 

driinae Stimpson, 1858. Only the first and the 

last are represented in the eastern Atlantic. 

Subfamily CAMPTANDRIINAE Stimpson, 1858 

REMARKS.—Stimpson (1858b: 106) erected a 

new family, Camptandriidae, to include a single 

genus, Camptandrium, which he was also newly 

describing. Later authors usually have synonymized 

the Camptandriidae with the subfamily 

Macrophthalminae Dana, 1851. However, the 

differences between the genera Macrophthalmus 

Desmarest, 1823, and Australoplax Barnes, 1966, 

on the one hand and the genera Camptandrium, 

Cleistostoma, Paracleistostoma, Tylodiplax, and Leipocten 

on the other are significant. It seems best, 

therefore, to separate the latter five genera as a 

distinct subfamily, for which the name Camptandriinae 

Stimpson, 1858, is available, confirming 

the views of Serene and Kumar (1971:75) and 

Serene (1974:59-68). Our reasons for recognizing 

Camptandriinae are practically the same as those 

noted by Serene and Kumar (1971) and Serene 

(1974). 

The characters that distinguish the Camptandriinae 

from the Macrophthalminae are the following: 

The most important is the shape of the 

Key to Genera of Camptandriinae 

193 

male gonopods, which are strongly recurved in 

the Camptandriinae, straight or slightly bent in 

the Macrophthalminae. The external maxillipeds 

in the Macrophthalminae have the merus shorter 

than the ischium and the last two segments thick; 

in the Camptandriinae the merus of the maxillipeds 

is as long as or longer than the ischium and 

the distal two segments are slender. 

The Camptandriinae are represented in West 

Africa by four genera, all of which are new. In 

order to ascertain their affiliations with the other 

genera of Camptandriinae, we had to study each 

genus within the subfamily Camptandriinae. 

These genera are very poorly defined and their 

limits are rather vague, so that a revision on the 

generic level was badly needed. Fortunately a fair 

number of species of Camptandriinae was available 

to us and made it possible to obtain a better 

idea of the various genera and their limits. At the 

same time the material showed the necessity to 

erect several new genera, as a number of species 

did not fit the older genera as redefined by us. 

For this reason we give here a diagnosis of the 

genera of Camptandriinae recognized by us with 

a discussion of the species that we assign to them, 

those of which we consider of dubious status and 

those that in our opinion should be eliminated 

from the Camptandriinae. 

1. Male abdomen constricted near fifth somite so that gonopods are partly 

visible when abdomen is pressed against thorax 2 

Male abdomen gradually tapering towards terminal somite, completely 

covering gonopods, which are not visible when abdomen is pressed 

against thorax 6 

2. Gonopod of male with 2 distinct appendages at the distal end 3 

Gonopod of male tapering to narrow pointed apex, without 

appendages 4 

3. Appendages at tip of male gonopod longer than distal recurved part of 

shaft. Carapace subhexagonal Camptandrium 

Appendages at tip of male gonopod short, at most half length of recurved 

part of shaft. Carapace transverse quadrangular Paratylodiplax 

4. Anterolateral margin of carapace with teeth. Meri of ambulatory legs 

with distinct subdistal dorsal tooth. Male gonopod drawn out into 

slender, narrow tip Calabarium, new genus


Anterolateral margin of carapace entire. Meri of ambulatory legs without 

subdistal dorsal tooth. Male gonopod with tip either short and triangular 

or with subdistal lobe 5 

5. Carapace only slightly wider than long, subhexagonal, without dorsal 

carina. Legs slender, merus of third (= longest) leg more than 3 times 

as long as wide. Male gonopod with broad subapical lobe. Fifth 

abdominal somite of male only slightly constricted 

Ecphantor, new genus 

Carapace distinctly wider than long, transversely quadrangular, with 

distinct transverse ridge. Legs broad, merus of third leg about twice as 

long as wide. Male gonopod regularly narrowing to triangular tip, 

without subapical lobe. Fifth male abdominal somite strongly constricted 

in basal part Deiratonotus, new genus 

6. First somite of male abdomen much wider than other somites and reaching 

coxae of fifth pereiopods 7 

First somite of male abdomen, though usually slightly wider than second, 

separated from coxae of fifth pereiopods by considerable distance .. 8 

7. Male gonopod ending in a pointed apex, which is not swollen. Chelipeds 

with strong sexual dimorphism Cleistostoma 

Male gonopod ending in blunt swollen apex. Males with chelipeds small, 

not different from those of females (dactylus of chelipeds without molarlike 

tooth) Ilyogynnis, new genus 

8. Merus of pereiopods with blunt spines on lower (flexor) 

surface Leipocten 

Merus of pereiopods without ventral spines 9 

9. Carapace without tubercles or transverse ridges dorsally. Anterolateral 

margins of carapace without teeth 10 

Carapace with transverse dorsal ridges and tubercles. Anterolateral margins 

of carapace with distinct or indistinct teeth. (Third maxilliped 

with small lobe at anterolateral angle of merus near base of carpus. 

Male gonopod without distinct appendages) 11 

10. Male gonopods swollen distally, provided with 1 or more distal appendages, 

which are at least half as long as recurved part of shaft. Apex of 

gonopod usually lying well free from basal part of shaft. Carapace with 

greatest width before middle Paracleistostoma 

Male gonopods tapering to narrow, pointed or blunt apex, not widened 

distally and without distal appendages. Gonopods so strongly recurved 

that tip lies over basal part of shaft. Carapace with greatest width 

behind middle Serenclla, new genus 

11. Anterolateral margin of carapace with 2 teeth, of which posterior very 

distinct, larger than anterior. Lateral margins of carapace converging 

posteriorly. Chelipeds showing strong sexual dimorphism. Propodi of 

second to fifth pereiopods longer than wide. Male gonopod not widened 

distally, distal margin deeply incised; morphological outer half of apex 

of gonopod slender and digitiform Telmatothrix, new genus


Anterolateral margin of carapace with 2 low teeth, posteriormost less 

distinct than anterior. Lateral margins of carapace diverging posteriorly. 

Chelae of male small, of same shape and size as those of female. 

Propodi of second to fifth pereiopods hardly longer than wide. Male 

gonopod slightly widened distally, without an appendage, apex of 

gonopod not incised; morphological outer half of apex of gonopod a 

wide and blunt lobe, wider than long Tylodiplax 

Genus Calabarium, new genus 

TYPE-SPECIES.—Calabarium crinodytes, new species. 

ETYMOLOGY.—Derived from the type-locality, 

the New Calabar River; the gender of the generic 

name is neuter. 

DIAGNOSIS.—A genus of Camptandriinae. Carapace 

flat and uneven, hardly wider than long, 

subhexagonal, with prominent front. Regions 

faintly indicated, with several elevations. Dorsal 

surface of carapace with sparse pubescence of 

short, curved, dark colored hairs, densest at elevations 

and at margins. Prominent front with 

wide V-shaped incision on anterior margin. Epigastric 

tubercles obscure, placed on base of front. 

Anterolateral margins of carapace with distinct 

teeth. Posterior margin of orbits lying in single 

transverse line, showing no indentations. 

Cornea well developed, but narrower than 

eyestalk. Antennules obliquely folded. Antennae 

entering orbit. Lower margin of orbit distinct in 

frontal view, suborbital ridge lying some distance 

below it; both showing only a few denticles. 

Epistome short and concave, with anterior and 

posterior margins elevated. Third maxillipeds not 

filling oral cavity, distinct space present between 

anterior margin of merus of maxilliped and posterior 

margin of epistome. Merus about as long 

as ischium and partly covering exopod. Anterior 

margin of merus sloping backwards towards median 

line of body. Inner anterior angle of ischium 

produced obliquely forwards. 

Chelipeds equal, those of males usually not 

enlarged, of about same size, or smaller, than 

those of females. Third and fourth pereiopods 

longer than second and fifth. Merus of second to 

fifth pereiopods with strong subdistal anterodorsal 

tooth, least pronounced in fifth leg. Dark 

colored, curved short hairs and longer plumose 

soft setae present on legs. 

Female abdomen with 7 somites, all somites 

free, abdomen broadly oval with apex slightly 

produced. Male abdomen elongate triangular, 

with the second, third, and fourth somites fused. 

First somite not wider than second, not reaching 

coxae of fifth pereiopods. Abdomen narrowing 

suddenly at end of fourth and beginning of fifth 

somite, with definite constriction, so much so that 

gonopods partly exposed, even when abdomen 

fully pressed against thorax. Male gonopods are 

strongly recurved, ending in simple, narrow, 

drawn out apex. 

REMARKS.—The shape of the carapace of Calabarium 

shows a remarkable resemblance to that 

of Camptandrium, being hexagonal with distinct 

anterolateral teeth and being almost as long as 

wide. Also the male abdomen, which is so strongly 

constricted that the gonopods become partly exposed, 

resembles that of Camptandrium. However, 

the fact that the male gonopods in Calabarium end 

in a single slender point and do not show distal 

appendages immediately distinguishes the present 

genus from the Indo-West Pacific Camptandrium. 

Furthermore, the walking legs in Camptandrium 

do not possess dorsal teeth, and in most 

species the chelipeds show a distinct sexual dimorphism. 

The male gonopod of Calabarium somewhat 

resembles that of Deiratonotus, but has the 

apex narrowly drawn out. Deiratonotus furthermore 

differs from Calabarium by the wide and 

ridged carapace, which is more quadrangular 

than hexagonal, by the unarmed anterolateral 

margins of the carapace, and by the walking legs, 

which do not show any dorsal teeth on the merus; 

also in Deiratonotus there is a strong sexual dimorphism 

in the chelipeds.

196 

Calabarium crinodytesy new species 

FIGURE 46 


Other Material: Nigeria: New Calabar River at Okpo 

waterside, Niger delta, 04°52'N, 06°54'E, 25 Jul 1978, C. B. 

Powell, 26, 3$ ov (one male is holotype) (L, W). Same 

locality, 7 Aug 1978, C. B. Powell, 26, 1 9 ov (W). Same 

locality, 15 Nov 1978, C. B. Powell, llcj, 5? (3 ov) (L, W). 

DESCRIPTION.—Carapace (Figure 46a) subhexagonal, 

slightly broader than long if lateral teeth 

included in width, slightly longer than broad 

exclusive of lateral teeth. Surface flat but uneven. 

Median area of carapace with following elevations 

from front to back: 2 indistinct epigastric, 2 

submedian mesogastric, 2 submedian cardiac, 1 

intestinal, latter five much stronger than epigastric, 

almost developed into tubercles. Each branchial 

region with 2 elevations, outermost placed 

slightly more posteriorly than innermost. Regions 

indistinctly indicated, cervical groove discernible. 

Lateral margins of carapace with strong tooth in 

anterior half, about twice as close to outer orbital 

angle as to posterolateral angle of carapace. This 

tooth about as strong as outer orbital tooth; 

between them are 2 smaller teeth on anterolateral 

margin. Posterior margin of orbit slightly sinuous, 

merging with lower orbital angle under broadly 

rounded curve. Neither orbital margin with incision, 

but lower orbital margin with distinct 

blunt tooth slightly before middle. Front horizontal, 

wide, anterior margin formed into 2 broad 

triangular teeth, each terminating in blunt point, 

separated by wide, V-shaped incision. Dorsal surface 

of carapace bearing short, stiff, dark brown, 

curved hairs, placed closest together on elevations 

and on margins. Few long, soft, plumose, uncolored 

hairs also present, especially on front. 

Eyes (Figure 46a,6) almost filling orbits, tapering 

distally, with distinct cornea. Some short, 

dark brown hairs placed on anterior surface of 

eyestalk. Lower margin of orbit with narrow denticulated 

process near base of antenna. Lateral to 

this process lower orbital margin with 1 or 2 small 

teeth on inner part, remainder of margin un- 


armed. Suborbital ridge lying distinctly behind 

and below orbit, with 2 denticles. 

Antennules covered by front, folding obliquely. 

Antennae short, entering orbit, latter widely open 

at its base. 

Epistome wide, somewhat concave, with anterior 

and posterior margins elevated, ridge-like; 

surface with some irregular pits. 

Third maxillipeds (Figure 466) not completely 

filling oral cavity, space present between anterior 

margin of merus and posterior margin of epistome. 

Outer margin of merus much longer than 

inner, anterior margin oblique. Outer anterior 

angle of merus produced, but not conspicuously 

so. Ischium about as long as merus, with inner 

anterior angle produced forward. Merus partly 

covering exopod. 

First pereiopods of male (Figure 46^) small, 

equal, much shorter than second pereiopods and 

not always larger than chelipeds of female. Fingers 

of male cheliped about as long as palm. Tips 

of chelipeds hoof-shaped and closing, proximal 

half of cutting edges gaping. Cutting edge of 

dactylus with blunt tooth in proximal third, proximal 

half of cutting edge of fixed finger with 

several small denticles. Palm about twice as long 

as broad when not swollen, slightly longer than 

broad when swollen. Carpus somewhat shorter 

than palm, unarmed. Merus slightly shorter than 

chela, with inner and outer lower anterior angles 

broadly rounded. Ischium short, with tooth on 

inner margin and several tubercles near distal 

margin. All segments of first pereiopod, up to 

palm, bearing stiff, short and curved, dark hairs 

similar to those on dorsal surface of carapace; a 

few long, soft, plumose hairs also present. Cheliped 

of female (Figure 46


a 

FIGURE 46.—Calabarium crinodytes, new genus, new species. Ovigerous female paratype: 

a, carapace; b, frontorbital region, ventral view; c, cheliped; d, abdomen. Male paratype: 

e, cheliped; /, second pereiopod; g, third pereiopod; h, fourth pereiopod; i, fifth pereiopod; 

j, abdomen in situ; k, basal part of abdomen. Holotype, male: /, gonopod. 

implanted on both upper and lower margins. 

Propodus slightly less than 1.5 times as long as 

dactylus, slightly widening proximally, bearing 

usual dark, short, and curved setae and some 

longer plumose hairs. Carpus about as long as 

dactylus. Merus, slightly longer than propodus, 

conspicuous by having large, sharp, forwardly 

curved, subdistal dorsal tooth. Pubescence of carpus 

and merus similar to that of propodus. Third 

leg (Figure 46g) longer and stronger than second, 

of about same structure and with same pubescence, 

except that lower margin of propodus and 

usually to lesser extent also that of merus, densely 

hairy, with numerous soft and long plumose hairs.

198 

Fourth leg (Figure 46/*) very similar in size, shape, 

and pubescence to third. Fifth leg (Figure 46?) 

distinctly shortest of all walking legs. Dactylus 

almost as long as propodus and longer than carpus. 

Subdistal short tooth of merus reduced to 

blunt lobe. In females legs longer and more slender 

than those of males, with short dark hairs 

more distinct and fewer long soft hairs. 

Female abdomen (Figure 46V) wide, reaching 

to coxae of pereiopods, broadly oval to almost 

circular, with all somites free. Seventh somite 

with apex somewhat broadly triangularly produced. 

Male abdomen (Figure 46/,A:) elongate triangular. 

First somite very short, about as wide as 

second, failing to reach base of fifth pereiopod, 

separated from coxa of that leg by considerable 

distance. Second, third, and fourth abdominal 

somites fused, but lines between them rather distinct. 

Second somite shortest of the three, but 

longer than first. Deep incision in lateral margin 

of abdomen indicating separation of second and 

third somites. Third and fourth somites of about 

equal length, fourth narrowing distinctly in distal 

part. Fifth and sixth somites free, fifth much 

narrower than third or basal part of fourth, narrowest 

at base, widening distally. Sixth somite 

about as wide as fifth but somewhat shorter, 

narrowing slightly distally. Seventh somite about 

as long as fifth but somewhat shorter, distinctly 

narrower, ending in almost semicircular distal 

margin. Outer margin of depression in thoracic 

sternum, which receives abdomen when latter 

completely folded against body, regularly triangular 

in shape; apex of depression rounded like 

seventh abdominal somite, lateral margins not 

showing any constriction; space present between 

margin of depression and outer margin of abdomen 

at level of constriction between fourth and 

fifth somites filled by part of ventral surface of 

gonopods, latter partly exposed even when abdomen 

completely pressed against thorax. 

Male gonopod (Figure 46/) with characteristic 

recurved shape of Camptandriinae. Anterior limit 

of curve just failing to reach separation between 

sternites of first and second pereiopods. Gonopod 


tapering regularly from base to top, widening 

somewhat in distal fourth, then narrowing rapidly 

to slender, sharp, drawn out point; latter 

simple, lacking appendages. Distal part of gonopod 

ornamented with few, scattered, minute tubercles, 

with some tubercles arranged in short 

row on inner margin proximal to tip. 

MEASUREMENTS.—Carapace lengths of males 

3.8 to 7.8 mm, of nonovigerous females 3.0 to 

5.0 mm, of ovigerous females 5.5 to 9 mm. Eggs 

numerous and small, diameter 0.3 to 0.35 mm. 

REMARKS.—The three new genera and new 

species, Calabrium crinodytes, Ecphantor modestus, and 

Telmatothrix powelli, are so far the only known 

West African Camptandriinae. Ecphantor can immediately 

be distinguished from the other two by 

the absence of anterolateral teeth on the carapace. 

The present new species is characterized by the 

shape of the front, which is more deeply incised 

anteriorly; furthermore, the chelipeds in Telmatothrix 

show a strong sexual dimorphism. Telmatothrix 

lacks the strong subdistal dorsal tooth on 

the merus of the walking legs. Also, the male 

abdomens and gonopods of the three species show 

significant differences, which necessitate recognition 

of a separate genus for each of the three West 

African species. 

TYPE-LOCALITY.—New Calabar River at Okpo 

waterside, Niger Delta, Nigeria, 04°52'N, 06°- 

54'E. 


D. 31950), a male, cl 5.7 mm, cb 6.3 mm, is in 


Paratypes have been deposited in that institution 

as well as in the National Museum of 


D.C. 


from the generic name of the aquatic lily, crinum, 

on which the species was found. 

BIOLOGY.—Mr. C. B. Powell (in litt., 26 July 

1978) informed us as follows: 

Specimens were collected from submerged leaves of the 

aquatic lily Crinum natans Baker in the zone of mixed Rhizophora 

and Pandanus in the New Calabar River. The water at 

the site is nearly fresh, but the tidal range is about 2 meters


and there is considerable disturbance to the littoral zone 

caused by the wash from motorboats and barges moving at 

top speed—when they pass by, one has to scramble to shore 

to avoid the large waves. The other Malacostraca present 

are Parorchestia n. sp., Cirolana, n. sp., Metastenasellus n. sp., 

Caridina africana Kingsley, Palaemonetes africanus Balss, Macrobrachium 

vollenhovenii (Herklots), M. felicinum Holthuis, Potamalpheops 

monodi (Sollaud), P. pylorus Powell (a single specimen), 

a new Potamalpheops, Telmatothrix powelli, Metagrapsus 

curvatus (H. Milne Edwards), Sesarma alberti Rathbun, S. 

angolense De Brito Capello, S. elegans Herklots, and S. buettikoferi 

De Man. 

Ovigerous females were collected in July, August, 

and November. 

DISTRIBUTION.—Known only from the typelocality. 

Genus Camptandrium Stimpson, 1858 

FIGURE 47 

Camptandrium Stimpson, 1858b: 106 [type-species: Camptandrium 

sexdentatum Stimpson, 1858, by monotypy; gender: 

neuter; name 289 on Official List]. 

DIAGNOSIS.—Carapace hexagonal rather than 

transversely quadrangular, with 2 or 3 anterolateral 

teeth, posteriormost reaching sideways. Surface 

of carapace flat, uneven, with elevations that 

may form longitudinal or transverse ridges. Epigastric 

lobes rather conspicuous, placed in basal 

part of front. Chelipeds equal, with conspicuous 

sexual dimorphism (except in the aberrant C. 

elongatum); dactylus in male with single tooth on 

cutting edge, tooth being absent from fixed finger; 

in female neither finger with tooth. Meri of 

pereiopods without spines, but lower distal angle 

may be produced. First somite of male abdomen 

only slightly wider than second and not extending 

to coxae of fifth pereiopods. Second to fifth somites 

of male abdomen fused, but lines between 

somites more or less distinctly visible. Abdomen 

gradually narrowing distally, but constricted at 

level of fifth somite so that gonopods partly visible, 

even when abdomen folded fully back against 

thorax. Male gonopods strongly recurved, each 

with apex somewhat swollen, ending in 2 long 

slender appendices, one practically straight, the 

FIGURE 47.—Camptandrium sexdentatum Stimpson: a, female, 

dorsal view; b, male abdomen; c, male chela; d, gonopod. 

(All from Shen, 1932, figs. 138, 140a,c,e.) 

other sinuous and longer of the two; appendages 

longer than recurved part of gonopod. 

REMARKS.—Shen (1935:30) assigned the following 

four species to the genus Camptandrium: C. 

sexdentatum Stimpson, 1858 (Figure 47); C. elongatum 

Rathbun, 1929; C. anomalum Shen, 1935; and 

C. paludicola Rathbun, 1909. Since 1935 the following 

three new species have been described in 

Camptandrium: C. starmuehlneri Pretzmann, 1968; 

C. ambonensis Serene and Moosa, 1971; and C. 

rathbunae Takeda, 1972. In our opinion four of 

these species are true Camptandrium, viz., C. sexdentatum, 

C. elongatum, C. ambonensis, and C. rathbunae. 

Camptandrium starmuehlneri Pretzmann (1968:16; 

see also Pretzmann, 1969:5, figs 1-4), which is 

known only from the female holotype, probably 

is a good Camptandrium near C. elongatum; but since 

the shape of the abdomen and the gonopods of 

the male are not known, we are not quite certain 

of its generic status. Serene and Moosa (1971:6)

200 

suggested that the species might belong either to 

the genera Shenius Serene or Ilyograpsus Barnard, 

which, however, seems unlikely. Camptandrium paludicola 

Rathbun belongs to the grapsid genus 

Ilyograpsus Barnard, 1955, being synonymous with 

the type and only known species of the genus, 

Ilyograpsus rhizophorae Barnard, 1955, the correct 

name of which thus is Ilyograpsus paludicola (Rathbun, 

1909) (see Crosnier, 1965:31). Camptandrium 

anomalum Shen was removed from Camptandrium 

by Serene (1971:916, 917) and made the type of 

the new genus Shenius Serene. Shenius should be 

removed from the Camptandriinae because the 

male gonopods of Shenius anomalus (Shen) are not 

recurved. Serene (1974:60) placed the genus Shenius 

in the subfamily Scopimerinae. Shenius also 

was left out of his (1974:62-66) lists of the genera 

and species of Camptandriinae. However, it was 

included in Serene's (1974:66) key to the genera 

of Camptandriinae. 

The range of the genus extends from India to 

Japan, Korea, the Malay Archipelago, and perhaps 

New Caledonia. 

Genus Cleistostoma de Haan, 1833 

FIGURE 48 

Cleistostoma de Haan, 1833:5, pi. B, pi. 7: fig. 3 [misspelled 

Cleistotoma on pi. 7] [type-species: Ocypode (Cleistostoma) 

dilatata de Haan, 1833, by monotypy; gender: neuter]. 

DIAGNOSIS.—Carapace convex both in longitudinal 

and transverse directions, not showing 

any appreciable ridges dorsally, but evenly hairy 

with short, dark pubescence. Carapace quadrangular 

in outline, being wider than long. Epigastric 

ridges not very distinct, extending forward far 

beyond base of front. Latter shallowly concave in 

middle, with anterior margin rounded, lacking 

anterolateral lobes. Suborbital ridge reaching far 

forward, protruding beyond distinct lower orbital 

margin. Upper orbital margin showing no incisions. 

Anterolateral margin of carapace lacking 

teeth apart from outer orbital angle, continuing 

posteriorly into posterolateral margin. No additional 

carina present on posterolateral part of 

a 


FIGURE 48.—Cleistostoma dilatatum (de Haan): a, female, dorsal 

view; b, male, frontal view; c, male abdomen; d, male 

chela; e, gonopod; /, tip of gonopod, enlarged. (From Shen, 

1932, figs. 145, 146, 147b,c, 148.) 

carapace. Chelipeds equal, left and right, but 

showing strong sexual dimorphism: much larger 

and stronger in male than in female and with 

different shape. Dactylus of male cheliped with 

distinct molariform tooth on cutting edge; such 

tooth absent on fixed finger of male cheliped or 

on either finger of that of female. Pereiopods 

lacking spines on merus. First somite of male 

abdomen very wide, reaching to coxae of fifth 

pereiopods; second somite much narrower. Second 

to fourth somites of male abdomen fused; 

fifth and sixth free. Lateral margins of abdomen 

somewhat sinuous, fitting snugly against thoracic 

sternum, covering gonopods completely. Strongly 

recurved male gonopods gradually narrowing toward 

apex, lacking distal appendages. Detailed 

figures of the type-species, C. dilatatum, are pro-


vided by Shen (1932:236, figs. 145-148, pi. 10; 

fig. 4). 

REMARKS.—Guinot and Crosnier (1963) listed 

the following species as belonging to the genus 

Cleistostoma: C. dilatatum (de Haan, 1833) (Figure 

48), C. wardi Rathbun, 1926, C. dotilliforme Alcock, 

1900, C. macneilli Ward, 1933, C. edwardsii Mac- 

Leay, 1838, and C. algoense Barnard, 1954. We 

have examined material of all these species except 

C. macneilli. In our opinion, only C. dilatatum, the 

type-species of the genus, which is known from 

Japan, North China, and Korea, is a true Cleistostoma; 

all the other species should be transferred 

to other genera. Cleistostoma wardi and C. dotilliforme, 

perhaps also C. macneilli, are placed in the 

genus Paracleistostoma (p. 208), and C. edwardsii 

and C. algoense are referred below to the genus 

Paratylodiplax (p. 209), as they differ in so many 

respects from Cleistostoma that it is impossible to 

assign them to that genus. 

Ocypode {Cleistostoma) dilatata de Haan, 1833, is 

a senior homonym of Ocypode (Macrophthalmus) 

dilatata de Haan (1835:55, pi. 15: fig. 2). We 

propose the replacement name, Macrophthalmus 

abbreviatus, new name, for the invalid Ocypode 

{Macrophthalmus) dilatata de Haan, 1835, for 

which, so far as we can ascertain, no other name 

has ever been proposed. The epithet abbreviatus is 

chosen because de Haan (1835:26) himself used 

it for the species, be it as a nomen nudum. 

In the original publication of the genus, de 

Haan (1833) used two different spellings: Cleistostoma 

in the text (p. 5) and on plate B, and 

Cleistotoma on plate 7: figure 3. In later parts of 

the Fauna Japonica de Haan was not consistent 

either; he mostly used Cleistostoma (part 2(1835): 

26, 27, 55, 56; part 3(1837), pi. 16: fig. 1), but at 

the end used again Cleistotoma (part 7(1849):233, 

234), namely in the index to the work. This 1849 

use of Cleistotoma was perhaps because H. Milne 

Edwards (1837:67, 68) had adopted that spelling; 

de Haan in his index (1849:233) referred to u Cleistotoma, 

n. Edw^ From de Haan's original publication 

it is not clear which spelling is the correct 

one; neither de Haan's (1835, 1837, 1849) later 

uses of the two spellings, nor H. Milne Edwards' 

(1837) use of Cleistotoma, indicates why a certain 

spelling is used. Thus none of these actions can 

be construed as first reviser actions. The actual 

first reviser is Dana (1852b:312, 313) who used 

the spelling Cleistostoma, and said in a footnote on 

p. 312 "not Cleistotoma." By Dana's action Cleistostoma 

is the correct spelling of the generic name; 

Cleistotoma, being an incorrect original spelling, 

has no nomenclatural standing. 

Agassiz (1846:89, 90) emended Cleistotoma to 

Clistotoma. Clistotoma Agassiz, 1846, thus is an 

unjustified emendation of the incorrect original 

spelling, but this notwithstanding is an available 

name, even though it is invalid as a junior objective 

synonym of Cleistostoma de Haan, 1833. 

Genus Deiratonotus, new genus 

FIGURE 49 

TYPE-SPECIES.—Paracleistostoma cristatum De 

Man, 1895. 

ETYMOLOGY.—From the Greek, deirado (ridge) 

and noton (back); gender of name is masculine. 

DIAGNOSIS.—Carapace about quadrangular in 

outline, distinctly broader than long with lateral 

FIGURE 49.—Deiratonotus cnstatus (De Man): a, male, dorsal 

view; b, male abdomen; c, abdomen in situ, male, cl. 7.5 

mm, Japan; d, male chela; e, gonopod; /, tip of gonopod, 

enlarged (a,M-/from Shen, 1932, figs. 141, 143b,c, 144).

202 

margins convex. Surface very flat, except for 3 

elevations forming a transverse ridge slightly behind 

middle, and short but distinct epigastric 

ridges. Ridges accentuated by covering of short 

dark brown hairs absent from rest of carapace 

except for marginal rim and additional ridge in 

posterolateral area. Inner dorsal orbital angle 

rounded. Front slightly concave in middle and 

with anterolateral angles angularly rounded but 

not produced. Suborbital ridge lying somewhat 

below distinct lower orbital margin, but reaching 

farther forward. Anterolateral margin of carapace 

unarmed. Sharp carina placed on posterolateral 

part of carapace, joining lateral margin anteriorly. 

Chelipeds with distinct sexual dimorphism. 

In male, chelipeds large, chelae with large molariform 

tooth on cutting edge of dactylus, but not 

on fixed finger. Meri of walking legs unarmed. 

Male abdomen with first somite somewhat wider 

than any of other somites, but failing to reach 

coxae of fifth legs by considerable distance. First 

somite of male abdomen free, second, third, 

fourth, and fifth fused, sutures only faintly indicated; 

incision in lateral margin marking separation 

between second and third somites. Outline 

of male abdomen narrowing gradually towards 

telson, but strongly constricted at level of the fifth 

somite, constriction revealing part of gonopods, 

even when abdomen is fully pressed against thoracic 

sternum. Exposed parts of gonopods, however, 

not flush with surface of abdomen. Sixth 

somite of male abdomen and telson are free. Male 

gonopod strongly recurved, ending in sharp point, 

neither swollen distally nor bearing appendages 

there. 

REMARKS.—This genus, in the shape of the 

abdomen, which leaves part of the gonopods 

exposed, resembles the genera Camptandrium and 

Paratylodiplax, and differs in this respect from all 

other Camptandriine genera. In Camptandrium 

and Paratylodiplax, however, the gonopods have 

two distinct appendages distally, while in Deiratonotus 

such appendages are entirely lacking. In 

the general shape of the carapace, with elevated 

hairy ridges the present new genus resembles the 

West African genus Telmatothrix. 


The type-species is the Indo-West Pacific 

Paracleistostoma cristatum De Man, 1895 (Figure 

49). A second species that we assign to this genus 

is Paracleistostoma japonicum Sakai. Although we 

saw no material, Sakai's (1934:321, fig. 26, pi. 18: 

fig. 1) good description indicates the main characters 

of Deiratonotus: ridged carapace, male abdomen 

revealing the gonopods, and the shape of 

the male gonopod. Deiratonotus cristatus is known 

from Japan, China and Korea, D. japonicus only 

from Japan. 

Genus Ecphantor, new genus 

TYPE-SPECIES.—Ecphantor modestus, new species. 

ETYMOLOGY.—From the Greek word ekphantor 

(revealer), referring to the partly exposed gonopods 

of the male; gender of name is masculine. 


subhexagonal, being only slightly (1.1 to 

1.2 times) wider than long. Surface of carapace 

flat but slightly uneven, with dense, uniform, 

short pubescence. Front directed obliquely down. 

Epigastric lobes strong, placed on base of front; 

no other distinct ridges present. Anterolateral 

margins without teeth. Orbits transverse, situated 

in common straight line. Eyes with distinct pigmented 

cornea. Antennules obliquely folded. Antennae 

entering orbit. Lower margin of orbit 

straight, without tubercles or teeth. Epistome 

short, concave. Third maxilliped filling entire 

oral field; merus and ischium covering palp and 

part of exopod. Merus shorter than ischium. Anterolateral 

angle of merus widely rounded, not 

produced. Inner anterior angle of ischium produced 

forward. Chelipeds equal, left and right; 

those of adult male much stronger and more 

robust than in female. Dactylus of cheliped in 

male with strong tooth in basal half of cutting 

edge, no other teeth present. No teeth on either 

finger in female. Tips of fingers of chelipeds 

spoon-shaped. Ambulatory legs longer than chelipeds; 

third and fourth pereiopods longer than 

second and fifth. No spines or teeth on any segments 

of ambulatory legs, latter covered by uniform 

short pubescence and scattered longer hairs.


Male abdomen narrow; all somites distinct, regularly 

tapering from first to sixth. First somite 

failing to reach coxa of fifth pereiopods. Fifth 

somite slightly constricted in basal part, first gonopods 

thereby slightly exposed. Female abdomen 

very wide, almost semicircular. Male gonopods 

strongly recurved; pointed apex with broad subdistal 

lobe, but without appendages. 

REMARKS.—The shape of the carapace of this 

new genus is somewhat similar in its hexagonal 

form to that of Calabarium, but it can immediately 

be distinguished by the total lack of anterolateral 

teeth. Like Camptandrium, Calabarium, Paratylodiplax, 

and Deiratonotus, Ecphantor has the abdomen 

constricted in such a way that the gonopods are 

partly visible even when the abdomen is firmly 

pressed against the sternum. The distal part of 

the gonopod, which carries a broad subdistal lobe 

but no appendages, distinguishes Ecphantor immediately 

from the other genera in the subfamily. 

Ecphantor modestus, new species 

FIGURE 50 


Other Material: Nigeria: Elechi Creek at College of Science 

and Technology, Port Harcourt, 04°47.3'N, 06°58.6'E, 19 

Oct 1979, C. B. Powell, Ic5 holotype (L). Same locality, 21 

Feb 1979, C. B. Powell, 1$ paratype (W). Same locality, 25 

Feb 1979, C. B. Powell, 1? paratype (L). 

DESCRIPTION.—Carapace (Figure 50a) subhexagonal, 

slightly (1.1 to 1.2 times) wider than long. 

Surface flat, margins slightly elevated; with pubescence 

removed, surface appearing slightly uneven, 

smooth, shining and pitted. Hair cover of 

carapace uniform, consisting of short, dark, 

curved, stiff hairs; cover dense enough to obscure 

surface, especially so since mud particles usually 

are caught between hairs. Central part of cervical 

groove rather distinct, with another median transverse 

depression some distance behind it; latter, 

however, very vague. Front directed obliquely 

down, separated from rest of carapace by distinct 

epigastric lobes, falling off precipitously anteriorly, 

separated from each other by short, distinct 

median groove; short longitudinal lateral groove, 

less distinct than median groove, present on either 

side of frontal lobes. Width of front at level of 

epigastric lobes slightly less than half frontorbital 

width; distal width of front 2/5 frontorbital 

width. Upper surface of front rather flat, slightly 

concave in middle. Anterior margin produced 

somewhat forward in middle, top slightly emarginate. 

Anterolateral angles of front broadly 

rounded. Lateral margin of front merging with 

orbital margin under wide curve. Orbital margin 

with indistinct notch in inner, posterior part, 

latter otherwise slightly convex. Outer orbital 

angle rectangularly rounded. Anterolateral margin 

of carapace unarmed. Angle between anterolateral 

and posterolateral margins blunt and 

wide. Posterolateral margin less sharply defined 

than anterolateral. Angle between anterolateral 

and posterolateral margins lying in anterior half 

of body, forming widest part of carapace. Faint 

but broad rim present on posterior margin; latter 

about 2/3 as wide as frontorbital width. 

Eyes with cornea distinct and globular but 

narrower than eyestalk. Lower margin of orbit 

straight, smooth, bearing several long hairs, lacking 

granules or teeth. Suborbital ridge similarly 

unarmed. 

Antennules folding obliquely, almost transversely. 

Antennae entering orbit. Flagellum short, 

consisting of 4 articles and terminating in long 

seta. 

Epistome wide, short, concave, anterior margin 

almost straight or slightly sinuous, posterior margin 

ending posteriorly in median carina separating 

the two halves of the oral field. No granules 

present. 

Third maxillipeds filling entire oral field. In 

ventral view merus and ischium covering larger 

part of both exopod and palp, of both only basal 

part being visible. Merus shorter than ischium, 

with anterolateral angle broadly rounded, not 

produced or auricular. Palp articulating in middle 

of anterior margin of merus. Exopod rather 

broad, with well-developed flagellum. 

First legs (Figure 50c,A) showing conspicuous 

sexual dimorphism. Chelipeds of male relatively 

small, shorter than any of other legs, but distinctly

204 


FIGURE 50.—Ecphantor modestus, new genus, new species. Male, holotype: a, dorsal view; 

b, anterior part of body, ventral view; c, cheliped; d, second pereiopod; e, abdomen;/, gonopod; 

g, apex of gonopod, enlarged. Smaller female, paratype: A, cheliped; i-l, second to fifth 

pereiopods.


more robust than those of female. Left and right 

chelipeds equal in both sexes. In adult male 

fingers of chela somewhat shorter than palm, 

gaping basally, tips spoon-shaped, with small 

horn-colored hoofs. Cutting edge of dactylus 

bearing strong tooth in gap, no other teeth present 

on it or on fixed finger. Palm slightly swollen, 

about 2/3 as high as long, highest in distal half. 

Carpus short, cup-shaped, slightly more than half 

as long as palm and distinctly less high. Merus 

elongate, slightly more than half as high as palm 

and about 2.5 times as long as wide. Chelipeds 

slenderer in female. Fingers narrow and elongate, 

somewhat shorter than palm, with spoon-shaped 

tips and fully unarmed cutting edges. Palm elongate, 

being about 3 times as long as high, increasing 

slightly in height distally. Carpus about as 

long as and as high as palm. Merus about 4 times 

as long as high, being slightly higher than carpus 

or palm. 

Of following legs (Figure 50d, i-l) second and 

third (= third and fourth pereiopods) longest. 

Second and fifth pereiopods (= first and fourth 

ambulatory legs) shorter, but still longer than 

cheliped. Second pereiopod slightly more slender 

than fifth, especially in shape of propodus. Dactylus 

of each ambulatory leg slender, about as 

long as propodus, unarmed. Carpus about as long 

or slightly shorter than propodus and about 2/3 

as long as merus. Merus totally lacking subdistal 

dorsal tooth, at most barely noticeable slight 

elevation there. All legs covered with same short, 

stiff, curved, dark hairs as are on carapace. Long, 

soft, somewhat plumose hairs present between 

short hairs, more distinct on anterior and posterior 

margins of segments. Longer hairs denser on 

posterior margins of dactylus and propodus than 

on anterior margins, and denser on posterior legs 

than on anterior ones. 


206 


type as follows: 

into anterolateral angles, latter triangularly pro- 

The specimen was taken from a bed of leaf fragments etc., 

with no sand and not much mud. At low tide seepage water 

drains through the bed, and the crab was in a handful of the 

duced. Posterior margin of orbit sinuous, with 

distinct incision at inner end. Suborbital ridge 

placed below distinct lower orbital margin, but 

litter I had scooped from the edge of the main trickle of reaching farther forward. Anterolateral margin of 

water. The most common macro-invertebrate present in the 

litter was the amphipod Quadrivisio, 

carapace showing 2 blunt lobes behind outer 

orbital angle. Sharp carina placed in posterolat- 

According to Powell (1979:126), Elechi Creek eral part of carapace above and mesial to postero- 

is a mangrove creek of 

lateral margin, forming pseudo-posterolateral 

margin, about parallel with true posterolateral 

about 6-12 meters wide and 1-2 meters deep, draining a 

large area of mangrove peat flats. The vegetation consists margin but not touching it. Chelipeds of male 

mainly of Avicemia africana and Lagunculana racemosa; also small, equal, similar to those of female; fingers 

present are Rhizophora spp. incl. R. mangle, the fern Acrostichumwith 

spoon-shaped tips, neither showing molari- 

aureum and the grass Paspalum vaginatum. The tidal range isform 

tooth. No sexual dimorphism shown by 

close to 2 meters so that at low tide the creek is dry except 

chelipeds. Meri of walking legs lacking spines. 

for a small shallow stream of drainage water running along 

the central sandy part of the creek bottom. . .. The maxi- Male abdomen about triangular in shape. First 

mum salinity of the creek water is probably about 20% . . . somite widest and shortest, reaching laterally as 

the normal dry-season salinity range is in the order of 5- far as coxae of fifth pereiopods. Second to fifth 

20%. In the rainy-season the maximum salinity is probably 

nearly as high . . . but the average and minimum salinities 

must be somewhat less. 

somites fused, separated by indistinct sutures 

only. Second somite much narrower than first 

and about as wide as third. Sixth somite and 

The type-locality of Ecphantor modestus is also telson free. Lateral margins of male abdomen 

the type-locality of Potamalpheops pylorus Powell, from third somite to telson about straight, being 

1979, the description of which is cited here. Pow- only slightly concave in sixth somite. Abdomen 

ell (1979:127) also provided a list of other inver- fitting closely against thoracic sternum, entirely 

tebrates present at the type locality. 

covering gonopods. Male gonopods recurved, 

Ovigerous females were collected in February. ending in widened bulbous apex, lacking distal 

DISTRIBUTION.—Known only from the type- appendage. 

locality. 

REMARKS.—The only species that we assign to 

this genus is Paracleistostoma microcheirum Tweedie 

(Figure 51), of which we examined material from 

Genus Ilyogynnis, new genus 

Singapore (USNM 137238). The genus Ilyogynnis 

FIGURE 51 

is primarily distinguished from Paracleistostoma by 

the very wide first segment of the male abdomen, 

TYPE-SPECIES.—Paracleistostoma microcheirum which reaches sideways as far as the coxae of the 

Tweedie, 1937. 

fifth pereiopods, the absence of sexual dimorph- 

ETYMOLOGY.—From the Greek ilyos (mud) and ism in the chelipeds, and by the shape of the male 

gynnis (a womanish man); gender of generic name gonopod, which lacks a distal appendage. In other 

is masculine. 

respects the male gonopod strongly resembles that 

DIAGNOSIS.—Carapace transversely quadran- of Paracleistostoma. Also the shape of the orbit with 

gular, only slightly wider than long, with lateral the distinct incision at the inner end of the pos- 

margins convex. Surface of carapace flat, but terodorsal margin, and the rather distinct regions 

regions rather well indicated by grooves. Dense of the carapace serve to distinguish the new genus. 

pubescence covering entire carapace. Epigastric The correct spelling of the name of the present 

ridges distinct. Front deeply grooved longitudi- species is Ilyogynnis microcheirum (Tweedie). Tweenally 

in middle, with oblique grooves extending die (1937) did not give the derivation of the


FIGURE 51.—Ilyogynnis microcheirum (Tweedie): a, anterior part 

of carapace of male; b, abdomen in situ, male, cl 6.5 mm, 

South China Sea; c, gonopod. (a,c, from Tweedie, 1937, fig. 

6a,d). 

specific epithet microcheirum, and since cheirum is 

not a known adjective, the word "microcheirum" 

must be considered either a noun or an arbitrary 

combination of letters; in either case the removal 

of the epithet from a neuter genus (Paracleistostoma) 

to one of which the name is masculine 

{Ilyogynnis) does not necessitate a change in spelling 

of the epithet. 

Genus Leipocten Kemp, 1915 

FIGURE 52 

Leipocten Kemp, 1915:243 [type-species: Leipocten sordidulum 

Kemp, 1915, by monotypy; gender: neuter]. 

DIAGNOSIS.—Carapace subquadrangular, 

slightly convex in both longitudinal and transverse 

directions. Upper surface with regions 

weakly indicated and bearing hairs and granules. 

Epigastric lobes faint, placed on base of front. 

Anterolateral teeth present. Third maxilliped 

with merus hardly auriculate. Part of exopod of 

third maxilliped visible. Chelipeds with distinct 

sexual dimorphism. Male chela with molariform 

tooth on cutting edge of dactylus, but not on 

fixed finger. Female with strong spines on chela, 

fingers about as long as palm. Walking legs short, 

robust, with spines on lower margin of merus. 

Male abdomen with first somite narrow, not 

reaching coxae of fifth pereiopods. Second and 

third abdominal somites of male fused. Male 

gonopod strongly recurved, ending in 2 processes, 

one narrow, pointed, the other lobiform, with 

several curved spines. 

REMARKS.—Only one species, the type, L. sordidulum 

(Figure 52), is known in this genus. It has 

an Indo-West Pacific distribution (India to Formosa 

and Australia). 

Serene (1974:62, 64, 66) justifiably intimated 

that Baruna Stebbing, 1904, probably is a senior 

synonym of Leipocten Kemp, 1915. Unfortunately, 

Stebbing's (1904:3, 4, pi. 1: fig. A) description 

and illustrations of his new genus Baruna and its 

single new species, Baruna socialis Stebbing, 1904, 

are insufficient by modern standards. For one 

thing the male pleopods have neither been described 

nor figured. On the other hand, the great 

similarity of the shapes of the male chelipeds, of 

the peculiar pereiopods, of the male abdomen, 

and of the anterolateral margin of the carapace 

of Leipocten and Baruna, as well as the sizes and 

the habitats of their species, strongly suggest that 

the two genera might be synonymous. The differences 

between the two, so far as can be judged by 

the available data, are minor: Baruna socialis has 

the carapace wider (1.4 times as wide as long) 

than Leipocten sordidulum (1.27-1.33 times); the 

exopod of the third maxilliped seems wider in 

Baruna than in Leipocten; the posterior anterolateral 

tooth of the carapace in Leipocten is granular, 

whereas in Baruna it is smooth, having been described 

by Stebbing as simple (i.e., not subdivided 

into small teeth); the fact that Stebbing's illustration 

of the front and the orbit of Baruna socialis 

show these without tubercles may be due to the 

inaccuracy of the drawing. Should the gonopods 

of Baruna and Leipocten prove to be very similar 

there would be no good reason not to synonymize 

the two names and the older of them, Baruna 

Stebbing, 1904, would have to be used. 

Baruna socialis Stebbing, 1904, so far the only 

species assigned to Baruna, is known only from the

208 


a d 

FIGURE 52.—Leipocten sordidulum Kemp: a, female, dorsal view; b, female, frontal view; c, male 

abdomen; d, male chela; e, female chela;/, fourth pereiopod. (From Kemp, 1915, figs. 16, 18, 

19, 20a, and pi. 12: fig. 8.) 

type-material, which originated from brackish 

water of Lake Negombo, Ceylon. 

Genus Paracleistostoma De Man, 1895 

FIGURE 53 

Paracleistostoma De Man, 1895a:580 [type-species: Paracleistostoma 

depression De Man, 1895, by selection by Guinot and 

Crosnier, 1963:608; gender: neuter]. 

DIAGNOSIS.—Carapace flat, being only slightly 

curved down at margins, lacking ridges, with only 

depressed H-shaped groove in center. Surface of 

carapace hairy or glabrous, outline quadrangular, 

being wider than long. Epigastric ridges indistinct, 

placed at base of front. Anterior margin of 

front with blunt or triangular projection at either 

lateral angle. Anterolateral margin of carapace 

lacking teeth. Suborbital ridge forming lower 

margin of orbit, true lower margin being barely 

visible as row of granules inside orbit. Shape and 

sexual dimorphism of chelipeds similar to Cleistostoma. 

Walking legs lacking spines on merus. 

First somite of male abdomen only slightly wider 

than second, failing by far to reach coxae of fifth 

pereiopods. Second to fifth somites of male abdomen 

fused, although lines separating these somites 

only faintly or more distinctly indicated by 

sutures. Male abdomen narrowing regularly distally, 

showing no constrictions; abdomen pressed 

against thorax, completely covering male gonopods. 

Latter recurved, strongly and bulbously 

widened apically, carrying distal appendage. 

REMARKS.—Guinot and Crosnier (1963) listed 

the following species as belonging to Paracleistostoma: 

P. depressum De Man, 1895 (Figure 53); P. 

leachii (Audouin, 1826); P. cristatum De Man, 

1895; P. eriophorum Nobili, 1903; P. dentatum Tesch, 

1918; P. longimanum Tweedie, 1937; P. fossulum 

Barnard, 1955; P. microcheirum Tweedie, 1937; and 

P. japonicum Sakai, 1934. In our opinion only the 

following species actually belong to Paracleistostoma: 

P. depressum, P. longimanum, P. wardi, and P. 

dotilliforme (we have examined material of all 

except P. longimanum). The last two species (P. 

wardi and P. dotilliforme) formerly had been assigned 

to Cleistostoma (p. 200). We doubtfully refer 

the following two species to Paracleistostoma: 

Cleistostoma mcneilli Ward, 1933: This species 

was placed by Barnes (1967:246) in Paracleistos-


FIGURE 53.—Paracleistostoma depression De Man: a, male, dorsal 

view; b, male abdomen; c, male chela; d, female chela; e, 

gonopods in situ; /, gonopod. (a-d from De Man, 1897, pi. 

14: figs. 13, I3d-f; ejfrom Gordon, 1931, fig. 26). 

toma. However, the anterolateral margin of the 

carapace has distinct teeth, and the male gonopod 

figured by Barnes (1967, fig. 16d) differs somewhat 

from that of the typical Paracleistostoma species. 

It is possible that C. mcneilli is a true Paracleistostoma, 

and that the definition of the genus as 

given above should be somewhat modified. 

Paracleistostoma eriophorum Nobili, 1903: Judging 

by Nobili's (1903:23) description of the carapace, 

the chelipeds, and the male abdomen, this species 

could well be a true Paracleistostoma. However, 

nothing is known about the shape of the male 

gonopods, while also some important characters 

of the abdomen are not mentioned by Nobili. Of 

this species, which so far has not been illustrated, 

we have not seen any material. 

The following five species are excluded from 

the genus Paracleistostoma: 

Paracleistostoma leachii (Audouin, 1826): The 

shape of the male gonopod, which ends in a 

narrow apex, shows that the species is not a 

d 

Paracleistostoma and we make it the type of a new 

genus Serenella (p. 211). 

Paracleistostoma cristatum De Man, 1895, and P. 

japonicum Sakai, 1934, are both placed here in the 

new genus Deiratonotus (p. 201). 

Paracleistostoma microcheirum Tweedie, 1937, is 

made the type of a new genus, Ilyogynnis (p. 206). 

Paracleistostoma dentatum Tesch, 1918: The general 

shape of the carapace is wholly unlike that of 

Paracleistostoma and resembles more that of Camptandrium. 

Since the only known specimen is a small 

female, very little can be said about the generic 

status of this species. 

Paracleistostoma fossulum Barnard, 1955: In this 

species the dorsal surface of the carapace shows 

transverse ridges. The anterolateral margins of 

the carapace have some feeble teeth. The third 

maxillipeds are separated by a wide gap and the 

peduncle of the exopod is fully exposed. The 

female chelipeds are larger than those in Paracleistostoma, 

have the fingers shorter than the palm 

and the cutting edges of both fingers show several 

teeth. The species certainly is no Paracleistostoma 

and possibly not even an ocypodid. Unfortunately 

it is only known from a female specimen. 

Genus Paratylodiplax Serene, 1974 

FIGURE 54 

Paratylodiplax Serene, 1974:62 [type-species: Cleistostoma blephariskios 

Stebbing, 1924, by original designation; gender: 

feminine]. 

DIAGNOSIS.—Carapace quadrangular or oval, 

wider than long, rather flat, somewhat more convex 

in longitudinal than in transverse direction. 

Dorsal surface smooth, showing only deep, Hshaped 

depression formed by central part of cervical 

groove and anterior part of cardiac grooves. 

Carapace pubescent or naked. Epigastric ridges 

present as rather indistinct rounded elevations at 

base of front. Latter somewhat concave in middle. 

Anterior margin of front with angularly rounded, 

but not produced, anterolateral angles. Inner orbital 

angle broadly rounded. Suborbital ridge 

lying distinctly below lower margin of orbit,


FIGURE 54.—Paratylodiplax derijardi (Guinot and Crosnier): a, carapace of male, dorsal view; b, 

male abdomen; c, male chela; d, female chela; e, gonopod. Paratylodiplax edwardsii (MacLeay): 

/, abdomen, male, cl 7.8 mm, South Africa; g, gonopod; h, tip of gonopod, enlarged, (a-e from 

Guinot and Crosnier, 1963, figs. 1-3, 8, 9; gji from Barnard, 1954, fig. 2e,f). 

reaching only slightly farther forward. Anterolateral 

teeth of carapace distinct or obscure. Anterolateral 

margin continuing into posterolateral; no 

additional ridge present posterolaterally on carapace. 

Chelipeds equal in female, subequal in 

male, showing strong sexual dimorphism, chelipeds 

of male being much stronger than those of 

female and having different shape. Dactylus of 

male cheliped with distinct molariform tooth in 

basal part of cutting edge; this tooth absent in 

female. Pereiopods lacking spines on merus. First 

somite of male abdomen widest of all abdominal 

somites, failing by considerable distance to reach 

coxae of fifth pereiopods. Second somite of male, 

abdomen slightly narrower than first, either free 

(P. derijardi, P. blephariskios) or fused with abdom- 

h 

inal somites 3 to 5. Latter fused, showing only 

traces of sutures between somites; sixth somite 

and telson free. Somites narrowing gradually 

from first to sixth, outline of male abdomen being 

bluntly triangular, except for constriction caused 

by concavity in lateral margin at level of fifth 

somite. Depressed area of thoracic sternum, which 

receives abdomen when latter is fully pressed 

against thorax, regularly triangular, leaving an 

opening between abdomen and thoracic sternite; 

through opening part of gonopod visible. Exposed 

part of gonopod completely filling gap between 

abdomen and sternum, exposed surface lying 

flush with lower surface of abdomen. Male gonopod 

strongly recurved, ending in 2 rather short 

appendages, inner slender, curved or sinuous,


outer lobiform; both shorter than recurved part 

of shaft. 

REMARKS.—The following four species are here 

assigned to this genus; material of the first three 

has been examined by us: 

Paratylodiplax edwardsii (MacLeay, 1838:64) 

(Figure 54/-A), originally described as Cleistotoma 

edwardsii, was included in this genus by Serene 

(1974:62). In it the second somite of the male 

abdomen is fused with the following three; in our 

specimen the suture between the second and third 

somites is rather distinct in the right half, but 

altogether absent in the left. Barnard's (1950:150) 

statement (in his definition of the genus Cleistostoma 

in which he placed the present species) that 

all somites of the abdomen are free certainly does 

not apply here. 

Paratylodiplax algoensis (Barnard, 1954:122, figs. 

1, 2a-d) originally described as Cleistostoma algoense, 

was correctly considered by Barnard to be 

close to P. edwardsii. Barnard (1954) described the 

chelipeds and the male gonopods, but gave no 

description or figure of the entire animal nor of 

the male abdomen. Examination of specimens (1 

male and 1 female) from Knysna estuary, South 

Africa (USNM collection, uncataloged), confirmed 

our idea that the species belongs in Paratylodiplax. 

It fully agrees with the above-cited 

definition of the genus. The male abdomen has 

the second somite either free or, if fused with the 

third, separated by a deep suture. The fifth somite 

is constricted, so that in ventral view the gonopods 

show next to the closed abdomen. Serene 

(1974:62) believed that this species probably was 

conspecific with P. edwardsii. 

Paratylodiplax blephariskios (Stebbing, 1924:3, pi. 

116), the type-species, was originally described as 

Cleistostoma blephariskios. Stebbing's (1924) description 

was rather poor, and Barnard (1950: 

816, 817) gave some additional details, e.g., of the 

chelipeds and the male gonopods, which agree 

with those of Paratylodiplax. A rather poorly preserved 

specimen at our disposal shows that the 

shape of the male abdomen was incorrectly described 

and figured by Stebbing as consisting of 

seven free somites. The male abdomen shows the 

shape typical for the present genus. There is a 

distinct suture between the second and third abdominal 

somites, but not between the third, 

fourth, and fifth somites. The male gonopods are 

visible lateral to the closed abdomen. The examined 

specimen came from Cape Town, South 

Africa, collected by J. Day, 1973, in USNM 

collection, uncataloged. 

Paratylodiplax derijardi (Guinot and Crosnier, 

1963:612, figs. 1-3, 5-11) (Figure bAa-e), was 

originally placed in the genus Tylodiplax. The 

species is well described and figured in the original 

publication, and shows all the characters 

listed in the above generic diagnosis of Paratylodiplax, 

in which genus we therefore do not hesitate 

to place it, following Serene (1974:62). 

The present genus differs from Cleistostoma (1) 

in the less strong development of the suborbital 

ridges, (2) by the fact that the first abdominal 

segment of the male does not reach the coxae of 

the fifth legs, (3) in showing part of the male 

gonopods when the abdomen is pressed against 

the thorax, and (4) by the presence of distal 

appendages at the end of the male gonopods. 

Of the four known species of Paratylodiplax, 

three (P. algoensis, P. blephariskios, and P. edwardsii) 

are known only from South Africa. The fourth, 

P. derijardi, has been reported from Madagascar. 

Genus Serenella, new genus 

FIGURE 55 

TYPE-SPECIES.—Macrophthalmus leachii Audouin, 

1826. 

ETYMOLOGY.—This genus is named for Dr. 

Raoul Serene in recognition of his many valuable 

contributions to the study of Brachyura; the gender 

of the generic name is feminine. 

DIAGNOSIS.—Carapace broadly quadrangular, 

being distinctly wider than long. Dorsal surface 

of carapace smooth, without ridges or tubercles, 

with only shallow grooves indicating regions. 

Front curved down, anterior margin at either end 

with 2 angular lobes, inner larger than outer and 

reaching farther forward. Epigastric ridges dis-

212 

FIGURE 55.—Serenella leachu (Audouin), male: a, dorsal view; 

b, gonopods in situ; c, tip of gonopod (L, Crust. D. 26887), 

Melita Bay, Red Sea; d, abdomen (L, Crust. D. 26888), 

Museri Island, Red Sea. (a, from Paulson, 1875, pi. 8: fig. 6; 

b, from Gordon, 1931, fig. 27 left.) 

tinct, placed before base of front. Lateral margins 

of carapace without teeth, converging posteriorly. 

Orbits transverse, situated in single straight line. 

Upper orbital border convex, lacking incisions. 

Eyes with cornea well developed although small. 

Antennules transversely folded. Antennae not entering 

orbit. Suborbital ridge fused with outer 

third of lower orbital margin, provided with granules. 

Epistome short, with transverse, smooth, 

sharp ridge; posterior margin produced, tonguelike, 

in middle between third maxillipeds, latter 

filling entire oral field. Merus and ischium covering 

part of exopod and entire flagellum; merus 

wider than ischium, somewhat auriculate, lacking 

lobe near base of carpus. Chelipeds equal, those 

of adult males larger than those of females and 

with quadrangular tooth on cutting edge of dactylus. 

Third pereiopod longer than other legs, 

fifth pereiopod shortest. No spines present on any 

pereiopod. Male abdomen narrow, not constricted 

at fifth abdominal somite. First somite 

only slightly wider than second and failing considerably 

to reach coxa of fifth pereiopod. Second, 

third, and fourth abdominal somites fused, only 

some faint grooves indicating borders between 

them. Fifth and sixth somites free. Female abdo- 


men almost circular, with all somites free. Male 

gonopods strongly recurved, so much so that apex 

overlaps base of shaft. Distal part of gonopod not 

widened, ending in narrow, bluntly rounded 

apex. Morphological inner side of apex bearing 

strong spines, curved at tip and on one side 

bearing some granules. Tip of gonopod bearing 

short, wide triangular lobe. 

REMARKS.—So far S. leachii is the only species 

known of this genus. It is possible that Tylodiplax 

indica Alcock, 1900, also should be placed here (p. 

217), but this species differs in several important 

respects from the type-species. In the first place 

Alcock's species evidently shows no sexual dimorphism 

in the chelipeds, while also the shape 

of the third maxilliped and the male gonopod are 

different. Stephensen's (1945, fig. 58D) illustration 

shows the male abdomen of T. indica slightly 

constricted at the fifth somite, but no information 

is available on whether or not this constriction is 

so strong that the gonopods become visible. A 

closer study of Alcock's species is necessary to 

solve its generic position. 

Genus Telmatothrix, new genus 

TYPE-SPECIES.— Telmatothrix powelli, new species. 

ETYMOLOGY.—The name is derived from the 

Greek words telma (mud of a pool) and thrix (hair) 

in allusion to the hairs of the animal, which are 

made quite conspicuous by the mud caught between 

them; the gender of the generic name is 

feminine. 

DIAGNOSIS.—Carapace transversely quadrangular, 

being wider than long. Surface of carapace 

flat, but regions rather distinct, some with elevations 

forming blunt transverse ridges. Short, dark 

pubescence visible on dorsal surface, especially 

dense and conspicuous on ridges. Front curved 

down. Epigastric ridges strong, placed on base of 

front. Anterolateral margins of carapace with 

distinct teeth. Orbits transverse, situated in common 

straight line; upper orbital border lacking 

incisions. Eyes with cornea well developed. Antennules 

obliquely folded. Antennae entering or-


bit. Lower orbital margin and suborbital ridge 

visible in front view, granulated. Epistome short, 

with smooth, sharp, transverse ridge. Third maxillipeds 

filling entire oral field; merus and ischium 

covering palp and part of exopod. Merus longer 

than ischium, and with anterolateral angle 

rounded or somewhat flattened, not produced, 

but with small lobe near base of carpus. Inner 

anterior angle of ischium bluntly triangularly 

produced. Chelipeds equal; those of adult males 

much larger than those of females and with different 

shape; dactylus in males with strong molariform 

tooth on cutting edge, no such tooth 

present on fixed finger of either sex or on dactylus 

of female. Third and fourth pereiopods longer 

than second and fifth. No spine on any of segment 

of pereiopods, but pubescence present on several. 

Male abdomen narrow, with second, third, and 

fourth somites fused. First somite only slightly 

wider than second, largely failing to reach coxae 

of fifth pereiopods. Female abdomen almost circular, 

consisting of 7 free somites. Male gonopods 

strongly recurved, ending in cleft apex, latter 

narrowing gradually, lacking distal appendages. 

The type and only species so far known of this 

genus is described below. 

Telmatothrix powelli, new species 


"a new ocypodid".—Powell, 1976:315. 


Other Material: Nigeria: Mayuku Creek at Ugbekoko (= 

Gbekoko), approximately 10 mi [16 km] W of Sapele, Midwest 

State, at about 05°54'N, O5°37'E, among mangroves 

(Rhizophora), oligohaline, 5 Oct 1975, C. B. Powell, 36, 3 juv 

9, 1 juv (L). Same locality, Oct-Dec 1975, C. B. Powell, 3

a 

FIGURE 56.—Telmatothrix powelli, new genus, new species: a, male, dorsal view; b, female, 

dorsal view. (Drawn by J. Wessendorp.)


distinctly granular, slightly sinuous, showing no 

indentations. Inner half of posterior orbital margin 

widened ventrally, with granules in lower 

part of widened area. Between widened area and 

base of eye short sharp carina present, branching 

off from orbital margin. Outer orbital tooth 

bluntly angular. Behind this tooth anterolateral 

margin of carapace bearing 2 distinct teeth, anteriormost 

lower, more rounded than second; latter 

longer, more triangular, directed more sideways; 

lateral 3 teeth separated from each other 

by deep triangular incisions, margins granular. 

Posterolateral margin longer than anterolateral, 

unarmed but granular; just before reaching posterior 

margin row of granules curving inward, not 

touching posterior margin. Second granular ridge 

present branching off from posterolateral ridge 

slightly behind posterior lateral tooth, lying more 

medially from posterolateral ridge. This second 

ridge, stronger than posterolateral, extending posteriorly, 

ending in sharp, elongate posterolateral 

tubercle, lying near margin of carapace above 

base of fifth pereiopod. Posterolateral margin of 

carapace with blunt tooth protruding between 

bases of fourth and fifth pereiopods. Posterior 

margin of carapace, like posterolateral, somewhat 

granular and elevated; another parallel granular 

ridge present, placed more anteriorly. 

Eyes (Figure 57a) elongate, cornea terminal, 

not surrounding stalk completely; lower margin 

of orbit finely granular, bearing long hairs that 

partly cover eye when latter retracted. Below 

lower orbital margin ridge present with about 11 

to 16 low but distinct tubercles, outer smaller, 

slightly more closely placed than inner. 

Antennules folding obliquely, almost transversely. 

Antenna entering orbit; flagellum short, 

consisting of about 6 segments, bearing long bristle, 

slightly longer than flagellum itself. 

Front (Figure 57a) touching epistome in middle. 

Epistome showing smooth transverse ridge in 

middle, posterior margin, strongly produced posteriorly 

in middle, distinctly granular. 

Third maxillipeds (Figure 57 d) fill entire oral 

cavity. In ventral view only part of exopod and 

part of palp visible, slender dactylus, somewhat 

FIGURE 57.—Telmatothrix powclli, new genus, new species: a, 

front; b, male abdomen; c, base of male abdomen in situ; d, 

third maxilliped; e, male chela; /, female chela; g, chela of 

juvenile male; h, gonopod. 

less slender propodus, part of carpus, and distal 

part of exopod hidden behind merus and ischium. 

Merus with anterolateral angle rounded or somewhat 

flattened, but not auricular, forming shallow, 

anteriorly directed lobe just outside base of 

palp. Latter articulating in middle of anterior 

margin of merus. Inner anterior angle of merus 

rectangularly rounded. Ischium almost as long as 

and as wide as merus. Inner margin of both 

segments elevated, rim-like, otherwise no longitudinal 

carinae or grooves present. 

First legs (Figure 51e-g) showing conspicuous 

sexual dimorphism. In adult males chelipeds 

large, robust, being far larger than following legs 

and more than twice as long as carapace. Fingers 

slightly more than half as long as palm, gaping 

distinctly. Finger tips slightly spoon-shaped, each

216 

bearing row of long hairs on inner margin. Cutting 

edges of fingers bearing small granules, 

distalmost larger than proximals. Cutting edge of 

dactylus with wide molariform tooth, occupying 

slightly more than 1/4 of length of edge; no such 

tooth present on cutting edge of fixed finger. 

Palm highest distally, narrowing proximally, 

bearing scattered small granules, densest dorsally. 

Carpus about half as long as palm, merus slightly 

shorter than palm. Both carpus and merus showing 

granules, but bearing no spines. Left and 

right chelipeds in male equal in shape and size. 

In female chelipeds much shorter than second 

pereiopods, of about same length as carapace, 

slender and equal. Fingers somewhat longer than 

palm, gaping and having spoon-shaped tips bordered 

with long hairs. Cutting edges entire. Carpus 

about as long as palm and less than 2/3 as 

long as merus. 

Dactyli of following legs simple, slightly shorter 

than propodi, provided on both dorsal and ventral 

surfaces with 3 longitudinal pubescent 

grooves separated by 2 ridges. Propodi about 

twice as long as wide and longer than carpi, 

neither bearing any granules. Merus about as 

long as propodus and carpus combined but wider, 

bearing distinct granules on lower surface, especially 

along margins; some granules visible also 

in upper part of outer (= posterior) surface. Third 

and fourth pereiopods distinctly longer and wider 

than second and fifth; third leg largest, fifth 

smallest of walking legs. Outer (= posterior) surface 

of these legs bearing numerous short black 

hairs, among which some longer hairs visible, 

especially on upper and lower margins. Third, 

fourth, and fifth pereiopods showing numerous 

soft, long, woolly hairs on lower surface of merus; 

such woolly hairs also visible in outer distal part 

of carpus and dorsal part of propodus of third 

and fifth legs, and on inner, upper, and lower 

surfaces of propodus and distal part of carpus of 

fourth legs. Woolly hairs more distinct in males 

than in females. No teeth or spines on any of 

segments of legs; anterodorsal and anteroventral 

angles of merus bluntly rounded. 

Male abdomen (Figure 57b,c) slender. First 


somite only slightly wider than second, largely 

failing to reach coxae of fifth pereiopods. Second 

somite slightly wider than third, of about same 

length as first and about half as long as third. 

Third somite about as long as but somewhat 

wider than fourth. Second, third, and fourth somites 

fused, but grooves indicating lines separating 

somites, while deep incisions in lateral margins of 

fused part also indicate limits of somites. Distal 3 

somites of about equal width. Fifth somite longest 

of three, also longer than fourth; differences 

slight. Sixth and seventh somites of about equal 

length. Sixth quadrangular, seventh with tip 

semicircularly rounded. Weak, low transverse carina, 

interrupted in middle, present in distal half 

of exposed surface of seventh somite. Other somites 

not showing any carinae, except first, traversed 

by distinct smooth carina. In adult female 

abdomen with all somites free, almost semicircular, 

reaching to, or overlapping, bases of the 

pereiopods. 

Male gonopods (Figure 57h) reaching to line 

between sternites of second and third pereiopods, 

strongly recurved, tapering gradually towards tip. 

Tip ending in 2 points, of which 1 minutely 

dentate on outer margin. Tip neither widened 

nor bearing any appendages. Male sexual openings 

placed on sternum. 

MEASUREMENTS.—The largest male examined 

(the holotype) has the carapace 10 mm long and 

14 mm wide, the largest females were of the same 

length and 14 and 15 mm wide. Females, with 

the abdomen wide and reaching to the bases of 

the pereiopods, ranged in length from 7 to 10 mm 

and in width from 9 to 15 mm. Ovigerous females 

had a carapace length of 9 and 9.5 mm and a 

carapace width of 12 and 13 mm. Juveniles (including 

females with a narrow abdomen and 

males in which the chelae still showed the female 

type) ranged from carapace width 2.5 to 8 mm. 

The eggs are numerous and have a diameter of 

about 0.35 mm, they are spherical. 

A male with a carapace length of 6 mm and a 

carapace width of 8 mm showed the chelipeds 

very similar to those of the females, the fingers 

(Figure big) were longer than the palm, with only


faint traces of the molariform tooth of the dactylus 

and of the granulation of the cutting edge 

of the fixed finger are visible. In still smaller 

males the chelipeds are indistinguishable from 

those of the female, e.g., in a male with cl 3.5 mm 

and cb 5 mm, in which the abdomen and the 

gonopods showed the typical male shape. 

TYPE-LOCALITY.—Mayuku Creek at Ugbekoko 

(= Gbekoko), approximately 10 miles [16km] west 

of Sapele, Midwest State, Nigeria, at 05°54'N, 

05°37'E. This locality was described in detail by 

Powell (1976:315), who also dealt with the various 

animal species found in it. 


D. 31508), a male collected 1-3 November 1975, 

forms part of the collection of the Rijksmuseum 

van Natuurlijke Historie, Leiden; the greater part 

of the paratypes are in the same museum. Series 

of paratypes are also deposited in the National 


Washington, D.C., and in the Koninklijk 

Museum voor Centraal Afrika, Tervuren, Belgium. 

ETYMOLOGY—The specific epithet is given in 

honor of the collector of the material, C. B. 

Powell, University of Port Harcourt, Nigeria, who 

by careful collecting activities, and even more by 

his investigations, has greatly furthered the 

knowledge of West African fresh and brackish 

water Decapoda. 

Genus Tylodiplax De Man, 1895 

FIGURE 58 

Tylodiplax De Man, 1895a:598 [type-species: Tylodiplax tetralylophora 

De Man, 1895, by monotypy; gender: feminine]. 

DIAGNOSIS.—Carapace almost semicircular, 

with anterolateral margins widely rounded and 

long, provided with 2 indistinct teeth of which 

anterior more pronounced than posterior. Posterolateral 

margins of carapace short. Dorsal surface 

of carapace somewhat convex, with 2 transverse 

ridges in anterior half and 4 tubercles in 

posterior half. Epigastric ridges not high, placed 

on base of front. Anterolateral angles of front 

distinct, little produced. Dorsal margin of orbit 

showing no incisions. Ischium and merus of third 

maxilliped covering part of exopod and part of 

palp; merus slightly longer than ischium and not 

auriculate, but with small lobe on anterior margin 

near base of carpus. Chelipeds of adult male 

small, not larger than those of female, and without 

teeth on cutting edges. Third and fourth 

pereiopods larger than second or fifth. First somite 

of male abdomen narrow, not reaching coxae 

of fifth pereiopods; second to fourth somites fused, 

fifth and sixth free. Abdomen of male not constricted 

at fifth somite and in reflexed position 

covering gonopods. Male gonopods strongly recurved, 

ending in somewhat swollen distal parts, 

the one on morphological inner half bearing 

broad lobe and that on morphological outer half 

several strong and some small spines. 

REMARKS.—So far three species have been described 

as belonging in the genus Tylodiplax: T. 

tetratylophora De Man, 1895; T. indica Alcock, 

1900; and 71 derijardi Guinot and Crosnier, 1963. 

Some authors also placed Cleistostoma blephariskios 

Stebbing, 1924, in this genus. As has been explained 

(p. 211), both T. derijardi and C. blephariskios 

are assigned to the genus Paratylodiplax, differing 

from Tylodiplax in several important characters. 

Also Tylodiplax indica has to be removed 

from Tylodiplax; it is here provisionally placed in 

Serenella (p. 211), but its generic position is still 

far from clear, and can only be decided upon 

after examination of additional material. 

The only species thus remaining in the genus 

Tylodiplax is its type-species, T. tetratylophora De 

Man, which is known only from Malaya. 

Subfamily OCYPODINAE Rafinesque, 1815 

Genus Ocypode Weber, 1795 

Ocypode Weber, 1795:92 [type-species: Cancer ceratophthalmus 

Pallas, 1772, by selection by Holthuis, 1962:244, 245; 


Ocypode Fabricius, 1798:312, 347 [invalid junior objective 

synonym and homonym of Ocypode Weber, 1795; typespecies: 

Cancer ceratophthalmus Pallas, 1772, by selection by 

Latreille, 1810:95, 422; gender: feminine; name 1738 on 

Official Index].

218 

a 


FIGURE 58—Tylodiplax tetratylophora De Man: a, male, dorsal view; b, male abdomen; c, carapace, 

dorsal view; d, third maxilliped; e, gonopod;/, tip of gonopod. (a,b, from De Man, 1897, pi. 14: 

fig. 15, 15d; c-f, from Serene and Kumar, 1971, figs. 1, 3-5.) 

Ocypoda Lamarck, 1801:149 [incorrect spelling of Ocypode 

Weber, 1795; name 1737 on Official Index]. 

Monolepis Say, 1817:155 [type-species: Monolepis inermis Say, 

1817, a subjective junior synonym of Cancer quadratic Fa- 

SYNONYM.—Ocypoda hexagonura Hilgendorf, 

1882. 


bricius, 1787, by selection by Fowler, 1912:457; gender: Other Material: Liberia: Grand Cape Mount, near Rob- 

feminine]. 

ertsport, 1880-1882, J. Biittikofer and J. H. Sala, 12 speci- 

Ceratophthalma MacLeay, 1838:64 [type-species: Cancer cursor mens (L). Monrovia, Apr 1894, O. F. Cook and G. N. 

Linnaeus, 1758, by monotypy; gender: feminine]. Collins, 1 juv (W). Ocean beach in front of Camp Johnson, 

Parocypoda Neumann, 1878:26 [type-species: Cancer ceratophMonrovia, 

G. C. Miller, 21 Jul 1952, \6, 19 (W). Mouth of 

thalmus Pallas, 1772, by monotypy; gender: feminine]. Mesurado River, Monrovia, O. F. Cook, 1


Angola: Musserra, P. Kamerman, lectotype, Id "from 

Congo coast" (L, reg. no. D. 235). Musserra, P. Kamerman, 

8 specimens (L). 




(Ghana). 



REMARKS.—De Man (1881:253-255), when describing 

the present species, based it on a male 

(Crust. D. 235) from "Congo" [almost certainly 

Musserra, Angola], collected by P. Kamerman. 

However, De Man also assigned to his new species 

a specimen from Liberia, not seen by him, but 

reported upon by Hilgendorf (1869:81), and 

noted as ZMB 3118. Both specimens are syntypes 

of De Man's species and the one from "Congo" 

is now selected to be the lectotype. Monod (1956: 

395) already indicated that specimen as the 

holotype of Ocypode africana. 

BIOLOGY.—Like all species of Ocypode, 0. africana 

is an inhabitant of sandy beaches, making its 

burrows above the tide line. Gauld and Buchanan 

(1956) and Gauld (1960) found the species "very 

common in sandy ground above high water 

marks, such as coconut groves; juveniles are 

found, with O. cursor, below high water mark but 

adults rarely so" (Gauld, 1960:71). According to 

Gauld and Buchanan (1956:295, 299), 0. africana 

occupies a habitat more distant from the sea than 

0. cursor with, in certain places, "no evidence of 

overlap of the two species . . . . O. africana is quite 

terrestrial in habit, emerging from its burrows at 

night." Longhurst (1958:53), on the other hand, 

found that "in Sierra Leone—at least on the 

beaches examined—both species occur together 

and excavate burrows above H.W. of springs; 

from these, isolated individuals of both species 

emerge during the daytime to feed along the lines 

of foam left by the surf. At night, the beaches are 

crowded with feeding individuals of both species, 

which appear to gather—as in the daytime—at 

the edge of the surf." 

DISTRIBUTION.—West Africa, from southern 

Mauritania to South-West Africa as far as 19° 

23'S. Monod (1956) enumerated the records of 

the species then known. To these the following 

can now be added: 

Senegal: Plage de Camberene, N of Dakar and Plage de 

Bargny Gouddou (as Bargny), E of Dakar (Sourie, 1957). 

Guinea: lies de Los (Uschakov, 1970). 


Liberia: No specific locality (Biittikofer, 1890). 

Ghana: No specific locality (Gauld, 1960). Denu and 

Labadi (Gauld and Buchanan, 1956). 

Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). 

Principe: No specific locality (Forest and Guinot, 1966). 

Congo: Baie de Pointe-Noire (Rossignol, 1957, 1962). 

Zaire: Banana to Vista (Dubois, 1957). 

Angola: Lobito (Bott, 1964). Moc.amedes (Guinot and 


South-West Africa: Rocky Point, 18°59'S, 12°29'E (Penrith 

and Kensley, 1970; Kensley, 1970). Near Kunene River 

mouth, 17°15'S, 11° 45'E; Dunedin Star wreck site, 18°13'S, 

11°56'E; False Cape Frio, 18°29'S, 12°01'E; Westies Mine 

Camp, 19°12'S, 12°37'E; and Mowe Point, 19° 23'S, 

12°42'E (all Kensley, 1970). 

* Ocypode cursor (Linnaeus, 1758) 

lOcypode rhomba.—Pechiiel-Loesche, 1882:287 [not Ocypode 

rhombea Fabricius, 1798]. 

Ocypode cursor.—Hilgendorf, 1879:802.—Biittikofer, 1890: 

465, 487.—Johnston, 1906:862.—Rossignol, 1957:86, pi. 

2: fig. 1.—Bott, 1964:31.—Forest and Guinot, 1966:89.— 

Voss, 1966:30.—Kensley, 1970:180.—Penrith and Kensley, 

1970b:252, 261— Hartmann-Schroder and Hartmann, 

1974:13, 23.—Sakai and Tiirkay, 1977:178. 

Ocypode ippeus.—Monod, 1933b: 548. 

Ocypoda cursor.—Capart, 1951:178, fig. 68.—Monod, 1956: 

391, 632, figs. 552-554.—Dubois, 1957:7, fig. 22.—Rossignol, 

1957:119 [key].—Sourie, 1957:14, 31, 43, 45.— 

Longhurst, 1958:53, 88.—Gauld, 1960:71.—Nicou, 1960: 


119—Ribeiro, 1964:14. 

Ocypoda hippeus.—Gauld and Buchanan, 1956:295, 296, 298, 

301; 1959:127. 

Ocypode.—Voss, 1966:52.—Bayer, 1966:98, 102. 

SYNONYM.—Ocypode ippeus Olivier, 1804. 


Sta 224, Lagos, sand beach, 1

220 

north of Dakar, 3 Dec 1975, W. Bohme, 1


Mesuca Bott, 1973b:316 [type-species: Cancer letragonon Herbst, 

1790, by original designation; gender: feminine]. 

Latuca Bott, 1973b:317 [type-species: Mesuca (Latuca) neocultnmana 

Bott, 1973, by original designation; gender: feminine]. 

Tubuca Bott, 1973b:322 [type-species: Gelasimus urvillei H. 

Milne Edwards, 1852, by original designation; gender: 

feminine]. 

Austruca Bott, 1973b:322 [type-species: Gelasimus annulipes H. 

Milne Edwards, 1837, by original designation; gender: 

feminine]. 

Paraleptuca Bott, 1973b:322 [type-species: Gelasimus chlorophthalmus 

H. Milne Edwards, 1837, by original designation; 


Heteruca Bott, 1973b:323 [type-species: Gelasimus heteropleurus 

Smith, 1870, by original designation and monotypy; gender: 

feminine]. 

Planuca Bott, 1973b:324 [type-species: Uca thayeri Rathbun, 


Leptuca Bott, 1973b:324 [type-species: Gelasimus stenodactylus 

H. Milne Edwards and Lucas, 1843, by original designation; 


Deltuca Crane, 1975:21 [type-species: Gelasimus forcipatus 

Adams and White, 1848, by original designation; gender: 

feminine]. 

Australuca Crane, 1975:62 [type-species: Gelasimus bellator 

Adams and White, 1848, by original designation; gender: 

feminine]. 

Thalassuca Crane, 1975:75 [type-species: Cancer tetragonon 

Herbst, 1790, by original designation; gender: feminine]. 

Amphiuca Crane, 1975:96 [type-species: Gelasimus chlorophthalmus 

H. Milne Edwards, 1837, by original designation; 


Borboruca Crane, 1975:109 [type-species: Uca thayeri'Rathbun, 

1900, by original designation and monotypy; gender: 

feminine]. 

Afruca Crane, 1975:116 [type-species: Gelasimus tangeri 

Eydoux, 1835, by original designation and monotypy; 


Celuca Crane, 1975:211 [type-species: Uca deichmanni Rathbun, 


REMARKS.—Latreille (1817b:5l7), when introducing 

the generic name Gelasimus, cited it as 

"Gelasimus (Buffon)." This caused Stebbing 

(1905:40) to remark: "Latreille at the first institution 

of Gelasimus attributed the genus to Buffon, 

though in 1820 he claims it as his own. He gave 

no reference for the name to any part of Buffon's 

works, and no such reference has since been discovered." 

Also Crane (1975:20) remarked: "I 

have been unable to trace the use of the word by 

Buffon." The solution of this problem may be 

that Latreille did not intend to refer to George 

Louis Leclerc, comte de Buffon, but that the word 

"Buffon" is merely cited as a translation of the 

latin word "Gelasimus," which means "buffoon," 

"jester," or "mocker." 

* Uca tangeri (Eydoux, 1835) 

Cancer Uka una, Brasiliensis Seba, 1759:44, pi. 18: fig. 8. 

Cancer vocans major Herbst, 1782:83, pi. 1: fig. 11. 

Ocypoda heterochelos Lamarck, 1801:150.—Bosc, 1802:197.— 

Desmarest, 1830:250. 

Cancer Uka Shaw and Nodder, 1803, pi. 588. 

Uca una Leach, 1814:430. 

Gelasimus Tangeri Eydoux, 1835, pi. 17. 

Gelasimus />«7a/uj.—Hilgendorf, 1879:806.—De Man, 1879: 

66.—Pechiiel-Loesche, 1882:288.—Biittikofer, 1890:464, 

487.—Aurivillius, 1893:31.—Johnston, 1906:862. 

Uca tangeri.—Maccagno, 1928:33, fig. 19.—Frade, 1950:11, 

26.—Capart, 1951:180, fig. 69.—Monod, 1956:399, figs. 

559, 560.—Dubois, 1957:7, fig. 23.—Rossignol, 1957:86, 

119 [key].—Sourie, 1957:14, 50.—Longhurst, 1958:53, 

88.—Gauld, 1960:71.—Guinot-Dumortier and Dumortier, 

1961:136.—Nicou, 1960:135-156, figs. 1-4.—Guinot 

and Ribeiro, 1962:67.—Rossignol, 1962:119.—Forest and 

Guinot, 1966:90.—Monod, 1967:180, pi. 17: fig. 4.—Uschakov, 

1970:448, 455, fig. 4.—Bright and Hogue, 1972: 

13.—Bott, 1973a:311-314, figs. 1, 3; 1973b:316, fig. 1.— 

Hartmann-Schroder and Hartmann, 1974:19.—Pauly, 

1975:57.—Powell, 1979:127. 

Gonoplax speciosus Monod, 1933b:548 [footnote; nomen nudum]. 

Gelasimus (Uca) tangeri.—Bruce-Chwatt and Fitz-John, 1951: 

117 [also Gelasimus tangeri, pp. 117 and 119; and Gelasimus 

Tangery, p. 119]. 

Uca tangeri typique Monod and Nicou, 1959:988, figs. 2, 4, 5. 

Uca tangeri matandensis Monod and Nicou, 1959:988, figs. 1, 

3,6. 

Uca (Minuca) tangeri.—Bott, 1968:168. 

Uca (Afruca) tangeri.—Crane, 1975:118, figs. 27D-F, 37E, 

45E-I, 45EE-II, 46F, 63D, 81E, 82F, 99, pi. 18A-D. 

SYNONYMS.—Gelasimus perlatus Herklots, 1851; 

Gelasimus cimatodus De Rochebrune, 1883. 

MATERIAL EXAMINED—Pillsbury Material: Nigeria: Sta 

1, Lagos harbor, shore, 26, 19 (W). 

Other Material: Morocco: Tangier, H. Milne Edwards, 

?type-material, lc?, 19 (dry, L). 

Senegal: Dakar, 1887, J. Biittikofer. 1$ (L). Dakar, 3 May 

1882, O. F. Cook, 26, 1? (W). 

Liberia: Grand Cape Mount near Robertsport, 1882, J.


Biittikofer, 4(5, 29 (L). Monrovia, Apr 1894, O. F. Cook and 

G. N. Collins, New York State Colonization Society, 16* (W). 

Rock Spring, Monrovia, Apr 1894, O. F. Cook and G. N. 

Collins, 1(5 (W). Mouth of Mesurado River, Monrovia; O. 

F. Cook: 3(5 (W). Tive's farm, Bushrod Island, Monrovia, 16 

Nov 1953, G. C. Miller, 2(5 (W). 

Ghana: Locality not specified, 1840-1855, H. S. Pel, 

paralectotypes of Gelasimus perlatus Herklots, 46* (L). Butre 

(04°50'N, 01°56'W), 1841-1850, H. S. Pel, types oiGelasimus 

perlatus Herklots, 16* lectotype, 3$ paralectotypes (L; lectotype 

Crust. D. 262). Elmina, 27 Nov 1889, U. S. Eclipse 

Expedition, 36*, 4$ (W). Accra, 1868-1869, M. Sintenis, 6(5, 

4$ (L). 

Nigeria: Lagos, Apr 1964, F. Kliige, 26*, 1? (L). Tarkwa 

Bay, Lagos, Oct 1957, J. Crane, 29(5, 179 (W). S bank of 

Escravos River near Ajudaibo, Niger delta, 05°34.5'N, 

05° 11.75'E, 20 Jul 1975, C. B. Powell, 26* (L). Wof Forcados 

near confluence of Odimodi Creek and Forcados River, 

05°22'N, 05°26'E, 28 Feb 1976, C. B. Powell, 15 specimens 

(L). Between Brass and Port Harcourt, Niger delta, May- 

Aug 1960, H. J. G. Beets, 12 specimens (L). 

Cabinda: Chinchoxo (= Quinchoxo, = Tschintschotscho, 

05°09.24'S, 12°03.75'E) (see Holthuis and Manning, 1970: 

250, 251), 1873-1876, J. Falkenstein, don. Mus. Berlin, 1(5 

(W). "Quilla Miindung" (= mouth of the Quila River, 

05°58'S, 14°47'E), 1873-1876, J. Falkenstein, don. Mus. 

Berlin, 1(5 (W). 

Zaire: Congo, Jan 1895, H. C. Kooiman, 19 (L). Banana, 

mouth of the Congo River, Aug 1915, H. Lang, American 

Museum Congo Expedition, 10(5, 99 (W). Banana, R. I. 

Meyer, 26*, 19 (L). 

Angola: Musserra, 1882, P. Kamerman, 20 specimens (L). 

Santo Antonio do Zaire, mouth of the Congo River, Aug 

1915, H. Lang, American Museum Congo Expedition, 1(5 

(W). Luanda (= St. Paul do Loanda), 11 Dec 1889, U. S. 

Eclipse Expedition, 16* (W). Samba, Luanda, Sep 1957, J. 

Crane, 686*, 349, 33 juv (W). Ilha de Cabo, Luanda, Sep 

1957, J. Crane, 946*, 989 (W). Luanda, 18 June 1967, G. 

Hartmann, 26*, 19 (L). Morro da Cruz, between Luanda and 

Cuanza, 23 km S of Luanda, 20 Jun 1967, G. Hartmann, 

26", 19 (L). Baia Farta, near Benguela, 3 Jul 1967, G. 

Hartmann, 16*, 19 (L). 

DESCRIPTION.—Crane, 1975:118-124. 

Figures: Capart, 1951, fig. 69; Monod, 1956, 

figs. 559, 560; Bott, 1973a, figs. 1, 3; Crane, 1975, 

figs. 27D-F, 37E, 45E-1,45EE-II, 46F, 63D, 8IE, 

82F, 99, pi. 18A-D. 

Male Pleopod: Bott, 1973b, fig. 1 (Zaire); Crane, 

1975, fig. 63D (no locality). 

MEASUREMENTS.—The examined males have 

the carapace width between 13 and 42 mm, the 

females between 12 and 28 mm. 

REMARKS.—Bott (1973a) very convincingly 

showed that the specimen which Seba (1759) 

showed on his figure 8 of plate 18 as "Cancer Uka 

una, Brasiliensis" belongs to the species that is best 

known as Uca tangeri (Eydoux, 1835), and not to 

the American species which Rathbun (1918:381) 

in her monograph of the American grapsoid crabs 

had indicated with the name Uca heterochelos (Lamarck, 

1801), Lamarck's species being based on 

Seba's figure. Holthuis (1959b:277; 1962:239, 

240) rejected the specific epithet heterochelos Lamarck, 

1801, and replaced it by its senior objective 

synonym major Herbst, 1782; he used the 

name Uca major (Herbst) for the American species, 

accepting Rathbun's interpretation of the identity 

of Ocypoda heterochelos Lamarck, and thus of 

Seba's specimen. Holthuis (1962:240) selected the 

specimen figured by Seba (1759, pi. 18: fig. 8) as 

the lectotype of Cancer vocans major Herbst, 1782, 

Ocypoda heterochelos Lamarck, 1801, and Uca una 

Leach, 1814, and thereby made these species 

objective synonyms of each other. As Uca una 

Leach, 1814, is the type-species (by monotypy) of 

the genus Uca Leach, 1814, the correct name of 

that type-species becomes Uca major (Herbst, 

1782). 

Bott's (1973a) discovery that the epithets major 

Herbst, 1782, heterochelos Lamarck, 1801, and una 

Leach, 1814, do not pertain to the American 

species (as Rathbun and most later authors 

thought), but to the European and West African 

Uca tangeri (Eydoux), would cause considerable 

confusion in the nomenclature of the group, unless 

action by the International Commission on 

Zoological Nomenclature prevents this. Bott 

(1973a) showed that the oldest available name 

for the American species is Gelasimus platydactylus 

H. Milne Edwards, 1837, and he suggested that 

henceforth this species be known as Uca platydactylus 

(H. Milne Edwards, 1837). This change in 

itself would not have been disturbing, as the 

species is not a very common one and had been 

known under several names before 1918 when 

Miss Rathbun's monograph stabilized the use of 

the name Uca heterochelos for it; not until 1959 was 

the epithet major reintroduced for it. However,


the name Uca major is now generally used for it 

and is also adopted in the recent monumental 

monograph of the genus Uca published by Crane 

(1975:136). 

If the change of the name Uca major sensu 

Crane to Uca platydactylus (H. Milne Edwards) is 

somewhat disturbing, this is very much so for the 

other nomenclatural change that results from 

Bott's discovery. The species which at present is 

best known as Uca tangeri (Eydoux, 1835) under 

the strict application of the International Code of 

Zoological Nomenclature should have to be given the 

name Uca major (Herbst, 1782). This species occurs 

on the extreme southwest coast of the Iberian 

peninsula and in West Africa as far south as 

Angola. Since 1835 the epithet tangeri (also spelled 

tangieri) has been mostly used for it and since 1900 

this usage has been unanimous. Uca tangeri is a 

far better known species than U. platydactylus, and, 

being the only species of Uca occurring in Europe, 

its behavior has been studied by many European 

zoologists and the ethological literature concerning 

it is extensive (see Crane, 1975:124, for a 

listing of this literature). The change of the epithet 

tangeri to major would be most undesirable, 

the more so as the epithet major at present is 

regularly in use for a different species. Also the 

fact that Crane adopted the epithet tangeri in her 

fundamental monograph, which will be the basis 

for all taxonomic and ethological studies of Uca 

by generations of biologists to come, is a most 

important argument for not changing this name. 

Dr. Bott (1973a:313, 314) was of the opinion that 

the names Cancer vocans major Herbst, Ocypoda heterochelos 

Lamarck, and Uca una Leach could be 

considered "nomina oblita" and should be conveniently 

ignored. This is not true, however. In 

the first place, only Uca una Leach under the Code 

before it was revised in 1972 (at the International 

Congress of Zoology held in Monaco) could qualify 

as a nomen oblitum, but certainly not the 

other two names, as those have been repeatedly 

used in the last 50 years (be it for a species 

different from U. tangeri, which, however, is irrelevant 

here). Secondly the present revised Code 

does not allow rejection of so-called "nomina 

oblita" without a definite action by the International 

Commission on Zoological Nomenclature. 

As far as we can see, there are two ways open 

to save the use of the name Uca tangeri for the 

present species: 

1. To request the International Commission 

on Zoological Nomenclature to use its plenary 

powers to suppress the names Cancer vocans major 

Herbst, 1782, Ocypoda heterochelos Lamarck, 1801, 

Cancer uka Shaw and Nodder, 1803, and Uca una 

Leach, 1814. This is a complicated procedure and 

is especially inadvisable as the names are those of 

the type-species of the genus Uca Leach. The 

results of such an action by the Commission 

would be that the valid name for the African 

species would remain Uca tangeri, while that of the 

American species would become U. platydactylus. 

The Commission then also has to decide which of 

these two species (or any other) should be made 

the type of the genus Uca. 

2. To request the International Commission to 

indicate under their plenary powers a specimen 

of Uca platydactylus (H. Milne Edwards) to be the 

neotype specimen of Cancer vocans major Herbst, 

1782. In this way the name Uca major (Herbst) 

would become the valid name for Uca platydactylus 

and the name Uca tangeri would become that of 

the African species. This action would preserve 

the current usage of both names and legalize the 

names used by Crane (1975) for the two species. 

Furthermore, no change in the name of the 

type-species of the genus Uca is then necessary. 

The second of these two actions seems to us to 

be preferable, and an application has been submitted 

on these lines to the International Commission 

on Zoological Nomenclature. In it the 

Commission is requested to use its plenary powers 

to validate the selection of the type-specimen of 

Uca platydactylus (H. Milne Edwards) to be the 

neotype of Cancer vocans major Herbst, 1782. This 

specimen, a full grown male from Cayenne, now 

forms part of the collection of the Museum national 

d'Histoire naturelle in Paris (see Crane, 

1975:601). In case H. Milne Edwards' (1837) 

original material of Gelasimus platydactylus consisted 

of more than one specimen, the type-spec-


imen examined by Crane (1975) is made the 

lectotype of that species. Of all the specimens of 

Uca platydactylus known at present, the neotype of 

Cancer vocans major Herbst originates from a locality 

(Cayenne) that is closest to the originally 

indicated type-locality of Herbst's species, viz. 

Brazil. By this neotype selection Cancer vocans major 

Herbst, 1782, Ocypoda heterochelos Lamarck, 1801, 

Cancer uka Shaw and Nodder, 1803, Uca una Leach, 

1814, and Gelasimus platydactylus H. Milne Edwards, 

1837, become objective synonyms. 

Bott's (1973a) discovery that Seba's specimen 

is identical with Uca tangeri also has consequences 

for the nomenclature of the subgenera of Uca. In 

1968 Bott placed Uca tangeri in the subgenus 

Minuca Bott, 1954, but in 1973 he (Bott, 1973b: 

316) split the old genus Uca into 10 genera (7 

new), one of which consisted of 2 subgenera (both 

new). The genus Uca was restricted to two species: 

Uca tangeri (Eydoux, 1835) and Uca marionis (Desmarest, 

1825) [= Uca vocans (Linnaeus, 1758)]. In 

Crane's (1975) monograph of Uca, all the fiddler 

crabs were still placed in a single genus, Uca, 

within which Crane recognized 9 subgenera (7 of 

which were given new names by her). Uca tangeri 

was placed by itself in a monotypic subgenus 

Afruca Crane, 1975. The limits of Crane's subgenera 

in many instances do not coincide at all 

with those of Bott's genera and subgenera. It 

furthermore is most unfortunate that Bott's 

(1973b) preliminary paper was published before 

Crane's (1975) monograph, the manuscript of 

which was in press too early to take either Bott's 

system into consideration or to adopt his new 

names. Bott's final revision of the genus Uca sensu 

lato was never published, as he died in 1974 

before finishing it. We have now the most unpleasant 

situation that Crane's subgenera are well 

defined and exhaustively treated in an ideal way, 

while Bott's names, published in a short, not too 

well-documented paper, have priority. Von 

Hagen (1976:223), in his review of Crane's monograph, 

has already pointed to this unfortunate 

nomenclatural situation and stated: 

Because of their being objective synonyms Crane's names 

Thalassuca, Amphiuca and Borboruca must be replaced by Bott's 

Mesuca, Paraleptuca and Planuca, respectively. When accepting 

Crane's limits of species groups, one has also to use Tubuca 

Bott instead of Deltuca Crane and should use Leptuca (not 

Austruca) Bott instead oiCeluca Crane. Only Australuca Crane 

and (provided the shift of the type species name in consequence 

of Bott, 1973a, is avoidable) Afruca Crane remain 

valid. 

Unfortunately, in practice (though not in theory) a chaos 

of names seems inevitable. Many non-taxonomists, taking 

Crane's monograph as the sound reference work that it 

actually is, will unsuspiciously use her new but invalid 

subgeneric names, while taxonomists would, of course, have 

to use Bott's names (regardless, whether on a subgeneric or 

generic level). 

If the International Code of Zoological Nomenclature 

is strictly applied, the name of the present species 

would be Uca (Uca) major (Herbst, 1782); if Bott's 

system is followed it should be named Uca tangeri 

(Eydoux, 1835); and under Crane's system it is 

Uca (Afruca) tangeri (Eydoux, 1835). Pending the 

decision of the International Commission on Zoological 

Nomenclature on this case, the well 

known name Uca tangeri (Eydoux) is used here. 

We also follow Von Hagen's (1976) suggestion 

not to use subgeneric names for the time being. 

Another change, in both Crane's (1975) and 

Bott's (1973b) nomenclature is needed. If Crane's 

system is adopted, then the name Thalassuca 

Crane, 1975, should not be replaced by Mesuca 

Bott, 1973, as Von Hagen (1976) thought, but by 

Gelasimus Latreille, 1817, because the type-species 

of Gelasimus, Cancer vocans Linnaeus, 1758, belongs 

in Crane's Thalassuca (see Crane, 1975:20). Under 

Bott's classification Gelasimus Latreille, 1817, must 

fall as a junior synonym of Uca Leach, 1814, and 

for the genus that Bott (1973b:323) indicated 

with the name Gelasimus, the generic name Acanthoplax 

H. Milne Edwards, 1852, would become 

available. 

DISTRIBUTION.—The species is found in the 

eastern Atlantic, its range extending from the 

south coast of Portugal to southern Angola; it 

does not occur in the Mediterranean. The records 

from America (West Indies, Brazil; cf. Rathbun, 

1918:388 and Bott, 1973a) in all likelihood are 

erroneous, due to incorrect labeling. Monod 

(1956) enumerated the localities from which the


species was then known. Since that time it has 

been reported from the following West African 

localities (including those overlooked by Monod; 

Crane's (1975) records are not duplicated here): 

Senegal: Saloum (Sourie, 1957). Ngor, near Dakar (Nicou, 

1960). 

Guinea-Bissau: Bissau (Maccagno, 1928; Frade, 1950). 

Guinea: lie Marara, mouth of Rio Pongo (Uschakov, 

1970). Boulbinet, Tanene, and Ralompa, all near Conakry 

(Monod and Nicou, 1959). 

Liberia: No specific locality (Hilgendorf, 1879; Biittikofer, 

1890; Johnston, 1906). 

Sierra Leone: Robene Point and Cline Bay (as Kline) 


Ivory Coast: Lagoon of Abidjan, 05°16'12"N, 04°00'- 

20"W (Forest and Guinot, 1966). 

Ghana: "Coast of Guinea" (De Man, 1879). Estuaries of 

Volta and Ankobra rivers (Gauld, 1960). Sakumo Lagoon 

(Pauly, 1975). 

Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). Elechi 

Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell, 

1979). 

Cameroon: No specific locality (Aurivillius, 1893). Douala 

(Monod and Nicou, 1959). 

Principe: No specific locality (Maccagno, 1928; Forest 

and Guinot, 1966). Praia Ponta da Mina and Santo Antonio 


Sao Tome: No specific locality (Maccagno, 1928; Forest 


Congo: No specific locality (Maccagno, 1928). Loango 

(Pechiiel-Loesche, 1882; Rossignol, 1957). Mouth of Songololo 

River (Rossignol, 1957, 1962). Djeno (Rossignol, 1957). 

Cabinda: Chinchoxo (Hilgendorf, 1879). 

Zaire: Banana (Dubois, 1957; Bott, 1973a,b). 

Angola: Luanda (Hilgendorf, 1879; Bott, 1968). Baia de 

Luanda, Lobito; Baia Farta, Benguela; and Baia dos Tigres 

(Guinot and Ribeiro, 1962). Between Cacuaco and Lobito- 

Benguela (Hartmann-Schroder and Hartmann, 1974). 

Family GRAPSIDAE MacLeay, 1838 

GRAPSIDAE MacLeay, 1838:63, 65. 

SESARMINAE Dana, 1851c:288. 

PLAGUSINAE Dana, 1851c:288 [corrected to Plagusiinae by 

Miers, 1878:147; name 377 on Official List]. 

VARUNACEA H. Milne Edwards, 1853:175 [corrected to Varuninae 

by Alcock, 1900:288, 400]. 

CYCLOGRAPSACEA H. Milne Edwards, 1853:191. 

EASTERN ATLANTIC GENERA.—Thirteen, 12 of 

which are represented by species occurring from 

localities between Mauritania and Angola. The 

other genus is: Eriocheir de Haan (1835:32), typespecies: 

Grapsus (Eriocheir) japonicus de Haan, 

1835, by selection by H. Milne Edwards (1854: 

146); gender: masculine. 

EASTERN ATLANTIC SPECIES.—Twenty-five, 19 

of which occur in tropical waters (the included 

Atlantic species of Brachynotus is not known to 

occur south of Mauritania so is not strictly tropical). 

The other species are as follows: 

Brachynotus foresti Zariquiey Alvarez, 1968. 

Mediterranean; littoral (Zariquiey Alvarez, 1968; 

Froglia and Manning, 1978). 

Brachynotus gemmellari (Rizza, 1839). Mediterranean; 

sublittoral (Froglia and Manning, 1978). 

Brachynotus sexdentatus (Risso, 1827). Mediterranean; 

introduced into Great Britain in artificially 

heated water at Swansea; northern part of 

Suez Canal and Black Sea (Zariquiey Alvarez, 

1968; Froglia and Manning, 1978). 

Eriocheir sinensis H. Milne Edwards, 1853. A 

native of China, introduced into NW Europe and 

now reported from Finland, Sweden, and Norway 

to England and the Atlantic coast of France, in 

rivers (Christiansen, 1969); and from Lake Erie, 

North America (Nepszy and Leach, 1973). 

Pachygrapsus marmoratus (Fabricius, 1787). Mediterranean 

and Atlantic coast of Europe from 

Brittany (France) southward to Morocco, including 

Madeira, the Azores, and the Canary Islands; 

intertidal (Zariquiey Alvarez, 1968). 

Pachygrapsus maurus (Lucas, 1846). Western 

Mediterranean, Madeira, the Azores and the Canary 

Islands; intertidal (Zariquiey Alvarez, 1968). 

Four species of Grapsidae at one time or another 

reported from West Africa in all probability 

do not belong to the West African fauna, and the 

material on which these records are based either 

is incorrectly identified or is labeled with a wrong 

locality. These species are as follows: 

Metopograpsus messor (Forskal, 1775). A species 

known with certainty from the Red Sea, the east 

coast of Africa, Madagascar and the Persian Gulf 

(Holthuis, 1977a). Monod (1956:422, 423) listed 

the records of this species from West Africa and 

indicated that most probably they were based on 

incorrect identifications. There was, however, a

226 

correctly identified specimen of Metopograpsus messor 

in the collection of the Museum national 

d'Histoire naturelle, Paris, labeled "Gabon," the 

correctness of which was doubted by Monod as 

this same lot included a specimen of another 

species, Pseudograpsus elongatus (see below), which 

also occurs only in the Indo-West Pacific region. 

Platychirograpsus spectabilis De Man, 1896. This 

species was originally described by De Man 

(1896:292, fig. 1) from Gabon. Apart from the 

types of this very characteristic species, no material 

has ever been reported from West Africa, 

whereas it has been found repeatedly in fresh 

water on the E coast of Mexico and also has been 

introduced into Florida (Marchand, 1946:93- 

100). As pointed out by Monod (1956:426-428) 

there is a definite possibility that De Man's material 

did not originate from West Africa but from 

Mexico. 

Pseudograpsus elongatus (A. Milne Edwards, 

1873). A well known species from the Indo-West 

Pacific (Red Sea, E Africa, Madagascar, the Seychelles, 

and New Caledonia) (Holthuis, 1977a), 

but not known from West Africa other than from 

the specimen labeled as originating from Gabon 

and found together with Metopograpsus messor (see 

above). 

Sesarma roberti H. Milne Edwards, 1853. Originally 

described from Goree, Senegal, but otherwise 

only reported from the West Indies and the 

Atlantic coasts of Central and South America. As 

stated by Monod (1956:443) and Chace and 

Hobbs (1969:184), the type-locality indication is 

very likely erroneous, so that this species, as well 


Subfamily GRAPSINAE MacLeay, 1838 

Genus Geograpsus Stimpson, 1858 

Geogmpsus Stimpson, 1858b: 101 [p. 47 on separate] [typespecies: 

Grapsus lividus H. Milne Edwards, 1837, by subsequent 

designation by Rathbun, 1918:231; gender: masculine]. 

Orthograpsus Kingsley, 1880a: 188, 194 [type-species: Orthograpsus 

hillii Kingsley, 1880, a subjective junior synonym 

of Grapsus lividus H. Milne Edwards, 1837, by present 

selection; gender: masculine]. 

* Geograpsus lividus (H. Milne Edwards, 1837) 

Geograpsus livtdus.—Monod, 1956:410, figs. 562, 563.—Dubois, 

1957:7.—Guinot and Ribeiro, 1962:69.—Ribeiro, 

1964:15.—Forest and Guinot, 1966:91. 

SYNONYMS.—Grapsus brevipes H. Milne Edwards, 

1853; Orthograpsus hillii Kingsley, 1880. 


Sta 1, Lagos harbor, shore, 2$ (L). 

Other Material: Zaire: Banana, mouth of the Congo River, 

Aug 1915, H. Lang, American Museum Congo Expedition, 

2c5, 2$ (W). 

Angola: Lobito, P. Kamerman, 1$ (L). 

DESCRIPTION.—Rathbun, 1921:442; Chace and 

Hobbs, 1969:157. 

Figures: Rathbun, 1921, pi. 15: fig. 1, pi. 22: 

figs. 2, 3; Monod, 1956, figs. 562, 563; Chace and 

Hobbs, 1969, fig. 48. 

Male Pleopod: Monod, 1956, fig. 563 (Senegal); 

Chace and Hobbs, 1969, fig. 52a-c (West Indies). 

MEASUREMENTS.—The females collected by the 

Pillsbury have the carapace width 12 and 13 mm. 

DISTRIBUTION.—The species inhabits the tropi- 

as Metopograpsus messor, Pseudograpsus elongatus, and cal Atlantic, and has also been reported from the 

Platychirograpsus spectabilis, had best be removed eastern Pacific. In the Indo-West Pacific area it 

from the list of the West African Brachyura. is represented by the closely related Geograpsus 

Other than the subgeneric names of the West 

stormi De Man. In West Africa the species is 

African Sesarma , our resurrection of Goniopsis pelii 

known from the Cape Verde Islands and Senegal 

to Angola and Sao Tome. Monod (1956) listed 

from the synonymy of Goniopsis cruentata, and our 

all West African records known to him, to which 

use of Metagrapsus instead of Sarmatium for S. 

now the following can be added: 

curvatum, the grapsid names used by Monod 

(1956) have not changed so we have not listed Cape Verde Islands: Bafa das Gatas, Sao Vicente (Guinot 

them separately for this family. 

and Ribeiro, 1962; Ribeiro, 1964). Sao Tiago (Forest and 

Guinot, 1966).


Fernando Poo: No specific locality (Forest and Guinot, 

1966). 

Annobon: No specific locality (Forest and Guinot, 1966). 

Zaire: No specific locality (as Belgian Congo) (Dubois, 

1957). 

Angola: Baia Farta, Benguela; Baia da Caota, Benguela; 

Baia de Santa Maria; and Praia Amelia, Mocamedes (all 


Genus Goniopsis de Haan, 1833 

Goniopsis de Haan, 1833:5 [genus established without included 

nominal species; the first nominal species to be 

assigned to the genus, by de Haan (1835:33), are Cancer 

strigosus Herbst, 1799, Grapsus pictus Latreille, 1803, and 

Grapsus cruentatus Latreille, 1803; type-species: Grapsus 

cruentatus Latreille, 1803, by selection by Rathbun, 1918: 


* Goniopsis pelii (Herklots, 1851) 

FIGURE 59 

Grapsus (Grapsus) Pelii Herklots, 1851:8, 23, pi. 1: figs. 6, 7. 

Grapsus (Grapsus) simplex Herklots, 1851:9, 23, pi. 1: fig. 8. 

Grapsus (Goniopsis) cruentatus.—Won Martens, 1872:105, 106 

[not Grapsus cruentatus Latreille, 1803]. 

Grapsus Pelii.—De Man, 1879:68; 1900, pi. 2: fig. 6 [Peli on 

pp. 43-46, 64]. 

Grapsus simplex.—Be Man, 1879:68.—Miers, 1886:255.—De 

Man, 1900:43, pi. 2: fig. 7. 

Goniopsis cruentatus.—Kingsley, 1880a: 190.—Studer, 1882: 

333 [listed].—Biittikofer, 1890:487.—Thallwitz, 1891: 

52.—Benedict, 1893:538.—Doflein, 1900:142.—Johnston, 

1906:862.—Vilela, 1949:63.—Frade, 1950:11, 26 [not 

Grapsus cruentatus Latreille, 1803]. 

Grapsus pelii.—Kingsley, 1880a: 190 [in synonymy]. 

Pachygrapsus simplex.—Kingsley, 1880a:201. 

Goniograpsus cruentatus.—Osorio, 1887:227; 1888:191; 1889: 

130, 134, 139; 1890:46; 1892:199; 1895a:54; 1895b:57, 58; 

1898:193 [not Grapsus cruentatus Latreille, 1803]. 

Grapsus Peli.—Dc Man, 1900:43, 64, pi. 2: fig. 6 [Pelii on 

plate]. 

Goniopsis cruentata.—Rathbun, 1900a:278.—Nobili, 1906b: 

311.—Rathbun, 1918:237; 1921:443, pi. 39.—Balss, 1922: 

80.—Monod, 1927:620; 1928:124.—Irvine, 1947:292, fig. 

197.—Bruce-Chwatt and Fitz-John, 1951:117.—Capart, 

1951:183, fig. 7.—Sourie, 1954a:84, 112, 293, 297, 306.— 

Monod, 1956:412, figs. 564-567.—Rossignol, 1957:89, 

121.—Longhurst, 1958:88.—Gauld, 1960:71.—Rossignol, 

1962:119.—Forest and Guinot, 1966:91—Uschakov, 

1970:443, 455 [listed].—Pauly, 1975:57. [Not Grapsus cruentatus 

Latreille, 1803.] 

Gonopsis cruentatus.—J. Roux, 1927:237 [not Grapsus cruentatus 

Latreille, 1803]. 

Goniopsis cruenta.—Hartmann-Schroder and Hartmann, 

1974:19 [erroneous spelling]. 

Goniopsis pelii.—Powell, 1979:127. 

Not Goniograpsus cruentatus.—Osorio, 1892:199 [=Grapsus grapsus 

(Linnaeus), see Forest and Guinot, 1966:90]. 

Not Pachygrapsus simplex.—Doflein, 1904:129.—Lenz, 1910: 

125.—Lenz and Strunck, 1914:283 [= P. gracilis (Saussure), 

see Balss, 1922:81]. 




specimens varied from 16 to 49 mm. There is no 

obvious size difference in males and females. 

Ovigerous females have carapace widths between 

25 and 47 mm. The male lectotype of G. pelii has 

a carapace width of 48 mm, the male lectotype of 

G. simplex one of 20 mm. 

REMARKS.—A comparison of the present West 

African material with specimens of Goniopsis cruentata 

from the north coast of South America and 

the Caribbean shows some differences, which, 

although small, seem to be constant and induce 

us to distinguish the two populations as distinct 

species. A closer study based on much more material 

and taking also the West American G. 

pulchra into account may establish that the three 

should be considered subspecies of a single species. 

Goniopsis pelii differs from G. cruentata in the 

following points. 

1. The front is relatively shorter and wider. 

The anterior margin of the front not only is 

emarginate in the middle, but in the middle of 

each half as well; this secondary emargination is 

less distinct in G. cruentata. The surface of the front 

bears coarse tubercles, some of which, near the 

anterior margin, form more or less distinct transverse 

ridge-like rows; in G. cruentata such transverse 

ridges, if present, are usually short and 

inconspicuous. This latter character, which was 

already described by Von Martens (1872), however, 

is not very constant. 

2. The pereiopods in G. pelii have the propodi 

more slender than in G. cruentata, and the dactyli 

are more robust. In G. pelii the length of the 

dactylus of the last pereiopod (Figure 59b) is 

distinctly less than that of the lower margin of 

the propodus, while in G. cruentata (Figure 59


FIGURE 59.—Goniopsispelii (Herklots), ovigerous female, Cameroon: a, dorsal view; b, propodus 

and dactylus of fifth pereiopod. Goniopsis cruentata (Latreille): c, propodus and dactylus of fifth 

pereiopod. 

the exception of the third maxillipeds, which are 

pale with a dark purple spot on the carpus and in 

the distal part of the merus. The distal four 

somites of the abdomen are whitish; the basal 

three are pale purple. 

The specimens from Nigeria in which the color 

is well preserved show the same color pattern, 

although the color itself is more reddish. Also the 

Liberian material agrees well with this description. 

In most other specimens the color has faded, 

but in a few some traces of the original pattern 

are visible, agreeing, as far as can be ascertained, 

with the above description. 

The color pattern of the West African speci- 

mens differs strikingly from that of G. cruentata, of 

which an excellent description is provided by 

Chace and Hobbs (1969:161), who also provided 

a beautiful figure of the color pattern. In Goniopsis 

cruentata (1) the carapace is lighter, instead of 

having small light spots on a purple background, 

it shows small purple or red spots on a pale 

background; (2) instead of having a pale streak 

along the posterolateral margin, it shows in the 

posterolateral area a number of characteristic 

white spots that are surrounded by a dark purple 

or red ring; (3) the eyestalks are marbled with 

purple or red instead of being uniformly colored; 

(4) the carpus of the cheliped is "scarlet with

230 

purple lines" (Chace and Hobbs, 1969:161) and 

does not show the bright white tubercles of G. 

pelii; (6) the merus of the chelipeds is of a uniform 

red color inside or orange red with darker marblings 

and shows on the outer surface conspicuous 

large pale spots surrounded by a dark red ring; 

(7) the pereiopods are paler than in G. pelii, the 

upper margins of merus, carpus, and propodus 

are blocked with dark and light color; the dactylus 

is not conspicuously paler than the propodus. 

The merus has conspicuous white spots with 

dark rings around them. 

Only very few authors give some information 

on the color of the present species. Herklots (1851: 

8) described the carapace as "ruber, viridi flavo 

marmoratus." Von Martens (1872:106) when 

dealing with a specimen from Liberia observed 

that the coloration of the carapace was more 

"kleinfleckig" than in his American material. De 

Man (1900:43) stated that the types of Grapsus 

pelii at that time had still "une belle couleur 

rouge jaunatre, moins intense sur les pattes anterieures." 

Rathbun (1921:443) published the following 

color description of a living animal "A 

very showy crab, distinguished by its colors; carapace 

brownish yellow or brick-red; legs red, with 

spots of a darker red, extremities yellow. Chelipeds 

red, except the palms, which are almost 

white, and the fingers, which are yellow." Irvine 

(1947:292) described the colors as follows: "The 

carapace is dark reddish black. . . . The eyes are 

black. The claws are pale, with yellowish tips, 

their upper segments being of a beautiful purple 

colour. The legs are dark red in colour and are 

flat and covered with long stiff bristles." 

Capart (1951:183) gave the following account 

"Couleurs vives, carapace brun-rouge; pattes 

rouges, de meme que les chelipedes, sauf les 

pinces, qui sont jaune clair." Unfortunately most 

of these authors paid more attention to the colors 

than to the color pattern of their animals, while 

the latter is far more important. The color description 

provided by Rossignol (1957:89) is far 

superior to most others. It is as follows: 

Face dorsale: Carapace noire avec des points jaunes tres 

petits et tres serres, sauf sur les regions branchiales. Une 


ligne mediane courte de couleur jaune sur la partie posterieure 

de la carapace (ligne continuee par une serie de points 

de meme couleur sur chaque segment abdominal). Une ligne 

blanche sur chaque bord lateral. Deux taches rouges entre la 

region gastrique et la region cardiaque. 

Pinces: face externe jaune clair a vert amande. Face 

interne: violacee. Sur le bord superieur de la main et du 

doigt, une serie de tubercules blancs ou jaunatres. 

Pattes: brun fonce, noir, avec une tache rouge a chaque 

articulation. Dactyle rouge. 

Face ventrale: jaunatre. Les deux premiers segments abdominaux 

violaces. Mxp. 3 blancs, tranchant sur le brun 

sale des regions pterygostomiales. [The species is also] tres 

facilement reconnaissable avec ses tubercules blancs sur le 

bord superieur des pinces. 

This description, obviously made after living 

specimens, agrees excellently with the one we 

made (quite independently from Rossignol's account, 

with which we were not aware at that 

time) after preserved specimens that had kept 

their coloration quite well. An illustration of our 

specimen is provided here to show the color pattern 

(Figure 59a). 

It is interesting to note that Dana (1855, atlas, 

pi. 21: fig. 7) published a colored figure ofGoniopsis 

cruentata from Brazil that shows a color pattern 

different from that of the West Indian form and 

in a few respects resembles that of G. pelii (presence 

of a white streak along the posterolateral 

margin of the carapace; pale dactyli of the pereiopods); 

however, it differs conspicuously from 

both types, and a more thorough study of Brazilian 

material of Goniopsis might yield interesting 

results. 

Herklots (1851:8, 9, pi. 1: figs. 6, 7, 8) described 

the present species under two different names, 

viz., Grapsus pelii and G. simplex. As pointed out 

by De Man (1900:43, pi. 2, figs. 6, 7), G. simplex 

is based on juvenile specimens of G. pelii. The 

types of both species are still extant in the Rijksmuseum 

van Natuurlijke Historie, Leiden, and 

after examining them we can fully confirm De 

Man's conclusions. We use here the specific name 

pelii in preference to simplex, as the former is based 

on fully adult, the latter on juvenile specimens. 

The two names are of the same data, and under 

the Law of Priority have equal rights. In an 

earlier paper De Man (1879:68) synonymized


Grapsus pelii with Goniopsis cruentata and was folcommon 

among stones and debris on the ground 

lowed in this by all subsequent authors. The (Rathbun, 1921:444; Uschakov, 1970:443), and 

specimens brought by Doflein (1904:129), Lenz Sourie (1954a:84, 293) mentioned that near 

(1910:125), and Lenz and Strunck (1914:283) to Dakar the species lives "dans les anfractuosites 

Pachygrapsus simplex, as Balss (1922:81) has des maconneries; abondant au voisinage des man- 

pointed out, are not Herklots' species, but Pachygroves," and "commun sur les falaises" near Congrapsus 

gracilis (De Saussure). 

akry. Pauly (1975:57) noted that it was not com- 

Forest and Guinot (1966:91) reported on a mon near the mouth of the Sakumo Lagoon, 

specimen of Geograpsus lividus from Fernando Poo Ghana. 

in the collection of the Museu Bocage, which was This species has been reported to live in bur- 

identified as "Goniograpsus cruentatus"; it is possible rows that it makes in the mud of mangrove areas. 

that this is one of the specimens reported by It is commonly seen climbing on the aerial roots 

Osorio (1895b:57) from that island. The same 

of mangrove trees and also on slender vertical 

authors also showed that the material that Osorio 

stems, up to 5 feet [1.5 m] over the surface 

(1892:199) reported as Goniograpsus cruentata from 

(Rathbun, 1921). Irvine (1947:293) described this 

Binda, Sao Tome, actually belongs to Grapsus 

as follows: 

grapsus. On the other hand, they could confirm 

Osorio's (1889:139) identification of this species It is often seen climbing up an aerial root of the red 

from Praia Salgada, Principe. 

BIOLOGY.—The Pillsbury specimens were taken 

on a muddy bottom near the sea wall of Lagos 

harbor in brackish water. Most records in the 

mangrove with four legs on either side of it. The claws are 

kept underneath as it climbs, and are used as legs, working 

alternately in pushing the animal up the aerial roots. When 

pursued it can climb rapidly, and if closely pressed it often 

drops into the water and escapes along submerged roots. 

literature indicate the species from a similar 

brackish muddy habitat. The best characterization 

is that given by Rathbun (1921:444): 

They are very agile and when disturbed seek 

shelter between mangrove roots, or under stones 

and debris (Rathbun, 1921). They feed on the 

These crabs avoid the seashore, and on finding them in 

good numbers in mangrove swamps one might at first 

consider these their favorite habitat. However, they are not 

found in any of these swamps far inland for they remain 

near the mouth of rivers, where the salinity of the water is 

ground. Irvine (1947:293) also remarked that "it 

appears to feed on the bark of dead roots and 

other rotting vegetation, but may live on minute 

animals living in the bark. When feeding it brings 

less than in the sea but still greater than about the creeks its claws up to its mouth alternately in a most 

up-river where mangroves are still able to flourish. amusing manner." 

Irvine (1947:292, 293) described the species as 

living "mainly in the wet mud underneath the 

tangled mass of aerial prop roots of the red mangrove 

(Rhizophora). It occupies small holes hollowed 

out in the mud and is often to be seen 

crawling on the surface." Furthermore, he stated 

that they "are also seen to a smaller extent in the 

Humes (1957) reported that this species was 

the host of the harpacticoid copepod Cancrincola 

longiseta Humes at several localities on the West 

African coast between Sierra Leone and the 

Congo. 

Ovigerous females have been reported during 

April, May, July, August, and November. 

slightly drier swamps round the banks of the The species seems to be of very little economic 

lagoons, where the land is covered with a growth value. Only Irvine (1947:293) remarked that it is 

of white mangrove (Avicennia nitida)." Several sold in the fish markets in Ghana. 

other authors also reported the species from man- VERNACULAR NAME.—Both Irvine (1947) and 

groves (e.g., Monod, 1927, 1956; Capart, 1951; Bruce-Chwatt and Fitz-John (1951) give this spe- 

Longhurst, 1958; Gauld, 1960; Rossignol, 1957, cies the vernacular name "Mangrove crab," a 

1962). However, Goniopsis pelii also is found to be name also found on the field label of the speci-

232 

mens from the mouth of the Junk River, Liberia, 

examined by us. 

DISTRIBUTION.—Goniopsis pelii is known from 

the West African coast from Dakar, Senegal, in 

the north to Mocamedes, Angola, in the south. 

Monod (1956:414, 415) listed many localities in 

Senegal, Guinea-Bissau, Guinea, Sierra Leone, 

Liberia, Ivory Coast, Ghana, Nigeria, Rio Muni, 

Gabon, and Congo, whence he had examined 

material of the present species. The other localities 

mentioned in the literature are as follows. 

Senegal: Dakar (Rathbun, 1900a, 1918; Sourie, 1954a). 

Casamance (Sourie, 1954a). 

Guinea-Bissau: No specific locality (Osorio, 1895a). Agongoon 

on Ilha Caravela, and Porto de Bissau on Ilha de 

Bissau (Vilela, 1949). 

Guinea: Conakry (Capart, 1951; Sourie, 1954a; Uschakov, 

1970). 


Liberia: No specific locality (Von Martens, 1872; Biittikofer, 

1890; Johnston, 1906). Monrovia (Rathbun, 1918). 

Rock Spring and mouth of Mesurado River, near Monrovia 

(Rathbun, 1900a, 1918). Cape Mesurado (Balss, 1922). 

Ivory Coast: Lagune Ebrie at Adiapo-doume (Humes, 

1957). 

Ghana: No specific locality (Gauld, 1960). Butre (typelocality; 

Herklots, 1851; De Man, 1879, 1900). Baya River 

near Elmina (Benedict, 1893, as Ashantee; Rathbun, 1918). 

Accra (Irvine, 1947). Sakumo lagoon, near mouth (Pauly, 

1975). 

Nigeria: Lagos (Balss, 1922; Bruce-Chwatt and Fitz-John, 

1951). Iru Fisheries Station near Lagos (Humes, 1957). 

Elechi Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell, 

1979). 

Cameroon: No specific locality (Doflein, 1900; Balss, 

1922; Monod, 1927, 1928). Douala (Monod, 1927, 1956). 

Rio Muni: Cabo San Juan (Nobili, 1906b). Rio Muni, 

20 km up from Elobey (Balss, 1922). Islas de Elobey (Osorio, 

1895a). 

Fernando Poo: San Carlos and Mongola (Osorio, 1895a; 

see "Remarks"). 

Principe: No specific locality (Frade, 1950).—Praia Salgada 

(Osorio, 1889; Forest and Guinot, 1966).—Rio Papagaio 


Sao Tome: No specific locality (Osorio, 1888, 1889; Balss, 

1922; Frade, 1950; Forest and Guinot, 1966). Praia das 

Conchas (Osorio, 1889). Iogoiogo (Osorio, 1890). Binda 

(Osorio, 1892). 

Gabon: No specific locality (Kingsley, 1880a). Cap Lopez 

(Balss, 1922). Port-Gentil (Roux, 1927). Ogooue (Thallwitz, 

1891). 

Congo: Songololo River near Pointe-Noire; Loeme River, 


about 18 km S of Pointe-Noire (Humes, 1957). Loango, and 

mouth of Songololo River (Rossignol, 1957, 1962). Djeno, 

near Pointe-Noire (Rossignol, 1962). 

Zaire: Banana (Rathbun, 1921; Capart, 1951). Moanda 

(Rathbun, 1921). 

Angola: No specific locality (Osorio, 1887). Santo Antonio 

do Zaire (as San Antonio) (Rathbun, 1921). Lobito (Osorio, 

1887). Mocamedes (Osorio, 1895a). Between Cacuaco and 

Lobito-Benguela (Hartmann-Schroder and Hartmann, 

1974). 

Genus Grapsus Lamarck, 1801 

Gmpsus Lamarck, 1801:150 [type-species: Cancer grapsus Linnaeus, 

1758, by tautonomy; gender: masculine]. 

* Grapsus grapsus (Linnaeus, 1758) 

Grapsus maculatus.—Biittikofer, 1890:465, 487.—Johnston, 

1906:862. 

Grapsus grapsus.—Bruce-Chwatt and Fitz-John, 1951:117.— 

Capart, 1951:181, fig. 70.—Monod, 1956:407, fig. 561.— 

Dubois, 1957:7.—Rossignol, 1957:87, 121 [key].—Gauld 

and Buchanan, 1959:124.—Forest and Games, 1960: 

353.—Gauld, 1960:71.—Guinot and Ribeiro, 1962:68.— 

Rossignol, 1962:119.—Ribeiro, 1964:14.—Chace, 1966: 

640.—Forest and Guinot, 1966:90.—Zariquiey Alvarez, 

1968:422, fig. 139a [Portugal].—Kensley, 1970:181.— 

Penrith and Kensley, 1970b:246, 250, 252, 260.—Hartmann-Schroder 

and Hartmann, 1974:24. 

SYNONYMS.—Grapsus pictus Latreille, 1803; 

Grapsus maculatus H. Milne Edwards, 1853; Grapsus 

webbi H. Milne Edwards, 1853; Grapsus pictus 

ocellatus Studer, 1883; Cancer jumpibus Swire, 1938. 


Poo: Sta 258, shore, 86, 3$, 3 juv (W). 

Annobon: Sta 271, shore, 36, 2$, 5 juv (L). Sta 273, shore, 

26, 3 juv (L). Sta 281, shore, 1 cast (L). 

Other Material: Azores: No specific locality, W. Trelease, 

3c5, 3$ ov (W). Near harbor of Ilha do Corvo, 4 Jun 1976, 

W. Backhuys, 6 spec (L). Ponta Delgada, Ilha de Sao Miguel, 

31 Aug 1949, G. J. Jacobs, lc? (W). Horta, Ilha do Faial, U. 

S. Exploring Expedition, 1$ (W). Pirn Bay, Horta, Ilha do 

Faial, L. Dexter, 1$ (W). 

Senegal: Dakar, in tide pool, 12 Feb 1969, D. E. Harvey, 

1


Ghana: Virgins Pool, Takoradi, 27-29 Jul 1961, Bane 

and Richards, 46\ 3$ (1 ov) (W). 

Cameroon: Kribi, common on rocky boulders, below and 

above low tide line, very fast, 8 Mar and 6 Aug 1964, B. de 

Wilde-Duyfjes, 19, 1 juv (L). 

Zaire: Banana, mouth of the Congo River, Jul 1915, H. 

Lang, American Museum Congo Expedition, 26, 19 ov (W). 

DESCRIPTION.—Capart, 1951:181; Chace and 

Hobbs, 1969:163. 


561; Chace and Hobbs, 1969, figs. 50, 52g-i. 

Male Pleopod: Chace and Hobbs, 1969, fig. 

52g-i (West Indies). 

MEASUREMENTS.—The carapace width of the 

examined males ranged between 16 and 70 mm, 

that of the non-ovigerous females between 23 and 

43 mm, that of the ovigerous females between 32 

and 71 mm, and that of the juveniles between 5 

and 14 mm. 

BIOLOGY.—The species can be found on rocks 

in the surf- or splash-zone. A good description of 

its habitat is given by Chace and Hobbs (1969: 

165, 166). 

The species is very agile and exceedingly difficult 

to catch in the daytime; at night it can be 

approached far easier. See also Chace and Hobbs 

(1969:166). Off West Africa ovigerous females 

have been collected in February, May, July, and 

August (Guinot and Ribeiro, 1962; Ribeiro, 1964; 

present paper). 

DISTRIBUTION.—The species has a wide range 

in the tropical Atlantic and East Pacific regions. 

In the East Atlantic it is found from the Azores 

and Morocco to South-West Africa; there is a 

single record from Setubal, Portugal (Osorio, 

1905), but this needs confirmation. Monod (1956) 

listed the West African records, to which the 

following can now be added. 

Morocco: Temara, S of Rabat (Forest and Gantes, 1960). 

Cape Verde Islands: Baia de Monte Trigo, Santo Antao; 

Matiota, Baia da Fateixa, and Baia da Calheta, Sao Vicente; 

Baia da Murdeira and Pedra Lurae, Sal; Baia do Tarrafal, 

Sao Tiago; and Ponta Rodrigo, Boavista (all Guinot and 



1906). 

Ghana: No specific locality (Gauld, 1960). Teshi, 8 mi [13 

km] E of Accra (Gauld and Buchanan, 1959). Takoradi and 

Tema harbors (Gauld, 1960). 

Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951). 

Principe: Praia Ponta da Mina (Forest and Guinot, 1966). 

Sao Tome: Praia Pantufo, Praia Melao, Ilheu das Cabras, 

Binda, and Ilheu Gago Coutinho (as Hot das Rollas) (Forest 


Annobon: San Antonio (Forest and Guinot, 1966). 

Congo: Djeno (Rossignol, 1957, 1962). Baie de Loango 

and Pointe-Noire (Rossignol, 1962). 

Zaire: No specific locality (as Belgian Congo) (Dubois, 

1957). 

Angola: Baia de Luanda; Lobito; Baia Farta, Baia da 

Caota, and Ponta da Carufta, Benguela; Bafa de Santa 

Marta; and Praia Amelia, Mocamedes (all Guinot and 

Ribeiro, 1962). Near Mocamedes (Hartmann-Schrbder and 

Hartmann, 1974). 

South-West Africa: Rocky Point, 18°59'S, 12°29'E (Kensley, 

1970; Penrith and Kensley, 1970b). Mbwe Point, 19°- 

23'S, 12°42'E (Kensley, 1970). 

Saint Helena: several localities (Chace, 1966). 

Genus Pachygrapsus Randall, 1840 

Pachygrapsus Randall, 1840:126 [type-species: Pachygrapsus 

crassipes Randall, 1840, by selection by Kingsley, 1880a: 

198; gender: masculine; name 1638 on Official List]. 

Goniograpsus Dana, 1851c:247, 249 [type species: Goniograpsus 

innotatus Dana, 1851, a subjective junior synonym of 

Pachygrapsus transversus (Gibbes, 1850), by present selection; 


* Pachygrapsus gracilis (De Saussure, 1858) 

Pachygrapsus gracilis.—Frade, 1950:11, 26.—Capart, 1951: 

187, fig. 74, pi. 3: fig. 19.—Monod, 1956:419, figs. 569, 

571, 574-577.—Rossignol, 1957:89, 122 [key].—Gauld, 

1960:71.—Guinot and Ribeiro, 1962:71.—Rossignol, 

1962:120.—Forest and Guinot, 1966:92.—Uschakov, 

1970:443, 444, 445.—Powell, 1979:127. 

Pachygrapsus "africanus".—Hartmann-Schrbder and Hartmann, 

1974:19 [error for P. gracilis]. 

SYNONYM.—Grapsus guadulpensis Desbonne and 

Schramm, 1867. 


Sta 1, Lagos harbor, shore, 156", 179 (8 ov) (L). Sta 227, 

Lagos harbor, shore, 36*, 2$ (W). 

Other Material: Liberia: No specific locality, 13 Mar 1953, 

G. C. Miller, 16*, 2$ (W). Freeport area, Monrovia, oyster 

cultch, 12 Mar 1953, G. C. Miller, 3c5, 8$ (4 ov) (W). 

Ghana: Virgins Pool, Takoradi, near sports club, 24 Jul 

1961, Bane and Richards, 19 (W).

234 

Dahomey: Cotonou, Point XI Lagune, 8 Apr 1964, H. 

Hoestlandt, 1$ ov (L). 

Nigeria: Ogudu River, 8 Aug 1926, A. S. Pearse, 2$ (W). 

Lagos, harbor, 18 Jul 1963, A. R. Longhurst, 1(5 (L). W of 

Forcados, near confluence of Odimodi Creek and Forcados 

River, 05°22'N, 05°26'E, 28 Feb 1976, C. B. Powell, 16, 

1 $ (L). Niger delta between Brass and Port Harcourt, May- 

Aug 1960, H. J. G. Beets, 3


stones on river shore, 15 Jul 1915, H. Lang, American 

Museum Congo Expedition, 10

236 

terial of this species from the Pacific Ocean between 

roughly 41 °N and 35°S latitudes and from 

the Indian Ocean; no reliable records of the 

species from the Atlantic Ocean were known to 

him. Later records showed that the species is 

rather common in the South Atlantic Ocean, 

being found there far more frequently than P. 

minutus; Juanico (1976:149) is of the opinion that 

it is the only species of Planes there (but see p. 

237). Juanico reported an extensive material (28 

specimens) of the species from various localities 

on the Uruguay coast. Chace (1966:646) examined 

11 specimens from various parts of the coast 

of St. Helena. The only certain record of the 

species from the coast of tropical West Africa is 

the one by Crosnier (1967), who examined a 

female from a buoy off Gabon, 02°50'S, 08°50'E. 

The species has once been reported from the 

North Atlantic, viz., by Shirley (1974) from North 

Padre Island, Texas, U.S.A. The unconfirmed 

records of Planes minutus from the West coast of 

Africa (Gambia, Liberia, Cameroon and "Gulph 

of Guinea"), discussed below, might pertain to 

the present species. 

Planes minutus (Linnaeus, 1758) 

Cancer minutus Linnaeus, 1758:625. 

Grapsus minutus.—Leach, 1818, in 1817-1818:414.—Monod, 

1970:66. 

Planes minutus.—Monod, 1956:425, fig. 583.—Christiansen, 

1969:94, fig. 39, map 32 [Netherlands]. 

SYNONYMS.—Cancer pusillus Fabricius, 1775; 

Grapsus pelagicus Say, 1818; Planes clypeatus Bowdich, 

1825; Grapsus testudinum Roux, 1828; Nautilograpsus 

major MacLeay, 1838; Nautilograpsus smithii 

MacLeay, 1838; Grapsus dirts Costa, 1838- 

1853; Planes linnaeana Bell, 1845. 


Other Material: Morocco: Off Cap Blanc du Nord, 33°- 

16'N, 09°10'W, surface, collected with hand net, 27 Mar 

1976, Orwersaagd Sta 140, 1


the Northside of Jamaica." Chace (1951:81) gave 

this as the first reliable record of Planes minutus. 

Finally Linnaeus' reference, "Osbeck.iter.," is to 

Osbeck (1757:307), who mentioned Cancer minutus 

from the Sargasso Sea (Gras-Sjb) at 22°N in the 

Atlantic north of Ascension. Osbeck gave no morphological 

description of this species, but did so 

of Cancer pelagicus, which showed the latter to be 

Portunus sayi Gibbes; therefore, there can be little 

doubt that Osbeck's Cancer minutus is a Planes, and 

Chace (1951:81) also considered it as such. 

LECTOTYPE.—So far as we know no lectotype 

has ever been selected for Cancer minutus L. and in 

order to legalize the continued use of the epithet 

minutus for the North Atlantic Planes species, we 

now select as the lectotype of Linnaeus' species 

the specimen from the north coast of Jamaica 

figured by Sloane (1725, pi. 245, fig. 1). 

BIOLOGY.—Monod (1927:621) reported on four 

specimens found in Douala Bay near Souellaba, 

Cameroon, on a floating trunk of a tree (Aucoumea 

sp., family Burseraceae). The same author later 

(1956:426) examined eight specimens found near 

Cotonou, Dahomey, on floating weeds "Fucus 

natans," by which the collector doubtless meant 

Sargassum. Outside the tropical West African region 

the species has been reported from floating 

Sargassum weed, from marine turtles, and from 

pieces of wood and other floating objects; it is a 

true oceanic form. 

DISTRIBUTION.—There are very few records of 

the species from West Africa. Monod (1956:425, 

426) listed only four: Gambia River (Gambia), 

Monrovia (Liberia), Cotonou (Dahomey), and 

Souellaba (Cameroon). Later, Monod (1970:66) 

drew attention to Leach's (1818, in 1817-1818: 

414) record of the species from the "Gulph of 

Guinea." No other West African records are 

known to us. 

Chace (1951:80, fig. 8) indicated the specimens 

from Gambia, Liberia, and Cameroon as of uncertain 

identity, being either Planes minutus or P. 

cyaneus Dana; no material from these localities 

was seen by Chace and the data provided in the 

literature were insufficient to identify them with 

certainty. The specimens mentioned by Leach 

(1818, in 1817-1818) and Monod (1970) as Grapsus 

minutus cannot be identified, because too little 

is known about them. 

Quite different is the case with the specimens 

from Cotonou mentioned and figured by Monod 

(1956:425, fig. 583). Monod based the identification 

of his material on Chace's (1951) revision 

of the genus Planes, in which the differences between 

P. minutus and P. cyaneus were extensively 

dealt with. Furthermore, the Cotonou specimen 

figured by Monod shows the slender second pereiopod 

indicated by Chace as characteristic for P. 

minutus, and resembles that species also in other 

respects, so that there is no good reason to doubt 

Monod's identification. It is rather difficult therefore 

to understand Juanico (1976:146), who tried 

to show that Monod's identification is not reliable 

by stating on this account: "Monod (1956), dispone 

de un material de 8 ejemplares, pero no 

hace ninguna referencia a haberlos analizado 

respecto a los parametros que permiten distinguir 

una especie de otra [= P. minutus from P. cyaneus]." 

Juanico reached the conclusion that P. 

minutus is restricted to the North Atlantic, but did 

not define what he considered the line between 

the North and South Atlantic. Judging by the 

fact that he evidently considered Surinam, Pernambuco, 

and Dahomey to be in the South Atlantic, 

this line is not the equator. Therefore, we 

consider for now that both species of Planes occur 

in West African waters, with Monod's (1956) 

record of Planes minutus from Cotonou constituting 

the only reliable record of that species for the 

area. 

Outside the tropical West African region Planes 

minutus is known with certainty only from the 

North Atlantic between 11° and 52°N (see also 

Chace, 1951, fig. 8). 

Subfamily PLAGUSIINAE Dana, 1851 

Genus Percnon Gistel, 1848 

Acanthopus de Haan, 1833:5, 6 [invalid junior homonym of 

Acanthopus Klug, 1807 (Hymenoptera); a genus established 

without included nominal species; the first species assigned 

to the genus, by de Haan, 1835:29, 30, were Plagusia


clavimana Latreille, 1806, Cancer planissimus Herbst, 1804 

(as a synonym of P. clavimana), and Plagusia serripes Lamarck, 

1818. Type-species: Cancer planissimus Herbst, 1804, 

by selection by Rathbun, 1918:337; gender: masculine; 

name 465 on Official Index]. 

Percnon Gistel, 1848:viii [substitute name for Acanthopus de 

Haan, 1833; type-species: Cancer planissimus Herbst, 1804; 

gender: neuter; name 345 on Official List}. 

Leiolophus Miers, 1876:46 [substitute name for Acanthopus de 

Haan, 1833; type-species: Cancer planissimus Herbst, 1804; 


Liolophus Alcock, 1900:439 [emendation of Leiolophus Miers, 


* Percnon gibbesi (H. Milne Edwards, 1853) 

Acanthopus Gibbesi H. Milne Edwards, 1853:180. 

Percnon planissimum.—Monod, 1956:453, fig. 613.—Gauld 

and Buchanan, 1959:128.—Forest and Games, 1960: 

355.—Gauld, 1960:71.—Guinot and Ribeiro, 1962:72.— 

Ribeiro, 1964:17.—Forest and Guinot, 1966:93.—Zariquiey 

Alvarez, 1968:436, fig. 146a [Portugal]. [Not Cancer 

planissimus Herbst, 1804.] 

Percnon gibbesi.—Chapman and Santler, 1955:375.—Figueira, 

1960:11. 

SYNONYM.—Plagusia Delaunayi De Rochebrune, 

1883. 


Poo: Sta 258, shore, \6, 1 juv (W). 

Annobon: Sta 271, shore, 3


*Plagusia depressa (Fabricius, 1775) 

Plagusia depressa.—Buttikofer, 1890:487.—Johnston, 1906: 

862.—Monod, 1956:455, figs. 614-617.—Rossignol, 1957: 

95, 121 [key], fig. 9.—Figueira, 1960:11.—Forest and 

Games, 1960:356.—Gauld, 1960:71.—Guinot and Ribeiro, 

1962:72.—Rossignol, 1962:121.—Ribeiro, 1964: 

17.—Chace, 1966:647.—Forest and Guinot, 1966:93.— 

Hartmann-Schroder and Hartmann, 1974:24. 

Plagusia squamosa.—Stimpson, 1907:122. 

SYNONYMS.—Cancer squamosus Herbst, 1790; Plagusia 

sayi De Kay, 1844; Plagusia gracilis De Saussure, 

1858. 


Sta 225, Lagos harbor, shore, 1$ ov (W). 

Annobon: Sta 271, shore, Id, 1$ ov (L). Sta 273, shore, 2$ 

(1 ov) (W). 

Other Material: Madeira: No specific locality; U. S. Exploring 

Expedition, 1(5, 1? (W). 

Liberia: No specific locality, 1882, J. Buttikofer, let, 1? 

(L). No specific locality, 1890, J. Demery, 1$ (L). 

Ghana: Butre, 1840-1855, H. S. Pel, 1$ (L). Sekondi, 

1853, H. S. Pel, several specimens (L). Baya River, Elmina, 

27 Nov 1889, W. H. Brown, U. S. Eclipse Expedition, 16* 

(W). Accra, 1868-1869, M. Sintenis, 26*, 1$ (L). 

Cameroon: Kribi, among algae on rocks on a sandy 

beach, 11 Mar 1964, B. de Wilde-Duyfjes, 8


Angola: Lobito, 1899, P. Kamerman, 1? (L). 

DESCRIPTION.—Rathbun, 1921:455; Chace and 

Hobbs, 1969:173. 

Figure: Monod, 1956, fig. 609. 

Male Pleopod: Monod, 1956, figs. 610-612 (Cape 

Verde Islands; Senegal); Ghace and Hobbs, 1969, 

fig. 58b-d (West Indies). 

Color: "Brun-rouge uniforme. Extremite des 

pattes brun fonce, extremites des pinces jaunatres" 

(Rossignol, 1957:94). 


widths ranging from 6 to 12 mm, those of 

ovigerous females being 8 and 9 mm. 

BIOLOGY.—This species is found under rocks in 

the intertidal zone. Off West Africa ovigerous 

females have been collected in March, May, and 

July- 

DISTRIBUTION.—The species inhabits both the 

eastern and the western Atlantic; it has also been 

reported from the Indo-West Pacific region, from 

East Africa to Polynesia (Campbell and Griffin, 

1966; Griffin, 1968b). In the western Atlantic it 

occurs from Bermuda and Florida to Brazil. In 

the eastern Atlantic its range extends from the 

Cape Verde Islands and Senegal to Angola. To 

the West African localities enumerated by Monod 

(1956) the following can be added: 

Cape Verde Islands: Baia da Murdeira, Sal (Guinot and 


Senegal: Dakar (Sourie, 1954a). 

Guinea: Conakry (Sourie, 1954a; Uschakov, 1970). 

Congo: Loango lagoon (Rossignol, 1957, 1962). Pointe- 

Noire (Rossignol, 1962). 

Angola: Lobito (Bott, 1964). Baia Farta and Baia de 

Santa Marta, Benguela (Guinot and Ribeiro, 1962). 

Genus Metagrapsus H. Milne Edwards, 1853 

Melagrapsus H. Milne Edwards, 1853:188 [type-species: Sesarma 

curvatum H. Milne Edwards, 1837, selection by Rathbun, 

1918:321; gender: masculine]. 

REMARKS.—Serene and Soh (1970:397) restricted 

the genus Sarmatium Dana, 1851, to include 

only two species: the type, Sarmatium crassum 

Dana, 1851, and Sesarma germaini A. Milne Edwards, 

1869. The West African species, which is 

best known as Sarmatium curvatum (H. Milne Edwards, 

1837), according to these authors belongs 

to a distinct genus, for which the name Metagrapsus 

H. Milne Edwards, 1853, is available. Serene 

and Soh (1970:398) placed a second species in the 

genus Metagrapsus, viz., Metagrapsus pectinatus H. 

Milne Edwards, 1853. According to Rathbun 

(1918:322), who examined the type of Metagrapsus 

pectinatus, this species is synonymous with M. curvatus. 

Some authors, however, doubted the correctness 

of Rathbun's identification, mainly on 

zoogeographic grounds. As the type of M. pectinatus 

(said to be from Martinique) so far is the only 

specimen of the genus ever reported from the 

western Atlantic, there seems to be more reason 

to doubt the correctness of the locality label 

"Martinique" than Rathbun's identification. If 

Rathbun is correct, M. curvatus at present is the 

only species known of the genus Metagrapsus. 

* Metagrapsus curvatus 

(H. Milne Edwards, 1837) 

Sarmatium curvatum.—Capart, 1951:193.—Monod, 1956:449, 

fig. 608.—Rossignol, 1957:93, 122 [key], fig. 7.—Jordan, 

1957:198.—Humes, 1957:184, 187, 189.—Longhurst, 

1958:88.—Gauld, 1960:71.—Rossignol, 1962:121.—Uschakov, 

1970:448,455 [listed].—Powell, 1979:129. 

Sesarma curvatum.—Humes, 1957:186. 

Sarmatium.—Humes, 1957:188, 189. 

SYNONYMS.—Sesarma violacea Herklots, 1851; 

?Metagrapsus pectinatus H. Milne Edwards, 1853. 




koko, about 10 km west of Sapele, 05°N, 05°34.5'E, Oct-Dec 

1975, C. B. Powell, Ic5 (L). Between Brass and Port Harcourt, 

Niger delta, May-Aug 1960, H. J. G. Beets, 26, 1? (L). 

Congo: No specific locality, 1894, Dybowski, legator, 16\ 

1$ (W). 

Zaire: Banana, mouth of the Congo River, Jul-Aug 1915, 

H. Lang, American Museum Congo Expedition, 4(5, 1$ (W). 

Malela, 8 Jul 1915, H. Lang, American Museum Congo 

Expedition, 5

242 


Episesarma De Man (1895b: 165). Type-species: Sesarmoides Serene and Soh (1970:392, 403). 

Sesarma taeniolatum Miers, 1877 (= Sesarma taeni- Type-species: Sesarma krausii De Man, 1887, by 

olata White, 1847, a nomen nudum), a subjective original designation; gender: masculine. 

junior synonym of Sesarma meden H. Milne Ed- Namlacium Serene and Soh (1970:392, 404). 

wards, 1853, selected by Holthuis (1978:24); gen- Type-species: Sesarma crepidatum Caiman, 1925, by 

der: neuter. 

Parasesarma De Man (1895b: 181). Type-species: 

original designation; gender: neuter. 

Cancer quadratus Fabricius, 1798 (not Fabricius, 

1787)(= Ocypode plicata Latreille, 1803), by selec- 

Subgenus Chiromantes Gistel, 1848 

tion by Rathbun, 1918:284); gender: neuter. 

REMARKS.—The name Chiromantes was pro- 

Perisesarma De Man (1895b: 208). Type-species: posed by Gistel (1848:x) as a replacement for the 

Sesarma {Perisesarma) eumolpe De Man, 1895, se- generic name Pachysoma de Haan, 1833, which is 

lected by Holthuis (1977a: 170); gender: neuter. preoccupied by Pachysoma MacLeay, 1821 (Co- 

Nanosesarma Tweedie (1950:310). Type-species: leoptera) and Pachysoma Geoffroy, 1828 (Mam- 

Sesarma andersonii De Man, 1887, by original desmalia).ignation; gender: neuter. 

De Haan (1833:5, circ. pi. 2, pi. 7: fig. 4, pi. 8: 

Beamum Serene and Soh (1970:389, 394). Type- fig. 3) introduced the name Pachysoma in the first 

species: Sesarma batavica Moreira, 1903, by original fascicle of his Crustacea volume of Ph. F. von 

designation; gender: neuter. 

Siebold's Fauna Japonica. In the text of this fascicle 

Neosesarma Serene and Soh (1970:389, 394). the name Pachysoma was given in a key to the 

Type-species: Sesarma gemmiferum Tweedie, 1936, subgenera of the genus Grapsus, and no species 

by original designation; gender: neuter. 

were mentioned there. However, on two of the 

Neoepisesarma Serene and Soh (1970:390, 395). plates of this first fascicle two species of the 

Type-species: Sesarma mederi H. Milne Edwards, subgenus Pachysoma were figured and mentioned 

1853, by original designation; gender: neuter. by name: Grapsus (Pachysoma) haematocheir de Haan 

Muradium Serene and Soh (1970:390, 396). (1833, pi. 7: fig. 4) and Grapsus (Pachysoma) quad- 

Type-species: Cancer tetragonus Fabricius, 1798, by ratus (Fabricius, 1798) (pi. 8: fig. 3). As these two 


species are the only nominal species referred to 

Selatium Serene and Soh (1970:390, 397). Type- Pachysoma in the original publication of that 

species: Sesarma brockii De Man, 1887, by original subgenus, it is from among them that a type may 

designation; gender: neuter. 

be selected for the subgenus. Therefore, as Hol- 

Tiomanum Serene and Soh (1970:391, 398). thuis (1977a: 170) pointed out, the selection of 

Type-species: Sesarma indica H. Milne Edwards, Grapsus (Pachysoma) bidens de Haan, 1835, as the 

1837, by original designation; gender: neuter. type of Pachysoma, made by Fowler (1912:439) is 

Bresedium Serene and Soh (1970:391, 399). invalid. The first valid type selection for Pachysoma 

Type-species: Sesarma edwardsii brevipes De Man, de Haan, 1833 (and thus automatically for Chi- 

1889, by original designation; gender: neuter. romantes Gistel, 1848) is the one by Holthuis 

Pseudosesarma Serene and Soh (1970:391, 399). (1977a: 170), who selected Grapsus (Pachysoma) hae- 

Type-species: Sesarma edwardsi De Man, 1888, by matocheir de Haan, 1833, as the type of that 


subgenus. 

Sesarmops Serene and Soh (1970:391, 400). As Grapsus (Pachysoma) haematocheir de Haan, 

Type-species: Sesarma impressa H. Milne Edwards, 1833, is also the type-species (by monotypy) of 

1837, by original designation; gender: masculine. the subgenus Holometopus H. Milne Edwards, 

Labuanium Serene and Soh (1970:392, 401). 1853, the name Chiromantes, being the senior of 

Type-species: Sesarma polita De Man, 1888, by the two, has to be used. 


Although the transfer of a name from one


taxon to another usually is harmful and to be 

avoided, the status of the taxonomy and nomenclature 

of the subgenera of Sesarma is at present 

in such a state of flux that it seems best to adhere 

strictly to the Code in the present case, at least 

until a revision of the Sesarma complex is achieved. 

The subgenus Chiromantes is represented in the 

eastern Atlantic by three species, all three of 

which occur in tropical West Africa. Sesarma roberti, 

erroneously reported from West Africa (p. 

226), also belongs in the present subgenus. 

The subgenus was referred to as Holometopus by 

Monod (1956) and all later authors, while they 

used the name Chiromantes for the subgenus here 

indicated with the name Perisesarma. 

Sesarma (Chiromantes) angolense De Brito 

Capello, 1864 

Sesarma angolensis.— Biittikofer, 1890:487.—Johnston, 1906: 

861.—Longhurst, 1958:88.—Gauld, 1960:71. 

Sesarma {Holometopus) angolensis.—Dartevelle, 1950:48.— 

Monod, 1956:445, fig. 605.—Rossignol, 1962:120. 

Sesarma {Holometopus) angolense.—Dartevelle, 1950:50.—Capart, 

1951:191, fig. 76.—Rossignol, 1957:92, 122 [key].— 

Jordan, 1957:198.—Humes, 1957:186, 187, 189. 


Other Material: Liberia: No specific locality, 1890, 1897, J. 

Demery, 2? (L). Grand Cape Mount near Robertsport, 1882, 

J. Biittikofer, 4(5, 3$ (L). 

Nigeria: Mayuku Creek near Ugbekoko, 10 km W of 

Sapele, Midwest State, ca. 05°54'N, 05°34.5'W, Oct, Oct- 

Dec, Dec 1975, Jan 1976, C. B. Powell, many specimens (L). 

Niger delta between Brass and Port Harcourt, May-Aug 

1960, H. J. G. Beets, 16, 29 (L). 

Angola: Lobito, 1899, P. Kamerman, 1

244 

of Sapele, 05°54'N, 05°37'E, in mangrove swamp, Oct-Dec 

1975, C. B. Powell, 28 specimens (1$ ov) (L). 

Cameroon: Kribi, among humid dead leaves on the bank 

of a forest stream not far from the sea, 8 Aug 1964, B. de 

Wilde-Duyfjes, Id, 2$ (L). 

Zaire: Banana, mouth of the Congo River, Jul-Aug 1915, 

H. Lang, American Museum Congo Expedition, Id, 19 (W). 

Malela, 8 Jun 1915, H. Lang, American Museum Congo 

Expedition, 226*, 109 (2 ov) (W). Zambi, 4 Jul 1915, H. 

Lang, American Museum Congo Expedition, 1 juv (W). 

DESCRIPTION.—Rathbun, 1921:449. 

Figures: Monod, 1956, figs. 606, 607; Rossignol, 

1957, fig. 6. 

Color: The specimens from Lagune Ebrie carried 

a note stating that in life the chelae are 

purplish blue with red tips. Additional color notes 

were provided by Rossignol (1957:91). 

BIOLOGY.—This species inhabits littoral estuarine 

areas, particularly mangroves and adjoining 

inland regions (see Rathbun, 1921:449, 450). 


March, June, July-August, and October through 

December (Rathbun, 1921; Rossignol, 1957; present 

paper). 

DISTRIBUTION.—Sesarma buettikoferi is found 

along the West African coast from Liberia to 

Angola. Monod (1956) listed the locality records 

known to him; to these the following can now be 

added: 


59 paralectotypes (L, lectotype Crust. D. 150, paralectotypes 

Crust. D. 151). 

Nigeria: Harbor of Lagos, 13 Jun 1963, A. R. Longhurst, 

1


West Africa (Guinea to Angola). To the localities 

listed by Monod (1956) the following can be 

added. 

Guinea: lie Marara, Rio Pongo (Uschakov, 1970). 

Sierra Leone: Kissy (Longhurst, 1958). 

Ghana: Near Butre (Gauld, 1960). 

Nigeria: Elechi Creek, Port Harcourt, 04°47'15"N, 06° 

48'45"E (Powell, 1979). 

Congo: Loya River and Djeno (Rossignol, 1957, 1962). 

Pointe-Noire (Rossignol, 1962). 

Zaire: Banana, mouth of the Congo River (Dartevelle, 

1950). 

Angola: Santo Antonio do Zaire (as Saint-Antoine) (Dartevelle, 

1950). 

Subgenus Perisesarma De Man, 1895 

REMARKS.—This subgenus was cited by Monod 

(1956) and most other authors as Chiromantes Gistel, 

1848. However, as shown by Holthuis (1977: 

170) the type-species of Chiromantes is Grapsus 

(Pachysomd) haematocheir De Haan, 1833; hereby 

Chiromantes Gistel, 1848, becomes a senior objective 

synonym of Holometopus H. Milne Edwards, 

1853, and should replace the latter name (p. 242). 

For the subgenus usually cited as Chiromantes the 

name Perisesarma is available. 

Three species of Perisesarma are known from the 

eastern Atlantic area, all three are restricted to 

tropical West Africa. 

Sesarma (Perisesarma) alberti Rathbun, 1921 

Sesarma (Chiromantes) alberti.—Dartevelle, 1950:48.—Capart, 

1951:189, fig. 75.—Monod, 1956:440, fig. 594.—Jordan, 

1957:198.—Rossignol, 1957:122 [key]; 1962:120.—Forest 

and Guinot, 1966:93. 

Sesarma (chiromantes) alberti.—Rossignol, 1957:90. 

Sesarma alberti.—Longhurst, 1958:88.—Powell, 1979:127. 

MATERIAL EXAMINED—Pillsbury Material: None. 

Other Material: Liberia: St. John River at upper Buchanan, 

mud in mangroves, collected at night, 24 Aug 1967, T. C. 

Rutherford, 1$ (W). Rock Spring, Monrovia, O. F. Cook 

and G. N. Collins, 9juv (W). St. Paul River lagoon at True 

Rubber Farm, 9 Oct 1953, G. C. Miller, 1

246 

mann-Schroder and Hartmann, 1974:19.—Powell, 1979: 

127. 

SYNONYM.—Sesarma africana H. Milne Edwards, 

1837. 


Other Material: Senegal: Senegal, legator M. Delambre, 

donor Mus. Paris, Ic3 syntype of Sesarma africanum H. Milne 

Edwards, dry (W). 

Liberia: Liberia, 1881, J. Buttikofer and J. A. Sala, 1


Elechi Creek, Port Harcourt, 04°47'15"N, 06°58'45"E (Powell, 

1979). 

Congo: Loango (Pechiiel-Loesche, 1882). Mouths of the 

Songololo and Loya rivers (Rossignol, 1957, 1962). Songololo 

River near Pointe-Noire; mouth of Loeme River near Pointe- 

Noire (Humes, 1957). 

Zaire: Banana, mouth of the Congo River (Dartevelle, 

1950). 

Angola: Santo Antonio do Zaire (as Saint Antoine) (Dartevelle, 

1950). Between Cacuaco and Lobito-Benguela (Hartmann-Schroder 

and Hartmann, 1974). 

Sesarma (Perisesarma) kamermani De Man, 

1883 

Sesarma (Chiromantes) kamermani.—Monod, 1956:441, figs. 

595-600.—Rossignol, 1957:122 [key]. 


Other Material: Angola: Musserra, 1882, P. Kamerman, Id 

holotype (L. Crust. D. 166). 

DESCRIPTION.—De Man, 1883:165; Balss, 

1936:1. 

Figures: Balss, 1936, figs. A, B; Monod, 1956, 

figs. 595-600. 

MEASUREMENTS.—The carapace length of the 

holotype is 27 mm, its carapace width is 32 mm. 

DISTRIBUTION.—So far only two specimens, 

both males, have become known of this species. 

The type-specimen (in the Leiden Museum) originated 

from Musserra, Angola. The second specimen 

was reported upon by Balss (1936) and 

Monod (1956) and was collected at Banana, 

Zaire. 

Subfamily VARUNINAE 

H. Milne Edwards, 1853 

Genus Brachynotus de Haan, 1833 

Brachynotus de Haan, 1833:5, 7 [subgenus established without 

included nominal species; type-species: Goneplax sexdentatus 

Risso, 1827, by subsequent monotypy by de Haan, 1835: 

34; gender: masculine; name 1610 on Official List]. 

Heterograpsus Lucas, 1846:18 [type-species: Heterograpsus 

sexdentatus Lucas, 1846, a subjective junior synonym and 

homonym of Goneplax sexdentatus Risso, 1827, by monotypy; 


Shurebus Verany, 1846:7 [type-species: Shurebus genuensis Verany, 

1846, an objective: junior 1 synonym of Cleistotomaigemmellan 

Rizza, 1839, by monotypy; gender: masculine]. 

Brachynotus atlanticus Forest, 1957 

Brachynotus sexdentatus.—Monod, 1933a:219, fig. 7.—Bouvier, 

1940, fig. 184A only [copied from Monod, 1933a, fig. 7]. 

[Not Goneplax sexdentatus Risso, 1827.] 

Brachynotus sexdentatus lucasi.—Monod, 1956:428, 631, figs. 

589-592, 881 [not Heterograpsus lucasi H. Milne Edwards, 

1853 = Goneplax sexdentatus Risso, 1827]. 

Brachynotus atlanticus Forest, 1957:505, figs. 2, 4, 6, 8, 10, 12, 

14.—Forest and Games, 1960:354, fig. 2. 

MATERIAL EXAMINED.—None. 

DESCRIPTION.—Forest, 1957:505. 

Figures: Monod, 1933a, fig. 7; Monod, 1956, 

figs. 589-592, 881; Forest, 1957, figs. 2, 4, 6, 8, 

10, 12, 14. 


(Mauritania); Forest, 1957, fig. 14 (Mauritania). 

DISTRIBUTION.—This species, which until recently 

was not distinguished from B. sexdentatus, 

is only known from the Atlantic coast of Morocco 

and from the Cap Blanc peninsula (Mauritania). 

The records are the following. 

Morocco: Numerous localities as far north as 34°53'N 

(Monod, 1933a; Forest and Gantes, 1960). 

Mauritania: Baie de l'Ouest, coast of Cap Blanc peninsula 

(Monod, 1956: Forest, 1957). Pointe de Cansado (Monod, 

1956). Baie de Cansado (Monod, 1956; Forest, 1957). Port- 

Etienne (Monod, 1956). Baie du Repos near Port-Etienne 

(Monod, 1956). 

The Mauritanian localities are all situated on 

the east coast of the Cap Blanc peninsula. The 

species is not known to occur farther south. 

Genus Euchirograpsus H. Milne Edwards, 1853 

Euchirograpsus H. Milne Edwards, 1853:175 [type-species: 

Euchirograpsus liguricus H. Milne Edwards, 1853, by monotypy; 


Euchirograpsus liguricus H. Milne Edwards, 

1853 

Euchirograpsus liguricus H. Milne Edwards, 1853:175.—Tvirkay, 

1975b: 105, figs. 1-3, 17, 23; 1976a:25 [listed], 39, figs. 

29-31. 

Euchirograpsus americanus.—A. Milne Edwards and Bouvier, 

1894:46, pi. 4: figs. 10-14; 1900:107.—Dollfus and

248 


bottoms of sand, sandy mud, mud, rock with 

gorgonians and/or sponges, and gravel at depths 

between 10 and 359 m, most of the records being 

from between 100 and 250 m. 

Few ovigerous females have been recorded 


Monod, 1927:216, figs. 1, 2.—Monod, 1933a:320; 1933b: from West Africa; the only record in the recent 

535.—Capart, 1951:184, fig. 72.—Monod, 1956:434, figs. 

592 bis, 882-884.—Rossignol, 1957:121 [key].—Forest and 

Games, 1960:355.—Rossignol, 1962:120.—Forest and 

Guinot, 1966:92.—Crosnier, 1970:1215 [listed], 1217. [Not 

Enchirograpsus americanus A. Milne Edwards, 1880]. 

literature is that by Crosnier (1970) based on a 

specimen taken by the Undaunted in March and 

also recorded herein. 

DISTRIBUTION.—The range of this species extends 

from west of Portugal and Madeira and the 

MATERIAL EXAMINED.—PUlsbury Material: None. Azores south to South-West Africa; it also occurs 

Geronimo Material: Gabon: Sta 211, 100 m, 1


EASTERN ATLANTIC GENERA.—Two, Cardisoma 

and Gecarcinus, each represented by one tropical 

species. 

EASTERN ATLANTIC SPECIES.—Two, both reported 

below. Neither species is represented in 

the Pillsbury collections. 

Genus Cardisoma Latreille, 1828 

Cardisoma Latreille, 1828b:685 [type-species: Cardisoma guanhumi 

Latreille, 1828, selected by H. Milne Edwards, 1838, 

in 1836-1844, pi. 20; gender: neuter]. 

Perigrapsus Heller, 1862:522 [type-species: Perigrapsus excelsus 

Heller, 1862, a subjective junior synonym ofCancer camifex 

Herbst, 1794, by monotypy; gender: masculine]. 

Discoplax A. Milne Edwards, 1867b:284 [type-species: Discoplax 

longipes A. Milne Edwards, 1867, by monotypy; gender: 

feminine]. 

Cardiosoma S.I. Smith, 1869b: 16, 36 [unjustified emendation 

of Cardisoma Latreille, 1828; type-species: Cardisoma guanhumi 

Latreille, 1828; gender: neuter]. 

Cardisoma armatum Herklots, 1851 

Cardisoma Guanhuni.—Studer, 1882:333, 353 [discussion] [not 

C. guanhumi Latreille, 1828]. 

Cardisoma armatum.—Pechiiel-Loesche, 1882:299.—Biittikofer, 

1890:464, 487, fig. on p. 465.—Johnston, 1906:861.— 

Gruvel, 1913:169 [listed].—Capart, 1951:196, fig. 79.— 

Monod, 1956:458, fig. 618.—Rossignol, 1957:95, 120 

[key], pi. 2: fig. 2.—Humes, 1957:181,189.—Dubois, 1957: 

7, fig. 24.—Gauld, 1960:72.—Rossignol, 1962:121.— 

Guinot and Ribeiro, 1962:73.—De Leersnyder and 

Hoestlandt, 1963:211 [physiology].—Ribeiro, 1964:18.— 

De Leersnyder and Hoestlandt, 1966:43 [physiology].— 

Forest and Guinot, 1966:94.—Monod, 1967:180, pi. 17: 

fig. 1.—Uschakov, 1970:450, 455 [listed].—Bright and 

Hogue, 1972:17.—Tiirkay, 1973:86, figs. 1, 3, 7-8, map 

1.—Hartmann-Schroder and Hartmann, 1974:19. 

Cardisoma Guanhumi var. armatum.—Bouvier, 1911:226. 

Cardisoma guanhumi.—Gruvel, 1913:169 [listed] [not C. guanhumi 

Latreille, 1828]. 

Cardiosoma armata.—Bruce-Chwatt and Fitz-John, 1951:117, 

118, 119. 

Gardiosoma.—Bruce-Chwatt and Fitz-John, 1951:120. 

Cardisoma.—Uschakov, 1970, fig. 4. 

Cardiosoma armatum.—Pauly, 1975:57. 


Other Material: Cape Verde Islands: Porto da Praia, Sao 

Tiago, coconut grove, North Pacific Exploring Expedition, 

16 (W). 

Senegal: Dakar, O. F. Cook, 16*, 1$ (W). 

Liberia: Grand Cape Mount, 1882, J. Biittikofer, 36", 

39 (L). 

Ghana: Accra, 1868-1869, M. Sintenis, lcJ (L). Elmina, 

1840-1855, H. S. Pel, 1$ holotype (L, Crust. D.58). 

Dahomey: Near Cotonou, Oct 1962, H. Hoestlandt, 16, 

19 (L). 

Nigeria: Lagos, 16 Aug 1926, A. S. Pearse, 19 (W). Adu, 

25 Aug 1926, A. S. Pearse, \6 fragmented (W). Niger delta 

between Brass and Port Harcourt, May-Aug 1960, H. J. G. 

Beets, 46, 59 (L). 

Cameroon: Batanga, A. J. Good, 13 Mar 1931, 16", 

19 (W). 

Zaire: Banana, Jul 1915, H. Lang, American Museum 

Congo Expedition, 18

250 

count of the species. To the references in Monod 

the following can be added: 

West Africa: No specific locality (Tiirkay, 1973). 

Cape Verde Islands: Baia de Sal-Rei, Boavista (Guinot 


Senegal: No specific locality (Tiirkay, 1973). Marigots de 

Hann (Bouvier, 1911). Saint-Louis (Monod, 1967; Tiirkay, 

1973). Dakar (Monod, 1967). Rtombo [?](Tiirkay, 1973). 

Joal; Ngor near Dakar (Humes, 1957). 

Guinea: Conakry (Uschakov, 1970). 

Sierra Leone: Bunce River, opposite Kamatare Island, 

near Freetown (Humes, 1957). 


1906). 

Ivory Coast: From market in Abidjan (Forest and Guinot, 

1966). Koumassi, near Abidjan (Humes, 1957). Abidjan 

(Tiirkay, 1973). 

Ghana: No specific locality (Gauld, 1960; Tiirkay, 1973). 

Kumasi and Sekondi (Tiirkay, 1973). Sakumo lagoon (Pauly 

1975). Accra; Osu Fisheries Station near Accra (Humes, 

1957). 

Dahomey: No specific locality (De Leersnyder and 

Hoestlandt, 1963, 1966). 

Nigeria: Lagos (Bruce-Chwatt and Fitz-John, 1951; Tiirkay, 

1973). Iru Fisheries Station near Lagos (Humes, 1957). 

Cameroon: Douala (Tiirkay, 1973). 

Fernando Poo: No specific locality (Tiirkay, 1973). 

Sao Tome: "environs de la ville" and Iogoiogo (Forest 


Gabon: No specific locality (Tiirkay, 1973). 

Congo: No specific locality (Rossignol, 1957). Baie de 

Pointe-Noire (Rossignol, 1962). Bahua [?] and Pointe-Noire 

(Tiirkay, 1973). Loango (Pechiiel-Loesche, 1882). Pointe- 

Noire; Songololo River near Pointe-Noire (Humes, 1957). 

Zaire: Banana (Dubois, 1957; Tiirkay, 1973). 

Angola: Luanda (Tiirkay, 1973). Santo Antonio do Zaire; 

Moâmedes (Guinot and Ribeiro, 1962). Between Cacuaco 

and Lobito-Benguela (Hartmann-Schroder and Hartmann, 

1974). 

Genus Gecarcinus Leach, 1814 

Gecarcinus Leach, 1814:430 [type-species: Cancer ruricola Lin- 

naeus, 1758, by selection by H. Milne Edwards, 1838, in 

1836-1844, pi. 21; gender: masculine]. 

Geocarcinus Miers, 1886:xiv, 216, 219 [invalid emendation of 

Gecarcinus Leach, 1814; type-species: Cancer ruricola Lin- 


EASTERN ATLANTIC GENERA.—One, Neotroglocarnaeus, 


cinus, represented in the tropical fauna. 

Johngarthia Tiirkay, 1970:335, 341, 343 [type-species: Gecarcinus 

planatus Stimpson, 1860, by original designation; EASTERN ATLANTIC SPECIES.—One, Neotroglocar- 


cinus balssi, reported below. It is not represented 

in the Pillsbury collections. 

Gecarcinus weileri (Sendler, 1912) 

Pelocarcinus weilen Sendler, 1912:191, figs, 4, 5. 

Gecarcinus lagostoma.—Capart, 1951:198, fig. 80.—Monod, 

1956:461 [not fig. 619 = G. lagostoma sensu stricto].— 

Rossignol, 1957:120 [key].—Forest and Guinot, 1966:94. 

[Not G. lagostoma H. Milne Edwards, 1837.] 

Gecarcinus (Johngarthia) weilen.—Tiirkay, 1973:96, figs. 15- 

17, map 2. 

Johngarthia weileri.—Tiirkay, 1976c:69, 70, 71 [lectotype designated]. 


Other Material: Fernando Poo: No specific locality, Feb 

1959, J. A. Warners, 1$ dry (L). 

DESCRIPTION.—Tiirkay, 1973:97. 

Figures: Tiirkay, 1973, figs. 15-17. 

MEASUREMENTS.—Our specimen has a carapace 

width of 60 mm. 

BIOLOGY.—This is a terrestrial species from the 

Gulf of Guinea, largely restricted to the offshore 

islands but also known from the mainland in 

Cameroon. Capart (1951) recorded an ovigerous 

female in January. 


taken from two localities in Cameroon and from 

the offshore islands of the Gulf of Guinea. Monod 

(1956) saw no material. Records in the literature 

since 1956 include: 

West Africa: No specific locality (Tiirkay, 1973). 

Cameroon: Bibundi (Tiirkay, 1973; 1976c). 

Fernando Poo: No specific locality (Tiirkay, 1973). 

Principe: Ilheu Caroô (Forest and Guinot, 1966). 

Sao Tome: No specific locality (Tiirkay, 1973). Ribeiro 

do Peixe (Forest and Guinot, 1966). Ilheu Gago Coutinho 

(as Hot das Rollas) (Forest and Guinot, 1966; Tiirkay, 1973). 

Annobon: No specific locality (Forest and Guinot, 1966; 

Tiirkay, 1973). 

Family HAPALOCARCINIDAE Caiman, 1900 

Lithoscaptes A. Milne Edwards, 1862b: F 10. 

HAPALOCARCINIDAE Caiman, 1900:3, 49. 

Genus Neotroglocarcinus Fize and Serene, 1957 

Neotroglocarcinus Fize and Serene, 1957:134 [type-species:


Troglocarcmus monodi Fize and Serene, 1955, by original 

designation; gender: masculine]. 

Neotroglocarcinus balssi (Monod, 1956) 

Troglocarcmus balssi Monod, 1956:463, 632, figs. 620-627.— 

Longhurst, 1958:88.—Gauld, 1960:72.—Monod, 1963, 

fig. 37 [no locality].—Crosnier, 1969:535. 

Neotroglocarcinus Balssi—Fize and Serene, 1957:135, 136, 141- 

143. 


Other Material: Canary Islands: Ponta de Matorra, Fuertaventura, 

14 km from Morro, 20-25 m, in Phyllangia mouchezi 

(Lacaze-Duthiers, 1897), 27 Jan 1975, M. Grasshoff and F. 

Engelhardt, 1 damaged specimen (MP). 

West Africa: No specific locality, in Asterosimilia marchadi 

(Chevalier, 1966), A. R. Longhurst, 1$ ov (BMNH). 

Ivory Coast: Off Abidjan, 62 m, 23 Aug 1968, C.R.O., 

Abidjan, 1? (MP). 



Male Pleopod: Monod, 1956, fig. 623 (Ghana). 


lengths of 3.2 to 3.8 mm; that of the ovigerous 

female is 3.8 mm. 

REMARKS.—As the single adult female from 

Sierra Leone listed by Monod (1956:632) was 

collected by A. R. Longhurst, and the single 

specimen reported upon by Longhurst (1958:88) 

was identified by Monod (see Longhurst, 1958: 

5), it seems likely that they are the same specimen 

and that an error crept into the ecological data 

reported for either: Monod reported the specimen 

as an epibiont of Cidaris cidaris (Linnaeus) from 

68 m depth, while Longhurst listed it as an 

epibiont of Eucidaris tribuloides (Lamarck) from 

200 m. (Under Eucidaris tribuloides, however, 

Longhurst (1958:101) only mentioned two specimens, 

both from 25-88 m depth, and under 

Cidaris cidaris 6 specimens from depths between 

88 and 132 m.) Longhurst (1958:7) reported that 

his collection finally was deposited in the British 

Museum (Natural History). Manning examined 

a specimen of this species of the British Museum 

collection, viz., the above-mentioned ovigerous 

female labeled "West Africa" and collected by 

Longhurst; this specimen is said to be taken from 

the coral Asterosimilia marchadi (Chevalier). The 

fact that both Cidaris and Eucidaris are very unlikely 

hosts for the species, and the conflicting 

information in the ecological data suggest that 

these should be regarded with the utmost reserve, 

and the possibility is not precluded that both the 

Monod (1956) and the Longhurst record pertain 

to the ovigerous female now in the British Museum, 

and that this specimen was collected in 

Asterosimilia marchadi in a depth of 68 m, at 

08°25'N, 14°18'W (off Freetown, Sierra Leone) 

on a sandy bottom on 22 Feb 1956 {Cape St. Mary 

StaMBl/A3). 

BIOLOGY.—This is a sublittoral species usually 

associated with corals. It has been taken with 

Dendrophyllia, in 44 m (Gauld, 1960); on rocks 

with gorgonians, probably associated with Phyllangia, 

in 10 m (Crosnier, 1969); on Eucidaris 

tribuloides (?) in 200 m (Longhurst, 1958); on 

Cidaris cidaris (?) in 68 m (Monod, 1956); and 

with Phyllangia mouchezi and Asterosimilia marchadi. 


(Crosnier, 1969; present paper); ova measured 

0.4 mm in the ovigerous female we examined. 

DISTRIBUTION.—West Africa, from localities in 

the Canary Islands, off West Africa proper from 

localities between Sierra Leone and the Congo, 

and from Ilheu Gago Coutinho (as I. das Rolas), 

near Sao Tome in the Gulf of Guinea (Balss, 

1922) in depths between 10 and 200 m. It has not 

previously been recorded from the Canary Islands 

or the Ivory Coast. Monod's specimens came from 

Ghana; since 1956 the species has been recorded 

from the following: 

Sierra Leone: No specific locality, 200 m (Longhurst, 

1958); 08°25'N, 14°18'W, 68 m (Monod, 1956). 


Congo: Pointe-Noire, ca. 10 m (Crosnier, 1969). 

Family HYMENOSOMATIDAE MacLeay, 1838 

HYMENOSOMIDAE MacLeay, 1838:68 [corrected to Hymenosomatidae 

by Stebbing, 1905:49]. 

HYMENICINAE Dana, 1851c:290. 

EASTERN ATLANTIC GENERA.—Two, Elamena

252 

and Hymenosoma, both represented by West African 

species. 

EASTERN ATLANTIC SPECIES.—Two, both listed 

below. This family was not represented in the 

Pillsbury collections. 

Genus Elamena H. Milne Edwards, 1837 

Elamena H. Milne Edwards, 1837:33 [type-species: Hymenosoma 

mathaei Desmarest, 1825, by monotypy; gender: feminine]. 

Trigonoplax H. Milne Edwards, 1853:224 [type-species: Ocypode 

(Elamene) unguiformis de Haan, 1839, by monotypy; 


Elamena (Trigonoplax) gordonae Monod, 1956 

Elamena (Trigonoplax) gordonae Monod, 1956:469, figs. 629- 

637 [between Conakry, Guinea and Monrovia, Liberia, 

30-40 m; Sierra Leone]; 1963, fig. 40 [no locality]. 

Elamena gordonae.—Uschakov, 1970:455 [listed; Guinea]. 

DISTRIBUTION.—West Africa, from the localities 

cited above. 

Genus Hymenosoma Desmarest, 1825 

Hymenosoma Desmarest, 1825:163 [type-species: Hymenosoma 

orbiculare Desmarest, 1825, by selection by H. Milne Edwards, 

1842, in 1836-1844, pi. 35: fig. 1; gender: neuter]. 

Leachium MacLeay, 1838:68 [type-species: Hymenosoma orbiculare 

Desmarest, 1825, by original designation; gender: 

neuter]. 

Centridion Gistel, 1848:viii [substitute name for Leachium 

MacLeay, 1838; type-species: Hymenosoma orbiculare Desmarest, 

1825; gender: neuter]. 

Hymenosoma orbiculare Desmarest, 1825 

Hymenosoma orbiculare.—Capart, 1951:61, fig. 18 [Angola; 

South-West Africa].—Monod, 1956:468, fig. 628 [Gabon 

(?); South-West Africa].—Penrith and Kensley, 1970a: 

209, 232 [South-West Africa]. 

DISTRIBUTION.—Known with certainty from localities 

in Angola, South-West Africa, and South 

Africa. The record from Gabon is questionable. 

Family MAJIDAE Samouelle, 1819 

MAIADAE Samouelle, 1819:88 [corrected to Majidae by Neumann, 

1878:5]. 


MACROPODIADAE Samouelle, 1819:90. 

INACHIDAE MacLeay, 1838:56 [given preference over Macropodiadae 

Samouelle, 1819, by International Commission 

on Zoological Nomenclature, Opinion 763, 1966; name 


EPIALTIDAE MacLeay, 1838:56. 

HUENIDAE MacLeay, 1838:56. 

EURYPODIDAE MacLeay, 1838:56. 

MITHRACIDAE MacLeay, 1838:56. 

LEPTOPODIADAE Bell, 1844:1. 

MACROCHEIRINAE Dana, 1851a:427. 

PISINAE Dana, 1851a:428 [name 368 on Official List, date 

and citation erroneous there]. 

LIBININAE Dana, 1851a:429. 

PRIONORHYNCHINAE Dana, 1851a:429. 

MICIPPINAE Dana, 1851a:429. 

CHORININAE Dana, 1851a:429. 

PYRINAE Dana, 1851a:430. 

OTHONINAE Dana, 1851a:430. 

SALACINAE Dana, 1851a:430. 

CYCLACINAE Dana, 185la:431. 

TYCHIDAE Dana, 1851a:431. 

CRIOCARCININAE Dana, 185 la:431. 

CAMPOSCINAE Dana, 1851a:431. 

AMATHINAE Dana, 1851a:431. 

STENORHYNCHINAE Dana, 1851a:432. 

ACHAEINAE Dana, 1851a:432. 

PERICERIDAE Dana, 1851a:432. 

PARAMICIPPINAE Dana, 1851a:432. 

INACHOIDINAE Dana, 1851a:432. 

MENAETHINAE Dana, 1851a:433. 

STENOCIOPINAE Dana, 1851a:433. 

ONCININEA Dana, 1852b:77. 

ONCINOPIDAE Stimpson, 1858d:222. 

LEPTOPINAE Stimpson, 1871a: 109. 

NAXIINAE Stimpson, 1871a: 114. 

COLLODINAE Stimpson, 1871a: 119. 

ANOMALOPINAE Stimpson, 1871a: 124. 

ACANTHONYCHINAE Stimpson, 1871a: 127. 

IXIONINAE Neumann, 1878:10. 

ACANTHOPHYRINAE Neumann, 1878:10. 

PICROCERINAE Neumann, 1878:12. 

PODOCHELINAE Neumann, 1878:13. 

CYPHOCARCININAE Neumann, 1878:15. 

EURYNOMINAE Neumann, 1878:17. 

MICRORHYNCHINAE Miers, 1879a:651. 

SCHIZOPHRYSINAE Miers, 1879a:659. 

LISSOIDA Alcock, 1895:161. 

BLASTIDAE Stebbing, 1902:2. 

MAMAIIDAE Stebbing, 1905:22. 

OPHTHALMIINAE Balss, 1929:6. 

HYASTENIINAE Balss, 1929:8, 14. 

MACROCOELOMINAE Balss, 1929:8, 16, 20. 

OREGONHNAE Garth, 1958:134.


EASTERN ATLANTIC GENERA.—Twenty, of which 

the following 15 are represented by tropical species: 

Acanthonyx, Achaeus, Apiomithrax, Calypsachaeus, 

new genus, Capartiella, new genus, Dorhynchus, 

Ergasticus, Eurynome, Herbstia, Inachus, Macropodia, 

Maja, Micropisa, Pisa, and Stenorhynchus. The 

other genera are as follows: 

Anamathia Smith (1885:493). Substitute name 

for Amathia Roux (1828, pi. 3), an invalid junior 

homonym of Amathia Lamouroux, 1812; type-species: 

Amathia rissoana Roux, 1828, by monotypy; 

gender: feminine; name 1606 on Official List. 

Hyas Leach (1814:431). Type-species: Cancer araneus 

Linnaeus, 1758, by monotypy; gender: masculine. 

Hyastenus White (1847b:56). Type-species: Hyastenus 

sebae White, 1847, by monotypy; gender: 

masculine. 

Lissa Leach (1815b:69). Type-species: Cancer 

chiragra Fabricius, 1775, by monotypy; gender: 


Rochinia A. Milne Edwards (1875, in 1873- 

1881:86). Type-species: Rochinia gracilipes A. 

Milne Edwards, 1875, by monotypy; gender: feminine; 


EASTERN ATLANTIC SPECIES.—Sixty-six, of 

which 49 occur in tropical waters. Monod (1956) 

recorded the following species. 

Name in Monod 

Maja squinado 

Maja verrucosa 

Maja goltziana 

Eurynome aspera 

Herbstia rubra 


Pisa tetraodon 

Pisa nodipes 

Pisa gibbsi 

Pisa carinimana 

Micropisa ovata 

Apiomithrax violaceus 

Apiomithrax bocagei 

Acanthonyx lunulatus 

Dorhynchus thomsoni 

Ergasticus clouei 

Inachus angolensis 

Inachus dorsettensis 

Inachus guentheri 

Current Name 

Maja squinado 

Maja crispata 

Maja goltziana 

Eurynome aspera* 


Herbstia condyliata* 

Pisa tetraodon 

Pisa nodipes 

Pisa armata* 

Pisa carinimana* 

Micropisa ovata 

Apiomithrax violaceus* 

Apiomithrax bocagei 

Acanthonyx lunulatus 

Dorhynchus thomsoni 

Ergasticus clouei 

Inachus angolensis 

Inachus nanus, new species* 

Inachus guentheri 

Inachus phalangium 

Inachus thoracicus 

Inachus aguiari 

Inachus leptochirus 

Physachaeus (?) longipes 

Achaeus crane hi 

Achaeus fores ti 

Achaeus sp. 

Achaeus monodi 

Macropodia gilsoni 

Macropodia macrocheles 

Macropodia rostrata 

Macropodia straeleni 

Stenorhynchus seticomis 

Inachus phalangium 

Inachus biceps, new species* 

Inachus aguiarii 

Inachus leptochirus 

Capartiella longipes* 

Achaeus cranchii 

Achaeus fores ti* 

Achaeus trifalcatus 

Achaeus monodi 

Macropodia gilsoni* 

Macropodia macrocheles* 

Macropodia spinulosa* 

Macropodia straeleni* 

Stenorhynchus lanceolatus* 

Representatives of 24 species were taken by the 

Pillsbury, eight of them previously undescribed. 

The following species occur outside of the tropical 

region: 

Achaeus gracilis O. Costa, 1839 (= Achaeus gordonae 

Forest and Zariquiey Alvarez, 1955). Forest 

and Zariquiey Alvarez (1955:68, figs. 2, 4, 6, 8) 

pointed out that two species of Achaeus could be 

recognized in the Mediterranean, A. cranchii 

Leach, 1817, and a second species that they 

named Achaeus gordonae. In their account Forest 

and Zariquiey Alvarez noted that O. Costa (1839, 

in Costa and Costa, 1838-1871:25) in his Fauna 

del Regno di Napoli, named a small crab, Macropodia 

gracilis, which has the general aspect of an Achaeus 

and that Costa also had recognized A. cranchii. 

Forest and Zariquiey Alvarez (1955:66) considered 

Costa's M. gracilis to be unidentifiable ("espece 

douteuse") and they did not synonymize it 

with A. cranchii. An examination of Costa's account 

and figure (pi. 3: fig. lA,b) suggests to us 

that he was dealing with an Achaeus, one in which 

the rostral teeth are almost contiguous and the 

hepatic lobes of the carapace are poorly developed, 

characters which Forest and Zariquiey Alvarez 

used to separate A. gordonae from A. cranchii. 

In addition, in Costa's figure 1A the length/width 

ratio of the illustrated specimen is 1.44; this ratio 

was given by Forest and Zariquiey Alvarez as 

1.16 to 1.38 in A. cranchii, 1.32 to 1.55 in A. 

gordonae. Thus this ratio in A. gracilis is larger than 

that of A. cranchii but fits well within the range 

reported for A. gordonae. We believe that Achaeus

254 


gordonae must be considered a synonym of Macro- Macropodia longirostris (Fabricius, 1775). Medipodia 

gracilis O. Costa, 1839. Achaeus gracilis occurs terranean; sublittoral, between 4 and 50 m 

in the Mediterranean and adjacent Atlantic; sub- (Zariquiey Alvarez, 1968). 

littoral, to about 20 m (Zariquiey Alvarez, 1968). Macropodia rostrata (Linnaeus, 1761). Northeast- 

Anamathia rissoana (Roux, 1828). Azores and ern Atlantic, from Norway southward at least to 

eastern Mediterranean; sublittoral to about 400 Mediterranean; sublittoral; tropical West Africa 

m (Zariquiey Alvarez, 1968). 

records are referable to M. spinulosa (Zariquiey 

Eurynome spinosa Hailstone, 1835. Eastern Atlan- Alvarez, 1968; Christiansen, 1969; Forest, 1978). 

tic, from Scandinavia and British Isles southward Macropodia tenuirostris (Leach, 1814). Northeast- 

to NW Spain, Azores, Mediterranean; sublittoral, ern Atlantic, from the Faroes southward to Por- 

from about 180 to about 400 m (Zariquiey Altugal; sublittoral, to more than 150 m (Zariquiey 

varez, 1968; Christiansen, 1969). 

Alvarez, 1968; Christiansen, 1969; Forest, 1978). 

Hyas araneus (Linnaeus, 1758). Eastern Atlantic Pisa corallina (Risso, 1816). Mediterranean; sub- 

from Barents Sea southward to NW France; also littoral, usually in less than 10 m (Zariquiey 

western Atlantic; sublittoral, shallow water to Alvarez, 1968). 

more than 300 m (Christiansen, 1969). 

Pisa muscosa (Linnaeus, 1758). Mediterranean; 

Hyas coarctatus Leach, 1815. Eastern Atlantic sublittoral, from 4-5 to about 40 m (Zariquiey 

from Barents Sea and Arctic southward to NW Alvarez, 1968). 

France; also western Atlantic, northern Pacific; Rochinia carpenteri (Thomson, 1873). Eastern At- 

sublittoral, between 1 and 500 m (Christiansen, lantic, from Iceland and the Faroes southward to 

1969). 

NW Africa, Azores; sublittoral, from about 180 

Hyastenus hilgendorfi De Man, 1887. An Indo- to about 1300 m (Zariquiey Alvarez, 1968; Chris- 

West Pacific immigrant into the eastern Meditertiansen, 1969). 

ranean, now known from the coasts of Israel We believe that the following species, tenta- 

(Lewinsohn and Holthuis, 1964) and Egypt (Ratively recorded by Monod (1956) from West Afmadan 


rica, are based on erroneously labeled specimens 

Inachus communissimus Rizza, 1839. Mediterra- (each is known from a single specimen thought to 

nean and adjacent Atlantic, at least as far as be West African) and should not be included in 

Portugal; sublittoral, 15 to 24 m (Zariquiey Al- the West African fauna: 

varez, 1968). 

Libinia erinacea A. Milne Edwards, 1879. A west- 

Lissa chiragra (Fabricius, 1775). Mediterranean, ern Atlantic species recorded from "West Africa" 

sublittoral, between 20 and 40 m (Zariquiey Al- as Libinia dubia by Streets (1870:105) and subsevarez, 

1968). 

quently recorded by Rathbun (1925:321). Monod 

Macropodia czernjawskii (Brandt, 1880). Medit- (1956:514, fig. 705), who examined the specimen, 

teranean; sublittoral, between 10 and 30 m (Zar- identified it with L. erinacea. 

iquiey Alvarez, 1968). 

Notolopas brasiliensis Miers, 1886. A western At- 

Macropodia deflexa Forest, 1978. Northeastern lantic species reported by Monod (1956:513, figs. 

Atlantic, from southern England to Portugal; 703, 704), based on a specimen collected off 

sublittoral, shallow water to 20 m (Forest, 1978). South-West Africa in 1900. 

Macropodia intermedia Bouvier, 1940. (See page Rochinia gracilipes A. Milne Edwards, 1875. A 

300). 

western Atlantic species included by Monod 

Macropodia linaresi Forest and Zariquiey Al- (1956:516, figs. 706-708), on the basis of a single 

varez, 1964. Eastern Atlantic, from southern England 

and France to Spain, Mediterranean; sublittoral, 

between 30 and 80 m (Zariquiey Alvarez, 

1968; Forest, 1978). 

specimen from Gabon collected by Heurtel. 

Much of the material from Gabon in this collection 

apparently has been erroneously labeled. 

REMARKS.—In our account of this family we


have departed from our usual format in presenting 

synonymies for several Mediterranean-North 

Atlantic species which may not be represented in 

the tropical fauna. The occurrence of three such 

both coasts of the Americas (Rathbun, 1925; 

Garth, 1958); A. lunulatus (Risso, 1816) (with A. 

brevifrons A. Milne Edwards, 1869, as a synonym), 

from the eastern Atlantic (see p. 256, and Zari- 

species, Pisa armata, Acanthonyx lunulatus, and Herbsquiey Alvarez, 1968 for Mediterranean refertia 

condyliata, is well documented, but in the cases ences); A. sanctaehelenae Chace, 1966, from Saint 

of the latter two species older records may well Helena; A. simplex Dana, 1852, from the Pacific; 

pertain to other species recognized herein. and three species from the western Indian Ocean: 

The occurrence of several other northern spe- A. consobrinus A. Milne Edwards, 1862, A. limbatus 

cies off tropical West Africa should be verified: A. Milne Edwards, 1862, and A. elongatus Miers, 

Achaeus cranchii, Inachus aguiarii, I. dorsettensis (trop1877 

(Stephensen, 1945; Barnard, 1950; Guinot, 

ical records of which are referred to /. norms, new 1967a). 

species, p. 291), /. leptochirus (tropical records are Acanthonyx closely resembles Dehaanius Macreferred 

to /. biceps, new species, p. 285), /. phal- Leay, 1838, the only difference mentioned in the 

angium, I. thoracicus, Macropodia intermedia, M. lonliterature 

being the presence of a seven-segmented 

gipes, and Pisa nodipes. 

abdomen in the male in the latter, and an abdomen 

with six or less somites in the former. Ste- 

Subfamily EPIALTINAE MacLeay, 1838 

phensen (1945) and Barnard (1950) both suggest 

that this character is unreliable. The status of the 

Genus Acanthonyx Latreille, 1828 two genera requires clarification. 

According to accounts in the literature (cited 

Acanthonyx Latreille, 1828a:698 [type-species: Maia lunulata 

Risso, 1816, by monotypy; gender: masculine; name 1603 

on Official List, there erroneously dated 1827]. 

Gonosoma Costa, 1844:69 [type-species: Gonosoma viridis Costa, 

1844 (= Acanthonyx viridis Costa, 1838), a subjective junior 

above), the four Indo-West Pacific species of Acanthonyx 

can be distinguished from those occurring 

in the Atlantic as follows: A. simplex lacks dorsal 

spines or tubercles on the carapace; A. dentatus 

synonym of Maia lunulata Risso, 1816, by monotypy; gen- has a strong tooth on the orbital margin; A. 

der: neuter]. 

consobrinus has a tooth on the orbital margin and 

Peltinia Dana, 1851d:272 [type-species: Peltinia scutiformis 

Dana, 1851, a subjective junior synonym of Acanthonyx 

petiverii H. Milne Edwards, 1834, by present selection; 


four lateral projections on the carapace; and A. 

limbatus has much stronger lateral projections on 

the carapace. 

In his account of A. sanctaehelenae, Chace (1966: 

REMARKS.—Acanthonyx is dated from 1827 in 13) provided good comparative illustrations of 

the Official List of Generic Names in Zoology. So far the three species then known from the Atlantic, 

as we can determine, this is an error that origi- A. lunulatus, A. petiverii, and A. sanctaehelenae. 

nated in Neave, Nomenclator Zoologicus, in which Material of Acanthonyx available to us from 

1827 is given. Other genera named in the same West African localities includes one new species 

article by Latreille are correctly dated 1828 in from the Pillsbury collections, a second new species 

Neave. 

from Point-Noire, and A. brevifrons from the Cape 

Until now Acanthonyx included seven species, as Verde Islands. We recognize four species from the 

follows: A. petiverii H. Milne Edwards, 1834, from eastern Atlantic. 

Key to Eastern Atlantic Species of Acanthonyx 

1. Carapace with only 2 lateral lobes. [Rostral teeth short. 8-13 tubercles 

ventrally on dactyli of pereiopods] A. brevifrons 

Carapace with 3 lateral lobes 2

256 


2. Orbital margin with blunt tooth or projection. [Size very small, adults with 

carapace lengths of less than 4 mm. 6-8 tubercles ventrally (usually 6; 3 

or 4 in juveniles) on dactyli of walking legs] . . . . A. minor, new species 

Orbital margin smooth, evenly concave 3 

3. Rostral teeth strongly depressed. Rostral sinus V-shaped. Merus of claw 

with 1 tubercle. [Size very small, adults with carapace lengths of 

less than 5 mm. 4-6 tubercles ventrally on dactyli of walking legs] 

A. depressifrons, new species 

Rostral teeth not markedly depressed. Rostral sinus U-shaped. Merus of 

claw with 2 tubercles. [Size moderate to large, adults with carapace 

lengths of 15 mm or more. Males with 11-17 and females with 8-12 

tubercles ventrally on dactyli of walking legs] A. lunulatus 

Acanthonyx brevifrons A. Milne Edwards, 

1869 

FIGURES 60a, 61 

Acanthonyx brevifrons A. Milne Edwards, 1869:353.—A. Milne 

Edwards and Bouvier, 1894:12; 1900:152.—Bouvier, 

1940:349. 

?Acanthonyx lunulatus.—Miers, 1886:43.—Barrois, 1888:9. [See 

p. 258, references to A. lunulatus from Cape Verde Islands; 

not Acanthonyx lunulatus (Risso, 1816)]. 

?Acanthonyx lunulatus var. brevifrons.—Chapman and Santler, 

1955:375. 


Other Material: Cape Verde Islands; between Ilheu Branco 

and Ilheu Raso, 110-180 m, sand and rock, Talisman, 27 Jul 

1883, 1?, ljuv(W). 

DESCRIPTION.—Carapace (Figure 61a) pearshaped, 

length in midline 1/5 longer than maximum 

breadth. Dorsal surface of carapace smooth, 

lacking distinct pubescence (in material studied). 

Hepatic lobe scarcely produced anteriorly. One 

prominent rounded lobe on branchial region, 

lateral margin between hepatic and branchial 

lobes straight, subparallel or slightly convergent 

posteriorly. Rostral teeth, preorbital, hepatic, and 

branchial lobes each with terminal tuft of stout 

setae. Rostral sinus (Figure 61£,


FIGURE 60.—Carapaces of West African species of Acanthonyx in lateral view: a, A. brevifrons A. 

Milne Edwards, female, cl 8.8 mm, Cape Verde Islands; b, A. depressifrons, new species, holotype, 

male, cl 4.0 mm, Pointe-Noire; c, A. lunulatus (Risso), female, cl 6.8 mm, Tunis; d, A. minor, new 

species, paratype, male, cl 2.6 mm, Pilhbury Sta 271. 

Cape Verde Islands by A. Milne Edwards (1869); 

no depth data were given. Miers (1886:43) recorded 

a small female of A. lunulatus from Sao 

Vicente and noted that it approached the "species 

or variety Acanthonyx brevifrons A. Milne Edwards, 

in the form of the front, but there are indications 

of three antero-lateral teeth, and the carapace, as 

in Acanthonyx lunulatus, bears several tufts of setae." 

Miers' material, which could not be located, may 

be referable to A. lunulatus. A. Milne Edwards and 

Bouvier (1894, 1900) gave additional differences 

between this species and A. lunulatus, but did not 

give figures. Balss (1914) identified material from 

Annobon with this species, but in 1922 he synonymized 

A. brevifrons with A. lunulatus. We suspect 

that the material reported by Balss from Annobon 

actually belongs to a new species, A. minor (p. 

261). Bouvier (1940:348) suggested that A. brevifrons 

should be considered as a race of A. lunulatus, 

and Chapman and Sander (1955) identified material 

from the Azores as A. lunulatus var. brevifrons. 

Monod (1956) synonymized A. brevifrons with A. 

lunulatus, and figured a male of the latter species 

from Dakar. In his list of material, Monod noted 

that his figured specimen, a large male (cl 15 

mm), had an obsolete intermediate tooth on the 

carapace and corresponded to "f. brevifrons". The 

figured specimen appears to be a typical lunulatus, 

as does the specimen from Mauritania figured 

earlier by Monod (1933b, fig. 7D). None of the 

material of A. lunulatus that we have examined 

approaches our material of A. brevifrons in short- 

FIGURE 61.—Acanthonyx brevifrons A. Milne Edwards, Cape 

Verde Islands. Female, cl 8.8 mm: a, carapace; b, front, 

ventral view; c, cheliped; d, fifth pereiopod. Juvenile, cl 5.4 

mm: e, front, ventral view;/, chela.

258 

ness of rostral teeth or in the total suppression of 

the second lateral tooth on the carapace. 

Bouvier (1940) noted that A. lunulatus occurred 

to a depth of about 20 m; it is usually taken in 

shallow water on algae. In the Mediterranean it 

often occurs on the algal genus Cystoseira. Two of 

the records for A. brevifrons given by A. Milne 

Edwards and Bouvier (1900) are from depths of 

75 m and 110-180 m (our material is from the 

latter collection). It seems likely that A. brevifrons 

occurs in much deeper water than does A. lunulatus, 

and it is possible that both occur together in 

the Cape Verde Islands. All of the material from 

the Cape Verde Islands, the Azores, and Madeira 

(below) should be reexamined to determine which 

species occurs in the localities. 

BIOLOGY.—Little information is available on 

the ecology of this species. Our specimens were 

taken by the Talisman on sand and rock in 110- 

180 m; no information was given for the type. A. 

Milne Edwards and Bouvier (1900) also recorded 

material from 75 m but did not record the habitat. 

In 1894 these same authors recorded the 

species from algae and rocks in 10 m. Shallower 

water records for this species, as well as those 

based on material collected in the alga Cystoseira, 

a favorite habitat for A. lunulatus, may be based 

on this latter species. 

DISTRIBUTION.—Acanthonyx brevifrons has been 

reported from localities in the Azores, Madeira, 

and the Cape Verde Islands. At least some of the 

following records may be referable to A. lunulatus; 

all should be verified. Records in the literature 

include: 

Azores: No specific locality (A. Milne Edwards and Bouvier, 

1900; Bouvier, 1940; Barrois, 1888). E coast of Ilha das 

Flores, 39°26'30"N, 33°29'15"W of Paris (= 31°09'15"W of 

Greenwich), intertidal, and Graciosa, Ilheu da Praia, 39°- 

03'15"N, 30°18'15"W of Paris (= 27°48'15"W of Greenwich), 

beach, 10 m (A. Milne Edwards and Bouvier, 1894). 

Pasteleiro, Feteira, and Almoxarife [?], Ilha do Faial; and 

Madalena, Ilha do Pico (Chapman and Santler, 1955). 

Madeira: No specific locality (Chapman and Santler, 

1955). 

Cape Verde Islands: Sao Vicente (A. Milne Edwards, 

1869; Miers, 1886). Channel between Sao Vicente and Santo 


Antao, 75 m, and between Ilheu Branco and Ilheu Raso, 

110-180 m (A. Milne Edwards and Bouvier, 1900). 

Acanthonyx depress!frons, new species 

FIGURES 60i, 62 

MATERIAL EXAMINED.—Pilkbwy Material. None. 

Other Material: Congo: Pointe-Noire, 5-10 m, in lobster 

nets, 24 Jan 1967, J. Marteau, 26 (larger male is holotype), 

4$ (L, W). 

DESCRIPTION.—Carapace (Figure 62a) smooth, 

length 1.20-1.35 times greatest width, greatest 

width at level of anterolateral tooth. Surface with 

2 protogastric, 1 gastric, and 1 cardiac tubercle 

in midline, lateral branchial tubercle also present. 

All dorsal tubercles with 1 or 2 broad setae. 

Rostral sinus (Figure 62a,b) V-shaped, rostral 

teeth noticeably depressed in lateral view (Figure 

606), teeth blunt. Carapace, posterior to base of 

rostrum, with usual 2 diverging rows of curved, 

hook-shaped hairs. Preorbital tooth blunt, 

rounded, much shorter than rostral teeth. Orbital 

margin curving evenly to anterior margin of first 

anterolateral tooth, which is not markedly set off 

as in A. lunulatus. No tooth present on orbital 

margin as in A. minor. First anterolateral tooth 

much larger than second and third, which are 

distinct and subequal in size. Carapace lacking 

subhepatic tubercle. Eyes small. 

Basal segment of antennal peduncle (Figure 

62b) unarmed. Antennal peduncle extending to 

or beyond apices of rostral teeth. 

Chelipeds (Figure 62c) small in both sexes. 

Fingers about as long as palm, cutting edges 

dentate, gape slight. Palm smooth. Carpus with 

low dorsal crest; carpus about as long as palm, 

much shorter than merus. Merus with single 

blunt tubercle on basal half. 

Dactyli of walking legs (Figure 62d) broad, 

opposable margins with 4-6 pairs of teeth. Second 

pereiopod with dactyl shorter than propodus. 

Propodus about 3 times as long as high, outer 

surface with disc-like projection extending over 

base of dactylus; posterior margin of propodus 

with broad tubercle, ornamented with large setae, 

near midline. Carpus about as long as propodus,


a 

FIGURE 62.—Acanthonyx depressifrons, new species. Female paratype, cl 3.6 mm, Pointe-Noire: a, 

carapace. Holotype, male, cl 4.0 mm, Pointe-Noire: b, front, dorsal view; c, cheliped; d, fifth 

pereiopod; e, abdomen;/, gonopod. 

much shorter than merus. Third and fourth pereiopods 

shorter than, but otherwise similar to, 

second. Fifth pereiopod shortest. Dactylus about 

as long as propodus. Propodus stout, slightly more 

than 2 times as long as wide; tubercles on posterior 

margin low, with slender setae and club-like 

hairs. Carpus shorter than propodus. Merus 

longer than propodus. 

Male abdomen (Figure 62*) narrow. Telson 

triangular, length and width subequal. Sixth somite 

expanded distally. Male pleopod as illustrated 

(Figure 62/). 


3.5 to 4.0 mm, of females 3.3 to 4.3 mm. 

REMARKS.—Acanthonyx depressifrons differs from 

the other three eastern Atlantic species of the 

genus in having the rostrum strongly depressed 

and in having only one tubercle on the merus of 

the cheliped. It further differs from A. brevifrons in 

having three distinct lateral lobes on the carapace 

and fewer tubercles on the dactyli of the walking 

legs; from A. lunulatus in having the rostral sinus 

V-shaped and fewer tubercles on the dactyls of 

the pereiopods; and from A. minor, new species, in 

lacking a projection on the orbital margin and in 

having its greatest width at the level of the anterolateral 

tooth of the carapace in both sexes and 

at all sizes. Like A. minor, and perhaps A. brevifrons, 

A. depressifrons is a much smaller species than A. 

lunulatus, which can attain 15 mm or more in 

carapace length. 

Acanthonyx depressifrons is a much smaller species 

than A. petiverii or A. sanctaehelenae, both of which 

as adults may attain a carapace length of 18 mm 

(Rathbun, 1925; Chace, 1966). It also differs from 

A. petiverii in having the rostral sinus V-shaped 

rather than broadly U-shaped and from A. sanctaehelenae 

in having one rather than two lobes or 

tubercles on the outer margin of the merus of the 

chelipeds. 

TYPE-LOCALITY.—Off Pointe-Noire, Congo. 

DISPOSITION OF TYPES.—The holotype is the


larger male, carapace length 4.0 mm (USNM 

127191); three female paratypes (USNM 169535) 

also have been deposited in the collections of the 

Smithsonian Institution. A male and a female 

paratype have been deposited in the Rijksmuseum 

van Natuurlijke Histoire, Leiden. 


Latin and refers to the characteristic depressed 

front of this species. 


Point-Noire, Congo, where it was taken 

at a depth of 5-10 m. 

Acanthonyx lunulatus (Risso, 1816) 

FIGURES 60C, 63 

Acanthonyx lunulatus.—Capart, 1951:84, fig. 26.—Monod, 

1956:517, figs. 709, 710.—Rossignol, 1957:78, 116 

[key].—Forest and Games, 1960:356.—Gauld, 1960:72.— 

Rossignol, 1962:122.—Guinot and Ribeiro, 1962:76.— 

Ribeiro, 1964:20.—Chace, 1966, fig. 13a-d [Mediterranean].—Forest 

and Guinot, 1966:106.—Zariquiey Alvarez, 

1968:466, figs. 7d, 153e,f, 154i,j [Spain; references].—Kensley, 

1970:181.—Penrith and Kensley, 

1970b:252, 260.—Uschakov, 1970:455 [listed]. 

SYNONYMS.—Maia glabra Latreille, 1836; Acanthonyx 

viridis O. Costa, 1838; Gonosoma viridis O. 

Costa, 1844. 


Other Material: Madeira: SE coast near Agua de Pena, 

32°41'N, 16°46'W, 0-25 m, diving, 9 Mar 1976, Onversaagd 

Sta 27, 1$ (L). 

REMARKS.—Monod (1956) summarized earlier 

records for this species and provided a good illustration 

of a large male from Dakar. He reported 

FIGURE 63.—Acanthonyx lunulatus (Risso), female, cl 6.8 mm, Tunis: a, carapace; b, front, dorsal 

view; c, fifth pereiopod; d, distal segments of fifth pereiopod, enlarged.


numerous specimens from localities in Mauritania, 

Senegal, Ghana, and Cameroon, and noted 

that its range in the Atlantic extended from 

Portugal to the equatorial region. 

We have examined material of A. lunulatus from 

the Gulf of Tunis, Messina and Trapani, Sicily, 

Naples, Nice, and Cadaques, Spain (all W) and 

have provided here some sketches of a young 

female from the Gulf of Tunis (Figures 60c, 63) 

to help distinguish this species from the three 

others occurring off West Africa. 

Our discovery of two previously undescribed 

species of Acanthonyx in West African waters and 

our recognition of A. brevifrons from the Cape 

Verde Islands and other localities indicate that 

earlier records of A. lunulatus may have been based 

on one or more of the four species now known to 

occur off West Africa. The specimens reported in 

Monod, the literature records cited by him, and 

the records given below all need to be verified. 


Atlantic, from Portugal to South-West Africa, 

generally in shallow water, littoral and subtidal. 

Records since 1956 include the following. 

Morocco: Rabat, Temara, Skhirat, Oued Cherrat, David, 

Tillet [?], Sidi Moussa, Agadir, and Foum Assaka (all Forest 

and Games, 1960). 

Cape Verde Islands: Matiota, Sao Vicente, beach and 

shore, and Tarrafal do Monte Trigo, Santo Antao (Guinot 



Ivory Coast: Lagoon of Abidjan, 05°16'12"N, 04°00'- 

20"W (Forest and Guinot, 1966). 

Ghana: Tenkpobo to Dixcove (Gauld, 1960). 

Sao Tome: Ilheu Macao (as Hot dos Cocos), 3-8 m, and 

Sant'Ana, shore (Forest and Guinot, 1966). 

Congo: Pointe Indienne (Rossignol, 1957). Baie de 

Loango and near Pointe-Noire (Rossignol, 1962). 

Angola: Baia do Lobito, shore; Praia das Conchas, Mocamedes, 

shore; and Baia das Vacas, Benguela, shore (all 


South-West Africa: Rocky Point, 18°59'S, 12°29'E (Kensley, 

1970; Penrith and Kensley, 1970b). Kunene River 

mouth, 17°15'S, 11°45'E, and Mowe Point, 19°23'S, 12°- 

42'E (Kensley, 1970). 

Mowe Point is the southernmost record for 

Acanthonyx on the West African coast. 

* Acanthonyx minor, new species 

FIGURES 60rf, 64 

?Acanthonyx brevifrons.—Balss, 1914:101 [not A. brevifrons A. 

Milne Edwards, 1869]. 


Sta 271, shore, 6c5 (includes holotype), 4$ (1 ov) (L, W). Sta 

273, shore, 1

262 

a 


FIGURE 64.—Acanthonyx minor, new species, Pillsbury Sta 271. Female paratype, cl 3.8 mm: a, 

carapace; b, front, ventral view; c, cheliped; d, second pereiopod; e, fifth pereiopod. Male 

paratype, cl 2.6 mm: /, carapace; g, abdomen; h, first pleopod. Male paratype, cl 1.7 mm: i, 

carapace. 

Fingers almost as long as palm (measured along 

upper surface). Cutting edges dentate, with only 

slight gape proximally. Palm smooth. Carpus 

globular, smooth, bearing no carinae or tubercles, 

shorter than palm, much shorter than merus. 

Latter smooth except for 2 blunt tubercles on 

basal half. 

Walking legs similar to those of other Acanthonyx 

species, but dactylus appearing more slender, 

with tip less strongly curved, terminating in slender 

apex, posterior margin with 7 or 8 small but 

distinct teeth. Second pereiopod (first walking 

leg) dactylus about as long as propodus (measured 

dorsally). Latter 4 times as long as high, 

h


outer surface ending in disc-like projection covering 

base of dactylus. Posterior margin of propodus 

with low tubercle or prominence near midlength 

ornamented with strong, stiff hairs. Carpus 

as long as propodus, much shorter and narrower 

than merus. First walking leg longer than remainder, 

latter of similar shape but decreasing in size 

posteriorly. Fifth pereiopod (Figure 64*) shortest. 

Dactylus slightly longer than propodus, more 

robust than that of second leg. Propodus about 

2.5 times as long as wide. Tubercle on posterior 

margin low but distinct because of tuft of hairs. 

Carpus distinctly shorter than propodus, merus 

about as long as propodus. 

Male abdomen (Figure 64g) narrow. Telson 

tongue-shaped. Sixth somite broad proximally, 

distinctly narrowing at telson. Abdomen of largest 

females very broad, reaching to bases of legs; 

sixth somite widest. 

First male pleopod as shown in Figure 64h. It 

does not differ significantly from that of A. sanctaehelenae 

Chace from Saint Helena (Chace, 1966, 

fig. 12f). 


1.7 to 3.4 mm, of non-ovigerous females 1.8 to 3.8 

mm, of the ovigerous female 3.7 mm. 

REMARKS.—Acanthonyx minor, like A. depressifrons, 

is a very small species, as the name implies. It can 

be distinguished from all of the Atlantic species 

now known by the distinct projection on the 

orbital margin; that margin is evenly concave in 

the other species. 

It is likely that the two specimens reported by 

Balss (1914:101) from Annobon under the name 

A. brevifrons belong to the present species; Balss 

indicated that they were juveniles. Later, Balss 

(1922:72) identified this material with A. lunulatus, 

synonymizing A. brevifrons with that species. 


05°38'E, equatorial Guinea. 


D. 31762), a male from Pillsbury Sta 271 with a 

carapace length of 3.4 mm, and seven paratypes 

are in the Rijksmuseum van Natuurlijke Historie, 

Leiden; two male and one female paratypes are 

in the collection of the Smithsonian Institution. 

ETYMOLOGY—The specific epithet is from the 

Latin and refers to the diminutive size of this 

species. 

BIOLOGY.—The present material all originates 

from Annobon. It was collected at two fish poison 

stations on the shore of Annobon. The first (Sta 

271) was situated on the NE shore between Punta 

Yoyo and Punta Pedrinha, the coast here was 

rocky with rubble and a small sandy beach; 

collecting was done in the surge where the water 

was rather turbid. The second station (Sta 273) 

was on the north shore of Annobon near Islote 

Piramide on a rocky and sandy shore. Because of 

the high surf, collecting was difficult, but most of 

the fishes and invertebrates killed at this station 

were gathered at the outer edge of the surf zone 

in about 5 to 8 feet (1.5 to 2.4 m) of water over 

rocks. 


Annobon Island, equatorial Guinea. 

Subfamily INACHINAE MacLeay, 1838 

REMARKS.—The extensive Pillsbury collections 

of inachine crabs has necessitated a much more 

detailed coverage of this subfamily than we had 

originally expected. Eight genera of this subfamily 

occur off West Africa. Two of these are characterized 

and named herein. Representatives of 

six genera are included in the Pillsbury material; 

Dorhynchus and Ergasticus, each represented in the 

eastern Atlantic by one species, were not collected. 

37 species of inachine crabs are now known 

to occur in the eastern Atlantic, a surprising 

increase in recognized species since 1956 when 

Monod recognized 22 species, 19 from tropical 

West African waters; we recognize 29 tropical 

species below. 

Because the Pillsbury collections included what 

appear to be representatives of two new genera, 

we have examined representatives of Achaeopsis, 

Dorhynchus, Ergasticus, Physachaeus, and several related 

genera in order to better understand generic 

characters and relationships. As a result, we believe 

that Dorhynchus Thomson, 1873, is distinct 

from Achaeopsis Stimpson, 1858. We have in-

264 

eluded a diagnosis of Dorhynchus (p. 279), under 

which our conclusions are discussed. 

We also believe that three interrelated "major" 

morphological features of inachine crabs are subject 

to misinterpretation: the supraorbital spine, 

the postorbital spine(s), and whether or not the 

eyes are retractile. 

The supraorbital spine of Dorhynchus (Barnard, 

1950, fig. 4d) or of Macropodia gilsoni probably 

should be interpreted as an outgrowth of the 

supraorbital eave rather than as being homologous 

with the intercalated spine or lobe in the 

Majinae. Balss (1929) stressed the importance of 

the intercalated spine in majid classification but 

his conclusions do not appear to have gained 

general acceptance. Garth (1958:37), however, 

includes in his key to American genera of Inachinae 

the genus Achaeopsis, which can be distinguished 

by the presence of a spine intercalated 

between pre- and postorbital spines. Griffin 

(1966b: 35) suggested that "if the supraorbital 

spine found in Achaeopsis species is regarded as 

homologous with the intercalated spine of other 

majids, Achaeopsis becomes widely separated from 

these genera ...." We agree with Griffin, for 

Achaeopsis and the closely related Dorhynchus 

(which Griffin considered a synonym of Achaeopsis), 

are otherwise similar to Macropodia, differing 

primarily in having the rostral spines divergent 

rather than appressed. Further, the West African 

Macropodia gilsoni and M. intermedia, which have a 

supraorbital spine but otherwise closely resemble 

the other species of the genus, would have to be 

separated into a distinct genus, a taxon which, to 

us, would be unnecessary and unnatural. We 

suspect that the supraorbital spine, like that of 

Dorhynchus and Macropodia gilsoni, which clearly 

arises from the supraorbital eave, should not be 

considered homologous to the intercalated spine. 

In the case of M. gilsoni, it is one of several specific 

characters that can be used to distinguish that 

species. 

The question of homologies of the postorbital 

or postocular spine in the subfamily Inachinae is 


perhaps more troublesome. The term has been 

used for a spine arising from the posterior margin 

of the orbit (Rathbun, 1925, fig. 1, for example), 

as well as for any spine that may be situated on 

the lateral margin of the carapace between the 

orbital margin and the hepatic lobe, i.e., posterior 

to the orbit (Griffin, 1966a, fig. 1, upper left 

figure). If this latter spine, which is situated between 

the orbit and the hepatic lobe, is considered 

to be a displaced, "true" postorbital spine like 

that of Inachus, then, again, Macropodia gilsoni 

would have to be generically distinct from M. 

rostrata, which lacks it. We believe that this latter 

spine should not be considered to be a true 

postorbital spine; we propose to call this the 

nuchal spine to differentiate it from the postorbital. 

Among the eastern Atlantic genera of Inachinae, 

a postorbital spine is found only in Inachus 

and Ergasticus; in Dorhynchus, Achaeopsis, Stenorhynchus, 

and in some species of Macropodia, there is a 

nuchal spine but no postorbital spine. Indeed, the 

eastern Atlantic species of Macropodia show a complete 

range of development of the nuchal spine. 

Finally, the question of whether the eye is 

retractile or not may be one of semantics but it is 

a feature that can be difficult to interpret. Among 

the eastern Atlantic genera of Inachinae, all of 

which have movable eyes, the eye in some appears 

to be retractile, that is, it can be moved into a 

position adjacent to the postocular spine. This 

occurs only in two genera, Inachus and Ergasticus. 

In Dorhynchus, in which the eye is said to be 

retractile to the sides of the carapace (Rathbun, 

1925:27, definition of Achaeopsis) (and which also 

has been described as having a postocular spine 

but which we interpret as a nuchal spine), the eye 

in material available to us is no more retractile 

than in our material of Macropodia. In Macropodia 

the eye is described as not retractile (Christiansen, 

1969:110), that is, the eye in representatives of 

the two genera folds backward about to the same 

level. The eye can approach the carapace in 

Dorhynchus, but, we suspect, only because the carapace 

is broader in that genus and the angle


formed by a line extending from the orbit to the 

hepatic lobe is quite different in Dorhynchus than 

it is in Macropodia. We would describe the eye as 

retractile only when it can be folded back into a 

postocular process like that of Inachus. 

Finally, although Ergasticus was assigned to the 

subfamily Pisinae by Bouvier (1940) and by Zariquiey 

Alvarez (1968), we can see no reason to 

Key to Eastern Atlantic Genera of Inachinae 

exclude it from the Inachinae, where it was placed 

by Balss (1957) and Monod (1956). The basal 

antennal article is extremely slender and the orbit 

is incomplete, with a supraorbital eave divided 

into spines, a large, postorbital spine, but no 

ventral orbital margin. We have not examined 

the male pleopod, which does not appear to have 

been illustrated. 

1. Rostrum undivided, elongate (as long as or longer than postrostral carapace), 

with spinules laterally. Basal antennal article convex ventrally, 

not longitudinally sulcate. [Abdomen of 6 somites in male, 5 in female] 

Stenorhynchus 

Rostrum distinctly bilobed or very short, anteriorly truncate. Basal antennal 

article flattened or longitudinally channelled ventrally 2 

2. Orbit with distinct spine or cupped process on posterior margin against 

which eye can be folded 3 

Orbit lacking distinct spine or process on posterior margin. [Carapace, 

between orbit and hepatic lobe, smooth, tuberculate, or with erect spine 

(nuchal). Abdomen of 6 somites in both sexes] 4 

3. Supraorbital spines present. Postorbital spine slender, not a cupped process. 

Rostrum composed of 2 long, slender, divergent spines. First walking leg 

not markedly longer than second. Abdomen of 7 somites in male, 6 in 

female Ergasticus 

Supraorbital margin unarmed dorsally. Postorbital spine a broad, cupped 

process. Rostrum short, of 2 blunt lobes or broad, triangular spines. First 

walking leg much longer than second, noticeably enlarged. Abdomen of 

6 somites in both sexes Inachus 

4. Rostral spines slender, divergent, upturned, extending anteriorly slightly 

beyond end of antennal peduncle. [Distinct supraorbital spine present. 

Antennular fossae grading into rostrum, lacking anterior rim. Nuchal 

spine present. Merus of pereiopods with distal spine(s). Dactyli of 

posterior 2 pereiopods not falciform] Dorhynchus 

Rostral spines usually short, separate, triangular or with spiniform apices, 

or composed of 2 appressed spines of varying length 5 

5. Front truncate, simple or obscurely bilobed, overreached slightly by interantennular 

septum. Carapace lacking spines dorsally. [Antennular fossae 

with anterior rim. Interantennular septum not produced ventrally into 

beak. None of dactyli falciform] Capartiella, new genus 

Front produced into 2 spines or spined lobes. Carapace with spines or 

tubercles dorsally. [Interantennular spine may be visible at base of rostral 

teeth] 6

266 


6. Interantennular septum distinct but not strongly produced ventrally, 

scarcely if at all visible in lateral view below level of basal antennal 

segment. [Rostrum bilobed. Antennular fossae lacking anterior rim, 

grading into rostrum. Chelae of male inflated. Dactyli of posterior 2 or 

3 pereiopods falciform] Achaeus 

Interantennular septum produced ventrally into distinct, prominent beak, 

visible in lateral view, extending well below level of basal antennal 

segment 

Rostrum produced into 2 distinctly separate spines, spinulous laterally, 

truncated and spinulous apically. Chelae compressed, palm cristate 

above and below in both sexes. Orbital margin spinulous above. [Basal 

article of antenna lined with spines on inner and outer margins. Dactyli 

of posterior 2 pereiopods falciform] Calypsachaeus, new genus 

Rostrum produced into 2 smooth, unarmed spines, contiguous throughout 

their length. Chelae inflated, especially in adult males. Orbital margins 

unarmed above or with 1 or 2 dorsal spines, not lined with spinules. 

[Basal article of antenna, if armed, with 1 line or irregular row of 

tubercles or spines] Macropodia 

Genus Achaeus Leach, 1817 

Achaeus Leach, 1817, in 1815-1875, legend of pi. 22C [typespecies: 

Achaeuscranchii'Leach, 1817,bymonotypy; gender: 


DIAGNOSIS.—Carapace pyriform to elongate triangular, 

length usually greater than width, somewhat 

narrowed behind orbit as a neck, smooth or 

ornamented with tubercles, spinules, or, rarely, 

with long dorsal spine on cardiac or gastric region 

or both (as in A. monodi sensu stricto). Rostrum 

very short, of 2 acute or rounded lobes or of 2 

spinules. Orbit dorsally with only narrow supraorbital 

eave, laterally spinulated or smooth; 

postorbital spine absent, nuchal lobe or spine, if 

present, not prominent. Eyestalks long, nonretractile, 

cornea obliquely subterminal, slightly 

ventral, large, ovoid. Interantennular septum not 

produced into ventrally directed beak. Basal antennal 

article extremely slender, smooth, channelled 

longitudinally, weakly tuberculate, spinulous, 

or armed with lateral spines, spines not 

arranged in 2 rows flanking longitudinal channel. 

Merus of third maxilliped ovate, not notched 

distally, narrower than ischium, palp articulating 

at summit. Chelipeds usually spinous, greatly 

enlarged with inflated palm in males. Walking 

legs long and slender, dactyli of posterior 2 or 3 

pairs usually strongly falcate and ventrally spinulated. 

Abdomen of 6 somites in both sexes, 

male abdomen widest at middle of third somite. 

Male first pleopod bluntly pointed, aperture located 

subterminally in groove (modified from 

Griffin, 1966b). 

REMARKS.—Achaeus now includes more than 30 

species (Serene, 1968, listed 25 Indo-West Pacific 

species) occurring in the eastern Atlantic and in 

the Indo-West Pacific, from South Africa to Japan 

(Griffin, 1966b, 1968a; Griffin and Yaldwyn, 

1965). 

The Pillsbury collections, comprising four species, 

bring to six the number of nominal species 

known from tropical West Africa, between Senegal 

and Angola. Two of the species taken by the 

Pillsbury are described here as new, and the other 

two are redescribed. One species, originally assigned 

to Achaeus and described from the Calypso 

collections in 1966 (Forest and Guinot, 1966), is 

transferred to another genus, Calypsachaeus. 

The seventh eastern Atlantic species, A. gracilis 

(O. Costa, 1839) (= A. gordonae Forest and Zariquiey 

Alvarez, 1955), occurs in the Mediterranean 

and so far has not been recorded from 

Atlantic localities. Monod (1956:542, figs. 767- 

770) supplemented the original description.


It seems very likely that several additional 

species of Achaeus from West Africa remain to be 

recognized and described. Monod (1956:539, figs. 

747-766) almost certainly included two species in 

his account of A. cranchii: A. cranchii sensu stricto 

from European waters and an undescribed species 

from Senegal. The male pleopods of the two 

species, as figured by Monod (1956, figs. 760-762 

from Senegal and figs. 763-766 from Spain, 

France, and the Canary Islands) are quite different. 

Monod's account of Achaeus monodi (1956:548, 

figs. 782-810), which he, as well as Forest and 

Guinot (1966:110), considered to be an extremely 

variable species, may be represented in Monod's 

account by three distinct species: (1) A. monodi 

sensu stricto (fig. 782, reproduced from Capart, 

1951, pi. 1: fig. 11); a broad, pyriform species 

from the Congo and Angola with spiniform rostral 

spines and slender, erect spines dorsally on 

the gastric and cardiac regions of the carapace; 

(2) a slender, elongate-triangular species from 

Senegal with broad, triangular rostral spines and 

no dorsal spines (figs. 783-785); and (3) a second 

species from Senegal with appressed rostral spines 

(figs. 786-789), similar to those of the South 

African A. spinosissimus Griffin, 1968(b). None of 

these species was represented in the Pillsbury collections. 

Our generic diagnosis, based on Griffin 

(1966b), has been modified to include A. monodi, 

sensu stricto, with its erect dorsal spines. That 

species should be restudied, however, because it 

is different enough from the other species of 

Achaeus for Capart (1951) to have assigned it to 

Podochela. 

The relative proportions of the carapace, even 

though they may vary within a species with size 

and sex, provide a helpful recognition character 

for species of Achaeus. They allow the immediate 

distinction between slender species, like A. turbator, 

new species (in which the carapace width varies 

from 0.64 to 0.72 times the carapace length), from 

the broader species, like A. foresti (in which the 

carapace width varies from 0.86 to 0.90 times the 

length). Key characters employed for species not 

in the collections have been extracted from the 

literature: Forest and Zariquiey Alvarez, 1955, 

for A. cranchii and A. gracilis (as A. gordonae); Forest 

and Guinot, 1966, for A. trifalcatus; and Forest, 

1968, for A. foresti. A similar feature to illustrate 

the relative proportions of the carapace, the relation 

of the width at the hepatic lobes to the 

distance between the summit of the gastric prominence 

and the anterior margin of the rostral 

spines, also proves to be helpful. In A. turbator, 

new species, for example, the width of the carapace 

at the hepatic lobes is less than the distance 

from the gastric prominence to the anterior margin 

of the rostral spines, whereas in A. buderes, new 

species, the hepatic width is far greater. The 

relative length of the neck also provides a helpful 

recognition character: the neck is almost absent 

in A. buderes, new species, whereas it is quite 

distinct and elongate in A. turbator, new species. 

The shape of the rostral spines or projections also 

is important and appears to be characteristic. In 

A. trifalcatus the rostrum is made up of two 

rounded, unarmed lobes; in A. turbator, new species, 

it is made up of two rounded lobes, each of 

which has an anterior spinule. There is a definite 

shoulder on each side of the apical spinule; and 

in A. cranchii and A. gracilis the rostrum is made 

up of two slender, separate spines, tapering evenly 

from the base to the apex and lacking a distinct 

shoulder posteriorly. Note that in A. cursor (A. 

Milne-Edwards and Bouvier, 1900:161, pi. 21: 

figs. 15, 16, and pi. 22: figs. 1-3), considered a 

synonym of A. cranchii by Forest and Zariquiey 

Alvarez, 1955, there may be a distinct shoulder 

at the base of the rostrum. The proportions of the 

segments of the pereiopods also appears to be 

important. Forest (1968), for example, pointed 

out that in A. foresti the dactylus of the fourth 

pereiopod, measured across its arc, is equal to the 

length of the ischium and merus together; in A. 

monodi, according to the figure of the holotype 

(Capart, 1951, fig. 31), the dactylus, measured 

across its arc, is less than half as long as the merus 

alone. The size of the eye (compare Figures 65 

and 66A of the eye in A. foresti and A. buderes, new 

species) appears to be important. Finally, the 

relative height of the gastric and cardiac promi-


nences and whether or not they are produced into The key to eastern Atlantic Achaeus given below 

erect spines seems to be important; in none of the should be used with caution, for it does not take 

eastern Atlantic species are these spines so well into account the "variants" identified with A. 

developed as in A. monodi sensu stricto. cranchii and A. monodi by Monod (1956). 

Key to Eastern Atlantic Species of Achaeus 

1. Rostrum unarmed, scarcely bilobed. Posterior 3 pereiopods with falciform 

dactyli. [Carapace lacking erect dorsal spines. Width between hepatic 

lobes greater than distance between gastric prominence and anterior 

margin. Carapace width 0.70 to 0.83 times carapace length] 

A. trifalcatus 

Rostrum with apical spinules or spines. At most posterior 2 pereiopods with 

falciform dactyli 2 

2. Rostrum consisting of 2 sharp spines, tapering evenly from base to 

apex 3 

Rostrum consisting of 2 lobes, each with apical spinule (i.e., rostrum 

posteriorly with distinct shoulder) 4 

3. Fissure between rostral spines narrow. Orbits usually unarmed dorsally, 

occasionally with tubercle or spinules. Posteromedian margin of carapace 

unarmed. Cardiac protuberance rounded, lower than gastric protuberance. 

Width between hepatic lobes equal to distance between gastric 

prominence and anterior margin. [Carapace width 0.65 to 0.79 times 

carapace length in males, 0.75 to 0.86 times carapace length in females] 

A. gracilis 

Fissure between rostral spines wide, U- or V-shaped. Orbits armed dorsally. 

Posteromedian margin of carapace usually tuberculate or lightly spinulose. 

Cardiac protuberance conical, as high as gastric protuberance. 

Width between hepatic lobes greater than distance between gastric 

prominence and anterior margin. [Carapace width 0.73 to 0.87 times 

carapace length in males, 0.75 to 0.95 times carapace length in females] 

A. cranchii 

4. Carapace with erect spines on both cardiac and gastric regions. [Width 

between hepatic lobes greater than distance from gastric protuberance 

to anterior margin. Carapace width greater than 0.8 times carapace 

length. Only 1 pereiopod with falciform dactylus (?)] A. monodi 

Carapace without erect spines on either cardiac and gastric regions .... 5 

5. Supraorbital margin spinulate. Posterior margin of carapace with strong 

spines posterolaterally, smaller spinules and tubercles posteriorly. Eyes 

slender, diameter of stalk about one-fourth length. [Width between 

hepatic lobes greater than distance from gastric protuberance to anterior 

margin. Carapace width 0.86 to 0.90 times carapace length] . A. foresti 

Supraorbital margin unarmed. Posterior margin of carapace unarmed, 

with at most low, inconspicuous tubercles posterolaterally. Eyes stout, 

diameter about half length 6

NUMBER 306 

6. Carapace stout, pyriform, greatest width 0.67 to 0.81 (mean 0.75) times 

carapace length. Width at hepatic lobes greater than distance from 

gastric protuberance to anterior margin. Hepatic lobe with apical spine 

and several subapical spinules. Neck very short, stout 

A. buderes, new species 

Carapace slender, elongate triangular, greatest width 0.64 to 0.72 (mean 

0.68) times carapace length. Width at hepatic lobes subequal to or less 

than distance from gastric protuberance to anterior margin. Hepatic 

lobe angled apically, unarmed. Neck very long, tapering anteriorly 

A. turbator, new species 

*Achaeus buderes, new species 

FIGURE 65 


Sta 17, 48 m, fine sand and green mud, 1

270 

spines distally and 1 or 2 tubercles or spinules 

proximally. Antennal segments unarmed. 

Antennular fossae large, longitudinally suboval, 

margins of fossae smooth. Basal antennular 

segment with line of tubercles ventrally. Interantennular 

process slender, terminating posteriorly 

in rounded right angle. Anterior process of 

epistome very thin. 

Epistome (Figure 65c) broader than long, 

widening posteriorly, most of surface smooth, 

with sharp spinule anterolateral to opening of 

antennal gland. Pterygostomian region with 

sharp spine along lateral border. 

Third maxillipeds not meeting in midline, 

hairy, not heavily spined. Ischium with 2 parallel 

lines of sharp denticles, mesial margin tuberculate 

or with small spines, lacking large spines. Merus 

with line of sharp denticles on surface, several 

small spines on mesial margin. Palp with slender 

dactylus, shorter than stout carpus and propodus 

combined. Carpus and propodus each with distal 

spine mesially. 

Chelipeds slender in adult female, merus not 

extending beyond orbit. Ischium with line of 

spines ventrally. Merus with line of spines, some 

large, on ventrolateral border, 2 widely separated 

and 1 distal spine dorsally, and 1 distal spine 

mesially. Carpus with proximal outer spine, 2 

spines on oblique dorsal line, and 1 inner distal 

spine, flanked proximally by sharp tubercle. 

Chela shorter than remainder of cheliped, fingers 

longer than palm. Palm with row of spines dorsally 

and ventrally, inner and outer surfaces 

smooth. Fingers flattened, lacking prominent 

gape (in $), cutting edges crenulate, lacking enlarged 

tooth on movable finger. Chela largely 

naked, with few long and short hairs scattered 

over surface. 

Walking legs short, slender, merus and propodus 

of first walking leg each shorter than carapace, 

with curled hairs arising singly on dorsal 

surface of carpus and propodus, longer hairs scattered 

over surfaces of both segments. First walking 

leg longest, remainder decreasing in length posteriorly. 

Proximal segments of walking legs unarmed, 

merus lacking conspicuous distal dorsal 


spine. Dactylus of first walking leg straight proximally, 

slightly curved distally, unarmed ventrally, 

more than half as long as propodus, latter 

slightly shorter than merus. Dactylus of second 

walking leg shorter than that of first, more evenly 

curved, with 1 subdistal tooth ventrally; dactylus, 

measured across arc, more than half as long as 

propodus, latter shorter than merus. Dactylus of 

third walking leg falciform, with 3 large distal 

and 3 smaller proximal ventral teeth, length, 

measured across arc, slightly more than half propodus 

length, latter 4/5 as long as merus. Dactylus 

of fourth walking leg (Figure 65d) falciform, 

with 5 or 6 large distal and several smaller proximal 

teeth ventrally, length, measured across arc, 

more than half that of propodus, latter shorter 

than merus. Sternum unarmed. Male too damaged 

to determine details of abdomen and gonopod. 

MEASUREMENTS.—Carapace length of male 4.7 

mm, of non-ovigerous females 3.4 to 4.0 mm, of 

ovigerous females 3.0 to 7.0 mm. 

REMARKS.—Achaeus buderes, with its stout, pyriform 

carapace with a very short neck, most closely 

resembles A. foresti. As in that species, there is a 

spinulose patch on the anterior margin of the eye, 

the hepatic lobes are armed, and there are spines 

and tubercles laterally on the carapace. This new 

species differs from A. foresti in having much 

stouter eyes, with the diameter of the stalk about 

half its length, having the supraorbital margin 

unarmed, having the posterior margin of the 

carapace unarmed, and in having a much shorter 

dactylus on the fourth pereiopods. In A. foresti 

that dactylus, measured across its arc, is longer 

than the ischium and merus combined, whereas 

in A. buderes it is slightly longer than half the 

length of the merus. 

TYPE-LOCALITY.—Off Ghana, 05°10'N, 00°- 

25'W to 05°08'N, OO°28'W, in 42 m (Pillsbury Sta 

23). 


D. 23903), an ovigerous female, is in the Rijksmuseum 

van Natuurlijke Historie, Leiden. One 

lot of paratypes is also deposited there and two 

lots of paratypes are in the National Museum of



D.C. 


from the Greek prefix bou-, large, and dere, neck. 

BIOLOGY.—This species was taken by the Pillsbury 

in depths between 35 and 51m, usually on 

soft bottom, fine sand and green mud, with bryozoans. 

Ovigerous females were taken in May. 

DISTRIBUTION.—Known only from type-locality, 

in 35 to 51 m. 

Achaeus cranchii Leach, 1817 

Achaeus cranchi.—Monod, 1956:539, 632, figs. 747-766 [Europe, 

Mediterranean, Canary Islands, Senegal, Sierra 

Leone].—Longhurst, 1958:89 [Sierra Leone].—Forest and 


Achaeus cranchii.—Zariquiey Alvarez, 1968:474, fig. 160e,f 


38 [North Atlantic]. 

SYNONYM.—?Achaeus cursor A. Milne Edwards 


DISTRIBUTION.—Eastern Atlantic from the British 

Isles southward to Sierra Leone, including the 

Azores and the Canary Islands; Mediterranean. 

Littoral to at least 68 m. 

* Achaeus foresti Monod, 1956 

FIGURE 66 

Achaeus foresti Monod, 1956:545, figs. 771-776.—Forest, 

1968:10%, figs. 1-6. 

MATERIAL EXAMINED.—PUlsbury Material: Liberia: 

Sta 68, 70 m, broken shell, 1$ (?). 

Ghana: Sta 17, 48 m, fine sand and green mud, 16* (?). 

DESCRIPTION.—Forest, 1968:1096. 

Male Pleopod: Forest, 1968, fig. 6 (Ghana). 

MEASUREMENTS.—Carapace length of male 2.8 

mm, of female 3.7 mm. The only other known 

specimen, the holotype, is an ovigerous female 

with a carapace length of 5 mm. 

REMARKS.—Forest (1968) redescribed this species 

from two specimens collected by the PUlsbury. 

His figures have been reproduced here (Figure 

66) for the sake of completeness and to facilitate 

the recognition of this species and the other species 

from the PUlsbury collections. 

The holotype, in the Museum national 

d'Histoire naturelle, Paris, is in poor condition. 

As Forest (1968:1098) pointed out, all that remains 

of the type is the carapace on which the 

apices of the rostrum have been broken. Thus, 

these rostral spinules are shown by Forest (1968, 

fig. 1) but the rostrum appears unarmed in the 

illustrations given by Monod (1956, figs. 771— 

773); this latter figure also does not show the 

marginal spinules on the basal portion of the 

rostrum. Monod (1956, fig. 776) illustrates one of 

the pereiopods of the type on which the merus is 

h 

FIGURE-66.—Achaeus foresti Monod. Female, cl 3.7 mm, PUlsbury 

Sta 68: a, carapace; b, eye; c, d, right cheliped, viewed 

from different angles. Male, cl 2.8 mm, PUlsbury Sta 17: e, 

right cheliped; /, fifth pereiopod; g,h, gonopod in anterior 

and posterior views. (All from Forest, 1968, figs. 1-6.)

272 

considerably longer than the propodus and the 

dactylus is subequal to the propodus; he may 

have shown the second pereiopod, for in the other 

species of Achaeus reported here that is the only 

pereiopod with the dactylus unarmed. Forest 

(1968) described the second pereiopod as having 

the merus shorter than the propodus and the 

dactylus 0.7 times as long as the propodus; his 

account of the third pereiopod indicates that the 

merus is slightly longer than the propodus and, 

as on the second pereiopod, the dactylus is 0.7 

times as long as the propodus. Finally, the strong 

spines near the opening of the antennal gland 

mentioned by Forest in the Pillsbury specimens 

are not shown in Monod's original illustration of 

the species. We have not had an opportunity to 

reexamine the two Pillsbury specimens. 

BIOLOGY.—Almost nothing is known of the 

biology of this species, represented by three specimens, 

one taken in 40 m off Senegal, one in 

70 m on broken shell off Liberia, and one from 

48 m on fine sand and green mud off Ghana. The 

only ovigerous female so far known was collected 

in March. 

DISTRIBUTION.—Known from the following 

three localities off West Africa. 


Achaeus trifalcatus Forest and Guinot, 1966 

Achaeus sp.—Monod, 1956:547, figs. 777-781 [Annobon]. 

Achaeus trifalcatus Forest and Guinot, 1966:110, figs. 15-17, 

[Principe, Sao Tome, Annobon]. 

DISTRIBUTION.—Known only from the offshore 

islands of the Gulf of Guinea, Principe, Sao Tome, 

and Annobon, in depths from 0-6 to 37 m.. 

* Achaeus turbator, new species 

FIGURE 67 


Sta 248, 33 m, 96* (includes holotype), 2$ ov (L, W). 

DESCRIPTION.—Carapace (Figure 67a,c) elongate 

triangular, width 0.64-0.72 (mean 0.68) 

times length, strongly narrowed anteriorly, 

slightly constricted behind orbits, neck appearing 

elongate, branchial regions swollen, regions well 

defined. Surface appearing smooth, not markedly 

granular. 

Rostrum (Figure 67a,c) consisting of 2 very 

short, slender spinules with distinct posterior or 

basal shoulder, arising from truncate lobe or from 

2 lobes separated by very shallow median emar- 

Senegal: S of lie de la Madeleine, near Dakar, 40 m gination. Rostral spines directed anteriorly, not 

(Monod, 1956). 

upturned in lateral view (Figure 676). 

Liberia: 04°23'N, 08°05.5'W to 04°24'N, 08°07.5'W, Supraorbital eave (Figure 67a,b) smooth or 

70 m (Forest, 1968). 

lightly tuberculate, not armed with spinules. 

Ghana: 05°35'N, 00°10'E to 05°36'N, 00°11.5'E, 48 m 

Postorbital margin smooth, unarmed. Eyestalks 

(not 42 m)(Forest, 1968). 

stout, width about half length, expanded distally, 

anterior margin sinuous, lacking well-marked 

Achaeus monodi (Capart, 1951) projection. Narrow process on anterior margin of 

Podochela monodi Capart, 1951:95, fig. 31, pi. 1: figs. 8, 11, pi. stalk extending to rounded tubercle over cornea. 

2: figs. 17, 18, 20 [Cabinda, Zaire, Angola]. 

Latter large, oval, obliquely terminal. 

Achaeus monodi—Monod, 1956:548, figs. 782-810 [Senegal, Hepatic region moderately expanded, pro- 

Guinea, Sierra Leone, Gabon].—Longhurst, 1958:89 duced into acute or subacute unarmed lobe, an- 

[Sierra Leone].—Rossignol, 1962:122 [Congo].—Forest terior margin almost perpendicular to body line, 

and Guinot, 1966:109 [Guinea, Ivory Coast, Nigeria, Principe, 

Sao Tome].—Le LoeufT and Intes, 1968, table 1 occasionally with lateral tubercle but lacking ap- 

[Ivory Coast]. 

ical spine. Width at hepatic lobes subequal to or 

less than distance from gastric protuberance to 

DISTRIBUTION.—West Africa, from Senegal to anterior margin. 

Angola, including the offshore islands of the Gulf Dorsal surface of carapace with 2 prominences 

of Guinea, Principe, and Sao Tome; sublittoral, on midline, gastric prominence an inflated tuber- 

0-4 to 100 m. 

cle, smaller and lower than cardiac prominence,


a ZL1 

FIGURE 67.—Achaeus turbator, new species. Ovigerous female paratype, cl 6.0 mm, Pillsbuty Sta 

248: a, carapace, dorsal view; b, carapace, lateral view. Male paratype, cl 6.6 mm, Pillsbury Sta 

248: c, carapace, dorsal view; d, epistome; e, chela; /, dactylus of fifth pereiopod; g, abdomen; 

h, gonopod. 

neither spined dorsally. Protogastric region with 

low, unarmed tubercle. Branchial region smooth, 

with low mesobranchial tubercle centered dorsally. 

Epibranchial swelling, if present, low, inconspicuous. 

Low swelling present on metabranchial 

region above posterior pereiopods. Posterior 

and posterolateral margins unarmed, tubercles 

present anteriorly on margin above chelipeds. 

Basal antennal article (Figure 67d) usually 

smooth, lightly tuberculate or with 1 or 2 low 

spinules in some specimens. Antennal segments 

unarmed. 

Antennular fossae large, longitudinally subovate, 

lateral margins of fossae irregularly crenulate 

along border with basal antennal segment. Basal 

antennular segment bearing irregular line of tubercles 

ventrally. Interantennular process slender, 

terminating posteriorly in thin, blunt, obtuse, 

triangular lobe. Anterior process of epistome very 

slender. 

Epistome (Figure 61d) appearing elongate, actually 

broader than long, widening posteriorly, 

most of surface smooth, with single sharp tubercle 

present anterolateral to opening of antennal

274 

gland. Pterygostomian region with sharp tubercle 

on lateral border. 


hairy, not heavily spined. Ischium with 2 parallel 

lines of small tubercles, mesial margin tuberculate, 

erect spines present on margin in some specimens. 

Merus largely smooth, with few surface 

tubercles, narrow, unarmed mesially or with tubercles 

or small spines. Palp with slender dactylus, 

shorter than stout carpus and propodus combined; 

carpus and propodus each with distal spine 

mesially. 

Chelipeds (Figure 67


ETYMOLOGY.—The name is from the Latin, 

turbator, meaning disturber or trouble-maker. 


off Nigeria. 

Genus Calypsachaeus, new genus 

TYPE-SPECIES.—Achaeus calypso Forest and 

Guinot, 1966. 

ETYMOLOGY.—The name is derived from the 

name of the vessel Calypso in combination with 

the generic name Achaeus, gender of the name is 

masculine. 

DIAGNOSIS.—Carapace subtriangular, longer 

than broad, narrowed behind orbit, largely 

smooth dorsally, margins spinulous. Rostrum of 

2 elongate, anteriorly truncated lobes, apex and 

margins spinulous. Orbit with only supraorbital 

eave above, lined with long spinules. Postorbital 

spine absent. Eyestalks long, nonretractile, cornea 

obliquely subterminal, slightly ventral, large, 

ovoid. Interantennular septum produced ventrally 

into cupped process with distal sharp spine, 

visible in dorsal view. Basal antennal article 

slender, channelled longitudinally, depression 

flanked on each side by line of tubercles, with 

strong distolateral spine. Merus of third maxilliped 

ovate, not notched distally, narrower than 

ischium, palp articulating at summit. Chelipeds 

spinous, slightly enlarged in adult male, palm 

strongly compressed, cristate dorsally and ventrally. 

Walking legs long and slender, dactyli of 

posterior 2 falciform, ventrally spinulated. Abdomen 

of 6 free somites in both sexes, male 

abdomen widest at third somite. Male first pleopod 

bluntly pointed, aperture terminal in groove. 

REMARKS.—Calypsachaeus superficially resembles 

Achaeus, differing in the form of the rostrum, 

which is much better developed, in the strong 

development of the interantennular septum, in 

the ornamentation of the basal antennal segment, 

and in the form of the chelae, on which the palms 

are strongly compressed and cristate dorsally and 

ventrally. The male pleopod of Calypsachaeus (Forest 

and Guinot, 1966, fig. \9c-d) is similar to that 

of Achaeus in overall shape, but opens terminally 

rather than subterminally. Achaeus calypso is the 

only species we assign to this genus. 

* Calypsachaeus calypso (Forest and Guinot, 

1966), new combination 

FIGURE 68 

Achaeus calypso Forest and Guinot, 1966:113, figs. 18, 19. 


Sta 68, 70 m, broken shell, 2

276 

orbital eave, completely lined with denticles or 

strong spinules. Distinct postorbital spine absent. 

Eyestalks slender, elongate, anterior or anteroventral 

margin with prominent, thin, blade-like, obtuse 

projection, as wide as or wider than stalk, 

with anterior spinules. Narrow process from anterior 

margin extending to rounded, setiferous 

tubercle over cornea. Latter broad, oval, 

obliquely terminal. 

Hepatic region well defined, strongly projecting 

laterally, smooth or with apical spine or with 

apical row of spinules flanked anteriorly by spinular 

crest on hepatic lobe. Carapace width at 

hepatic lobes subequal to or greater than distance 

from gastric prominence to anterior margin. 

Dorsal surface of carapace with 2 prominences 

in midline, gastric slender, usually an erect spine, 

often obsolete or broken, much the higher, cardiac 

prominence a low, obtusely rounded, conical lobe. 

Branchial regions smooth, unarmed, subregions 

not well marked. Lateral and posterior margins 

of carapace completely lined with tubercles or 

erect spinules. 

Basal antennal article longitudinally sulcate, 

with inner and outer lines of spinules, fewer in 

smaller specimens, outer line terminating in enlarged 

distolateral tooth. Free segments of antennal 

peduncle variably tuberculate or spinulose. 

Antennular fossae elongate-triangular, margins 

unarmed, lacking anterior rim, each fossa grading 

anteriorly into rostral spine. Basal antennular 

segment with patch or line of spiniform tubercles. 

Interantennular process strongly produced (Figure 

68/>), projecting well beyond level of basal 

antennal segment, expanded laterally to form 

broad, anteriorly cupped lobe, curved anteriorly, 

apically acute or spiniform. Inner, dorsal margin 

of cupped lobe grading evenly into inner surfaces 

of rostral horns. Anterior process of epistome thin, 

rounded, blade-like, projecting ventrally. 

Epistome subrectangular, broader than long, 

most of surface smooth, with spinulose crest or 

patch of spinules lateral to opening of antennal 

gland. Pterygostomian region with stalked projection, 

apically spinulose, on lateral border. 



hairy, heavily spined. Exopod lined with spinules. 

Ischium with 2 divergent rows of sharp denticles, 

mesial margin spinulose, tuberculate. Merus with 

line of sharp denticles on surface, spined mesially, 

with spinules laterally and distally. Palp with 

slender dactylus, shorter than stout carpus and 

propodus combined. Carpus and propodus each 

with distal spine mesially. 

Chelipeds slender in both sexes, longer and 

slightly inflated in males, merus extending beyond 

eye in males, not extending to eye in females 

and juveniles. Ischium very spinous, largest spine 

set distoventrally. Merus with line of erect spines 

on sharp ventral border, outer surface smooth, 

upper border cristate, smooth or with line of 

tubercles, inner surface with some lines of tubercles. 

Palm strongly compressed, cristiform dorsally 

and ventrally, upper margin with spaced 

spines and tubercles, lower margin smooth, inner 

surface of lower margin with line of spines and 

denticles, outer surface smooth, inner surface tuberculate. 

Fingers longer than palm, flattened, 

lacking prominent gape, cutting edges crenulate, 

lacking enlarged proximal tooth on movable finger. 

Chelae largely naked, with few hooked hairs 

on outer surface, upper and lower borders with 

few long, stiff hairs interspersed between spines. 

Walking legs long and slender (merus of second 

pereiopod longer than carapace, propodus subequal 

to or slightly longer than carapace), with 

curled hairs arising in groups on dorsal surface of 

distal 4 segments and longer hairs scattered over 

surface, especially on propodus and dactylus. Second 

pereiopod longest, remainder decreasing in 

length posteriorly. Merus of walking legs with 

distal arc of spinules or low tubercles dorsally. 

Dactylus of second pereiopod straight proximally, 

curved distally, unarmed ventrally, more than 

half as long as propodus, latter shorter than 

merus. Dactylus of third pereiopod as long as that 

of second, slightly more curved distally, unarmed 

ventrally, more than half as long as propodus, 

latter subequal to merus. Dactylus of fourth pereiopod 

falciform, ventral margin with numerous 

triangular teeth, some swollen, proximally tuberculate, 

length across arc subequal to that of


propodus, latter shorter than merus. Dactylus of 

fifth leg falciform, armature similar to that of 

fourth, length across arc subequal to that of 

propodus, latter shorter than merus. 

Male abdomen narrowed distally, apex 

broadly rounded, with median tubercle on each 

somite, sixth somite with 2 subapical tubercles or 

spinules. Male pleopod as figured by Forest and 

Guinot (1966, fig. 19c,d). 

Sternum granular in both sexes, in male ornamented 

with anteriorly convex arc of strong 

spines and tubercles, with 1 enlarged spine near 

base of each cheliped. 


5.7 to 9.0 mm, of females 4.8 to 9.3 mm, of 

ovigerous females 8.6 to 9.3 mm. The male holotype 

has a carapace length of 10.0 mm (Forest 


REMARKS.—This striking species can readily be 

recognized by its relatively smooth carapace, eyes 

with their flattened anterior projection, broad 

rostral spines, well-developed interantennular 

spine, flattened chelae, and the spinous ventral 

surface. 

BIOLOGY.—This species occurs on the shelf in 

the Gulf of Guinea in depths between 30 and 70 

m. The Pillsbury specimens were taken in 48 m on 

fine sand and green mud, in 51 m on rough 

bottom, and in 70 m on broken shell. The holotype 

was taken by the Calypso in 32 m on mud 

with Area, and Forest and Guinot (1966) also 

recorded material taken on mud in 30, 50, and 

70 m. Ovigerous females were collected in May. 

DISTRIBUTION.—Gulf of Guinea, where it has 

been recorded from localities between Liberia 

and Cameroon in depths between 30 and 70 m. 

Records in the literature, all from Forest and 

Guinot (1966), include the following: 

Ivory Coast: No specific locality, 30, 50, and 70 m. 


06°12'E, 32 m. 

It has not been recorded previously from Liberia, 

Ghana, Dahomey, or Cameroon. 

Genus Capartiella, new genus 

TYPE-SPECIES.—Achaeus longipes Capart, 1951. 

ETYMOLOGY.—We take great pleasure in dedi- 

cating this genus to A. Capart. His pioneering 

work on West African crabs published in 1951 

added much to our knowledge of the brachyuran 

fauna there. Gender of the generic name is masculine. 

DIAGNOSIS.—Carapace pyriform, length 

slightly greater than width, scarcely narrowed 

behind the orbit as a neck, lacking long spines or 

sharp tubercles. Rostrum a single short rounded 

projection, depressed anteriorly, overreached by 

strong interantennular spine. Orbit consisting 

above of narrow supraorbital eave, postorbital 

spines absent. Eyestalks long, nonretractile, movable. 

Cornea terminal, slightly ventral. Interantennular 

septum produced into distinct spine. 

Basal article of antennule slender, longitudinally 

sulcate, ornamented with low, rounded tubercles 

and distal blunt angular projection. Merus of 

third maxilliped subovate, not notched distally, 

narrower than ischium, palp articulating anterointernally. 

Chelipeds short, slender in both sexes. 

Walking legs long, slender, dactyli elongate, 

arched, with triangular ventral spines. Abdomen 

of 6 somites in both sexes. Male first pleopod with 

blunt apex, aperture terminal. 

REMARKS.—In his original description, based 

on a single non-ovigerous female, Capart (1951: 

63) noted that "e'est avec grande hesitation que 

je place cette curieuse espece dans le genre 

Achaeus...." He further noted that C. longipes 

could be separated from other species of Achaeus 

by its simple rostrum and by the shape of the 

dactyls of the walking legs. Monod (1956), who 

had no additional material, transferred the species 

to Physachaeus Alcock, with reservations, noting 

the superficial resemblance of C. longipes to P. 

ctenurus Alcock. Subsequent authors recording the 

species have followed Monod in placing the species 

in Physachaeus. 

Forest and Guinot (1966:108, 109), in recording 

the first males to be collected, interpreted the 

rostrum as being formed of two very short lobes 

which were overreached ventrally by the interantennular 

septum, i.e., the rostrum comprised 

two fused lobes rather than a single one. They 

also pointed out differences in the eyes and the

278 

basal antennal segment of C. longipes and those of 

species of Physachaeus, based on accounts in the 

literature, and noted (p. 109) "il est possible que 

Ton soit amene par la suite a creer pour elle un 

nouveau genre." 

We have been able to compare our material 

with a male and female Physachaeus ctenurus (13°- 

17'15"N, 93°10'25"E, 185 fm (339 m), Investigator, 

USNM 42757) and from this comparison conclude 

that the resemblance between C. longipes 

and P. ctenurus is superficial. In Physachaeus the 

rostrum is clearly bifid, the eyes are short and 

fixed, with a terminal cornea, there are large 

gastric and cardiac spines on the carapace, the 

basal article of the antenna bears a long spine, 

and there are but 5 somites in the abdomen of 

the female. The condition of the eyes in C. longipes, 

long and movable, versus those of P. ctenurus, 

would be enough to exclude C. longipes from 

Physachaeus. 

Capartiella most closely resembles Achaeus 

Leach, Achaeopsis Stimpson, Dorhynchus Studer, 

and the American Podochela Stimpson (definition 

of the latter genus in Rathbun, 1925). It resembles 

Achaeus, Achaeopsis, and Dorhynchus and differs 

from Podochela in having the abdomens of each 

sex composed of 6 somites. In Podochela, which 

also has an unarmed carapace and usually has a 

simple but anteriorly produced rostrum, the abdomen 

is composed of 6 somites in the male, 5 in 

the female. The legs of Capartiella, like those of 

Achaeopsis, are very long, slender, setose, with the 

dactyl slender, not strongly recurved, but Capartiella 

lacks the spines on the carapace. Capartiella 

shares many features with Achaeus, but differs in 

having a single rostrum, slender chelipeds that 

are not inflated or enlarged in adult males, and 

slender dactyli on the walking legs. The male 

pleopod of Capartiella longipes (figured by Forest 

and Guinot, 1966, fig. 14), resembles that of some 

species of Achaeus figured by Griffin (1970, figs. 

13-15) in general configuration, and differs 

strongly from the pleopods of Podochela, which are 

apically acute with a subterminal opening 

flanked by a rounded flap, as shown by Garth 

(1958, pi. H). 


* Capartiella longipes (Capart, 1951), 



Achaeus ? longipes Capart, 1951:62, fig. 19, pi. 1: fig. 1, pl. 2: 

figs. 21,22. 

Physachaeus (?) longipes.—Monod, 1956:537, fig. 746.—Forest, 

1959:15.—Rossignol, 1962:122.—Forest and Guinot, 

1966:108, fig. 14.—Crosnier, 1967:340. 


Sta 68, 70 m, broken shell, 29 (L). 

Ivory Coast: Sta 42,62-75 m, mud with brown, branched 

Foraminifera, 16\ 19 (L, W). Sta 46, 38-42 m, mud with 

dense Julltenella, 1$ ov (L). Sta 60, 79-82 m, coral or rock, 26 

(W). 

Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 16*, 

29 ov (L). Sta 23, 42 m, foliate brown to orange bryozoans, 

39 ov (L, W). Sta 24, 35-37 m, dark red bryozoans, 19 (W). 

Sta 28, 49-53 m, 16\ 19 ov (L). 

Nigeria: Sta 239, 73 m, 19 (W). Sta 240, 37 m, 16* (L). 

Sta 241, 59-63 m, mud and shell, 19 (L). Sta 248, 33 m, ld\ 

19 (W). 

Other Material: Congo: Off Pointe-Noire, 04°48'S, 11°- 

39'E, 54-56 m, sandy mud, 23 Sep 1965, Ombango, A. Crosnier, 

26, 49 (1 ov) (W). 

. Cabinda: SW of Landana, 50 m, 23 May 1959, Ombango, 

A. Crosnier, 19 ov (W). 


Male Pleopod: Forest and Guinot, 1966, fig. 14 

(Ghana). 


6.0 to 11.1 mm, of non-ovigerous females 4.0 to 

13.9 mm, of ovigerous females 6.0 to 13.0 mm. 

REMARKS.—We have provided here sketches of 

C. longipes, as well as Capart's original figures 

(Figures 69, 70) of it, apparently one of the more 

common species on the shallow part of the shelf 

in the Gulf of Guinea. All of our specimens were 

completely covered with a dense coat of algae 

and mud, so dense that their shape was scarcely 

discernible. This may have caused the species to 

be overlooked in earlier collections, for the shelf 

from Ghana to Nigeria has been well sampled in 

the last 25 years. 

BIOLOGY.—This species apparently lives on soft 

bottoms in depths of no more than 82 m. The 

Pillsbury specimens were taken in depths of 33 to 

82 m, usually on mud bottom with foraminifera


FIGURE 69.—Capartiella longipes (Capart) (from Capart, 1951, fig. 19.) 

or Bryozoa; one sample was taken on broken shell 

and one on coral or rock (which terminated the 

haul; a rock outcrop may have been encountered 

on an otherwise soft bottom). Capart (1951) reported 

the species from a depth of 60 m in green 

mud. The specimens reported by Forest and 

Guinot (1966) came from mud, sand, and compact 

sand (sable construit) in 67-75 m and from 

mud in 50 m. Crosnier (1967) found the species 

on mud and sandy mud in depths between 35 

and 58 m. 


May, June, and September. 

DISTRIBUTION.—Tropical West Africa, where it 

has been taken at a few localities between Senegal 

and Angola; sublittoral, in depths between 33 

and 82 m. Records include the following: 

Senegal: 12°55.5'N, 17°33'W, 65-75 m (Forest, 1959; 


Ivory Coast: No specific locality (Forest and Guinot, 

1966). 

Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m 


Dahomey: No specific locality, 35-38 m (Crosnier, 1967). 

Cabinda: W of Landana, 50 m (Rossignol, 1962). 

Angola: 10 mi NxW of Luanda, 08°37'S, 13°12'E, 60 m 

(Capart, 1951). 

Genus Dorhynchus Thomson, 1873 

Dorhynchus Thomson, 1873:175 [type-species: Dorhynchus thomsoni 

Thomson, 1873, by monotypy; gender: masculine; 


Lispognathus A. Milne Edwards, 1881, in 1873-1881:349 

[type-species: Lispognathus furcillatus A. Milne Edwards, 

1881, by monotypy; gender: masculine]. 

DIAGNOSIS.—Carapace triangular, pyriform, 

longer than broad, expanded behind orbit, with 

dorsal spines, margins spinulous. Rostrum of 2

280 

a 


FIGURE 70.—Capartiella longipes (Capart). Male, cl 10.0 mm, Pillsbury Sta 248: a, front, dorsal 

view; b, front, oblique anterior view; c, front, ventral view; d, abdomen. Female, cl 12.0 mm, 

Pillsbury Sta 248: e, third maxilliped. Male, cl 11.1 mm, Congo:/, propodus and dactylus of 

fifth pereiopod. 

slender, sharp spines usually divergent anteriorly. 

Orbits with supraorbital eave provided with supraorbital 

spine(s), preorbital spine absent. 

Postorbital spine absent, nuchal spine present 

between hepatic lobe and orbit. Eyestalks short, 

movable but nonretractile, cornea terminal, large, 

subglobular. Interantennular septum produced 

into sharp spine, usually visible in dorsal view. 

Basal antennal article slender, channelled longitudinally, 

channel flanked by inner and outer 

rows of spines, with enlarged terminal spine distolaterally. 

Merus of third maxilliped spinous, 

ovate or subrectangular, not notched distally, 

narrower than ischium, palp articulating at summit. 

Chelipeds spinous, slenderer in female, palm 

inflated. Walking legs long and slender, dactyli 

elongate, not falciform, posterior 2 with low teeth 

ventrally. Abdomen of 6 somites in both sexes. 

Male first pleopod bluntly pointed, with distinct 

subterminal lappet. 

REMARKS.—Although the Pillsbury collections 

included no representatives of this genus, our 

recognition herein of some new genera in this 

subfamily necessitated examination of representatives 

of all of the genera of Inachinae in the 

eastern Atlantic. In the case of Dorhynchus, considered 

by some (Rathbun, 1925; Barnard, 1950; 

Griffin, 1966b) but not others (Bouvier, 1940; 

Monod, 1956; Zariquiey Alvarez, 1968; and 

Christiansen, 1969) to be synonymous with 

Achaeopsis Stimpson (1858d:218; type-species: A. 

spinulosa Stimpson, 1858, by monotypy; gender: 

feminine; name 1604 on Official List), comparison 

of material of D. thomsoni (off Morocco, 1105 m, 

Talisman, USNM 22974, 3d, cl 7.3-7.5 mm, 3$, cl 

5.8-7.5 mm) with a specimen of A. spinulosa 

(South Africa, 36°40'S, 21°26'E, 200 m, USNM: 

Id, cl 12 mm) leads us to suggest that the two 

should be regarded as distinct but closely related 

genera. 

Dorhynchus differs from Achaeopsis in several features: 

the rostral spines are much longer; the 

interantennular spine is much larger, projects 

further ventrally, and its apex usually is visible 

between the rostral spines in dorsal view; the 

walking legs are longer and slenderer: in D. thorn-


soni the merus of the second pereiopod is longer 

than the carapace whereas in A. spinulosa it is 

shorter than the carapace; and the dactyli of the 

pereiopods are slender and similar on all legs in 

D. thomsoni, whereas they are falciform in the last 

two legs of A. spinulosa. Also, as Barnard (1950: 

25) pointed out, A. spinulosa generally occurs in 

shallower water than does D. thomsoni. 

If this interpretation is correct and Achaeopsis 

and Dorhynchus are retained as separate genera, 

then two other species assigned to Achaeopsis sensu 

lato by Griffin (1966b), Achaeopsis rostrata Sakai, 

1932, from Japan and Stenorhynchus ramusculus 

Baker, 1906, from South Australia and New Zealand, 

both of which have slender legs, should be 

transferred to Dorhynchus, and Achaeopsis should 

be restricted to include only the type-species, A. 

spinulosa. 

Although Dorhynchus thomsoni is now considered 

to be widely distributed ["virtually cosmopolitan," 

Griffin (1966b:35)], we agree with Bouvier 

(1940:351), Monod (1956:523), and Guinot 

(1967a:289, footnote), all of whom question 

whether the populations from different areas are 

conspecific. In the collection of the Smithsonian 

Institution, for example, there is one lot from 

Chiloe Island, off Chile (USNM 156225), in 

which the rostrum is spinulose ventrally, rather 

than smooth, and in which there are three distal 

spines on the meri of the pereiopods, rather than 

one as shown for D. thomsoni by Christiansen 

(1969:106, fig. 44). Material of Dorhynchus thomsoni 

should be examined from throughout its range to 

determine the relationship of the various populations. 

Dorhynchus thomsoni Thomson, 1873 

Dorynchus thomsoni Thomson, 1873:174, 175, fig. 34. 

Dorynchus Thomsoni.—Filhol, 1885a: 56. 

Lispognathus Thompsoni.—Filhol, 1885a, fig. 1. 

Dorhynchus thomsoni.—Monod, 1956:522 [references].—Zariquiey 

Alvarez, 1968:467 [Spain; references].—Christiansen, 

1969:106, fig. 44, map 37 [North Atlantic]. 

DISTRIBUTION.—Eastern Atlantic, W and S of 

Iceland and the Faroes southward to the Cape 

Verde Islands; deep water species, in depths between 

100 and 2100 m. The occurrence of this 

species outside of the eastern Atlantic requires 

verification. 

Genus Ergasticus Studer, 1883 

Ergasticus Studer, 1883:7 [type-species: Ergasticus clouei Studer, 



Ergasticus clouei Studer, 1883 

Ergasticus Clouei Studer, 1882:335 [nomen nudum]; 1883:8. 

Ergasticus clouei.—Monod, 1956:523.—Peres, 1964:20.—Zariquiey 

Alvarez, 1968:463, fig. 155b [Spain; references].— 

Tiirkay, 1976a:25 [listed], 39, fig. 32 [Portugal, in part]. 




151, 1$ (L). 

Cape Verde Islands: 16°52'N, 27°30'W of Paris (25° 10'W 

of Greenwich), 400-580 m, sand and rock, 29 Jul 1883, 

Talisman Sta 111, 16", 1? ov (W). 


1900:140. 

Figure: A. Milne Edwards and Bouvier, 1900, 

pi. 21: figs. 1-7. 

MEASUREMENTS.—Carapace length of nonovigerous 

females 8-10 mm, of ovigerous female 

8 mm. 

BIOLOGY.—Ergasticus clouei is a deepwater species, 

occurring in depths between 70 m and 

1241 m; more than 90% of the depth records in 

the literature are from depths between 300 and 

750 m. 

Ovigerous females have been recorded in July 

and August (A. Milne Edwards and Bouvier, 

1899, 1900; Bouvier, 1922). 

DISTRIBUTION.—Eastern Atlantic from the Bay 

of Biscay to the Cape Verde Islands, including 

Madeira and the Azores; Mediterranean. Deepwater, 

in depths between 70 and 1241 m. Monod 

(1956) reported no specimens. Other West African 

records include the following: 

Morocco: Bane de Spartel, 35°54'N, 06°14'W, 210 m

282 

(Peres, 1964). 33°12.6'N, 09°15.2'W, 500 m, and 33°10.5'N, 

19°17.5'W, 170-345 m (Tiirkay, 1976a). 

Cape Verde Islands: 15°40'N, 23°06'W, 38 fm (70 m) 

(Studer, 1882). 

Genus Inachus Weber, 1795 

Inachus Weber, 1795:93 [type-species: Cancer scorpio Fabricius, 

1779, a subjective junior synonym of Cancer dorsettensis 

Pennant, 1777, by subsequent designation by H. Milne 

Edwards, 1840, in 1836-1844, pi. 34: fig. 2; gender: masculine; 


Macropus Latreille, 1803a:27 [an invalid junior homonym of 

Macropus Shaw, 1790; type-species: Cancer phalangium Fabricius, 

1775, by monotypy; gender: masculine; name 

1777 on Official Index]. 

Leptopodia Leach, 1814:431 [type-species: Cancer phalangium 

Fabricius, 1775, by monotypy; gender: feminine]. 

Pseudocollodes Rathbun, 1911:247 [type-species: Pseudocollodes 

complectens Rathbun, 1911, by monotypy; gender: masculine]. 

REMARKS.—Griffin (1974:18) correctly synonymized 

Pseudocollodes Rathbun, 1911 with Inachus 

Weber, 1795, but, as pointed out later (p. 

290), we suspect that Inachus complectens (Rathbun), 

known from the western Indian Ocean and 

South Africa (Barnard, 1950; Griffin, 1974) is not 

conspecific with /. dorsettensis (Pennant). 

As is the case with most of the West African 

spider crabs, the species of Inachus appear to 

require much further study. Four species are represented 

in the Pillsbury collections and three of 

these are undescribed; two of the species previously 

were identified with European species. 

Monod (1956) recorded seven species from 

West Africa: Inachus angolensis Capart (the only 

endemic species to be listed), /. dorsettensis (Pennant), 

/. guentheri (Miers), /. phalangium (Fabricius), 

/. thoracicus (Roux), /. aguiarii De Brito 

Capello, and /. leptochirus Leach. Monod's accounts 

of /. dorsettensis and /. leptochirus, both 

relatively well-known European species, appear 

to be based on material of the two new species 

described herein. Monod's account of /. guentheri 


(Miers), which unfortunately is not represented 

in the Pillsbury collections, does not agree with the 

account of that species in Barnard (1950:27): the 

rostral teeth and the spination of the carapace 

appear to be quite different in the Gulf of Guinea 

and South African populations now referred to 

this species. Our studies of the Pillsbury collections 

have suggested that relatively few West African 

species identified by earlier workers with European 

or Mediterranean species actually are conspecific 

with those species. We suspect, therefore, 

that available West African material referred to 

/. phalangium, I. thoracicus, and /. aguiarii, should 

be carefully reexamined. 

The identity of the dwarf forms of Inachus 

dorsettensis reported by Doflein (1904) and Odhner 

(1923) from deep water off West Africa cannot 

be determined with certainty without examining 

their material; they could have been dealing with 

the new species /. grallator or /. nanus, both of 

which occur in relatively deep water and are 

smaller than European Atlantic specimens of /. 

dorsettensis sensu stricto. We have tentatively referred 

them to /. grallator. 

The identity of the juvenile female from Angola 

reported by Odhner (1923:19) as Inachus sp. 

also cannot be determined with certainty on the 

basis of his account. He may have been dealing 

with /. biceps, new species, or with /. guentheri sensu 

lato. 

The following key distinguishes the West African 

species of Inachus. The key would be complete 

for the eastern Atlantic by adding the Mediterranean 

/. communissimus (Rizza), which would key 

out with /. nanus, new species, /. dorsettensis, which 

would key out with /. angolensis and /. grallator, 

new species, and /. leptochirus, which would key 

out with /. biceps, new species. We decided against 

adding these species to the key because of the 

uncertain status of the Mediterranean population 

of /. dorsettensis (see Inachus grallator, new species, 

p. 287). 

Key to Tropical West African Species of Inachus 

1. Gastric areas of carapace with transverse row of 4 tubercles anterior to 

gastric spine 2 

Gastric area of carapace with at most 2 tubercles anterior to gastric 

spine 4


2. Interantennular spine short, not visible between rostral spines in dorsal 

view /. nanus, new species 

Interantennular spine large, well developed, visible between rostral spines 

in dorsal view [often overreaching rostral spines] 3 

3. Cardiac and branchial regions of carapace with erect spines 

/. grallator, new species 

Cardiac and branchial regions of carapace lacking erect spines 

/. angolensis 

4. Rostral spines appressed for most of their length [white sternal callosity 

absent in male] /. phalangium 

Rostral spines separate for all of their length 5 

5. Branchial regions of carapace each with erect dorsal spine 6 

Branchial regions of carapace each with obtuse swelling or rounded tubercle 

dorsally, lacking erect dorsal spine 7 

6. Male lacking white sternal callosity /. guentheri 

Male with prominent, simple sternal callosity /. biceps, new species 

7. Walking legs elongate, first (= second pereiopod) more than 4 times as 

long as carapace in males. Sternal callosity in males only .. /. thoracicus 

Walking legs short, first (= second pereiopod) less than 4 times as long as 

carapace in males. Sternal callosity present in both sexes .... /. aguiarii 

Inachus aguiarii De Brito Capello, 1876 

Inachus aguiari.—Monod, 1956:533, figs. 736-739 [Guinea]. 

—Forest and Guinot, 1966:107 [Spanish Sahara]. 

Inachus aguiarii.—Zariquiey Alvarez, 1968:473, fig. 158c,d 

[Spain; references]. 


Madeira, and Guinea; Mediterranean. Sublittoral, 


* Inachus angolensis Capart, 1951 

Inachus angolensis Capart, 1951:72, fig. 22, pi. 1: fig. 7, pi. 2: 

fig. 10.—Monod, 1956:524, figs. 711-714.—Rossignol, 

1957:77, 116 [key], pi. 1: fig. 1.—Gauld, 1960:72.— Rossignol, 

1962:122.—Crosnier, 1964:34.—Forest, 1965b:394 

[discussion].—Forest and Guinot, 1966:106.—Le Loeuff 

and Intes, 1968:31, 46, table 1, figs. 51, 63.—Maurin, 

1968a:62; 1968b:486, 489, 491.—Le Loeuff and Intes, 

1969:66.—Crosnier, 1970:1215 [listed], 1218. 

Inachus.—Maurin, 1968b, figs. 4, 9. 

?Inachus mauritanicus.—Maurin, 1968b:484 [not Inachus mauritanicus 

Lucas, 1846 = /. communissimus Rizza, 1839]. 


Coast: Sta 42, 62-75 m, mud with brown, branched Foraminifera, 

2(5, 2$ (L). Sta 49, 73-77 m, 1$ (L). Sta 50, 128- 

192 m, 19 (L). Sta 62, 46 m, brown, branched and foliate 

Foraminifera, 5(5, 1? (L). Sta 63, 64 m, sandy mud with 

shells, 3c5, 1$ ov (L). Sta 64, 68 m, 1(5 (L). 

Ghana: Sta 16, 46 m, mud with Foraminifera, shells, 19 

(L). Sta 22, 51 m, rough bottom, 19 (W). Sta 28, 49-53 m, 

2(5, 49 (L). 

Nigeria: Sta 239,73 m, 2(5 (L). Sta 241, 59-63 m, mud 

and shell, 79 (6 ov) (L). 

Cameroon: Sta 259, 59 m, mud and broken shell, 26(5, 219 

(3 ov) (W). 

Undaunted Material: Angola: Sta 94, 90 m, 1(5, 19 (L). Sta 

96, 162 m, 1(5, 19 (L). Sta 103, 90 m, 19 (L). 

Other Material: Congo: Off Pointe-Noire, 150 m, slightly 

sandy mud, 27 Feb 1967, A. Crosnier, 1(5 (W). 



Monod, 1956, figs. 711-714. 




lengths of 7 to 23 mm; the ovigerous females 

have carapace lengths of 12 to 16 mm. The largest

284 

specimen recorded by Capart (1951) was 23 mm 

long. 

REMARKS.—Inachus angolensis, one of five Atlantic 

species now known to have a transverse line of 

four tubercles on the gastric region of the carapace, 

is one of three of those species in which the 

interantennular spine is strongly developed, being 

clearly visible in dorsal view. It is the only one in 

this group of species lacking erect spines on the 

cardiac and branchial regions. 

In /. angolensis the branchial margin of the 

carapace is continuous with the free epineral 

margin as described by Forest (1965a) for /. dorsettensis 

(Pennant). The dactylus of the chela is 

provided with 1 or 2 large basal teeth. The second 

pereiopod may be as much as 5 times as long as 

the carapace in adult males, and the merus of 

that leg may be up to 1.5 times as long as the 

carapace. The dactylus of the fifth pereiopod 

lacks subapical spinules as in the Mediterranean 

/. communissimus Rizza (Forest, 1965a:393). 

Monod (1956) suggested that /. angolensis might 

be a southern form of /. mauritanicus Lucas [a 

junior synonym of/, communissimus Rizza, according 

to Forest (1965a) who examined the types of 

/. mauritanicus], but /. angolensis differs in numerous 

features and, as pointed out by Forest (1965a) 

and Forest and Guinot (1966), can be distinguished 

by the long interantennular spine alone. 

BIOLOGY.—This species apparently prefers soft 

bottoms in moderate depths on the shelf and 

upper slope. Most of the specimens collected by 

the Pillsbury were taken at depths between 46 and 

77 m, but one sample came from 128-192 m; the 

samples were taken on mud with Foraminifera 

and shells and brown, branching and foliate Foraminifera 

in 46 m, rough bottom in 51 m, mud 

with brown, branched Foraminifera in 62-75 m, 

mud and broken shell in 59 m, and sandy mud 

with shells in 64 m. The specimen recorded here 

from Pointe-Noire was taken in slightly sandy 

mud in 150 m. Capart (1951) recorded material 

on green mud in 60-110 m, on black, brown mud 

in 100 m, on sand and mud in 240 m, and on 


green sandy mud in 250-300 m. Monod (1956) 

did not specify bottom type, but recorded material 


Crosnier (1964) characterized /. angolensis as a 

cold water species, living at depths greater than 

50 m. Maurin (1968b) found this species off 

Spanish Sahara on muddy detritic bottom in 50- 

90 m and off Mauritania on mud or sandy mud 

in 90-100 m and on fine detritic sand, occasionally 

muddy, in 200-400 m. Le Loeuff and Intes 

(1968:46) note that "c'est une espece d'eaux 

froides (19°5 a 15°2) et salees (35,4 %o a 35,72 

%o)"; they found the species in 80 to 200 m. 

Capart (1951) recorded it in 240 m with a bottom 

temperature of 10.85°C. Forest and Guinot 

(1966) found /. angolensis on sand and compact 

sand [sables construit] in 65-75 m and on mud in 

50 m. 

Ovigerous females have been recorded in February, 

March, April, May, June, September, and 

October (Capart, 1951; Monod, 1956; Crosnier, 



Spanish Sahara and southern Angola in 

depths between 46 and at least 350 m. Capart's 

(1951) material came from off the Congo and 

Angola and Monod (1956) reported material 

from Senegal; since 1956 it has been recorded 

from the following localities. 


Blanco, 50-90 m (Maurin, 1968b). 

Mauritania: Bane d'Arguin, 90-150 m (Maurin, 1968a), 

and 40-60, 60-70, and 90-100 m (Maurin, 1968b). Off 

Tamzak (as Tamxat), 200-400 m (Maurin, 1968b). 

Senegal: Saint-Louis, 75-85 m (Maurin, 1968b). 

12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 1966). 

Ivory Coast: No specific locality, 60+ m (Le Loeuff and 

Intes, 1969), 80-200 m (Le Loeuff and Intes, 1968). 

Ghana: Off Accra, 60 m (Gauld, 1960). 04°40'N, 02°- 

08'W to 04°39'N, 02°05'W, 50 m (Forest and Guinot, 1966). 

Cameroon: No specific locality, depths greater than 50 m 

(Crosnier, 1964). 

Congo: Off Pointe-Noire (Rossignol, 1957). W of Pointe- 


Angola: 16°27'S, 11°35'E, 90 m; 16°41'S, 11°21'E, 162 

m; and 17°06'S, 11°35'E, 90 m (Crosnier, 1970).


*Inachus biceps, new species 


Inachus leptochirus.—Monod, 1956:535, 632, figs. 740-745.— 

Longhurst, 1958:89.—Gauld, 1960:72.—Forest and 

Guinot, 1966:107. [Not Inachus leptochirus Leach, 1817.] 


Sta 17, 48 m, fine sand and green mud, 16* (L). Sta 23, 42 m, 

foliate brown to orange bryozoans, 6c5, 39 (2 ov) (L). Sta 24, 

35-37 m, dark red bryozoans, 14(5 (includes holotype), 14$ 

(13 ov) (L, W). Sta 26, 27m, shell bottom (scallops), 16\ 2$ 

(1 ov) (L). 

Nigeria: Sta 248, 33 m, Ic5, 2$ ov (L). 

Other Material: Dahomey: OffGrand-Popo, 30 m, Petersen 

grab, 23 Feb 1964, Guinean Trawling Survey, Tr 34, Sta 2: 

16\l$ov(L). 

DESCRIPTION.—Carapace (Figure l\a,d) 

broader than long. Gastric region with pair of 

tubercles anterior to erect gastric spine (Figure 

72a). Cardiac region with low, rounded or obtuse 

dorsal projection, lower than gastric spine, 

FIGURE 71.—Inachus biceps, new species. Ovigerous female, 

Ghana: a, dorsal view; b, carapace, lateral view; c, front, 

ventral view. Male, Ghana: d, dorsal view; e, first pleopod; 

/, apex of first pleopod; g, sternum. (From Monod, 1956, figs 

740-745.) 

flanked posteriorly on midline by lower prominence. 

Branchial regions each with dorsal tubercle, 

lateral branchial margin tuberculate. Hepatic 

region irregularly tuberculate anteriorly. Rostrum 

very short, teeth obtusely rounded, each 

usually with minute apical tubercle. Eyes large, 

cornea often extending laterally beyond broad 

postocular spine. Interantennular spine a very 

small, obtuse lobe, not visible in dorsal view, 

scarcely or not at all visible in lateral view (Figure 

72a). Posterolateral border of carapace continuous 

with free epimeral margin. Basal antennal 

segment (Figure 12a,b) with row of tubercles, 

some sharp, on mesial and lateral margins, anterolateral 

angle of basal segment lacking prominent 

spine. 

Chelipeds (Figure 12c) enlarged in adult males, 

about 1.5 times as long as carapace, merus and 

propodus markedly inflated in adult. Chelipeds 

in females slender, subequal to or slightly longer 

than carapace. Merus and carpus spiny in both 

sexes. Palm with row of large tubercles proximally 

on outer face in males. Palm depth 1/2 length in 

females, 2/3 length in males. Fingers subequal to 

or slightly longer than palm. Cutting edges of 

fingers unarmed basally in females, dactylus with 

2 prominent proximal teeth in males. 

First walking leg 4 or slightly more than 4 times 

as long as carapace, merus, propodus, and dactylus 

each longer than carapace in males, relatively 

shorter in females. Dactylus unarmed, sinuous 

distally, apex gently curved, length subequal 

to or slightly greater than that of propodus. Second 

to fourth walking legs decreasing in size 

posteriorly, fourth about twice carapace length in 

females, slightly longer in males. Second walking 

leg scarcely overreaching propodus of first by tip 

of dactylus. Latter sinuous, gently curved distally, 

shorter than propodus, with 1 or 2 subdistal teeth 

ventrally. Third walking leg not extending to end 

of propodus of second, overreaching carpus of 

second by distal third of propodus. Dactylus more 

curved than that of preceding leg, shorter than 

propodus, with 2 subdistal teeth ventrally. Fourth 

walking leg rather short, not extending to end of 

propodus of third. Dactylus short, more strongly

286 


FIGURE 72.—Inachus biceps, new species, paratype, male, cl 8.3 mm, Pillsbury Sta 24: a, front, 

lateral view; b, front, ventral view; c, chela; d, abdomen. 

curved than those of preceding legs but not sickleshaped, 

length across arc subequal to that of 

propodus, armed ventrally with 1 or 2 large, 

subdistal teeth and several teeth proximally. 



Inachus biceps can be distinguished readily from 

the other West African species which have two 

gastric tubercles and a sternal callosity in the 

male. The new species differs from /. aguiarii De 

Brito Capello in several features: the interantennular 

spine is smaller; there is a dorsal tubercle 

rather than a spine on the branchial region; and 

the sternal callosity is subcircular in shape. In /. 

aguiarii the sternal callosity is developed in both 

sexes, rather than only in the males as in /. biceps. 

Neither of the other West African species with 

two gastric tubercles, /. phalangium (Fabricius) or 

/. guentheri (Miers), have the sternal callosity developed 

in either sex. 

TYPE-LOCALITY.—Off Ghana, 04°56'N, 00°- 

47.5'W to 04°56'N, 00°50'W, in 35 to 37 m 

(Pillsbury Sta 24). 


D. 31770), a male, cl 9.4 mm, cb 10.3 mm, from 

Pillsbury Sta 24, is in the Rijksmuseum van Natuurlijke 

Historie, Leiden. Paratypes are in the 

collections of the National Museum of Natural 

History, Smithsonian Institution, and the Rijksmuseum 

van Natuurlijke Historie at Leiden. 


English derivative of the Latin musculus biceps, 

alluding to the inflated cheliped. 

BIOLOGY.—Inachus biceps is a sublittoral species, 

occurring in depths between 27 and 300 m. Only 

two of the 16 depth records available are from 

relatively deep water, namely off Sierra Leone in 

220-240 m (Monod, 1956) and 300 m (Longhurst, 

1958); the remainder of the records are from 

depths of less than 60 m. Material from the two 

deepest records should be reexamined. The Pillsbury 

collected the species off Ghana, where it 

appears to be relatively common (28 specimens 

at Sta 24), on fine sand and green mud and on 

bottom with bryozoans or shell. It was taken by 

the Calypso on shells, and mud with Area. 


May, and November (Monod, 1956; Forest 

and Guinot, 1966; present paper). 

DISTRIBUTION—Inachus biceps is a tropical West 

African species, known from a few localities between 

Senegal and Nigeria in depths between 27 

and 300 m. The following records are in the 

literature: 

Senegal: S of Goree, 40-41 and 46-50 m (Monod, 1956). 

Sierra Leone: No specific locality, 220-240 m (Monod, 

1956), 300 m (Longhurst, 1958). 

Ghana: Off Accra, 43, 44, and 51 m (Monod, 1956) and 

44-51 m (Gauld, 1960). 



It has not been recorded previously from Dahomey, 

although that is well within its known 

range. 

Inachus dorsettensis (Pennant, 1777) 

Cancer Dorsettensis Pennant, 1777:8, pi. 9A: fig. 18. 

Inachus dorsettensis.—A. Milne Edwards and Bouvier, 1899:45 

[key], 46 [Azores]; 1900:143 [?part; Ilhas Desertas; Spanish 

Sahara].—Balss, 1922:72 [listed].—Bouvier, 1922:79 [Canary 

Islands].—Monod, 1933b:503 [p. 48 on separate; 

listed].—Capart, 1951:70 [part; Spanish Sahara].— 

Maurin, 1968a:31 [Morocco]; 1968b:489 [Mauritania].— 

Zariquiey Alvarez, 1968:472, figs, 157f, 159b, 160b [Spain; 

references].—Christiansen, 1969:100, fig. 41, map 34 

[North Atlantic ].—Tiirkay, 1976a:26 [listed], 40, fig. 34 

[Portugal, Morocco]. 

SYNONYMS.—ICancer dodecos Linnaeus, 1767; 

Macropus parvirostris Risso, 1816; Doclea fabriciana 

Risso, 1827. 

DISTRIBUTION.—Eastern Atlantic, from the Hebrides 

southward to Mauritania (?), including the 

Azores and the Canary Islands; Mediterranean; 

?South Africa. Sublittoral, from a few meters to 

about 110 m. All of these records require verification. 

* Inachus grallator, new species 


PInachus dorsettensis.—A. Milne Edwards and Bouvier, 1900: 

143 [part?: Cape Verde Islands, 75-90 and 318 m].— 

Doflein, 1904:72 [Bane de la Seine; Spanish Sahara].— 

Odhner, 1923:19 [Angola, 72-108 m].—Monod, 1956:526 

[? part].—Longhurst, 1958:89 [Sierra Leone, 72-118 

m].—Guinot and Ribeiro, 1962:77 [Angola, 150-220 m]. 

Inachus dorsettensis.—Capart, 1951:70, fig. 21, pi. 1: fig. 6, pi. 

2: fig. 11 [part, not specimens from Spanish Sahara?].—

288 

Crosnier, 1970:1218. [Not Inachus dorsettensis (Pennant, 

1777).] 


Sta 255, 264-269 m, 36 (includes holotype), 1$ (L, W). 

Undaunted Material: Angola: Sta 96, 162 m, 1? (L). 

Other Material: Congo: 05°03'S, 11°23'E, 247-250 m, 

sandy mud, 23 Jan 1968, A. Crosnier, 3$ ov (W). 

DESCRIPTION.—Carapace (Figure 73a) distinctly 

longer than broad, length 1.06 to 1.12 

times width. Gastric region of carapace with 4 

tubercles in transverse row anterior to erect gastric 

spine. Cardiac region of carapace with erect 

dorsal spine. Each branchial region with shorter 

erect spine, not markedly recurved anteriorly, 

and with dorsal tubercle anterior to each spine. 

Hepatic lobe with 2 distinct tubercles, and 

smaller, less conspicuous tubercles scattered on 

surface. Rostrum (Figures 73b, 744) short, spines 

broad, lateral margins convergent anteriorly. 

Eyes large, but not extending laterally beyond 

strong postorbital spine. Interantennular spine 

well developed, clearly visible in dorsal view (Figure 

73a,b) overreaching rostral spines in some 

specimens. Branchial margin of carapace tuberculate, 

anteriorly continuous with free epimeral 

margin. Basal antennal segment (Figure 73b) 

with row of tubercles, posteriormost largest, anterolateral 

angle lacking prominent spine. 

Chelipeds subequal to or slightly longer than 

carapace, slender, not markedly inflated (slightly 

inflated in some males), equal and similar in both 

FIGURE 73.—Inachus grallator, new species: a, dorsal view; b, 

ventral view of front; c, male pleopod. (All from Capart, 

1951, fig. 21, pi. 1: fig. 6, pi. 2: fig. 11.) 


sexes; ischium with inner row of 4 or 5 erect 

tubercles; merus rounded, with low, blunt spines 

and tubercles, primarily on ventral surface; carpus 

with scattered small tubercles and spinules; 

palm covered with small, blunt spines; dactylus 

much longer than palm, cutting edge lacking 

enlarged proximal tooth or teeth. 

Second pereiopod (Figure 74c) about 4 (3.8 to 

4.5, mean 4.1) times as long as carapace, very 

slender, merus, propodus, and dactylus each 

longer than carapace; dactylus unarmed, subequal 

to or slightly shorter than propodus. Third 

pereiopod not extending to end of propodus of 

second, overreaching carpus of second by half of 

propodus and all of dactylus; latter shorter than 

propodus, lacking subdistal spines ventrally. 

Fourth pereiopod extending to or slightly overreaching 

propodus of third; dactylus slightly 

curved, with 2 low, subdistal tubercles, subequal 

to propodus. Fifth pereiopod (Figure 74d,e) 

scarcely overreaching propodus of fourth by tip 

of dactylus; latter similar to those of third and 

fourth pereiopods, subequal to propodus, with 2 

subapical tubercles apically. Male pleopod shown 

in Figure 73c. 


14.2 to 15.5 mm, of non-ovigerous females 12.7 to 


REMARKS.—Inachus grallator is the southern 

counterpart of Inachus dorsettensis, which it closely 

resembles. It may be a smaller species than /. 

dorsettensis, Atlantic populations of which may 

attain a carapace length of 30 mm (Christiansen, 

1969), almost twice as large as adult specimens of 

/. grallator available for study. It also appears to 

live in deeper water, occurring off West Africa in 

depths of 162-269 m. Zariquiey Alvarez (1968) 

noted that /. dorsettensis is usually found at depths 

of 30-40 m in Spain, and Christiansen (1969) 

recorded it from 6-10 m to about 110 m, and, 

because Bouvier (1940) noted that it occurred 

from the shore to 550 m, it seems possible that he 

was dealing with more than one species. Material 

of /. dorsettensis from Spain in the collections of 

the Smithsonian Institution comprises more than 

30 lots from the Ria de Arosa, where it is common


FIGURE 74.—Inachus grallator, new species, male paratype, cl 14.2 mm, Pillsbury Sta 255: a, 

carapace, lateral view; b, front, dorsal view; c, second pereiopod; d, fifth pereiopod; e, dactylus 

of fifth pereiopod. Inachus dorsettensis (Pennant), male, cl 15.1 mm, Spain:/, front, dorsal view; 

g, second pereiopod; h, fifth pereiopod; i, dactylus of fifth pereiopod. 

in depths of less than 40 m, although 8 lots were 

taken in depths between 45 and 70 m. 

The best way to distinguish /. grallator from /. 

dorsettensis is by the slenderness of the pereiopods 

and the shape of the dactyli of the fifth pereio- 

pods. In /. grallator, the pereiopods are slenderer, 

appearing longer, and the dactyli of the fifth 

pereiopods are much less curved, being subsimilar 

to those of the fourth pereiopods (compare Figure 

74d,e with 746,i), as well as comparatively longer.

290 


In adult females of /. grallator the dactylus of the Griffin was misled in using the accounts of Monod 

fifth pereiopod is longer than the carapace, (1956) and Barnard (1950) to help identify his 

whereas in /. dorsettensis it is shorter. In females of material with /. dorsettensis. Monod was not deal- 

the new species the pereiopods are slightly stouter ing with /. dorsettensis but with an undescribed 

than in males, but they are noticeably slenderer species, /. nanus, new species (p. 291). Although 

than the same pereiopods of /. dorsettensis. we have seen no material from South Africa, 

Although Forest (1965a:392-393), in differen- Barnard's (1950) account of/, dorsettensis suggests 

tiating the Mediterranean /. communissimus Rizza that he was dealing with /. complectens, a much 

from /. dorsettensis, stated that in the latter species spinier species with sharper rostral teeth, in which 

the carapace is longer than broad (mean length/ the four dorsal spines of the carapace all attain 

width ratio of 1.1 to 1), that the second pereiopod 

was three times as long as the carapace, and that 

the merus of the second leg was equal to the 

carapace, our material from Spain suggests that 

/. dorsettensis may be somewhat variable in these 

features. Our specimens have the carapace 

length/width ratio varying from 0.97 in ovigerous 

females to 1.04 in males; the carapace of adult 

females is noticeably broader than long. In our 

specimens the second pereiopods are 3.29 to 4.09 

(mean 3.65) times as long as the carapace, being 

comparatively longer in males, and, in all of the 

specimens measured, the merus is perceptibly 

longer than the carapace. 

As Forest (I965a:391) noted, Atlantic specimens 

of /. dorsettensis appear to be much larger 

than do Mediterranean specimens; the former 

may attain a carapace length of 30 mm, whereas 

the latter rarely exceed 20 mm in length. Our 

material suggests that in the Mediterranean population 

the dorsal spines of the carapace are much 

slenderer and higher than they are in material 

the same height (in /. dorsettensis, I. grallator, and 

/. nanus, the median spines are higher than the 

branchials). A redescription of /. complectens 

(Rathbun) is in preparation. 

Specimens identified with /. dorsettensis from 

West African localities by A. Milne Edwards and 

Bouvier (1900), Odhner (1923), Longhurst 

(1958), Guinot and Ribeiro (1962), and, possibly, 

part of the material referred to /. dorsettensis by 

Monod (1956) may be referable to /. grallator; all 

of these records have been questioned in the 

synonymy, above. Both Doflein and Odhner commented 

on the small size of specimens they identified 

with /. dorsettensis from deep water off West 

Africa. All of their material, as well as all of the 

material listed above under /. dorsettensis from 

NW African localities, should be restudied. 

Some of the deeper records for /. dorsettensis, 

assigned to /. nanus, new species (p. 291), may 

well be based on /. grallator. The specimen from 

Senegal illustrated by Monod (1956, fig. 720; 

from Atlantic localities except in very large fe- Figure 75*/ herein) with long dorsal spines resemmales 

in which the median spines usually are bles /. grallator in that feature, but the interanten- 

worn down. If the Mediterranean population nular spine appears to be small as in typical /. 

proves to be distinct, the name Inachus parvirostris nanus. 

(Risso, 1816), which Risso originally proposed as ETYMOLOGY.—The specific epithet is derived 

Macropus parvirostris, is available. 

from the Latin and means "one who goes on 

Although Griffin (1974) synonymized Pseudo- stilts," alluding to the long, slender legs of this 

collodes complectens Rathbun, 1911, with /. dorsetten- species. 

sis (Pennant, 1777), reexamination of Griffin's TYPE-LOCALITY.—Gulf of Guinea off Nigeria, 

material from the western Indian Ocean, as well 03°49'N, 07°38'E to 03°48'N, 07°42'E, in 264as 

the type of P. complectens, leads us to believe 269 m (Pillsbury Sta 255). 

that the Indian Ocean form is a distinct species. DISPOSITION OF TYPES.—The holotype (Crust. 

As noted above Griffin was correct in synonymiz- D. 27154) a male from Pillsbury Sta 255, is in the 

ing Pseudocollodes with Inachus. We suspect that Rijksmuseum van Natuurlijke Historie, Leiden;


paratypes also are in that collection and in the 



BIOLOGY.—Inachus grallator is a deep water species, 

known to occur in depths between 100 m 

and 250-300 m on soft bottom; Capart (1951) 

reported specimens from green mud, brown sandy 

mud, muddy sand and green muddy sand. The 

specimens recorded herein from the Congo were 

taken on sandy mud in 247-250 m. 


and November (Capart, 1951; present paper). 

DISTRIBUTION.—West Africa, from a few localities 

between Nigeria and Angola. Records in the 

literature include the following: 

Gabon: 45 mi [72.5 km] NxE of Port-Gentil, 00°S, 08°- 

58'E, 250-300 m (Capart, 1951). 

Angola: 25 mi [40 km] WNW of Ambriz, 07°39'S, 

12°47'40"E, 100 m; 8 mi [12.8 km] NxE of Baia dos Elefantes, 

13°05'S, 12°46'E, 100-110 m; 18 mi [29 km] WSW 

of Baia dos Tigres, 16°36'S, 11°27'E, 110 m (all Capart, 

1951). 16°37'S, 11°22'E, 126 m (Crosnier, 1970). 

Inachus guentheri (Miers, 1879) 

Inachus guentheri.—Monod, 1956:529, figs 723-730 [Senegal, 

Gabon, Angola]. 

SYNONYM.—Inachus antarcticus Doflein, 1904. 

DISTRIBUTION.—Off West Africa and South 

Africa, sublittoral, from about 18 to more than 

180 m. 

Inachus leptochirus Leach, 1817 

Inachus leptochirus Leach, 1817, in 1815-1875, pi. 22B.—A. 

Milne Edwards and Bouvier, 1894:7 [Azores]; 1899:45 

[key]; 1900:145 [part; Spain; Azores; Mauritania].—Doflein, 

1904:73 [Seine Seamount].—Bouvier, 1922:79, pi. 2: 

fig. 5 [color] [Josephine Seamount; Princesse Alice 

Bank; S of Almadena].—Balss, 1922:72 [listed].—Monod, 

1933b:503 [p. 48 on separate] [listed].—Chapman and 

Santler, 1955:376 [Azores].—Maurin, 1968b:484 [Spanish 

Sahara].—Zariquiey Alvarez, 1968:472, fig. 157a,b 


36 [North Atlantic ].—Tiirkay, 1976a:26 [listed], 40, fig. 

35 [Morocco]. 

SYNONYM.—Inachus affinis Rizza, 1839. 

DISTRIBUTION.—Eastern Atlantic, from the He- 

brides southward to Mauritania, including the 

Azores; Mediterranean. Sublittoral, from about 

30 m to more than 500 m. 

*Inachus nanus, new species 

FIGURE 75 

Inachus dorsettensis.—Monod, 1956:526, 632, figs. 715-722 

[?part].—Rossignol, 1957:116 [key]. [Not Inachus dorsettensis 

(Pennant, 1777).] 


Sta 68, 70 m, broken shell, 36, 29 ov (L). Sta 69, 29 m, coral 

or rock, 1(5 (W). 

Ivory Coast: Sta 42,62-75 m, mud with brown, branched 

Foraminifera, 1

292 

a 


FIGURE 75.—Inachus nanus, new species: a, dorsal view; b, front, ventral view; c, carapace, lateral 

view; d, carapace of different specimen, lateral view (this could be based on /. grallator, new 

species); e, male pleopod. (All from Monod, 1956, figs. 715-717, 720, 722.) 

half the carpus beyond front. Chelipeds in females 

slenderer than in males, about 1.1 times as long 

as carapace, carpus not extending to front. Chelipeds 

in both sexes with lower margin of merus 

and inner border of carpus very spiny. Palm 

tuberculate in both sexes, length about 1.7 times 

height in females, 1.3-1.4 times greatest height in 

males. Dactylus of chela with 2 rounded teeth 

proximally on cutting edge in males. 

Second pereiopod about 4 times carapace


length in males, about 3.5 times carapace length 

in females (most second pereiopods detached); 

dactylus with minute tubercles ventrally, shorter 

than propodus, latter slightly shorter than merus. 

Dactylus of third pereiopod not overreaching propodus 

of second. Second to fifth pereiopods decreasing 

in size posteriorly, fifth about twice as 

long as carapace in males, about 1.5 times as long 

as carapace in females. Dactylus of fifth pereiopod 

curved but not strongly falcate, with 1 or 2 

large, subdistal spinules flanked proximally by 

line of smaller spinules and tubercles. 

Terminal somite of male abdomen rounded 

apically. Male pleopod as shown (Figure I5e). 


4.9 to 8.9 mm, of ovigerous females 4.8 to 

8.3 mm. 

REMARKS.—Inachus nanus resembles the Mediterranean 

/. communissimus Rizza and differs from 

the other Atlantic species of Inachus, which have 

a transverse row of four tubercles on the gastric 

region. It differs from /. dorsettensis (Pennant) and 

/. grallator, new species, in having a very small 

interantennular spine, which is not visible between 

the teeth of the rostrum in dorsal view. 

Inachus nanus, which has a maximum carapace 

length of less than 10 mm, is a much smaller 

species than /. communissimus; in the latter species 

Forest (1965a) recorded specimens with carapace 

lengths up to 26 mm. The carapace in /. nanus is 

longer than broad or about as broad as long; in 

/. communissimus it is much broader than long. In 

/. nanus the posterolateral margin of the carapace 

is continuous with the free epimeral margin 

whereas in /. communissimus it is not continuous, 

being interrupted on the anterior part of the 

branchial region. 

Some of the West African records of/, dorsettensis, 

referred above to /. grallator, new species, may 

well be based on /. nanus; this can only be determined 

by examining the material. 

TYPE-LOCALITY.—Gulf of Guinea off the Ivory 

Coast, 05°02.5'N, 03°49.5'W to 05°05'N, 03°- 

52*W, in 62-75 m {Pillsbury Sta 42). 


D. 27155), a male, cl 8.3 mm, cb 7.8 mm, is in 

the Rijksmuseum van Natuurlijke Historie, Leiden; 

paratypes also are deposited there and in the 


Institution. 


Latin and refers to the diminutive size of this 

species. 

BIOLOGY.—Inachus nanus is a sublittoral species, 

occurring in depths between 29 and 118 m. Only 

three of the 21 depth records available are from 

100 m or more and only one is from less than 40 

m. Although three of the lots taken by the Pillsbury 

were from rough bottom, coral or rock, most of 

the specimens were taken on mud, either with 

Jullienella or bryozoans. Some of the deeper records 

in Monod (1956) may be based on specimens 

of/, grallator. 

Ovigerous females have been taken in February, 

May, June, July, September, and October, 

suggesting that the species spawns all year 

(Monod, 1956; Pillsbury). 

DISTRIBUTION.—Inachus nanus is a sublittoral 

West African species, known from a few localities 

between Senegal and Gabon. The following records 

are all from Monod (1956): 

Senegal: S of Cap Vert, 97-98 m; Fosse de Kayar, 100 m; 

S of Fosse de Kayar, 50 m; SE of lie de la Madeleine, Dakar, 

48 m; off Goree, 95, 96, and 132 m. 

Sierra Leone: No specific locality, 118 m. 

Gabon: Libreville, 60 m. 

It has not previously been recorded from Liberia, 

Ivory Coast, Ghana, Nigeria, or Cameroon, 

but all of these lie within its known range. 

Inachus phalangium (Fabricius, 1775) 

Inachus phalangium.—Monod, 1956:531 [references].—Zariquiey 

Alvarez, 1968:472, fig. 159c [Spain; references].— 

Christiansen, 1969:102, fig. 42, map 35 [North Atlantic]. 

Inachus dorhynchus.—Rossignol, 1957:115 [key].—Forest and 


SYNONYMS.—ICancer tribulus Linnaeus, 1767 

[name suppressed by ICZN]; Inachus dorynchus 

Leach, 1814; Macropus aracnides Risso, 1816. 

DISTRIBUTION.—Eastern Atlantic, from about 

60°N latitude southward to the Cape Verde Is-

294 

lands; Mediterranean. Sublittoral, from about 10 

to at least 150 m. 

Inachus thoracicus Roux, 1830 

lnachus thoracicus.—Capart, 1951:68 [Congo].—Monod, 1956: 

532, figs. 731-735 [Senegal, Gambia].—Rossignol, 1957: 

115 [key].—Longhurst, 1958:89 [Sierra Leone].—Maurin, 

1968a:48, 59; 1968b:480, 484, 486, fig. 5 [both Spanish 

Sahara, Mauritania].—Zariquiey Alvarez, 1968:473, figs. 

157c,d, 158a,b [Spain; references]. 

SYNONYM.—Inachus cocco Rizza, 1839. 

DISTRIBUTION.—Eastern Atlantic, from southern 

Europe to the Congo, including the Canary 

Islands; Mediterranean. Sublittoral, to at least 

100 m. 

Genus Macropodia Leach, 1814 

Macropodia Leach, 1814:395 [type-species: Cancer longirostris 

Fabricius, 1775, by monotypy; gender: feminine; name 


Peridromus Gistel, 1848:ix [replacement name for Macropodia 

Leach, 1814 (as Macropus); type species: Cancer longirostris 

Fabricius, 1775; gender: masculine]. 

REMARKS.—Specific identification of the eastern 

Atlantic species of Macropodia has remained 

particularly difficult until recently when Forest 

and Zariquiey Alvarez (1964) reviewed the Mediterranean 

species and their nomenclature (Forest, 

1965b). They pointed out that in the past 

identification often was based on keys provided 

by A. Milne Edwards and Bouvier (1899:48) or 

Bouvier (1940:362) and that results obtained 

through using these keys generally were unsatisfactory. 

Forest and Zariquiey Alvarez (1964) recognized 

five species in the Mediterranean: M. rostrata 

(Linnaeus, 1761); M. longipes (A. Milne Edwards 

and Bouvier, 1899); M. longirostris (Fabricius, 

1775); M. czernjawskii (Brandt, 1880); and a 

new species they named M. linaresi. 

In discussing the literature relating to these 

species, Forest (1965b:349) pointed out that the 

epithet longirostris in the past had been applied to 

three distinct taxa: M. longirostris proper, M. longipes, 

and a third species from areas in the north- 


eastern Atlantic north of the Mediterranean, M. 

tenuirostris Leach, 1814. Thus many earlier records 

of M. longirostris from the Mediterranean were 

based on that species or M. longipes. 

Monod (1956:559) included M. longirostris as a 

member of the West African fauna, based on 

earlier records by A. Milne Edwards and Bouvier 

(1900:156, 157) who reported two males and a 

female from the Bane d'Arguin, one male from 

Senegal, and one female from the Cape Verde 

Islands, as well as material from Morocco. Forest 

and Zariquiey Alvarez (1964:228) considered M. 

longirostris to be a Mediterranean endemic and 

identified many of A. Milne Edwards and Bouvier's 

records of M. longirostris with M. longipes, 

noting (1964:226, footnote) that material reported 

by these authors from three localities, 

including Morocco and Senegal, no longer was in 

the collection at Paris. Apparently there are no 

substantiated records for either M. longipes or M. 

longirostris from tropical African waters, and we 

have omitted these species from our accounts 

below. 

Material from the third locality, the Strait of 

Bonifaccio (between Corsica and Sardinia), is in 

the collection of the National Museum of Natural 

History, Smithsonian Institution, under catalog 

number USNM 22978. It consists of a single 

male, cl 15.9 mm, clearly identifiable with M 

longirostris. Outside of the Mediterranean, M. longipes 

is known with certainty from the Bay of 

Biscay (Forest and Zariquiey Alvarez, 1964:226). 

The eastern Atlantic species of Macropodia still 

require a great deal of work, especially the status 

of the species occurring on the NW African coast, 

including M. longirostris auctorum (including 

Peres, 1964:20, Morocco), M. longipes auctorum 

(Forest and Gantes, 1960:357, Morocco; Bas, Arias, 

and Guerra, 1976, table 3, Spanish Sahara; 

Tiirkay, 1976a:40, fig. 33, Portugal and Morocco), 

M. egyptia sensu A. Milne Edwards and 

Bouvier, 1900, and M. intermedia Bouvier, 1940 

(see p. 300). We believe that the status of Mediterranean 

material identified with M. rostrata 

(Linnaeus, 1761) also requires investigation.


Key to Tropical West African Species of Macropodia 

1. Dactylus of fifth leg slightly curved 2 

Dactylus of fifth leg strongly arcuate 4 

2. Rostrum longer than antennal peduncle, more than 2/3 postrostral length 

of carapace. [Nuchal spine present. Basal article of antenna with strong 

spines ventrally] M. longipes 

Rostrum shorter than antennal peduncle, less than half postrostral length 

of carapace 3 

3. Rostrum extending beyond midlength of distal article of antennal peduncle. 

Nuchal spine present. Orbital margin with dorsal spines or tubercles. 

Basal article of antenna with strong spines ventrally M. gilsoni 

Rostrum falling short of midlength of distal article of antennal peduncle. 

Nuchal spine absent. Orbital margin smooth. Basal article of antenna 

smooth, unarmed M. spinulosa 

4. Epistome with longitudinal ridge on each side extending from pore of 

antennnal gland to basal article of antenna M. macrocheles 

Epistome lacking longitudinal ridge on each side 5 

5. Rostrum longer than antennal peduncle. Basal segment of antenna 

unarmed 6 

Rostrum not reaching end of antennal peduncle. Basal segment of antenna 

armed with spinules and tubercles 7 

6. Rostrum more than 2/3 postrostral length of carapace. Basal article of 

antenna smooth M. longicornis 

Rostrum less than half postrostral length of carapace. Basal article of 

antenna irregularly tuberculate M. straeleni 

7. Rostrum extending beyond midlength of distal segment of antennal peduncle. 

Gastric region with sharp, erect spine ... M. doracis, new species 

Rostrum not extending to midlength of distal segment of antennal peduncle. 

Gastric region with low, conical prominence 

Af. hesperiae, new species 

Macropodia doracis, new species Rostrum (Figure 76a) moderately long, extending 

almost to but falling short of end of distal 

FIGURE 76 segment of antennal peduncle, ornamented with 

„ , . , „ , , , „ . curved hairs. Apices of rostral teeth appressed. 

Stenorhynchus aeeyptius.—A. Milne Edwards and Bouvier, r \ r . 

1900:155 [part] [not Sunorynchus tgyptius H. Milne Ed- Rostrum upturned dorsally. Nuchal region with 

wards, 1834 = Cancer longirostns Fabricius, 1775]. lateral spinule, visible in dorsal view (Figure 76a). 

MATERIAL EXAMINED.—PilUbury Material: None. 

Other Material: Cape Verde Islands: between Ilheu Branco 

and Ilheu Raso, 110-180 m, sand, rocks, Talisman No. 105, 

Protogastric region smooth, lacking dorsal tubercies 

medially. Hepatic regions each with low, 

conical prominence dorsally. Hepatic lobes each 

- i i i i J J L I n r> 

,, ¥ , " „ „ nk, , , s 

27Jul 1883, l9ov(W, holotype). 

with bluntly rounded tubercle apically. Gastric 

r 7 

. . J . 

region with high, erect spine dorsally. Cardiac 

DESCRIPTION.—Size small, carapace length of region with broad, obtuse prominence dorsally, 

adult less than 10 mm (7.8 mm). lower than gastric spine. Branchial regions each

296 


FIGURE 76.—Macropodia doracis, new species, holotype, ovigerous female, cl 7.8 mm, Cape Verde 

Islands: a, carapace, dorsal view; b, carapace, lateral view; c, chela; d, third pereiopod; e, fourth 

pereiopod;/, fifth pereiopod. (All pereiopods slightly damaged.) 

with 1 low dorsal epibranchial tubercle, 1 broadly 

rounded anterolateral epibranchial tubercle, 1 

dorsal mesobranchial tubercle, and 1 metabranchial 

tubercle situated almost on posterior margin. 

Anterolateral margins with rounded tubercle 

posterior to hepatic lobe. 

Basal antennal segment (Figure 76b) with 2 

tubercles on each side, larger, almost spiniform, 

anteriormost. Distal article of antennal peduncle 

about twice as long as penultimate. Epistome 

with rounded lobule near opening of antennal 

gland and with tubercle at base of antenna. 

Ocular peduncle lacking distinct anterior lobe. 

Cornea with distal dorsal tubercle. 

Cheliped (Figure 76V) of adult female slender, 

fingers, measured dorsally, longer than palm, latter 

with row of spinules dorsally. Merus with 2 

prominent dorsal tubercles. 

Dactylus of second pereiopod slender, slightly 

curved distally, apex broken; merus shorter than 

carapace. Merus of each walking leg with blunt, 

unarmed dorsal projection (Figure 76d). Dactyli 

of fourth and fifth pereiopods (Figure 76eJ) 

strongly arcuate, ventral margin toothed along 

entire margin. 

MEASUREMENTS.—Carapace length of ovigerous 

female 7.8 mm. 

REMARKS.—Macropodia doracis approaches M. 

hesperiae, (p. 298), in general facies, but differs in 

having a longer rostrum, which extends almost to


the end of the antennal peduncle, in having a 

shorter distal segment on the antennal peduncle, 

and in having a sharp, erect gastric spine. The 

present species may be closest to Macropodia aegyptia 

sensu Bouvier (1940:364, fig. 220), from the 

Canary Islands, but in that species the epistome 

has more spines and the basal segment of the 

antennal peduncle is armed with three large 

sharp spines rather than a spinule and a tubercle. 

TYPE-LOCALITY.—Cape Verde Islands, between 

Ilheu Branco (16°39'N, 24°41'W) and IIheu 

Raso (16°37'N, 24°36 / W) in 110-180 m. 

DISPOSITION OF TYPES.—The unique holotype 

(USNM 22980), an ovigerous female, cl 7.8 mm, 

is in the National Museum of Natural History, 

Smithsonian Institution. Apparently it was sent 

to the Smithsonian on exchange early in the 

1900's. 


from a Latin name for the Cape Verde Islands, 

Doraces insulae. 

DISTRIBUTION.—Known only from the typelocality 

in the Cape Verde Islands. 

Nigeria: Sta 237, 101 m, 16\ 29 (L). Sta 239, 73 m, 86, 79 

(L). Sta 241, 59-63 m, mud and shell, 116\ 89 (5 ov) (W). 

Cameroon: Sta 259, 59 m, mud and broken shell, 36, 39 

(W). Sta 260, 46 m, Id (L). 

Geronimo Material: Gabon: Sta 235, 100 m, 16* (W). 

Undaunted Material: Angola: Sta 95, 126 m, 26, 19 ov (L). 

Other Material: Cameroon: 03°54'N, 08°53'E, 64 m, Aug 

1963, A. Crosnier, 56, 29 (1 ov) (W). 03°55.5'N, 08°52.5'E, 

60 m, Ombango, A. Crosnier, 5 Jan 1963, 19 ov (W). 

* Macropodia gilsoni (Capart, 1951) 


Figures: Capart, 1951, fig. 20, pi. 1: figs. 4, 10, 

pi. 2: fig. 3; Monod, 1956, figs. 811-822. 

Male Pleopod: Capart, 1951, pi. 2: fig. 3 (Cabinda); 



6.5 to 19 mm, of non-ovigerous females 7.5 to 

14 mm, of ovigerous females 11 to 16.7 mm. 

REMARKS.—Macropodia gilsoni, M. hesperiae, new 

species, M. intermedia, and M. longipes are the only 

West African species of the genus with strong, 

well-developed nuchal spines; these spines are 

also present in the northern species, M. tenuirostris 

(Leach) (Christiansen, 1969, fig. 47). Further, M. 

gilsoni and M. intermedia are the only species in the 

genus with spines on the supraorbital margin. 

Differences between these two species are pointed 

Achaeopsis gilsoni Capart, 1951:65, fig. 20, pi. 1: figs. 4, 10, pi. out (p. 300) under the account of M. intermedia. 

2: fig. 3.—Rossignol, 1957:115 [key]. 

Macropodia gilsoni appears to be quite variable 

Macropodia gilsoni.—Monod, 1956:555, figs. 811-822.— in some features, especially in the extent of spi- 

Longhurst, 1958:89.—Rossignol, 1962:123.—Crosnier, nation: the supraorbital spines may be reduced 

1964:34. 

to tubercles; the walking legs often have but one 

Macropodia intermedia.—Guinot and Ribeiro, 1962:78.—Forest 

and Guinot, 1966:115.—Crosnier, 1970:1215 [listed], spine or even up to five spines on the merus of 

1218. [Not M. intermedia Bouvier, 1940.] 

the fourth leg rather than three; the hairiness of 

Macropodia.—Voss, 1966:22. 

the legs is variable; and in some specimens the 



298 


bottoms, including bottoms with shells and bry- dorsal view. Protogastric region smooth, lacking 

ozoans or foliate Foraminifera. 

dorsal tubercles medially. Hepatic regions each 

Ovigerous females have been found in all with low, conical prominence dorsally. Hepatic 

months but July, indicating that the species lobes each with apical spinule, flanked mesially 

spawns all year. 

with spine on anterior margin in some specimens. 

DISTRIBUTION.—West Africa, from localities be- Gastric and cardiac regions each with low, conical 

tween Senegal to Angola (16°37'S); sublittoral, prominence. Branchial regions with poorly 

in depths between 37 and about 200 m. Capart marked prominences: 1 low, rounded epibran- 

(1951) studied numerous specimens from localichial, 1 broad dorsal mesobranchial, 1 low metaties 

between Gabon and Angola, and Monod branchial near posterior margin, and sharp tu- 

(1956) had material from Senegal and Guinea. bercles laterally. Anterolateral margins with 

Records since 1956 include the following: sharp, anteroventrally directed spine posterior to 





hepatic lobe. 

Basal antennal segment (Figure lib) with 1-3 

spinules or sharp tubercles, anteriormost largest. 

Distal article of antennal peduncle about 3 times 


1958). 

Ivory Coast: 04°35'N, 06°40'W to 04°35'N, 06°41'W, 64 

m (Voss, 1966). 

Cameroon: No specific locality, deeper than 50 m (Crosnier, 

1964). 

as long as penultimate. Epistome with spinule 

near opening of antennal gland, occasionally with 

spinule anteriorly near base of antenna. 

Ocular peduncles lacking distinct anterior lobe. 

Cornea with distinct dorsal tubercle. 

Gabon: W of Barre des Portugais; W of Mayumba, 110 

m (Rossignol, 1962). 

Angola: Luanda, 102-122 m (Guinot and Ribeiro, 1962). 

16°37'S, 11°22'E, 126 m (Crosnier, 1970). 

Chelipeds of adult males larger and more robust 

than in females. Fingers as long as palm, 

latter with row of spinules dorsally. Merus with 

3 erect dorsal spines or lobes. 

* Macropodia hesperiae, new species 

Dactylus of second pereiopod (Figure lie) slender, 

slightly curved distally, 7/9 as long as merus, 

FIGURE 77 

latter longer than carapace and antennal peduncles. 

Meri of second through fifth pereiopods each 

Macropodia linaresi.—Forest and Guinot, 1966:117 [not Macrowith 

single large distal dorsal spine, latter lower 

podia linaresi Forest and Zariquiey Alvarez, 1964]. 


on merus of fifth pereiopod. Dactyli of fourth and 

fifth (Figure lid) pereiopods strongly arched, 


Coast: Sta 60, 79-82 m, coral or rock, 16\ 1$ (L). Sta 65, 46ventral 

margin toothed along entire margin. 


49 m, 16* (holotype), 1? (L). 

Ghana: Sta 22, 51 m, rough bottom, 16* (W). 

Nigeria: Sta 230, 82-97 m, hard ground, with gorgonians, 

coral, rock, 1(5 (W). 

5.2 to 9.4 mm, of females 4.2 and 7.3 mm. 

REMARKS.—Macropodia hesperiae resembles M. 

linaresi Forest and Zariquiey Alvarez, 1964, but 

differs as follows: The rostrum is longer, with the 

DESCRIPTION.—Size small, carapace lengths of apices of the spines divergent rather than ap- 

adults less than 10 mm. Rostrum (Figure 77a,*) pressed; the dorsal prominences of the carapace 

short, overreaching base but falling short of mid- are much less prominent; the carapace is 

dle of fifth article of antennal peduncle, orna- smoother and less hairy; and the legs are longer, 

mented with curved hairs. Apices of rostral teeth 

divergent in some specimens. Rostrum slightly 

upturned dorsally (Figure Tib). Nuchal region 

usually with lateral spinule or tubercle, visible in 

the merus of the second pereiopod being longer 

than the carapace and antennal peduncles combined. 

If our interpretation of the original account of


a 

FIGURE 77.—Macropodia hesperiae, new species. Male paratype, cl 6.0 mm, Pillsbwy Sta 60: a, 

carapace; b, front, lateral view; c, second pereiopod; d, fifth pereiopod. Female paratype, cl 4.2 

mm, Pillsbury Sta 60: e, carapace. 

M. linaresi is correct, the basal antennal segment 

in that species lacks distinct spines; it was described 

as "avec des granules ou des courtes spinules" 

(Forest and Zariquiey Alvarez, 1964:241). 

Also, in that species, the merus of the second 

pereiopod was described as being slightly longer 

than the carapace. We have examined a male cl 

7 mm, and a female, cl 7 mm (RMNH Crust. D. 

20133), from Cadaques, Spain, and found that 

both specimens had three spinules ventrally on 

the basal segment of the antenna, the anteriormost 

largest, and, in the female, the merus of the 

second pereiopod is longer than the carapace; all 

the pereiopods of the male were detached. 

We have no hesitation in identifying material 

identified with M. linaresi from Senegal by Forest 

and Guinot (1966:117) with M. hesperiae as they 

noted that "elle ne differe guere des exemplaires 

typiques .. . que par un rostre legerement plus 

grele et une carapace plus lisse et plus glabre." 

Macropodia hesperiae is a much smoother species 

than M. linaresi. 

TYPE-LOCALITY.—Off the Ivory Coast, 

04°15'N, 07°32 / W to 04°12'N, 07°35.5'W, 46-49 

m, Pillsbury Sta 65. 


D. 23891), a male from Pillsbury Sta 65, is in the 

Rijksmuseum van Natuurlijke Historie, Leiden; 

paratypes also are at Leiden and in the National 


Washington, D.C. 

ETYMOLOGY.—The specific name is derived 

from the Latin name for the Gulf of Guinea, 

Hesperium mare. 

BIOLOGY.—Macropodia hesperiae is a sublittoral 

species known from depths of 46-49 m, 51 m, 79- 

82 m, and 82-97 m (Pillsbury) and 65-75 m 

(Calypso; Forest and Guinot, 1966). Our specimens


were taken on rough bottom at three or four 

stations. The Calypso specimens were taken on 

mud, sand, and compact sand (sable construit). 

An ovigerous female was taken in May by the 

Calypso (Forest and Guinot, 1966). 

DISTRIBUTION.—Gulf of Guinea, from off Senegal, 

the Ivory Coast, Ghana, and Nigeria, in 

depths between 46-49 and 82-97 m. Records in 

the literature include the following: 



Nigeria: 06° 1 l'N, 03°36'E to 06° 10'N, 03°38'E, 82-97 m 

(Voss, 1966). 

Macropodia intermedia Bouvier, 1940 

Macropodia longirostris var. intermedia Bouvier, 1940:366.—Capart, 

1951:76. 

?Macropodia gilsoni.—Maurin, 1968a:48, 62; 1968b:484, 486, 

489, fig. 5 [not Macropodia gilsoni (Capart, 1951)]. 


Other Material: Morocco: No specific locality, holotype of 

Macropodia longirostris var. intermedia, 16* (MP). 

Remarks.—Although several authors have synonymized 

Macropodia gilsoni Capart, 1951, with 

Macropodia intermedia Bouvier, 1940 (p. 297), we 

believe that two taxa may be involved, M. intermedia 

from areas to the north of the tropical 

region and M. gilsoni from the Gulf of Guinea 

proper, and, until more material from the two 

regions can be studied together, we prefer to 

recognize two species. 

The holotype of M. intermedia, a male, cl 19 

mm, cb 13.5 mm, was examined by Manning at 

the Museum national d'Histoire naturelle in Paris 

in 1971. Several differences between it and our 

material of M. gilsoni were observed: (1) The 

rostrum is longer than the antennal peduncle; it 

is shorter in M. gilsoni (Monod, 1956, fig. 814). 

(2) The posterior margins of the antennular fossae 

as well as those portions that overlap the basal 

segment of the antenna are smooth; they usually 

are spinulose in M. gilsoni (Monod, 1956, figs. 

814, 815). (3) The ventral spine on the ischium of 

the fifth pereiopod appears to be much less prom- 

inent and appears to be directed laterally rather 

than posteriorly as in M. gilsoni. 

Although these differences are relatively minor, 

they suggest to us that M. intermedia should be 

considered distinct from M. gilsoni, at least for the 

time being. 

DISTRIBUTION.—Eastern Atlantic, from the NW 

coast of Africa, sublittoral. All but the first of the 

following records require verification: 

Morocco: Atlantic coast (Bouvier, 1940). 

Spanish Sahara: Pulpito Bay, 20-30 m (Capart, 1951). 

Between Cabo Corbeiro and Cabo Blanco, 60-80 m 

(Maurin, 1968a). Between Cabo Barbas and Cabo Blanco, 

50-90 m (Maurin, 1968b). 

Mauritania: S Bane d'Arguin, 90 150 m (Maurin, 

1968a). Bane d'Arguin, 40-60, 60-70, 90-100 m (Maurin, 

1968b). 

Macropodia longicornis (A. Milne Edwards 

and Bouvier, 1899) 

Macropodia longicornis.—Monod, 1956:562 [references]. 

DISTRIBUTION.—Cape Verde Islands; sublittoral, 

to at least 275 m. 

Macropodia longipes (A. Milne Edwards and 

Bouvier, 1899) 

Stenorhynchus longipes A. Milne Edwards and Bouvier, 1899: 

48. 

IMocropodia longirostris.—Monod, 1956:559 [references].— 

Rossignol, 1957:115 [key].—Peres, 1964:20 [Morocco]. 

[Not Cancer longirostris Fabricius, 1775.] 

Macropodia longipes.—Forest and Games, 1960:357 [Morocco].—Forest 

and Zariquiey Alvarez, 1964:226, 241 

[key], figs. 2, 6, 13 [review of Mediterranean species].— 

Forest, 1965b:349, 350 [discussion].—Bas, Arias, and 

Guerra, 1976, table 3 [Spanish Sahara].—Tiirkay, 1976a: 

25 [listed], 40, fig. 33 [Portugal, Morocco]. 

Macropodia tenuirostris longipes.—Forest, 1978:337, figs. 9, 16. 

REMARKS.—Forest (1978:337) considers M. longipes 

to be the southern subspecies of M. tenuirostris 

(Leach, 1814). 


Gulf of Gascogne to Mauritania, Cape Verde 

Islands, and western Mediterranean; sublittoral, 

in 50-445 m (Forest, 1978).


*Macropodia macrocheles (A. Milne Edwards 

and Bouvier, 1898) 

Macropodia macrocheles.—Capart, 1951:77, fig. 23.—Monod, 

1956:560, 632, figs. 823-827.—Rossignol, 1957:115 

[key].—Longhurst, 1958:89.—Guinot and Ribeiro, 1962: 




Sta 82, 146-150 m, 1(5, 19 ov (L). 

Nigeria: Sta 255, 264-269 m, 36, 2$ (1 ov) (L, W). 

Geronimo Material: Gabon: Sta 197, 200 m, 1$ (W). 

Other Material: Mauritania: Off Cabo Blanco, 240 m, 

Talisman, 13 Jul 1883, syntype, 1$ ov (W, USNM 22979). 



figs. 823-827. 

Male Pleopod: Monod, 1956, fig. 827 (Senegal). 


12.3 to 21 mm, of non-ovigerous females 11.8 to 


BIOLOGY.—Macropodia macrocheles is a deep water 

species, occurring between 96 and 300 m on 

soft bottoms (mud and sand, muddy sand), generally 

in depths greater than 150 m. 

Ovigerous females have been taken in January, 

February, May, June, and December (Capart, 

1951; Monod, 1956; Pillsbury). 


Mauritania and Angola, in depths between 

96 and 300 m. Monod (1956) reported the species 

from Senegal and Sierra Leone; since 1956 it has 

been taken from the following localities: 

Guinea-Bissau: 10°32'N, 16°53.5'W, 174 m (Forest and 


Sierra Leone: 08°45'N, 14°38'W, 220-240 m (Longhurst, 

1958). 

Liberia: 04°57'N, 09°30'W to 04°58'N, 09°32'W, 146- 

150 m (Voss, 1966). 

Angola: Baia dos Tigres, 150-200 m (Guinot and Ribeiro, 

1962). 

* Macropodia spinulosa (Miers, 1881) 

FIGURE 78a 

Stenorhynchus rostratus var. spinulosus Miers, 1881a:206. 

Stenorhynchusphalangium.—Studer, 1882:335; 1883:7 [not Cancer 

phalangium Fabricius, 1775]. 

Macropodia rostrata spinulosa.—Rathbun, 1900a:293 [listed]. 

Macropodia rostrata.—Rathbun, 1900a:293 [listed].—Balss, 

1922:71.— Odhner, 1923:18.—Monod, 1933b:501 

[listed].—Capart, 1951:74, pi. 1: fig. 3.—Sourie, 1954b: 

147.—Monod, 1956:562, figs. 828-836.—Rossignol, 1957: 

115 [key].—Longhurst, 1958:89.—Gauld, 1960:72.— 

Guinot and Ribeiro, 1962:78, pi. 4: figs. 2, 3.—Forest and 

Guinot, 1966:116.—Le Loeuff and Intes, 1968:46, table 

1.—Crosnier, 1970:1218. [Not Macropodia rostrata (Linnaeus, 

1761).] 

Stenorhynchus rostratus.—Doflein, 1904:69.—Lenz and 

Strunck, 1914:272 [part, not South African specimens?]. 

[Not Macropodia rostrata (Linnaeus, 1761).] 

?Stenorhynchus phalangium.—Stimpson, 1907:22 [Madeira] 

[not Cancer phalangium Fabricius, 1775]. 


Coast: Sta 46, 38-42 m, mud with dense Jullienella, 56, 10$ 

(7 ov) (W). Sta 47, 37 m, bottom with Jullienella, 16 (L). Sta 

48, 22 m, Id, 3$ ov (L). Sta 65, 46-49 m, 16* (L). 

Ghana: Sta 16, 46 m, mud with Foraminifera, shells, ld\ 

1$ (W). Sta 17, 48 m, fine sand and green mud, 16 (L). Sta 

23, 42 m, foliate brown to orange bryozoans, 26, 2$ ov (L). 

Sta 24, 35-37 m, dark red bryozoans, 26, 19 ov (W). Sta 28, 

49-53 m, 10

302 

a 

FIGURE 78.—a, Macropodia spinulosa (Miers) (from Monod, 

1956, fig. 828); b, Stenorhynchus lanceolatus (Brulle) (from 

Monod, 1956, fig. 838). 

rostrata (a character mentioned by Forest and 

Zariquiey Alvarez, 1964:241, key); only the proximal 

third or fourth of the dactylus of the fifth 

pereiopod is armed with spinules or low tubercles 

whereas in M. rostrata the proximal two-thirds to 

three-fourths is armed (most of the dactylus of 

that leg is naked in M. spinulosa, lacking even 

setae); and the dorsal spines of the carapace are 

much longer (Monod, 1956, fig. 830). In addition 

the pereiopods are longer, with the third leg 

almost five times rather than three times as long 

as the carapace (Christiansen, 1969:112) and, as 

noted by Miers (188la:206) the chelipeds are very 

spinulose. Some of these differences already had 

been noted by Guinot and Ribeiro (1962:79) and 

by Crosnier (1970:1218). 

We have compared our material of M. spinulosa 

with M. rostrata from the North Sea, as well as 

from the Mediterranean, and we found some 


differences in material of M. rostrata from those 

two areas. The Mediterranean form is more slender 

(compare carapace shape in Forest and Zariquiey 

Alvarez, 1964, fig. 1, and Christiansen, 

1969, fig. 46), a feature reflected in the shape of 

the epistome, which in Mediterranean specimens 

is as long as wide but in northern specimens is 

much broader than long. Also, the carapace in 

Mediterranean specimens is much smoother and 

less hairy. Finally, in Mediterranean specimens 

the dactylus of the fifth leg is longer, more slender, 

less curved, less setose, and has low denticles only 

on the proximal third; in northern specimens the 

dactylus is short, more strongly curved, much 

more setose, and has erect spinules on the proximal 

two-thirds. These differences strongly suggest 

that two species may be involved. If they prove 

to be constant and the Mediterranean form is a 

distinct species, the name Macropodia inermis (Heller, 

1856) is available. It was named originally as 

Stenorhynchus inermis from the Adriatic Sea (Heller, 

1856:3). 

Material from South Africa identified with M. 

rostrata by Lenz and Strunck (1914) and Barnard 

(1950, 1954) resembles M. spinulosa rather than 

M. rostrata in having a short rostrum, with the 

margins of the antennular fossae visible at the 

base of the rostrum. Barnard's material, at least, 

differs from both of these more northern species 

in having distal spinules on the basal segment of 

the antenna, an apical seta on the eye, and in 

having spinules along the entire margin of the 

dactylus of the fifth pereiopod. 

BIOLOGY.—Macropodia spinulosa is a sublittoral 

species, having been taken in depths between ca 

1 and 126 m; of 55 depth records available, only 

3 (5%) are from depths of more than 100 m (108, 

110, and 126 m), 11 (20%) are from between 50 

and 100 m, 11 (20%) are from depths of 20 m or 

less, and the remainder (55%) are from depths 

between 20 and 50 m. The deeper material identified 

here with this species probably should be 

restudied. 

The species generally occurs on soft bottoms, 

mud or muddy sand, but it was taken by the 

Calypso (Forest and Guinot, 1966) on mud, rocks,


calcareous algae, sand, and Foraminifera; mud, 

sand, and compacted sand (sable construit); mud; 

mud with Area; and on calcareous algae. Our 

specimens were taken on mud, mud with Foraminifera, 

fine sand and green mud, and on bottom 

with bryozoans. Sourie (1954b) found it on 

coarse, shelly sand, bottom with Area and Pyura, 

in 10-12 m in the Baie de Dakar. Longhurst 

(1958) found it on shelly mud in 60 m, and Le 

Loeuff and Intes (1968) considered it a eurythermic 

species, living on muddy sand in 25-40 

m. 

Ovigerous females have been collected from 

January through June, and August (Capart, 

1951; Monod, 1956; Guinot and Ribeiro, 1962; 

Forest and Guinot, 1966; Crosnier, 1970; herein 

p. 301). 


Verde Islands and at least Senegal on the African 

mainland S to Angola; sublittoral, in depths between 

1 and 126 m, generally in less than 50 m. 

Records of M. rostrata from areas N of Senegal, 

such as those by Capart (1951) from Spanish 

Sahara and Stimpson (1907) from Madeira, especially 

the latter, may be based on either this 

species or M. rostrata proper. They require verification. 


Cape Verde Islands: 15°40'N, 23°06'W) 38 fm [70 m] 

(Studer, 1882, 1883). 

Senegal: Off Dakar, 14°40'-41'N, 17°18'30"-20'30"W, 

20-25 m; 14°38'-41'N, 17°2O'-23'W, 22-34 m; 14°38'- 

40'N, ^H'-iaW, 22-27 m; off Dakar; Anse Bernard, 8- 

10 m; Baie de Hann; between Goree and Thiaroye-sur-Mer 

(as Tiaroye), 15 m; and between Dakar and Goree, 16 m (all 

Monod, 1956). Baie de Dakar, 10-12 m (Sourie, 1954b). 

Goree (the type-locality) (Miers, 1881a; Lenz and Strunck, 

1914; Monod, 1956); in 20 m (Balss, 1922). Around Goree, 

ca. 1, 5-11, 7-8, 13, 15-20, 19, 20, 25, 40-41, and 46-50 m; 

between Thiaroye-sur-Mer and Bel-Air, 9 m; Mbao; SE lie 

de la Madeleine, 35 and 48 m; and off Joal, 15-17 m 

(Monod, 1956). 13°01'N, 17°24'W, 51-55 m, and 12°55.5'N, 


Guinea: Off lie Kassa, lies de Los (Capart, 1951). 09°- 

36'N, 13°57'W, 18-30 m (Forest and Guinot, 1966). 

Sierra Leone: No specific locality, 60 m (Longhurst, 1958). 

Ivory Coast: Off Grand Bassam, 25-40 m (Le Loeuff and 

Intes, 1968). 

Ghana: Off Accra, 26 and 44 m (Monod, 1956; Gauld, 

1960). 



Principe: 01°43'10"N, O7°28'2O"E, 73 m, and 01°43'N, 

07°28'55'E, 37m (Forest and Guinot, 1966). 

Annobon: 2 mi [3.2 km] off Annobon, 18-20 m (Capart, 

1951). 

Gabon: Libreville, 60 m (Monod, 1956). 

Congo: 11 mi [17.7 km] WSW Pointe-Noire, 04°52'S, 

ll°39'30"E, 58-60 m (Capart, 1951). 

Cabinda: Off the lighthouse (Capart, 1951). 

Zaire: Off the mouth of the Congo River, 44 m (Doflein, 

1904). 28 mi [32 km] WNW of Banana, 05°54'S, 11°58'- 

30"E, 50 m, and 05°56'S, 12°E, 50-60 m (Capart, 1951). 

Angola: Benguela, 13-26 m; Baia da Caota, 13, 16, and 

18 m; and Baia Farta, 5 m (Guinot and Ribeiro, 1962). 

Porto Alexandre, 72 and 108 m (Odhner, 1923). 18 mi 

[29 km] WSW Baia dos Tigres, 16°36'S, 11°27'E, 110 m 

(Capart, 1951). 16°37'S, 11°22'E, 126 m (Crosnier, 1970). 

* Macropodia straeleni Capart, 1951 

Macropodia straeleni Capart, 1951:79, fig. 24, pi. 1: fig. 5.— 

Monod, 1956:566, fig. 837.—Rossignol, 1957:115 

[key].—Forest and Guinot, 1966:117. 

MATERIAL EXAMINED.—Pilbbury Material: Liberia: 

Sta 68, 70 m, broken shell, 2c? (L). 


Foraminifera, 26* (L). Sta 60, 79-82 m, coral or rock, 26, 29 

ov (W). Sta 65, 46-49 m, 1$ ov (L). 

Nigeria: Sta 239, 73 m, 1$ ov (W). 

Other Material: Congo: 05°20'S, 11°37'E, 200 m, 10 Dec 

1963, A. Crosnier, 36, 1? ov (W). 


Figures: Capart, 1951, fig. 24, pi. 1: fig. 5. 


5.1 to 8.7 mm, of ovigerous females 5.2 to 6.5 

mm. 

BIOLOGY.—Macropodia straeleni occurs sublittorally 

in moderately deep water. Capart (1951) 

took it in 73 to 140-150 m on sandy mud and it 

was taken by the Calypso in 65-75 m on mud, 

sand, and compacted sand (sable construit) (Forest 

and Guinot, 1966). The Pillsbury specimens, 

for which data are available, were taken on mud

304 

with brown, branched Foraminifera in 62-75 m, 

broken shell in 70 m, and on coral or rock in 79- 

82 m. The deepest recorded occurrence of the 

species is in 200 m off the Congo and the Ivory 

Coast (Forest and Guinot, 1966). 

Ovigerous females have been taken in May, 

June, October, November, and December (Capart, 

1951; Forest and Guinot, 1966; present paper). 


Senegal (12°55.5'N) and Angola (07°39'S), 

in depths between 46-49 m and 200 m. Capart 

(1951) based his account on material from off the 

Congo, Cabinda, and Angola; Monod (1956) 

added no material. Other records in the literature 




Ivory Coast: No specific locality, 200 m (Forest and 


Genus Stenorhynchus Lamarck, 1818 

Pactolus Leach, 1815b: 19 [type-species: Pactolus boscii Leach, 

1815, a subjective junior synonym of Cancer seticomis 

Herbst, 1788, by monotypy; gender: masculine; suppressed 

by the International Commission on Zoological 

Nomenclature under its plenary powers (Opinion 763, 

1966); name 1778 on Official Index]. 

Stenotynckus Lamarck, 1818:236 [invalid original spelling of 

Stenorhynchus; name 1779 on Official Index]. 

Stenorhynchus Lamarck, 1818:236 [emendation for Stenorynchus 

by the International Commission of Zoological Nomenclature 

under its plenary powers (Opinion 763, 1966); typespecies: 

Cancer seticomis Herbst, 1788, by designation by 

the International Commission on Zoological Nomenclature 

under its plenary powers (Opinion 763, 1966:19); 


* Stenorhynchus lanceolatus (Brulle, 1837) 

FIGURE 784 

Leptopodia lanceolata Brulle, 1837, unnumbered pi.: fig. la,b. 

Leptopodia Canariensis Brulle, 1839:15. 

Leptopodia sagittaria.—Bru\le, 1839:15.—White, 1847a: 1 

[part].—Stimpson, 1858d:219.—Kingsley, 1880b:383 

[part].—Miers, 1886:3, 4.—Osorio, 1887:221; 1888:187, 

191.—Koelbel, 1892:114.—Osorio, 1898:187, 192.—A. 

Milne Edwards and Bouvier, 1900:153—Stimpson, 1907: 


23.—Balss, 1922:72. [Not Cancer Sagittarius Fabricius, 1793 

= Cancer seticomis Herbst, 1788.] 

"une nouvelle espece de ce genre" [Leptopodia].—Guerin- 

Meneville, 1844:10. 

Leptopodia Sagittarius.—Herklots, 1861:136 [part]. 

Leptopodia vittata (Guerin MS).—Kingsley, 1880b:384. 

Leptopodia sagitaria.—Osorio, 1889:130; 1898:185. 

Stenorynchus Sagittarius.—Rathbun, 1900a:293.—Stimpson, 

1907:23. 

Stenorhynchus Sagittarius.—Odhner, 1923:19. 

Stenorynchus seticomis.—Rathbun, 1925:13 [part, not pi. 2, 3]. 

[Not Stenorhynchus seticomis (Herbst, 1788).] 

Stenorhynchus seticomis.—Monod, 1933b:503.—Capart, 1951: 

81, fig. 25.—Sourie, 1954b: 147.—Monod, 1956:567, figs. 

838, 839.—Rossignol, 1957:78, 115 [key].—Longhurst, 

1958:89.—Gauld, 1960:72.—Guinot and Ribeiro, 1962: 

79.—Rossignol, 1962:123.—Ribeiro, 1964:21.—Crosnier, 

1964:38, fig. on pi. A.—Forest and Guinot, 1966:117.— 

Maurin, 1968b:484, 486.—Le Loeuff and Intes, 1968, 

table 1; 1969:65.—Uschakov, 1970:455 [listed]. 

Stenorynchus.—Voss, 1966:17, 19. 

Stenorhynchus lanceolatus.—Yang, 1967:220.—Barr, 1975:47 

[discussion]. 




Foraminifera, 1


Stenorhynchus (S. seticornis, S. lanceolatus, and Stenorhynchus 

species A), gave the differences between 

the latter two species in the following tabular 

form: 

Stenorhynchus species A 

Propodite of female chela 

smooth 

Hiatus of female chela 

narrow 

Meropodite of male chela 

strongly granulated 

S. lanceolatus 

Propodite of female chela 

covered by fine spines 

Hiatus of female chela 

round; similar to S. 

seticomis 

Meropodite of male chela 

smooth 

The differences between S. seticornis (Herbst) 

and S. lanceolatus were discussed by Yang (1967: 

221) as follows: 

S. seticomis is also different from S. lanceolatus in having 

three spines at the distal margin of the meropodite (rather 

than two). Also S. seticomis is distinguished from 5. lanceolatus 

by having smooth propodite of the female chelae, and 

strongly granulated meropodite of the male chelae. The tip 

of the male first pleopod of S. lanceolatus is round without a 

tooth-like projection and resembles that of S. species A 

(based on one large specimen). However, the hiatus of the 

female chela and the spination of pereiopod carpodite of 

Stenorhynchus lanceolatus is very similar to that of 5. seticomis. 

The future larval study of S. lanceolatus will show the affinity 

of this species to Stenorhynchus sp. A or 5. seticomis. 


figs. 838, 839. 

Color: Koelbel (1892:115) mentioned "tief 

purpurbraunen, manchmal auch lauchgriinen 

Riickenstreifen" and noted in a footnote 

"Vermoge eines ausgepragt griinlichen 

Schimmers entspricht die Farbung derartiger 

Streifen vollstandig jener der Fliigeldecken der 

Mannchen von Dyticus marginalise He described 

the background of the stripes as "Elfenbeinweiss." 

A. Milne Edwards and Bouvier (1900: 

154) described the color of the species as follows: 

Les couleurs de ce Crabe sont disposees d'une facon tres 

remarquable. Une serie de bandes brunes, jaunatres ou 

presque blanches, s'etendent longitudinalement; elles se reunissent 

en avant sur la ligne mediane, puis se dirigent 

en arriere en suivant les contours de la carapace, les plus 

longues etant en dehors, les plus courtes au centre et la 

region cardiaque pouvant etre consideree comme le centre 

de ces lignes concentriques. Les pattes sont brunes, relevees 

de taches et de bandes longitudinales d'un jaune vif. Les 

doigts des pinces sont d'un bleu violace. 

Capart (1951:82) described the color as follows: 

"La carapace de couleur orange, avec bande 

longitudinale violette, les pattes et pinces marquees 

de bandes de memes couleurs." 

Male Pleopod: Monod, 1956, fig. 839 (Sierra 

Leone). 

MEASUREMENTS.—The carapace lengths of the 

present male specimens ranged from 18 to 24 

mm, in the females from 19 to 28 mm (the same 

range was shown by the ovigerous females). In 

the literature males have been reported with cl 

18.2-54.5 mm (Guinot and Ribeiro, 1962), 62.5 

mm (Miers, 1886), 71 and 75 mm (Rossignol, 

1957) and 72 mm (Capart, 1951); A. Milne Edwards 

and Bouvier (1900) gave the span of their 

males as 200 to 250 mm. Non-ovigerous females 

with cl 14.4-49.5 mm and ovigerous females with 

cl 36.9, 40.2, and 44.2 mm were listed by Guinot 

and Ribeiro (1962). Monod (1956) gave the diameter 

of the eggs as 0.6 to 0.7 mm. 

REMARKS.—Brulle (1837) gave a figure of the 

present species and on the plate used the name 

Leptopodia lanceolata for it. Later the same author 

(Brulle, 1839), now using the name Leptopodia 

canariensis sank it as a synonym of Leptopodia sagittaria 

(Fabricius, 1793) [= Stenorhynchus seticomis 

(Herbst, 1788)]. Practically all later authors accepted 

this synonymy. Only recently Yang (1967) 

showed that not one but three distinct species of 

the present genus have to be distinguished in the 

Atlantic region, one West African and two western 

Atlantic. Yang showed that the correct name 

for the West African species is Stenorhynchus lanceolatus 

(Brulle, 1837) and he indicated the differences 

between the three species. He based the 

West African form largely on part of the Pillsbury 

material listed above. Dr. Won Tack Yang intends 

to publish a more extensive paper on this 

problem, for which reason we only deal briefly 

with it here. 

BIOLOGY.—The species has been collected at 

depths between 6 and 96 m (extremes 2-6 m, 5- 

7 m, 80 m, 75-90 m, and 96 m); more than 85% 

of the catches were from between 20 and 80 m.

306 

The bottom on which the species was found was 

variously reported: sand and rock; sand and shells 

(A. Milne Edwards and Bouvier, 1900). Mud; 

green mud; brown and green mud; sand, mud, 

and rock (Capart, 1951). Coarse shelly sand, bottom 

with Area and Pyura (Sourie, 1954b). Mud 

and sand; shell and sand (Longhurst, 1958). 

Stones; rock (Guinot and Ribeiro, 1962). Mud; 

mud and sand; mud and shells; sand and rock; 

sand, rock, coral, and calcareous algae; calcareous 

algae; calcareous and other algae; sand, calcareous 

and other algae; rock (Forest and Guinot, 

1966). 

Ovigerous females have been reported from all 

months except February (Capart, 1951; Monod, 

1956; Guinot and Ribeiro, 1962; Forest and 

Guinot, 1966; present paper). 

Monod (1956) reported upon a specimen carrying 

a branchial isopod parasite. 

DISTRIBUTION.—Stenorhynchus lanceolatus is an 

eastern Atlantic species, known from Madeira, 

the Canary Islands, the Cape Verde Islands, and 

from numerous localities on the African mainland 

between Spanish Sahara and Angola. The following 

records are in the literature: 

Madeira: No specific locality (White, 1847a; Miens, 1886). 

Off the S side of Madeira 15 fm [27 m] (Stimpson, 1858d, 

1907). 

Canary Islands: E coast of Isla de Gran Canaria and near 

the harbor of Puerto de Cabras, Isla de Fuertaventura 

(Koelbel, 1892). Near Isla de Tenerife, 28°28'N, 18°32'W 

[of Paris, 16°12'W of Greenwich], 80 m, and Estrecho de la 

Bocaina, 28°49'N, 16°13'W [of Paris, 13°53'W of Greenwich], 


Cape Verde Islands: No specific locality (Osorio, 1898). 

Sao Vicente (Miers, 1886; Osorio, 1888). Channel between 

Santo Antao and Sao Vicente, 75-90 m (A. Milne Edwards 

and Bouvier, 1900). Faja di Agua, Brava, 8 and 50 m 

(Guinot and Ribeiro, 1962; Ribeiro, 1964). 


Guinot, 1966). Between Cabo Barbas and Cabo Blanco and 


1968b). 

Mauritania: Bane d'Arguin, 50 m (Maurin, 1968b). 

Senegal: No specific locality (Herklots, 1861; Kingsley, 

1880b). S of He de la Madeleine, 40 m; near Dakar, 

14°41'30"-41"N, 17°18'30"-20'30"W,20-25m; 14°39'30"- 

4C30"N, 17°16'-18°30'W, 18-19 m; 14°38'-39'N, 17° 16.5'n.S'W, 

26 m; between Goree and Dakar, 16 m; area around 


Goree, 33-35 and 41 m; Bane du Seminole, Baie de Goree, 

ca. 38 m; between Cap Manuel and S point of Goree, 24-27 

m; and off Ngaparou (all Monod, 1956). Baie de Dakar, 10- 

12 m (Sourie, 1954b). Rufisque, 14°30'N, 17°25'W, 24 m 

(Capart, 1951). 12°55.5'N, 17°33'W, 65-75 m (Forest and 




Guinea: No specific locality (Uschakov, 1970). 09°44'N, 

13°56'W, 10 m; 09°40'N, 14°21'W, 25 m; 09°16'N, 

13°42'W, 20 m; 09°16'N, 13°34'W, 10 m; 09°07'N, 

13°41'W, 25 m; and 09°N, 13°50'W, 30 m (Monod, 1956). 

09°36'N, 13°57'W, 18-30 m (Forest and Guinot, 1966). Off 

Conakry, 10 m; Matakong, ca. 15 m (Monod, 1956). Between 

lie Tamara and lie Roume, lies de Los, 10-12 m 

(Capart, 1951). 


1958). 08°38'-08°42'N, 8-12 m (Monod, 1956). 

Ivory Coast: 05°05'N, 04°59.5'W, 22 m (Voss, 1966; 

Yang, 1967). Lagoon at Abidjan, 05°16'N, 04°01'20"W 

(Forest and Guinot, 1966). Off Port-Bouet (Monod, 1956). 

Off Grand-Bassam, 15-60 m (Le Loeuff and Intes, 1968). 

Over all of the continental shelf (Le Loeuff and Intes, 1969). 

Ghana: 04°46'N, 02°30'W to 04°45'N, 02°33'W, 61- 

64 m (Voss, 1966; Yang, 1967). Off Takoradi (Gauld, 1960). 

Off Accra, 32-51 m (Monod, 1956; Gauld, 1960). 

Nigeria: No specific locality (Monod, 1956). Off the 

mouths of the Niger River, 04°03'N, 06°12'E, 32 m (Forest 

and Guinot, 1966). Off the Niger delta, 04°03'N, 05°41'E to 

04°07'N, 05°40'E, 33 m, and 04°04'N, 06°18'E to 04°05'N, 

06°22'E, 30 m (Yang, 1967). 

Cameroon: No specific locality, in 30-50 m (Crosnier, 

1964). 

Principe: 01°43'10"N, 07°28'20"E, 73 m; 01°43'N, 

07°28'55"E, 37 m; 01°38'35"N, 07°21'35"E, 35 m; 

01°35'N, 07°28'E, 45 m (Forest and Guinot, 1966). 

Sao Tome: No specific locality (Osorio, 1887, 1889, 

1898; Balss, 1922). Praia da Fernao Dias (Osorio, 1888). 

00°25'40"N, 06°40'10"E, 50 m; OO°25'15"N, 06°43'05"E, 

8-30 m; Morro Peixe, 2-6 m (Forest and Guinot, 1966). 

Annobon: No specific locality, 18-20 m (Capart, 1951). 

N of San Antonio, 23 m (Forest and Guinot, 1966). 

Congo: Off Pointe-Noire (Monod, 1956; Rossignol, 1957). 

W of Pointe-Noire, 40 m (Rossignol, 1962). 32 mi [51.5 km] 

W of Pointe-Noire, 04°48'S, 1 l°30'E, 50 m; 11 mi [17.7 km] 

WSW of Pointe-Noire, 04°52'S, 1 l°39'30"E, 58-60 m; 8 mi 

[12.9 km] WSW of Pointe-Noire, 04°53'S, 11°53'E, 50-70 m 

(Capart, 1951). 

Cabinda: 05°03'S, 11°24'E, 36-48 m (Capart, 1951). 

Angola: No specific locality (Miers, 1886; Guinot and 

Ribeiro, 1962). 11 mi [17.7 km] W of Cabo Ledo, Luanda, 

09°40'S, 13°02'E, 80 m (Capart, 1951). Benguela, 13-26 m; 

Baia Farta, 30 m; Baia da Caota, 13.5 and 30 m (Guinot


and Ribeiro, 1962). Porto Alexandre, Mocamedes (Odhner, 

1923). 

Subfamily MAJINAE Samouelle, 1819 

Genus Maja Lamarck, 1801 

Maja Lamarck, 1801:154 [type-species: Cancer squinado 

Herbst, 1788, by subsequent designation under the plenary 

powers of the International Commission on Zoological 

Nomenclature in Opinion 511; gender: feminine; name 


Meria Griffith and Pidgeon, 1833:165 [erroneous spelling of 

Maja]. 

Paramya de Haan, 1837, pi. 24 [type-species: Pisa {Paramya) 

spinigera de Haan, 1837, by monotypy; gender: feminine]. 

Mamaia Stebbing, 1905:23 [substitute name for Maja; typespecies: 

Cancer squinado Herbst, 1788; gender: feminine]. 

REMARKS.—Guinot and Ribeiro (1962:76, footnote), 

in their account of M. squinado, remarked 

on the status of West African species of Maja as 

follows: "Nous identifions provisoirement ces 

specimens a Maja squinado, mais la systematique 

des Maja ouest-africaines n'est pas entierement 

satisfaisante et fera l'objet d'une mise au point 

ulterieure." To our knowledge these authors have 

not pursued this problem, and, inasmuch as we 

lack material of Maja from tropical localities, we 

have not been able to investigate it ourselves. 

Those working with West African Maja in the 

future should take this into account. 

Maja crispata Risso, 1827 

Maia crispata Risso, 1827:23. 

Maja verrucosa.—Monod, 1956:477 [references].—Zariquiey 

Alvarez, 1968:447, figs. 149b, 150a-f, 156a [Spain; references]. 

SYNONYMS.—Maia verrucosa H. Milne Edwards, 

1834 (see Holthuis, 1977b:72); Cancer majodes 

Nardo, 1847. 

DISTRIBUTION.—Eastern Atlantic, from Portugal 

to Cabo Blanco and the Cape Verde Islands, 

Mediterranean; shallow water, littoral and sublittoral. 

Maja goltziana d'Oliveira, 1888 

Maja goltziana.—Capart, 1951:100, fig. 33 [Gabon].— 

Monod, 1956:478, 632, figs. 644, 645 [Guinea, Sierra 

Leone, Nigeria, Principe].—Rossignol, 1957:116 [key]; 

1962:121 [Congo].—Crosnier, 1967:339, figs. 31, 32 

[Congo, Annobon].—Longhurst, 1958:88 [Sierra Leone]. 

—Zariquiey Alvarez, 1968:447 [Portugal; references].— 

Massuti, 1970:127 [Rio Muni]. 

Maia goltziana.—Le Loeuff and Intes, 1968, table 1 [Ivory 

Coast]. 

Maia goltziana.—Maurin, 1968a: 30, 48 [Morocco, Spanish 

Sahara]; 1968b:484 [Spanish Sahara]. 


Mediterranean and Portugal S to the Congo, 

including Annobon and Principe islands in the 

Gulf of Guinea; sublittoral, in 15-200 m. 

Maja squinado (Herbst, 1788) 

Maja squinado.—Capart, 1951:98, fig. 32 [Spanish Sahara, 

Mauritania].—Monod, 1956:474, figs. 638-643 [Spanish 

Sahara, Mauritania, Senegal, Guinea].—Figueira, 1960: 

11 [Azores].—Guinot and Ribeiro, 1962:75 [Cape Verde 

Islands, Angola].—Ribeiro, 1964:18 [Cape Verde Islands].—Forest 

and Guinot, 1966:95 [Guinea-Bissau].— 

Crosnier, 1967:339 [Gabon; Congo or Gabon].—Monod, 

1967:182, pi. 16: fig. 2 [no localities].—Zariquiey Alvarez, 

1968:446, figs. 149a, 150g,h [Spain; references].—Christiansen, 

1969:131, fig. 54, map 47 [North Atlantic].— 

Kensley, 1970:180 [South-West Africa].—Ribeiro, 1973:6 

[Cape Verde Islands]. 

Maia squinado.—Sourie, 1954b: 147 [Senegal]. 

Maia squinado.—Chapman and Santler, 1955:375 

[Azores].—Forest and Gantes, 1960:356 [Morocco].— 

Gauld, 1960:72 [Ghana].—Maurin, 1968a:48; 1968b:484 

[both Spanish Sahara].—Bas, Arias, and Guerra, 1976: 

169 [Spanish Sahara]. 

Maja squinada.—Massuti, 1970:127 [Rio Muni]. 

SYNONYM.—Maia squinado var. brachydactyla 

Balss, 1922. 


North Sea southward to South-West Africa, including 

the Mediterranean; sublittoral to about 

75 m. 

Subfamily PISINAE Dana, 1851 

Genus Apiomithrax Rathbun, 1897 

Phycodes A. Milne Edwards, 1869:374 [invalid junior 

homonym of Phycodes Guenee, 1852 (Lepidoptera); type-

308 

species: Phycodes antennarius A. Milne Edwards, 1869, a 

subjective junior synonym of Micropisa violacea A. Milne 

Edwards, 1867, by monotypy; gender: masculine]. 

Apiomithrax Rathbun, 1897b: 164 [substitute name for Phycodes 

A. Milne Edwards, 1869; type-species: Phycodes antennarius 

A. Milne Edwards, 1869, by monotypy; gender: 

masculine]. 

Apiomithrax bocagei (Osorio, 1887) 

Micropisa violacea.—A. Milne Edwards and Bouvier, 1900: 

130 [part].—Rathbun, 1925:303, pi. 241: figs. 5-8 

[part].—Capart, 1951:93, fig. 30, pi. 2: figs. 12, 13 [part, 

all but one specimen from Dakar (Monod, 1956:509)]. 

[Not Micropisa violacea A. Milne Edwards, 1867.] 

Apiomithrax bocagei.—Monod, 1956:508, figs. 692-702.— 

Longhurst, 1958:89.—Gauld, 1960:72.—Guinot and Ribeiro, 

1962:76.—Forest and Guinot, 1966:105.—LeLoeuff 

and hues, 1968:46, 70, table 1, figs. 52, 61; 1969:63, 64, 

65. 

Micropisa bocagei.—Rossignol, 1957:78, 116 [key], pi. 1: figs. 

6-9, pi. 3: fig. 2. 

Micropisa (Apiomithras) bocagei.—Rossignol, 1962:122. 

Micropisa spinosa Forest and Guinot, 1966:105 [nomen nudum; 

discussion of manuscript name]. 


Other Material: West Africa: 1906, F. P. Vermeulen, 1


recorded many specimens from localities between 

Senegal and the Congo. Since 1956 it has been 

reported from the following: 


1958). 



off Grand-Bassam, 8-40 m (Le Loeuff and Intes, 

1968). 

Ghana: No specific locality, shore to 20 m (Gauld, 1960). 

Principe: Between Ponta da Mina and Ponta Novo Destino, 


Congo: Pointe-Noire (Rossignol, 1957). Baie de Pointe- 


Angola: No specific locality; Benguela (Forest and 

Guinot, 1966). Luanda, beach seine; Porto Amboim, 40 m; 

Baia de Benguela, shore (Guinot and Ribeiro, 1962). 

* Apiomkhrax violaceus 


Micropisa eryophora De Rochebrune, 1883:167.—Rathbun, 

1900a:294 [listed].—Balss, 1922:73 [listed].—Monod, 

1956:513. 

Herbstia eryophora.—Miers, 1886:49 [listed].—Rathbun, 

1893a:93 [listed]. 

Micropisa violacea.—Capart, 1951:93 [part, not illustrated 

specimen].—Sourie, 1954b: 147, 152.—Rossignol, 1957:78, 

116 [key], fig. 2, pi. 1: figs. 2-5, pi. 3: fig. 1.—Buchanan, 

1958:20, 24.—Crosnier, 1964:31, 32. 

Apiomithrax violaceus.—Monod, 1956:502, figs. 682-691.— 

Longhurst, 1958:89.—Gauld, 1960:72.—Monod, 1963, 

fig. 41 [no material].—Forest and Guinot, 1966:104.—Le 

Loeuff and Intes, 1969:64 [discussion]. 

Micropisa (Apiomithras) violacea.—Rossignol, 1962:122 [erroneous 

spelling]. 

SYNONYMS.—Phycodes antennarius A. Milne Edwards, 

1869; Micropisa eryophora De Rochebrune, 

1883. 


Sta 70, 33 m, branched Foraminifera, Ic5 (L). 

Other Material: Dahomey: Harbor of Cotonou, 26 Mar 

1964, H. Hoestlandt, 1 juv 9 (L). 

Congo: Off Pointe-Noire, 04°46'15"S, 11°48'15"E, 18 m, 

25 Jan 1966, Ombango, A. Crosnier, \6 (W). 

DESCRIPTION.—Monod, 1956:500-501 (comparison 

with A. violaceus). 


Male Pleopod: Monod, 1956, figs. 687-689 (Senegal); 

Rossignol, 1957, pi. 3: fig. 1 (Congo). 

MEASUREMENTS.—Our specimens are all quite 

small, having carapace lengths of 10.5 mm or less. 

Monod (1956) recorded a specimen 50 mm long. 

REMARKS.—The identity of the species described 

by De Rochebrune (1883) as Micropisa 

eryophora, from the mouths of the Casamance and 

Gambia rivers, has remained unsolved until now. 

Miers (1886) placed it in Herbstia and Balss (1922) 

retained it in Micropisa but suggested it could be 

identified with A. violaceus. Monod (1956) listed it 

as ''''Micropisa eryophora" and discussed it after his 

account of Apiomithrax, but did not identify it 

with any West African species then known. 

De Rochebrune's longer French account (1883: 

167, 168), based on a small specimen, 18 mm 

long, 15 mm wide, is as follows: 

Carapace plus longue que large, couverte ainsi que les 

pattes de longs poils grisatres enchevetres; les parties [p. 168] 

inferieures garnies de poils courts, rigides; front bifide, a 

comes courtes et obtuses; bord externe de 1'orbite arme d'une 

dent obtuse, dirigee en dehors; bords lateraux garnis d'epines 

assez longues, molles; trois tubercules saillants disposes sur 

une ligne transversale au niveau de la region gastrique; 

quatre autres epines au centre de la carapace; regions branchiales 

et hepatiques supportant des epines assez fortes, 

terminees par une etranglement surmonte d'un bouton de 

consistance molle; pinces faibles, a doigts ecartes, se touchant 

seulement a leur extremite; pattes ambulatoires assez fortes, 

laineuses. 

Couleur generale brun rougeatre, visibles a travers le 

feutrage des poils; pointe des epines rose, pattes rosees. 

The body proportions of M. eryophora, length 

6/5 width, suggest that it is based upon an Apiomithrax 

rather than a Herbstia or Micropisa; small 

specimens of Herbstia are relatively slender 

(Monod, 1956, figs. 649, 650) and Micropisa ovata 

(Monod, 1956, fig. 669) lacks dorsal spines on the 

midline of the carapace. That the branchial and 

hepatic spines were described as terminating in 

spongy buttons suggests that the specimen was 

covered with a sponge, not that the spines were 

enlarged distally. The mention of long setae on 

the carapace and legs suggests that De Rochebrune 

was dealing with the relatively hairy A. 

violaceus rather than A. bocagei. Finally, the de-

310 

scription of three prominent tubercles arranged 

in a transverse row across the level of the gastric 

region plus four other spines in the center of the 

carapace (not described by De Rochebrune as 

being in a longitudinal row) is precisely the condition 

figured by Monod (1956, fig. 682) for A. 

violaceus. We have no hesitation in identifying 

Micropisa eryophora De Rochebrune, 1883, with 

Micropisa violacea A. Milne Edwards, 1867. 

Although Monod (1956:502, synonymy) indicated 

that the type-locality of Micropisa violacea 

was "St. Vincent du Cap Vert," A. Milne Edwards 

(1867a: 33-35) actually had material from 

two localities, Sao Vicente and "la cote 

d'Angola," and specimens from these two localities 

are syntypes. Monod (1933b:506 and 1956: 

508) examined the specimen from Angola and, 

like A. Milne Edwards, considered it to be conspecific 

with the material from the Cape Verde 

Islands. However, the type-locality of M. bocagei 

also is Angola. In order to insure that the name 

Micropisa violacea will continue to be used in the 

sense originally employed by A. Milne Edwards 

(1867a), we select the specimen figured by him 

on the plate (pi. 21: fig. 1) that accompanied his 

account of the species in volume 1 of Les Fonds de 

la Mer. Apparently the plates that accompanied 

this volume were not issued with it but were 

issued after livraison 24 of volume 2, published in 

1872, i.e., with one of the livraisons of that volume 

issued between 1873 and 1876 (see Rehder, 1946: 

74, for an account of the dates of publication of 

the individual livraisons and some of the figures). 

BIOLOGY.—Like A. bocagei, this species apparently 

prefers soft bottoms in relatively shallow 

water. Although it has been recorded from a 

depth of 110-180 m from the Talisman collections, 

it usually occurs in depths of 35 m or less. The 

single specimen taken by the Pillsbury was collected 

from branched Foraminifera in 33 m. 

Sourie (1954b) in a study of sand habitats, found 

it in Senegal in 10-12 m on bottom with Area and 

Pyura, in coarse shelly sand and on fine shelly 

sand with mud and with Molgula hannensis (Peres), 

2-7 m. In a study of the rocky shore fauna of 

Senegal, Sourie (1954a) included it with the "hy- 


pobiotes lapidicoles mobiles," the "hypobioses detricicoles" 

where it was associated with Sidonops 

senegalensis (Topsent) and Eurythoe complanata (Pallas), 

and as a representative of the "horizons 

moyens et inferieurs" on rocky shores. Buchanan 

(1958) reported that it was abundant in the sandy 

silt community in 8-20 fin (15-36 m) and also in 

the inshore fine sand community in 3-8 fm (6-15 

m) off Accra, Ghana. Monod (1956) recorded a 

few notes on habitat: bottom with Palythoa and 

Molgula on sand in 15-20 m, muddy sand in 10 

m, and specimens collected at the shore, beach, 

or reef. Longhurst (1958) found the species off 

Sierra Leone in sandy mud in estuaries or on 

shelly sand offshore, in depths between 6 and 

25 m. Crosnier (1964) found it off Cameroon in 

0-32 m, in sand with Foraminifera on rocky 

bottom with gorgonians, and characterized it as 

a warm water crustacean. Gauld (1960) noted 

that it was common on rocky shores throughout 

Ghana and was occasionally taken sublittorally. 

Forest and Guinot (1966) recorded the following 

habitats: from mud with Area in 32 m off the 

Niger delta; from rocks, coral, sand, and calcareous 

algae in 3-10 to 35 m off Principe; from 

rocks on shore and from calcareous algae, rocks, 

coral, sand, on an anchor and in a sponge, in 2 to 

10 m off Sao Tome; and from similar bottoms in 

7 to 60 m off Annobon. 

Ovigerous females have been taken in January 

through June and July through September 

(Monod, 1956; Forest and Guinot, 1966). 

DISTRIBUTION.—This species, like A. bocagei, occurs 

off West Africa from numerous localities 

between Cabo Blanco and Angola; in depths to 

110-180 m, generally in less than 35 m; unlike A. 

bocagei, it also occurs off Brazil (Rathbun, 1925). 

Monod (1956) reported on many specimens from 

localities between Mauritania and Gabon; in addition 

to Monod's records, A. violaceus has been 


Senegal: Dakar (Sourie, 1954b). Mouth of the Casamance 

River (De Rochebrune, 1883). 

Gambia: Mouth of the Gambia River (De Rochebrune, 

1883).



1958). 

Ghana: Off Accra, 6-36 m (Buchanan, 1958). No specific 

locality (rocky shores throughout Ghana) (Gauld, 1960). 




Principe: Baia das Agulhas, 4-8 m; 01°38'35"N, O7°2l'- 

35"E, 35 m; Ilheus dos Mosteiros, 3-10 m (Forest and 


Sao Tome: In front of Sao Tome, 8 m; 00°20'N, 06°46'E, 

10 m; Ponta Diogo Vaz; Praia Santa Catarina; Ilheu Macaco; 

off Ponta Diogo Nunes, shore and 4-5 m; in front of 

Ponta Oquedelrei; Morro Peixe; and Ilheu das Cabras, shore 


Annobon: 01°24'04"S, 05°36'45"E, 7-10 m; 01°26'15"S, 

05°35'40"E, 60 m; NW coast, Isla Tortuga, 15-40 m (all 


Congo: Pointe-Noire (Rossignol, 1957). Baie de Pointe- 


REMARKS.—Apparently this species is rare off 

West Africa. Monod (1956) recorded six specimens 

and only two were taken by the Pillsbury. 

Barnard (1950) has recorded this species from 

South Africa and suggested that E. longimana 

Stimpson, 1858, should be considered a synonym 

of/s. aspera. Capart (1951) considered E. longimana 

to be a distinct species, but Griffin (1964, 1974) 

left it in synonymy. Crosnier (1970) agreed with 

Barnard and Griffin and pointed out additional 

similarities between material available to him 

and that reported by Barnard. If those considering 

E. longimana to be a synonym of E. aspera are 

correct, the latter species occurs from Norway 

southward to South Africa. 

BIOLOGY.—Little information is available for 

West African specimens of E. aspera, but apparently 

it prefers a firm substrate with relatively 

Genus Eurynome Leach, 1814 

large particles in it. Chapman and Santler (1955) 

Eurynome Leach, 1814:431 [type-species: Cancer asper Pennant, reported it from rocks in a harbor. Longhurst 



(1958) took one specimen on shelly sand in 100 m 

and Forest and Guinot (1966) also reported one 

specimen from algae and calcareous algae in 

60 m. Our specimens were taken on broken shell 

* Eurynome aspera (Pennant, 1777) 

Eurynome aspera.—Capart, 1951:86, pi. 2: fig. 4.—Chapmanin 

70 m. The overall depth range is wide, from 

and Santler, 1955:375.—Monod, 1956:480, figs. 646- 10 m or less to 550 m (Monod, 1956). 

648.—Longhurst, 1958:89.—Forest and Guinot, 1966: DISTRIBUTION.—Eurynome aspera is widely dis- 

95.—Zariquiey Alvarez, 1968:462, figs. 14g, 153a, 154f 

tributed in the eastern Atlantic, from Norway to 


45 [North Atlantic].—Crosnier, 1970:1217, pi. 1: figs. 1-4 Angola, including the Mediterranean, and, pos- 

[Mediterranean in part].—Tiirkay, 1976a:25 [listed], 39 sibly, from South Africa, in depths between 10 

[Portugal in part]. 

and 550 m. Monod (1956) reported on material 

from Senegal and Principe Island; in addition to 

SYNONYMS.—Eurynome scutellata Risso, 1827; Eu- 

Monod's records the species has been recorded 

rynome boletifera Costa, 1838; tEurynome longimana 

from the following: 

Stimpson, 1858; Eurynome aspera var. acuta A. 

Milne Edwards and Bouvier, 1900. 


Azores: Horta, Faial (Chapman and Santler, 1955). 

Sta 68, 70 m, broken shell, 16", 1 juv (L). 

Morocco: 33°27.7'N, 08°50.8'W, 161-168 m; and 

Undaunted Material: Angola: Sta 96, 162 m, 1

312 

*Eurynome parvirostris Forest and Guinot, 

1966 

Eurynome parvirostris Forest and Guinot, 1966:95, fig. 9. 


Sta 16, 46 m, mud with Foraminifera, shells, 1$ (W). Sta 23, 

42 m, foliate brown to orange bryozoans, 1$ ov (L). 

Other Material: Dahomey: Off Grand-Popo, 30 m, Petersen 

grab, 23 Feb 1964, Guinean Trawling Survey, Tr 34, Sta 2, 

1(5 (L). 

DESCRIPTION.—Forest and Guinot, 1966:95. 

Figure: Forest and Guinot, 1966, fig. 9. 


lengths of 6 mm. The two type-specimens 

had carapace lengths of 6.3 mm each. 

BIOLOGY.—This species may have different 

habitat requirements than E. aspera, for most 

specimens have been taken on soft bottoms: mud 

with Area in 32 m (Forest and Guinot, 1966), and 

mud with Foraminifera and shells in 46 m, or 


mud with foliate bryozoans in 42 m. The known 

depth range is relatively narrow, from 30 to 46 

m. 

DISTRIBUTION.—Off tropical West Africa, 

where it has been recorded from off Dahomey, 

Ghana, and off the mouths of the Niger River, 

04°03'N, 06°12'E, 32 m (Forest and Guinot, 

1966). 

Genus Herbstia H. Milne Edwards, 1834 

Key to Species of Adult Herbstia from West Africa 

Herbstia H. Milne Edwards, 1834:301 [type-species: Cancer 

condyliatus Fabricius, 1787, by monotypy; gender: feminine; 


Rhodia Bell, 1835:169 [type-species: Rhodia pyriformis Bell, 

1835, by monotypy; gender: feminine]. 

Herbstiella Stimpson, 1871b:93 [type-species: Herbstia depressa 

Stimpson, 1860, by original designation; gender: feminine]. 

Fisheria Lockington, 1877:72 [type-species: Fisheria depressa 

Lockington, 1877, a subjective junior synonym of Herbstiella 

camptacantha Stimpson, 1871, by monotypy; gender: 

feminine]. 

1. Posterior median margin of carapace with trilobed projection. [Opposable 

margins of dactyli of walking legs with fixed, triangular teeth] 

H. condyliata 

Posterior median margin of carapace with single projection 2 

2. Carapace, smooth, regions poorly defined. Upper surface of palm in males 

smooth. Opposable margins of dactyli of walking legs with fixed triangular 

teeth H. nitida, new species 

Carapace rugose, regions well defined. Upper surface of palm in males with 

row of tubercles. Opposable margins of dactyli of walking legs lacking 

fixed triangular teeth H. rubra 

* Herbstia condyliata (Fabricius, 1787) 

FIGURE 79 

Cancer condyliatus Fabricius, 1787:324. 

Herbstia condyliata.—Monod, 1933a:212.—Forest and Games, 

1960:356.—Zariquiey Alvarez, 1968:455, fig. 153c,d 

[Spain; references]. 

Herbstia rubra.—Sourie, 1954a: 113, 254, 256, 294; 1954b: 

147.—Chapman and Sander, 1955:375.—Monod, 1956: 

482, figs. 650-653 [part, not fig. 649]. [Not Herbstia rubra 

A. Milne Edwards, 1869.] 

SYNONYMS.—Mithrax herbsti Risso, 1827; Mithrax 

scaber Costa, 1840. 


Sta 22, 51 m, rough bottom, Ic5 (L). 

Other Material: Madeira: SE coast, near Canical, 32°44'N, 

16°44'W, 0-22 m, shore collecting, snorkeling, and diving, 

11 Mar 1976, Orwersaagd Sta 48, \S (L). S coast Ponta da 

Garajau, 32°38'N, lG^l'W, 5-26 m, diving, 17 Mar 1976, 

Orwersaagd Sta 111, 1$ (L). 

DESCRIPTION.—Carapace (Figure 79a,c,d) pyriform, 

irregular dorsally, ornamented with numerous 

spines, regions well marked; surface covered 

with thin coat of low, dark setae. Gastric, cardiac, 

and intestinal regions each elevated in midline.


a 

FIGURE 79.—Herbstia condyliata (Fabricius). Juvenile, cl 4.2 mm, Sicily: a, carapace; b, dactylus 

of fifth pereiopod. Male, cl 10.8 mm, Spain: c, carapace. Male, cl 29.6 mm, Pillsbwy Sta 22: d, 

carapace; e, dactylus of fifth pereiopod; /, abdomen; g, apex of gonopod. 

Gastric region with 3 raised protogastric tubercles 

in transverse line anteriorly (Figure 79

314 

podus, ventral margin with slender spines and 

usually with 1 or 2 triangular teeth (Figure 19b,e), 

large in juvenile, smaller in adult. 

Male gonopod and abdomen as shown (Figure 

79/*). 


7.8 to 29.6 mm, of female 18 mm. 

REMARKS.—Herbstia condyliata can readily be 

distinguished from other West African species of 

the genus, H. rubra and H. nitida, new species, by 

its larger size, much more spinulose carapace, and 

presence of a trilobed projection on the posterior 

margin of the carapace (as shown in Monod, 

1956, fig. 650, and herein, Figure *19d), where it 

is almost quadrilobed. The chelipeds in adult 

males of H. condyliata are greatly enlarged, being 

much longer than the carapace; in the two 

smaller West African species the chelipeds in 

adult males are subsimilar to those of females. 

Herbstia condyliata agrees with H. nitida, new species, 

and differs from H. rubra in having fixed, 

triangular teeth on the opposable margin of the 

dactyli of the walking legs. 

Because smaller specimens of this species have 

been confused with the adults of the two smaller 

West African species, we have sketched the carapace 

shape in a juvenile, cl. 4.2 mm (W), from 

Sicily (Figure 79a) and a larger specimen from 

Spain, cl. 10.8 mm (L) (Figure 79c), as well as an 

outline of our adult from Ghana (Figure 79d). In 

the smallest specimen, the carapace is very slender, 

its dorsal armature is reduced, and there is 

but one posterior spine. In the specimen from 

Spain the characteristic three posterior spines of 

the adult are developed, although the spinulation 

of the carapace is still reduced. Monod (1956, fig. 

650) figured a female 15.5 mm long, in which the 

carapace is still more slender than in the adult 

but in which the posterior spines of the carapace 

are well developed. In Monod's figure, the transverse 

line of three protogastric tubercles is not 

shown. 

A. Milne Edwards and Bouvier (1900:128) suggested 

that H. rubra might only be a race of H. 

condyliata and Monod (1956:485) more or less 

agreed with this: 


II n'est nullement impossible que l'etude de materiaux 

plus nombreux, et provenant en particulier des regions marocaine 

ou saharienne, n'oblige a tenir un jour rubra pour 

synonyme de condyliata, ou tout au moins comme une forme 

subordonnee, moins tuberculee, et aussi plus petite. 

Monod was working with juveniles of H. condyliata 

as well as adults of both other species, and, 

in view of the similarities between specimens of 

similar size of all three species, it is not surprising 

that they were not separated by him. 

Although it is impossible to be certain in view 

of the relatively small number ofHerbstia recorded 

from West Africa and the relatively unreliable 

literature records, it seems likely that H. rubra is 

restricted to the Cape Verde Islands, H. nitida, 

new species, occurs only on the offshore islands of 

the Gulf of Guinea, and H. condyliata occurs on 

the continental shelf. 

The male pleopod of H. condyliata, the typespecies 

of the genus, is shown in Figure 19g. It 

resembles that shown by Garth (1958, pi. S: figs. 

7, 8) for the eastern Pacific species H. pubescens 

Stimpson and H. pyriformis (Bell) in that there is 

a subapical rounded, spinulose knob flanked by 

a distinct notch; this knob is present on the 

gonopods of all three species reported here. As 

noted by Garth (1958:300) in his key to West 

American Herbstia, the other western American 

species lack the subapical notch on the male 

pleopod. Further study may well indicate that 

Herbstia, as currently recognized, comprises representatives 

of more than one genus. 

BIOLOGY.—Sourie (1954b) found this species 

on coarse, shelly sand, with mud, bottom with 

Area and Pyura, in 2-7 m, at Anse de Hann, Baie 

de Dakar, and subsequently included the species 

as a component of the rocky shore fauna of 

Senegal. Monod (1933a:212) reported two specimens 

associated with sponges, and, in 1956, reported 

a specimen collected during a spring low 

tide. The single specimen taken by the Pillsbury 

was taken on rough bottom at a depth of 51 m. 


Atlantic south to Ghana. Since 1956 it has 

been recorded from Morocco: Temara, Skhirat, 

and David (Forest and Games, 1960).


Barrois (1888) recorded the species from the low tubercles and several larger spines arranged 

Azores, and also indicated, without citing a ref- in dorsal and ventral rows, spination variable, 

erence, that the species occurred in the Canary absent in some specimens; carpus with numerous 

Islands. The specimens reported from Horta and tubercles and 1 larger spine on outer margin. 

Pasteleiro, Faial, Azores, by Chapman and San- Palm not markedly inflated in males, with 1 or 2 

der (1955) as H. rubra may be referable to H. (usually 1) posteriorly directed spines on dorsal 

condyliata; H. rubra is known with certainty onlyarticular 

knob, remainder of surface smooth but 

from the Cape Verde Islands. 

setose. Fingers shorter than palm, cutting edges 

crenulate, not strongly toothed or gaping. 

*Herbstia nitida, new species 

Walking legs slender, long, fifth leg longer than 

carapace, ornamented with corneous bristles and 

FIGURE 80 

longer simple setae. Merus with row of corneous 

bristles dorsally, occasionally with dorsal and pos- 

?Herbstia rubra.—Monod, 1956:482 [part].—Forest and 

Guinot, 1966:97. [Not Herbstia rubra A. Milne-Edwards, 

1869.] 

terior spines or with irregular tubercles posteriorly, 

not heavily armed. Dactylus (Figure 80/) 

shorter than propodus, ventral margin with 1-4 

MATERIAL EXAMINED.—Pillsbury Material: Annobon: prominent, triangular teeth and slender spines. 

Sta 275, 9-69 m, rubble of coralline algae, 3c5, 3$ (2 ov), 1 Male abdomen and gonopods as shown (Figure 

juv (W). Sta 282, 18-37 m, nodular coralline algae, 3

316 

a 


FIGURE 80.—Herbstia nitida, new species, paratype, male, cl 9.0 mm, Pillsbury Sta 282: a, dorsal 

view; b, carapace; c, front, dorsal view; d, front, ventral view; e, chela; /, dactylus of fifth 

pereiopod; g, abdomen; h, apex of gonopod. 


D.31780), a male with a carapace length of 8.9 

mm from Pillsbury Sta 282, is in the Rijksmuseum 

van Natuurlijke Historie, Leiden. Paratypes have 

been deposited in the Smithsonian Institution, 

Washington, D.C., and in Leiden. 

BIOLOGY.—All of our material came from Annobon 

Island, in depths between 9 and 55 m, in 

the beds of calcareous algae described by Forest 

(1959) and Voss (1966). If our assumption is 

correct that the material from the offshore islands 

of the Gulf of Guinea previously reported as H. 

rubra actually is identifiable with H. nitida, the 

latter species lives on rough bottom in moderate 

depths on rock, coral, algae, and calcareous algae. 

Forest and Guinot (1966) recorded the following 

habitats and depths (all under H. rubra): Principe, 

calcareous algae in 37-73 m; Sao Tome (6 localities), 

in rocks and coral in 0-6 and at 30 m, in 

calcareous algae in 6 and 10 m, in a sponge from 

rocks and calcareous algae in 2-6 m; and Annobon 

(5 localities), in rocks and coral, in calcareous 

algae, sand, and coral in 7-10 m, and in calcareous 

algae in 23-60 m. 

Ovigerous females have been recorded in May 

and June (Forest and Guinot, 1966; Pillsbury). 


Latin nitidus, shining, neat, referring to the 

smooth carapace of this species. 

DISTRIBUTION.—Gulf of Guinea, where it is 

known to occur off Annobon in 15-55 m. We 

9


suspect that all of the records of H. rubra given 

below are based on H. nitida: 

Principe: No specific locality, 15 m (Monod, 1956). 01°- 

43'10"N, 07°28'20"E, 73 m, and 01°43'N, 07°28'55'E, 37 m 


Sao Tome: Off Sao Tome, 8 m; 00°20'N, 06°46'E, 10 m; 

Ponta Diogo Vaz, 0-6 m and 30 m; in front of Ponta 

Oquedelrei, 6 m; and Morro Peixe, 2-6 m (all Forest and 


Annobon: No specific locality, 12 m (Monod, 1956). 

01°24'04"S, 05°36'45"E, 7-10 m; O1°27.5'S, 05°36.5'E, 

35 m; N of San Antonio, 23 m; 01°26'15"S, 05°35'40"E, 60 

m; and Isla Tortuga, NW coast (all Forest and Guinot, 

1966). 

Herbstia rubra A. Milne Edwards, 1869 

FIGURE 81 

Herbstia rubra A. Milne Edwards, 1869:354.—Miers, 1886:49, 

pi. 7: fig. 1,1a.—Rathbun, 1893a:93 [listed].—A. Milne 

Edwards and Bouvier, 1900:128, pi. 19: fig. 17 [fig. 16 in 

text].—Balss, 1922:73 [listed].—Monod, 1956:482, fig. 649 

[part, not fig. 650].—Guinot and Ribeiro, 1962:73.—Ribeiro, 

1964:19. 


Other Material: Cape Verde Islands: Sao Vicente, 20 m, 26 

Jul 1883, Talisman, \6, 1$ (L, W). Sao Vicente, Challenger, 

BMNH 84.31, 1


which the chelipeds of the male are not greatly 

enlarged and there is but one posterior projection 

on the carapace. This species differs from H. 

nitida, new species, in having the regions of the 

carapace better developed, more dorsal tubercles 

and spines on the carapace, a narrower posterior 

projection on the carapace, more spines on stouter 

walking legs, a row of spines on the upper surface 

of the palm in males, and no triangular teeth on 

the opposable margin of the dactylus of the walking 

legs. 

An outline of the carapace and a sketch of the 

dactylus of a walking leg are shown in Figure 81. 

BIOLOGY.—There is little information available 

on the ecology of this species. It was reported 

from coralline algae in 75 m (A. Milne Edwards 

and Bouvier, 1900), and near shore and in 6 to 

20 m (Guinot and Ribeiro, 1962; Ribeiro, 1964). 


Verde Islands (the type-locality) and possibly the 

Azores; most records in the literature require 

verification. They include the following: 

Cape Verde Islands: No specific locality (A. Milne Edwards, 

1869). Sao Vicente (Miers, 1886). Praia Matiota, Sao 

Vicente, shore, and Porta da Furna, Brava, 6-20 m (Guinot 

and Ribeiro, 1962; Ribeiro, 1964). Between Ilheu Branco 

and Ilheu Raso, 75 m (A. Milne Edwards and Bouvier, 

1900). Ponta do Sol, Ilha de Santo Antao (Monod, 1956). 

Genus Micropisa Stimpson, 1858 

Micropisa Stimpson, 1858a:218 [type-species: Micropisa ovala 

Stimpson, 1858, by monotypy; gender: feminine]. 

Micropisa ovata Stimpson, 1858 

Micropisa ovata.—Monod, 1956:495, figs. 669-681 [Cape 

Verde Islands, Senegal, Gabon(?)].—Guinot and Ribeiro, 

1962:74 [Cape Verde Islands].—Ribeiro, 1964:19 [Cape 


DISTRIBUTION.—Off tropical West Africa, from 

the Cape Verde Islands and Senegal, with one 

doubtful record from Gabon; sublittoral, from 

less than 10 to more than 110 m. 

Genus Pisa Leach, 1814 

Arctopsis Lamarck, 1801:155 [type-species: Arctopsis lanata 

Lamarck, 1801, by monotypy; gender: feminine; sup- 

pressed for purposes of the Law of Priority but not for 

those of the Law of Homonymy by the International 

Commission on Zoological Nomenclature in opinion 708 

(1964); name 1701 on Official Index]. 

Pisa Leach, 1814:431 [type-species: Cancer biaculeatus Montagu, 

1813, a subjective junior synonym of Maja armata 

Latreille, 1803, by monotypy; gender: feminine; name 

1597 on Official List}. 

Blastus Leach, 1814:431 [type-species: Cancer tetraodon Pennant, 

1777, by monotypy; gender: masculine]. 

*Pisa armata (Latreille, 1803) 

Maja armata Latreille, 1803c:98. 

Pisa Gibbsii.—Studer, 1882:335. 

Pisagibbsi.—Capart, 1951:90, fig. 28, pi. 2: fig. 23.—Monod, 

1956:486, fig. 654.—Rossignol, 1957:116 [key].—Figueira, 

1960:12.—Crosnier, 1964:34. 

Pisa armata.—Forest and Gantes, 1960:356.—Forest and 

Guinot, 1966:98.—Zariquiey Alvarez, 1968:454, figs. 

151d, 152e, 154d [Spain; references].—Christiansen, 1969: 

124, fig. 51, map 44 [North Atlantic].—Crosnier, 1970: 

1215 [listed], 1218.—Turkay, 1976a:25 [listed], 39 [Portugal]. 

Pisa.—Voss, 1966:31. 

Pisa gibbi.—Maurin, 1968b:486 [erroneous spelling]. 

SYNONYMS.—Cancer biaculeata Montagu, 1813; 

Pisa gibbsii Leach, 1815. 


Sta 230, 82-97 m, hard ground with gorgonians, coral, and 

rock, 1


habits moderate depths on rough ground on the 

shelf. The Pillsbury specimen was taken on hard 

ground with gorgonians, coral, and rock in 82- 

97 m. Capart (1951) reported material collected 

at depths from 20-30 to 110 m, on sandy mud, 

that from 20-30 m being the shallowest record 

off West Africa. Crosnier (1964) characterized it 

as a cold-water species, occasionally occurring 

deeper than 50 m off Cameroon. Forest and 

Guinot (1966) recorded it from mud, sand, and 

"sable construit" in 65-75 m off Senegal and on 

mud, shell, and Cidaris in 60-73 m off Guinea- 

Bissau. Maurin (1968b) found it off Morocco in 

40-60 m on bottom with sponge, Suberites, and 

asteroid, Allopatiria. 

Ovigerous females have been recorded from 

West African localities in March, May, August, 

October, November (Capart, 1951; Monod, 

1956), and summer (Figueira, 1960). 


southern North Sea and England to Angola, 

Mediterranean; sublittoral, to at least 162 m. 

Monod (1956) reported material from Senegal, 

Guinea, and Angola; other records include the 


Azores: Horta (Figueira, 1960). 

Morocco: Temara (Forest and Games, 1960). 

Mauritania: Bane d'Arguin, 40-60 and 90-100 m 


Cape Verde Islands: 15°40'N, 23°06'W, 38 fm [70 m] 

(Studer, 1882). 





Nigeria: 06° 1 l'N, 03°36'E to 06° 10'N, 03°38'E, 82-97 m 

(Voss, 1966). 

Cameroon: No specific locality, more than 50 m (Crosnier, 

1964). 

Angola: 16°41'S, 11°21'E, 162 m (Crosnier, 1970). 

The latter record appears to be the southernmost 

as well as the deepest for the species. 

*Pisa calva Forest and Guinot, 1966 

Une espece nouvelle de Pisa.—Forest, 1959, pi. 3: fig. 1. Pisa 

(n. sp.).—Forest, 1959:19. 

Pisa calva Forest and Guinot, 1966:99, figs. 10, lla-f, 13. 


Sta 275, 9-69 m, rubble of coralline algae, 66, 4$ (3 ov) (L). 

Sta 282, 18-37 m, nodular coralline algae, 16*, 7$ (5 ov), 2 

juv (L, W). Sta 283, 51-55 m, nodular coralline algae, 

116\ 89 (5ov), 3juv(W). 

Other Material: Annobon: 01°24'S, 05°37'30"E, 7-8 m, 11 

Dec 1965, Ombango, A. Crosnier, 29 (W). 

DESCRIPTION.—Forest and Guinot, 1966:100. 

Figures: Forest and Guinot, 1966, figs. 10, 

lla-f, 13. 

Male Pleopod: Forest and Guinot, 1966, fig. 13. 

MEASUREMENTS.—Carapace lengths of males 5 

to 11 mm, of females 5 to 8 mm, of ovigerous 

females 5 to 10 mm, of juveniles 3 to 5 mm. The 

largest specimens recorded by Forest and Guinot 

(1966) were a male 14 mm long and a female 10.5 

mm long. 

REMARKS.—As Chace (1966:653, footnote) 

pointed out, this species is very close to Pisa 

sanctaehelenae Chace (1966:651, fig. 14) from Saint 

Helena. However, P. calva has less divergent rostral 

spines and much more slender pereiopods 

(Figure 82), with 10 or 11 rather than 5-7 fixed, 

triangular teeth on the flexor margin of the dactylus. 

In our specimens, the propodus of the sec- 

FIGURE 82 

FIGURE 82.—Pisa calva Forest and Guinot, male, cl 11.0 mm, 

Pisa carinimana.—Monod, 1956:488 [part] [not Pisa carinimana Pillsbury Sta 283: a, second pereiopod; b, dactylus of second 

Miers, 1879]. 

pereiopod, enlarged.

320 

ond pereiopod in the males is not markedly swollen 

distally as in P. sanctaehelenae. 

Apparently Monod's (1956) record of P. carinimana 

from Annobon was based on this species 

(Forest and Guinot, 1966:104) and his record of 

P. carinimana from Principe may also be based on 

P. calva. However, Forest and Guinot, who reexamined 

Monod's material from Annobon, could 

not locate the specimen from Principe and they 

noted (1966:104) that both species were taken at 

the same station off Principe by the Calypso. 

BIOLOGY.—The Pillsbury representatives of this 

species all were taken off Annobon Island in beds 

of coralline algae in 9-69 m; the Ombango specimens 

were taken at Annobon in 7-8 m. Forest 

and Guinot (1966) recorded the following habitats 

for their material: Principe (5 stations): on 

mud, calcareous algae, shell, rocks, and coral in 

31-73 m; Sao Tome (12 stations): from a beach 

seine in 0-4 m and in rocks, coral, calcareous 

algae, "epave," algae, sand, and mud in 5-50 m; 

Annobon (6 stations): calcareous algae, sand, 

coral, algae, and rocks in 7-60 m. 


June, and July. 

DISTRIBUTION.—West Africa, from the offshore 

islands of the Gulf of Guinea; sublittoral, from 0- 

4 to 73 m. Records in the literature include the 

following: 

Principe: 01°38'25"N, 07 o 22 / 05 w E, 31 m; 01°43'10"N, 

07°28'20"E, 73 m; 01°43'N, O7°28'55"E, 37 m; in front of 

Baia de Santo Antonio, 50 m; Tinhosa Grande Island, 12 mi 

[19.3 km] SSW of Principe, 01°20'45"N, 07°17'37"E, 25-40 

m (all Forest and Guinot, 1966). 

Sao Tome: In front of Sao Tome, 8 m; 00°20'N, 06°46'E, 

10 m; N of Ilheu das Cabras, 32 m; Ponta Diogo Vaz, 30 m; 

Praia Santa Catarina, 15-18 m; 00°25'40"N, 06°40'10"E, 

50 m; 00°25'15"N, 06°43'05"E, 8-30 m; Baia de Ana de 

Chaves, 5 m; in front of Ponta Oquedelrei, 6 m; Morro 

Peixe, 0-4 m; in front of Ponta Sao Sebastiao, 11 m (all 


Annobon: No specific locality, 12 m (Monod, 1956).— 

NW coast, Isla Tortuga, 15-40 m; N of Santo Antonio, 

23 m; 01°27.5'S, 05°36.5'E, 35 m; 01°26'15 w S, O5°35'4O"E, 

60 m; 01°25'10"S, 05°36'10"E, 20-25 m; 01°24'04"S, 

05°36'45"E, 7-10 m (all Forest and Guinot, 1966). 


*Pisa carinimana Miers, 1879 

Pisa carinimana.—Capart, 1951:87, fig. 27, pi. 2: fig. 8.— 

Sourie, 1954b: 147.—Monod, 1956:488, figs. 655-668 

[part].—Rossignol, 1957:116 [key].—Longhurst, 1958: 

89.—Forest, 1959:19 [discussion].—Gauld, 1960:72.— 

Rossignol, 1962:122.—Guinot and Ribeiro, 1962:74.— 

Forest and Guinot, 1966:99, figs, llg, 12.—Zariquiey 

Alvarez, 1968:452, fig. 155a [Spain; references].—Uschakov, 

1970:439, 455 [listed].—Tiirkay, 1957a: 71 

[listed], 74. 


Sta 69, 29 m, coral or rock, 1$ ov (L). Sta 70, 33 m, branched 

Foraminifera, 86, 8$ (7 ov) (W). 


Foraminifera, 16 (W). Sta 46, 38-42 m, mud with dense 

Jullienella, 4


100 m off Ghana. The Pillsbury collected this 

species in depths between 29 and 82 m, usually 

on relatively soft bottom, sand and mud, or mud 

with bryozoans or Foraminifera; two stations 

were made on rock and coral. Monod (1956) 

recorded material collected on bottom with Palythoa 

and Molgula and sand in 15-20 m, on sand in 

10-45 m, muddy sand in 10-15 m, and on pebbles 

in 14.5 m. Guinot and Ribeiro (1962) reported 

material from Angola in depths between 5 and 

90 m on sand and muddy sand. Forest and Guinot 

(1966) noted the following habitats for the species: 

mud, shell, gorgonians, and ascidians in 43- 

45 m; mud, sand, and compact sand [sable construit] 

in 65-75 m; mud and shells in 18-30 m; 

muddy sand and Foraminifera in 21-27 m; 

gravel, shells, and Foraminifera in 50 m; mud 

with Area in 32 m, and mud, calcareous algae, 

and shells in 31 m. This species was taken with P. 

calva by the Calypso at this latter station off Principe 

Island. Uschakov (1970) found the species in 

clear water on hard sand in depths greater than 

10 m off Guinea. Apparently this species prefers 

habitats on softer bottoms than does P. calva, 

which the Pillsbury took only in coralline algae. 

Ovigerous females have been recorded in all 

months but July (Monod, 1956; Forest and 


DISTRIBUTION.—Pisa carinimana is an eastern Atlantic 

species, known from localities between the 

Canary Islands (the type-locality) and Spanish 

Sahara southward to Angola (12°36'S), including 

Melilla in the Mediterranean (Zariquiey Alvarez, 

1968); littoral and sublittoral to a depth of about 

100 m. Monod (1956) recorded it from numerous 

localities between Mauritania and Gabon; other 

records include the following: 



Mauritania: Off Cap Blanc, 20°37.3'N, 17°24.4'W, 57 m 

(Tiirkay, 1975a) 

Senegal: Baie de Dakar, 10-12 m (Sourie, 1954b). 

12°55.5'N, 17°33'W, 65-75 m (Forest and Guinot, 1966). 

Guinea: No specific locality, depth greater than 20 m 

(Uschakov, 1970). 09°40'N, 14°05'W, 18 m, and 09°36'N, 

13°57VJ, 18-30 m (Forest and Guinot, 1966). 


1958). 

Ivory Coast: 05°02.5'N, 05°25'W, 21-27 m (Forest and 

Guinot, 1966) 

Ghana: Off Accra, 14-100 m (Gauld, 1960). 04°36.5'N, 

0l°31'W, 50 m (Forest and Guinot, 1966). 



Principe: No specific locality (Forest, 1959). 01°38'25"N, 


Gabon: W of Pointe Claire (as Pointe Clara) and mouth 

of Gabon River, 20-40 m (Rossignol, 1962). 


Cabinda: No specific locality, in 38-39 m (Guinot and 


Angola: Baia da Caota, Benguela, 11-12 m, 13 m, 16 m, 

18 m, and 30 m; Baia Farta, 22 m, 22-28 m, and 90 m; 

Sombreiro, 5 m; between Sombreiro and Ponta da Caruita, 

23 m (all Guinot and Ribeiro, 1962). 

Pisa nodipes (Leach, 1815) 

Pisa nodipes.—Monod, 1956:486 [references].—Forest and 


454, figs. 151e, 152f, 154e, [Spain; references].—Maurin, 

1968a:59, 107 [Mauritania, Mediterranean]; 1968b:486, 

489 [Mauritania]. 

SYNONYM.—Inachus musivus Otto, 1821. 

DISTRIBUTION.—Eastern Atlantic, from the Canary 

Islands, the Cape Verde Islands, Morocco, 

Mauritania, and the Mediterranean; sublittoral, 

to about 75 m. 

Pisa tetraodon (Pennant, 1777) 

Pisa Mraodon.—Caparx, 1951:92, fig. 29, pi. 2: fig. 16 [Mauritania].—Monod, 

1956:485 [references].—Forest and 


452, figs. 6d, 151a, 152a, 154a [Spain; references]. 

SYNONYMS.—Cancer hircus Fabricius, 1781 (see 

Rathbun, 1925:195); Cancer praedo Herbst, 1796; 

Pisa convexa Brandt, 1880. 


to Cabo Blanco, Mauritania, Mediterranean; 

sublittoral, shallow water to about 50 m 

(100 m in Monod, 1956).

322 

Family MIMILAMBRIDAE Williams, 1979 

MIMILAMBRIDAE Williams, 1979:399. 

This family, comprising one genus and species, 

does not occur within the study area. 

Family PARTHENOPIDAE MacLeay, 1838 

PARTHENOPINA MacLeay, 1838:55, 58 [corrected to Parthenopidae 

by Bell, 1844:45; name 362 on Official List]. 

AETHRINAE Dana, 1851b: 127 [originally cited as Oethrinae]. 

CRYPTOPODIINAE Stimpson, 1871a: 137. 

HEPATINAE Stimpson, 1871a: 154 [see p. 325]. 

EUMEDONINAE Neumann, 1878:17. 

LAMBRINAE Neumann, 1878:17. 

EASTERN ATLANTIC GENERA.—Five, Daldorjia, 

Heterocrypta, Parthenope, Sakaila, new genus, and 

Solenolambrus, each represented by species occurring 

off tropical West Africa. 

EASTERN ATLANTIC SPECIES.—Eleven, of which 

eight occur off West Africa. Several name changes 

have been made since this family was studied by 

Monod (1956), as follows: 

Name in Monod 

Osachila stimpsoni 

(Calappidae) 

Lambrus massena (including 

various forms) 

Lambrus miersi 

Lambrus macrochelos 

Lambrus expansus 

Heterocrypta maltzani 

Solenolambrus noordendei 

Parthenope bouvieri 

Current Name 

Sakaila africana, 

new genus, new species* 

(Parthenopidae) 

Parthenope massena* 

Parthenope miersii* 

Parthenope notialis, 

new species* 

Parthenope expansa* 

Heterocrypta maltzami* 

Solenolambrus noordendei* 

Daldorjia bouvieri 

Three nominal species occur to the north of the 

tropical region: 

Heterocrypta marionis A. Milne Edwards, 1881. 

In deep water off the Azores, Toulon, France, and 

the Bay of Biscay (Bouvier, 1940). 

Parthenope angulifrons Latreille, 1828. Mediterranean 

(Zariquiey Alvarez, 1968), with one questionable 

record from Senegal (Monod, 1956:587). 

Parthenope macrochelos (Herbst, 1790). Mediter- 

ranean and adjacent Atlantic (Zariquiey Alvarez, 


1968). See under P. notialis, new species (p. 335), 

for possible records along the NW African coast. 

The status of Lambrus spinosissimus Osorio, 1923 

(see Nobre, 1936:85) remains to be determined. 

One specimen originated from the latitude of 

Lisbon, the other from Morocco. Osorio's account 

is not available to us. According to Nobre's quote 

of the original, L. spinosissimus (cl 33 mm, cb 37 

mm) is a larger species than the West African P. 

notialis (cl 5-21 mm, cb 11-23 m) (see p. 334). 

Osorio described the upper surface of the palm of 

the cheliped as follows (our translation): "The 

upper surface of the hand also shows spines, 

rather widely spaced one from the other, and the 

lower surface is covered with granules, which do 

not stop at the base of the fingers, but, on the 

contrary, are widely distributed on them." This 

description applies well to P. macrochelos; in P. 

notialis the upper surface of the palm of the 

cheliped is almost smooth. We believe that L. 

spinosissimus Osorio, 1923 should be identified 

with the northern Parthenope macrochelos (Herbst, 

1790) rather than the smaller tropical species, P. 

notialis. 

Subfamily AETHRINAE Dana, 1851 

Genus Heterocrypta Stimpson, 1871 

Heterocrypta Stimpson, 1871b: 102 [type-species: Cryptopodia 

granulata Gibbes, 1850, by original designation; gender: 

feminine; name 1626 on Official List]. 

* Heterocrypta maltzami Miers, 1881 

Heterocrypta Maltzami Miers, 1881a:209, pi. 13: fig.l. 

Heterocrypta Maltzani.—Miers, 1881a:364, 374.—A. Milne Edwards 

and Bouvier, 1900:121, pi. 19: fig.6 [part ?].—Balss, 

1921:54.—Bouvier, 1940:315 [part ?]. 

Heterocrypta maltzani.—Ortmann, 1893b:417.—Rathbun, 

1900a:296.—Sourie, 1954b: 150.—Monod, 1956:589, figs. 

862-867.—Longhurst, 1958:89.—Gauld, 1960:72.— 

Guinot and Ribeiro, 1962:80.—Rossignol, 1962:123.—Ribeiro, 

1964:21.—Forest and Guinot, 1966:120.—Zariquiey 

Alvarez, 1968:442 [Spain; references].—Maurin, 

1968b:486.—Tiirkay, 1975a:71 [listed], 74, fig. 6. 




Ivory Coast: Sta 46, 38-42 m, mud with denseJullienella, 

29 (1 ov) (L). 

Ghana: Sta 23, 42 m, foliate brown to orange bryozoans, 

16* (L). Sta 24, 35-37 m, dark red bryozoans, 3

324 

tion to consider that "le specimen de H. maltzani 

Miers signale par T. Odhner (1923 p. 20) de Port 

Alexandre (Angola), appartient vraisemblablement 

a cette espece nouvelle [= Solenolambrus noordendei]" 

Monod (1956:590, 595) seemed to accept 

Capart's conclusions. Guinot and Ribeiro (1962: 

80, 81) showed that H. maltzami does occur rather 

commonly in Angola and concluded that "on 

peut done supposer que 1 'echantillon d'Odhner 

appartient bien a H. maltzani." This conclusion 

seems to be accepted by Grosnier (1967:340, 1970: 

1219). As long as Odhner's specimen has not been 

reexamined, speculation on its identity is futile: 

The size of the specimen may fit either species, 

the locality of it lies within the known range of 

H. maltzami, but outside that of Solenolambrus noordendei, 

while the depth at which Odhner's specimen 

was collected lies well within the depth range 

of S. noordendei, but is much greater than at present 

known for tropical West African H. maltzami. 

BIOLOGY.—Monod (1956:593) gave the depth 

range of the species as "sublittoral, jusqu'a 400 

metres." Our specimens were obtained at depths 

of 33 to 70 m. All previous West African records 

also are from far less than 100 m depth: 18-28 m 

(Miers, 1881a: type-locality), 10 m, 20-22 m 

(Balss, 1921), 10, 14-15,15, 15, 15-20, 15-20, 16- 

18, 19, 19, 20, 20, 20, 22, 23-24, 25, 25, 27-28, 

30, 30, 31, 32, 32, 35, 36, 37, 38, 38, 40-41, 41, 

46-50, 50 m (Monod, 1956), 0-10, 3-17 m 

(Sourie, 1954b), 18-60 m (Longhurst, 1958), 19- 

37 m (Gauld, 1960), 5, 8, 11-12, 13, 16, 22 m 

(Guinot and Ribeiro, 1962), 6-8 m (Rossignol, 

1962), 8 m (Ribeiro, 1964), 18 m (Forest and 

Guinot, 1966), 30-40 m (Maurin, 1968b), 40, 

57 m (Tiirkay, 1975a). On the other hand all the 

records of the species from the Mediterranean, 

the Bay of Biscay, the Azores, and north of the 

Gape Verde Islands are much deeper: 100 to 

550 m. The West African specimens were found 

on the following types of bottom: shelly and 

muddy (Miers, 1881a), sand with Palythoa and 

Molgula (Monod, 1956), coarse shelly sand with 

Area and Pyura (Sourie, 1954b), muddy sand, 

shelly sand, and shelly mud (Longhurst, 1958), 

sand (Guinot and Ribeiro, 1962), mud and shells 


(Forest and Guinot, 1966), and sand or shelly 

sand (Maurin, 1968b). 

DISTRIBUTION.—Heterocrypta maltzami has been 

reported from the Bay of Biscay, the Mediterranean 

(as far as the Adriatic Sea), the Azores, N of 

the Cape Verde Islands (16°55'-16°51'N, 27°27'- 

27°29'W of Paris = 25°07'-25°09'W of Greenwich), 

and from tropical West Africa. Monod 

(1956) summarized earlier records and reported 

material from Senegal, Ghana, and Gabon; records 

not in Monod include the following: 

Spanish Sahara: OffCabo Blanco, 20°43.5'N, 17° 10.8'W, 

40 m, and 2O°37.3'N, 17°24.4'W, 57 m (Tiirkay, 1975a). 

Mauritania: Bane d'Arguin, 30-40 m (Maurin, 1968b). 

Cape Verde Islands: Baia de Porto Grande, Sao Vicente 

(Guinot and Ribeiro, 1962; Ribeiro, 1964). 

Senegal: Anse de Hann and Anse Bernard near Dakar 

(Sourie, 1954b). 


1966). 


1958). 

Ghana: Off Accra, 18-37 m (Gauld, 1960). 

Congo: Baie de Pointe-Noire, 6-8 m (Rossignol, 1962). 

Angola: Baia da Caota, 11-12, 13, and 16m; Sombreiro, 

5 m; and Baia Farta, 22 m (all Benguela) (Guinot and 


Genus Sakaila, new genus 

TYPE-SPECIES.—Sakaila africana, new species. 

ETYMOLOGY.—It is with great pleasure that we 

dedicate this genus to Professor Tune Sakai, the 

eminent Japanese carcinologist, who pointed out 

the differences between the species assigned here 

to the new genus and the species of Osachila sensu 

stricto, here restricted to the American and central 

Atlantic species (Rathbun, 1937:248). The 

gender of the generic name is feminine. 

DEFINITION.—Carapace broad, suboval in 

shape, narrowing anteriorly, regularly arcuate 

laterally, margins thin, scalloped or lobed. Front 

narrow, produced anteriorly, bilobed. Orbits 

small. Eyes scarcely or not at all visible in dorsal 

view. Antennules oblique. Antennae at inner angle 

of orbit. Buccal cavity broad anteriorly. Efferent 

branchial orifices separated, not meeting at 

midline. Merus of third maxilliped shorter than


ischium, anterior margin of merus divided into 2 

lobes. Chelipeds symmetrical. Walking legs compressed, 

dorsally toothed or with large, irregular 

tubercles. 

REMARKS.—Sakaila can readily be distinguished 

from Osachila Stimpson, 1871, by the 

position of the orbits, the distinctly separated 

efferent branchial channels, by the ischium of the 

third maxillipeds being shorter than the merus 

and being notched anteriorly, and by the spined 

or tuberculate walking legs. 

Guinot (1966:754) noted "pour des raisons 

d'ordre taxonomique, la separation d'O. stimpsoni, 

0. japonica, et 0. imperialis dans un genre ou sousgenre 

nouveau pourrait se justifier car plusieurs 

caracteres concomitants les distinguent des autres 

Osachila." Guinot (1968b) referred to 0. stimpsoni 

[sensu Monod, 1956] as a primitive Osachila. As a 

result of her study of the affinities of Osachila 

sensu lato, she assigned it, Aethra, Hepatus, Hepatella, 

and Actaeomorpha to a group that she called 

"parthenoxystomienne," and tentatively assigned 

this group to the Parthenopinae (Guinot, 1967b: 

841). This group was retained in the parthenopids 

by Sakai (1976). Pending further studies, we retain 

Sakaila in the Parthenopidae, subfamily 

Aethrinae, following Sakai (1976:288). 

In addition to the type-species, Sakaila africana, 

new species, we assign two other species to this 

genus, both from Japanese waters: Osachila imperialis 

Sakai, 1963, and Osachila japonica Sakai, 1963 

(Sakai, 1963, 1965). Osachila expansa Takeda 

(1977), from the Ogasawara Islands, in which the 

maxillipeds are strongly narrowed anteriorly, 

may be retained in Osachila sensu stricto. 

* Sakaila africana, new species 

FIGURE 83 

Osachila stimpsoni.—Monod, 1956:100, 624, figs. 874-876.— 

Forest, 1959:15.—Forest and Guinot, 1966:51.—Guinot, 

1966:747, 750, 752-755, figs. 2, 6, 14; 1967b:828-830, 

832-838, figs. 26, 29, 32, 33; 1968b: 165, fig. 15 [discussion]. 

[Not Osachila stimpsonii Studer, 1883.] 


Sta 284, 73 m, black basaltic rocks, 1

326 


FIGURE 83.—Sakaila africana, new genus, new species, holotype, male cl 23 mm, Pillsbury Sta 284: 

a, carapace, dorsal view; b, carapace, ventral view; c, chela; d, fifth pereiopod; e, abdomen. 

buccal area shows correctly that the infraorbital 

tooth, which lies against the outer margin of the 

antennal peduncle, continues posteriorly and medially 

as a flattened lobe, which partly overlaps 

the area along the anterolateral margin of the 

mouthfield. In our specimen this lobe ends in the 

anterior and largest of the three deep pits in the 

subhepatic region. These pits are not shown in 

Guinot's figure. 

The outer surface of the merus of the cheliped 

(Figure 83r) is reticulate by the presence of shorter 

and longer ridges and tubercles, rather than being 

"granuleuse" as described by Monod. The upper 

surface of the merus, not described by Monod, is


very narrow, widening somewhat anteriorly. The 

inner margin shows three high teeth, which are 

anteroposteriorly compressed; the outer margin 

shows a sharp high rim-like crest, the anterior 

part of which is highest and is fused with the 

anterior inner tooth to a high screen-like structure 

on the anterodorsal margin of the merus. The 

inner surface of the carpus has an anterodorsal 

and an anteroventral tooth connected with a 

ridge-like row of smaller teeth. The ridge on the 

inner surface of the palm bears a row of tubercles, 

one near the middle being high and sharp. The 

fingers have some of the tubercles rather strong, 

especially near the upper margin (in the dactylus), 

the lower margin (in the fixed finger), and 

near the base. The fingers of both chelae are 

inflected downward (Figure 83c), making a blunt 

angle with the longitudinal axis of the palm. In 

Monod's (1956) figure 875 the fingers are shown 

directed straight forward. 

Through the kindness of H.-E. Gruner, Zoologisches 

Museum, Berlin, we were able to compare 

our specimens with syntypes of Osachila stimpsonii 

Studer, 1883, from Ascension Island. Studer's 

species is correctly placed in Osachila. A redescription 

of 0. stimpsonii is in preparation by Manning 

in collaboration with Fenner A. Chace, Jr., in a 

report on the marine decapods of Ascension Island. 


05°36'E, in 73 m. 


D. 31541), a male taken at Pillsbury Sta 284, is in 


The male paratype (USNM 139766) taken 

at Geronimo Sta 235 is in the National Museum of 


D.C. 

BIOLOGY.—Sakaila africana has been collected in 

depths between 65-75 and 132 m. The type of 

bottom on which it was found has been indicated 

as mud and sand, 65 to 75 m (Forest and Guinot, 

1966) and black basaltic rocks, 73 m {Pillsbury). 

Ovigerous females were collected in July 

(Monod, 1956). 


from Latin and refers to the occurrence of the 

species off the West African coast. 

DISTRIBUTION.—Off tropical West Africa. It has 

not previously been recorded from either Gabon 

or Annobon. Records in the literature include the 

following: 

Senegal: S of Goree, 96 m; off Goree, 132 m (Monod, 

1956). 12°55.5'N, 17°33'W, 65-75 m (Forest, 1959; Forest 

and Guinot, 1966; Guinot, 1966, 1967b, 1968b). 

Subfamily PARTHENOPINAE MacLeay, 1838 

Genus Daldorfia Rathbun, 1904 

Parlhenope Fabricius, 1798:315, 352 [type-species: Cancer horridus 

Linnaeus, 1758, by selection by H. Milne Edwards, 

1838, in 1836-1844, pi. 26: fig. 2; a junior homonym of 

Parthenope Weber, 1795; gender: feminine; name 1679 on 

Official Index]. 

Daldorfia Rathbun, 1904:171 [type-species: Cancer horridus 

Linnaeus, 1758, by monotypy; gender: feminine; name 


Daldorfia bouvieri (A. Milne Edwards, 1869) 

Parthenope bouvieri A. Milne Edwards, 1869:350.—Capart, 

1951:106, fig. 36 [Cape Verde Islands].—Monod, 1956: 

595, figs. 871, 872 [Cape Verde Islands; references].— 

Forest and Guinot, 1966:121 [Principe, Sao Tome].— 

Crosnier, 1967:340 [Congo]; 1969:535 [Congo]. 


Verde Islands and the Gulf of Guinea at the 

localities listed above, in 4-5 to 91 m. 

Genus Parthenope Weber, 1795 

Parthenope Weber, 1795:92 [type-species: Cancer longimanus 

Linnaeus, 1758, by subsequent designation by Rathbun, 

1904:171; gender: feminine; name 1581 on Official List]. 

Lambrus Leach, 1815a:308 [type-species: Cancer longimanus 

Linnaeus, 1758, by monotypy; gender: masculine; name 

1678 on Official Index). 

Platylambnts Stimpson, 1871a: 129 [type-species: Lambrus crenulatus 

Saussure, 1858, a subjective junior synonym of 

Lambrus serratus H. Milne Edwards, 1834, by subsequent 

designation by Rathbun, 1925:516; gender: masculine]. 

Aulacolambrus Paulson, 1875:9 [type-species: Lambrus pisoides 

Adams and White, 1848, by monotypy; gender: masculine]. 

Pseudolambrus Paulson, 1875:9 [type-species: Parthenope calap-

328 

poides Adams and White, 1848, by monotypy; gender: 

masculine]. 

Enoplolambrus A. Milne Edwards, 1878, in 1873-1881:147 

[type-species: Lambrus carenatus H. Milne Edwards, 1834, 

by monotypy; gender: masculine]. 

Parthenolambrus A. Milne Edwards, 1878, in 1873-1881:148 

[type-species: Parthenope tarpeius Adams and White, 1849, 

by subsequent designation by Rathbun, 1925:528; gender: 

masculine]. 

Rhinolambrus A. Milne Edwards, 1878, in 1873-1881:148 

[type-species: Cancer contrarius Herbst, 1804, by original 

designation; gender: masculine]. 

Parthenopoides Miers, 1879a:672 [type-species: Lambrus massena 

Roux, 1830, by monotypy; gender: masculine]. 

Oncodolambrus De Man, 1906:400 [type-species: Lambrus (Oncodolambrus) 

praedator De Man, 1906, by monotypy; gender: 

masculine]. 

* Parthenope expansa (Miers, 1879) 

FIGURE 84 

Lambrus (Parthenopoides) expansus Miers, 1879b:25, pi. 5: 

fig. 9. 

Parthenolambrus expansus.—Adensamer, 1898:611 [Mediterranean]. 

Lambrus expansus.—Monod, 1956:588 [references].—Holthuis 

and Gottlieb, 1958:119 [listed].—Forest and Guinot, 1966: 

120. 

Parthenope expansus.—Pastore, 1975:145, 147, figs. 1-3 [Mediterranean]. 


Sta 283, 51-55 m, nodular coralline algae, 2$ (L, W). 

DESCRIPTION.—Carapace triangular in outline. 

Front bluntly rounded, its sides forming a straight 

line with sides of carapace. Orbits visible above 

as small cavities in lateral margin of carapace, 

but eyes, when retracted, exactly fill cavity and 

lateral margin appears unbroken. In posterior 

half, lateral margin slightly widened and showing 

3 shallow teeth just before posterolateral angle. 

Posterior margin slightly and evenly convex, 

showing few broad, inconspicuous lobes in median 

part. Dorsal surface of carapace uneven, but 

showing hardly any tubercles. Front somewhat 

concave, sunk in middle. Transverse broad ridge 

or elevated area present in metagastric region 

behind sunken area. Cervical groove indicated by 

wide depression. Oblique ridge present over each 

branchial region ending in posterolateral angle 


and running parallel to lateral margin of carapace. 

Posterior part of carapace, between branchial 

ridges, bearing median and 2 submedian 

elevations, median strongest. Surface of carapace 

rather uniformly and finely pitted, with few tubercles 

visible on elevated portions, but not very 

distinct. General shape of carapace in many respects 

resembling more that of Heterocrypta than 

that of Parthenope. 

Eyes small, completely retractable in their orbits, 

visible in dorsal view. 

Antennules with basal segment very wide, 

reaching beyond antennal peduncle and forming 

greater part of the lower inner margin of orbit. 

Antennular sockets not sharply delimited distally, 

ending in wide groove in ventral surface of front, 

groove reaching margin of front, but less distinct 

distally than proximally. 

Basal antennal segment (Figure 84a) short, distinctly 

failing to reach orbit and separated from 

orbit by almost entire length of second segment. 

Outer angle of merus of third maxilliped somewhat 

triangularly produced laterally. 

Chelipeds markedly different, resembling those 

of Parthenope massena. Right cheliped heaviest, 

somewhat swollen. Upper surface of palm slightly 

convex, almost smooth; outer margin bearing 

ridge with 3 large blunt teeth, more pronounced 

in smaller than in larger cheliped. Inner margin 

of the upper surface of palm bearing about 5 

FIGURE 84.—Parthenope expansa (Miers), female, cl 7.5 mm, 

Pillsbury Sta 283: a, basal antennal segment; b, second pereiopod; 

c, fifth pereiopod.


blunt teeth, middle largest. Outer surface of palm 

with some longitudinal rows of tubercles. Inner 

surface of palm smooth, slightly concave. Fingers 

somewhat more than half as long as palm, upper 

surface of dactylus bearing few large and several 

smaller tubercles proximally. Cutting edge of 

fixed finger of large cheliped with single large 

molariform tooth, occupying greater part of edge, 

flanked by small denticle distally. Cutting edge 

of fixed finger in small chela with 3 large distal 

and 3 small proximal teeth. Edges of dactylus of 

both chelipeds bearing 3 or 4 low small teeth. 

Carpus short, cup-shaped, with small irregular 

tubercles, forming more or less distinct longitudinal 

rows. Merus short and wide, inner margin 

bearing distinct larger and smaller teeth, on outer 

margin teeth less conspicuous. Lower margin of 

merus with longitudinal row of large tubercles, 

inner surface bearing some scattered tubercles, 

outer surface almost smooth. 

In following legs (Figure 84b,c) lower surface of 

merus bearing 2 rows of granules, 1 on either 

margin; upper margin of merus armed with tubercles 

in extreme proximal part only. Carpus 

with lower surface unarmed, upper carrying angular 

tubercle; in addition, last leg with few distal 

tubercles on upper margin. Propodus with some 

ventral and dorsal denticles; denticles and tubercles 

far more distinct in posterior than in anterior 

legs. Dactylus of second pereiopod (= first walking 

leg) about as long as, but narrower than, 

propodus, somewhat longer than carpus; merus 

about twice as long as, and broader than, carpus. 

Third and fourth pereiopods similar to second. 

Fifth pereiopod (Figure 84c) shortest of all legs. 

Dactylus longer than, but half as high as, propodus; 

carpus about as long as propodus, and half 

or less than half as long as merus. 

Female abdomen with all somites free. 

MEASUREMENTS.—The two examined females 

have cl 6 and 7.5 mm and cb 7 and 10 mm. The 

holotype had cb 11 mm (Miers, 1879b). The 

measurements given in the literature are: cl 10 

mm, cb 11 mm (Miers, 1886); cl 8 mm, cb 10 

mm (A. Milne Edwards and Bouvier, 1894); for 

123 cl 4.0-9.5 mm, cb 5.0-12.0 mm, for 2 oviger- 

ous 9 cl 5.0 and 6.0 mm, cb 5.5 and 7.5 mm, and 

2 non-ovigerous $ cl 5.0 and 6.5 mm, cb 5.8 and 

7.5 mm (Pastore, 1975:150; in Pastore's table, 

length and width have evidently been interchanged 

by accident). The known size range for 

the species thus is cl 4-10 mm, cb 5-12 mm. 

REMARKS.—Capart (1951:106, fig. 36) described 

and figured a female of what he considered 

to be a juvenile Daldorfia bouvieri (A. Milne 

Edwards). This specimen (cl 9 mm, cb 12 mm) 

originated from Boa Vista, Cape Verde Islands. 

The description and figure given by Capart show 

a great similarity to the present species, and a 

reexamination of Capart's specimen might be of 

interest. Our specimens differ from that described 

by Capart in having the carapace uniformly and 

evenly minutely pitted and not eroded. Capart's 

specimen might be abnormal in having two equal 

chelipeds, both corresponding with the smaller 

cheliped in the present species. 

BIOLOGY.—The species has been found in 

depths between 30 and 185 m. The bottom at the 

localities where it was found was noted in some 

instances: gravel, sand and broken shell (A. Milne 

Edwards and Bouvier, 1894, 1899); coral, rock 

and sand, sand and rock, sand, shell and coral, 

sand and shell (A. Milne Edwards and Bouvier, 

1900); calcareous and other algae (Forest and 

Guinot, 1966); nullipores and coarse sand (Adensamer, 

1898). 

Ovigerous females have been collected in June 

and August (Monod, 1956; Pastore, 1975). 


Mediterranean, Madeira, Seine Seamount, the 

Azores, the Canary Islands, and West Africa from 

Spanish Sahara to Sao Tome and Annobon islands 

in the Gulf of Guinea. This species has not 

been recorded previously from Annobon. The 

Mediterranean records are: NW of Crete, 36°- 

03'N, 23°06'E (Adensamer, 1898; Pastore, 1975) 

and Acitrezza, Bay of Catania, Sicily (Pastore, 

1975). Monod (1956) summarized earlier records 

and reported material from Madeira, the Canary 

Islands, and the Cape Verde Islands. West African 

records since 1956 include the following:

330 

Canary Islands: No specific locality (Pastore, 1975). 

Sao Tome: OO°25'4O"N, 06°40'10"E, 50 m (Forest and 


* Parthenope massena (Roux, 1830) 

?Lambrus massena.—Capart, 1951:105, fig. 35. 

Lambrus massena. —Sourie, 1954b: 147, 150.—Monod, 1956: 

572, 632, figs. 840-856.—Gauld, 1960:72.—Rossignol, 

1962:123.—Crosnier, 1964:31.—Forest and Guinot, 1966: 

118— Zariquiey Alvarez, 1968:441, fig. 147 [Spain; references]. 

Lambrus sp.—Monod, 1956:583, figs. 857, 858. 

Lambrus massenae.—Longhurst, 1958:89 [erroneous spelling]. 

SYNONYMS.—PParthenope contracta Costa, 1840; 

?Parthenope hexacantha Costa, 1840; Lambrus pumilus 

Costa, 1851; Lambrus rugosus Stimpson, 1857; Lambrus 

setubalensis De Brito Capello, 1866; Lambrus 

pulchellus A. Milne Edwards, 1868; Lambrus massena 

var. atlanticus Miers, 1881; Lambrus massena 

var. spinifer Miers, 1881; Lambrus massena var. 

goreensis Miers, 1881; Lambrus bicarinatus Miers, 

1881. 



Ghana: Sta 22, 51 m, rough bottom, 2c5, 1 juv (L). Sta 23, 

42 m, foliate brown to orange bryozoans, 1


material reported upon here was taken between 

24 and 51 m. The species was taken on the 

following types of bottom: sand with Palythoa and 

Molgula (Monod, 1956); coarse shelly sand, bottom 

with Area and Pyura (Sourie, 1954b); sand 

and shells, sand and mud (Longhurst, 1958); mud 

and shells (Longhurst, 1958; Forest and Guinot, 

1966); sand with Foraminifera on rocky bottom 

with gorgonians (Crosnier, 1964); rock and shells, 

mud, calcareous algae and shells, calcareous algae, 

calcareous algae, sand and shells, sand and 

calcareous and other algae, mud and calcareous 

algae (Forest and Guinot, 1966). 

Ovigerous females have been recorded in all 

months but January, April, and August, suggesting 

that off West Africa the species spawns all 

year (Monod, 1956; Forest and Guinot, 1966; 

Pillsbury). 

DISTRIBUTION.—Eastern Atlantic, from Brittany, 

Atlantic coast of France, southward to the 

Congo, including the Mediterranean. Monod 

(1956) summarized the literature and reported 

material from Senegal, Guinea, Sierra Leone, 

Ghana and Principe. Other records in the literature 


Senegal: Anse Bernard and Anse de Hann, Baie de Dakar, 

0-17 m (Sourie, 1954b). 


1958). 

Ghana: Off Accra, 20-40 m (Gauld, 1960). 04°37'N, 




Cameroon: No specific locality, in 10-50 m (Crosnier, 

1964). 

Principe: 01°38'25"N, O7°22'O5"E, 31 m; 01°43'10"N, 

07°28'20"E, 73 m;.01°43'N, 07°28'55"E, 37 m; [Cais de] 

Santana, 11 m; Praia Grande, 3-12 m (all Forest and Guinot, 

1966). 

Sao Tome: 00°25'15"N, 06°43'05"E, 8-30 m; Baia de 

Ana de Chaves, 5 m (Forest and Guinot, 1966). 

Annobon: 01°27.5'S, 05°36.5'E, 35 m (Forest and Guinot, 

1966). 

Gabon: W of Pointe Gombe, 40 m (Rossignol, 1962). 

Congo: W of Pointe-Noire, 10 m (Rossignol, 1962). 

Parthenope miersii (A. Milne Edwards and 

Bouvier, 1898) 

Lambrus miersi— Capart, 1951:105 [discussion].—Monod, 

1956:583 [references]. 

Parthenope miersi.—Zariquiey Alvarez, 1968:439 [Spain; references]. 


the Bay of Cadiz, Seine Seamount, and the 

Cape Verde Islands; sublittoral, in depths between 

91 and 240 m. 

* Parthenope notialis, new species 

FIGURES 85, 86a,* 

Lambrus mediterraneus.—Studer, 1882:335 [not Lambrus mediterraneus 

Roux, 1828 = Cancer macrochelos Herbst, 1790]. 

Lambrus Mediterraneus.—Studer, 1883:9. 

Lambrus macrochelos.— Rathbun, 1900a:295.—Monod, 1956: 

585, figs. 859-861.—Longhurst, 1958:89.—Gauld, 1960: 


123.—Crosnier, 1964:34. [Not Cancer macrochelos Herbst, 

1790.] 

Lambrus macrocheles.— Doflein, 1904:87 [part], pi. 32: fig. 5.— 

Balss, 1921:54.—Odhner, 1923:20.—Capart, 1951:102, 

fig. 34, pi. 2: figs. 5, 6.—Forest and Guinot, 1966:119.— 

?Maurin, 1968a:59, 62; 1968b:480, 486, 489.—Crosnier, 

1970:1215 [listed], 1219. [Not Cancer macrochelos Herbst, 

1790.] 

Lambrus (Lambrus) macrocheles.—Monod, 1933b:498 [not Cancer 

macrochelos Herbst, 1790]. 

Parthenope.—Vass, 1966:19, 22. 

Lambrus.— ?Maurin, 1968b, fig. 9. 


Sta 68, 70 m, broken shell, 66, 12? (2 ov) (L). 

Ivory Coast: Sta 42, 62-75 m, mud with brown branched 

Foraminifera, 66*, 4$ (W). Sta 46, 38-42 m, mud with dense 

Jullienella, 1(5, 29 (W). Sta 50, 128-192 m, 1$ ov (L). Sta 59, 

55-64 m, mud with dense branched Foraminifera, 1

332 

it was not distinguished by the authors that have 

dealt with it. We found, however, a number of 

characters that made it impossible to consider the 

present West African specimens and typical Med- 

iterranean specimens of P. macrochelos as belonging 

to the same species or subspecies. 

The shape of the carapace (Figure 85), which 

in adult specimens is distinctly wider than long 

(proportion length:width being about 5:6), has 

the same general shape as in P. macrochelos. The 

rostrum is narrow and bears a tooth at either side. 

The inner orbital angle bears two teeth and a 

strong tubercle. The upper margin of the orbit is 

provided with a strong tubercle (Figure 86a), 

behind which sometimes a trace of a smaller 

tubercle may be visible; in P. macrochelos (Figure 

86c) this second tubercle is quite distinct. Some 

distance behind the orbits, in the median area of 

the mesogastric region, there are four tubercles 

placed in a quadrangle. The anterior pair stands 

close to the posterior pair and its tubercles are 

placed slightly wider apart than those of the 

posterior pair. The four tubercles thus form a 

trapezium with the widest margin anteriorly. In 

P. macrochelos, the posterior tubercles are wider 

apart than the anterior, so that the trapezium 

formed by the four tubercles has the narrowest 


margin anteriorly; in P. macrochelos the two pairs 

of tubercles are separated from each other by a 

greater distance than in P. notialis. Behind the 

group of four tubercles there is a strong median 

gastric spine. Behind the cervical groove there is 

a median row of 3 strong spines (1 urogastric and 

2 cardiac); the posterior of these is the largest. In 

P. notialis these spines are relatively longer than 

in P. macrochelos; the arrangement of the spines is 

the same in the two species. The median gastric 

region is swollen and so is the branchial region. 

In the depressed area between these two swollen 

regions there is a row of three blunt tubercles 

close to and parallel with the median row of 

spines. The branchial region ends posterolaterally 

in three large teeth, between which there are 

small tubercles. In P. macrochelos, instead of these 

tubercles, there are teeth that often are only 

slightly smaller than the large teeth. The middle 

of the three large teeth is placed at the posterior 

end of a blunt oblique ridge, which carries another 

spine more anteriorly. The anterolateral 

margin of the carapace bears a row of about 7 

teeth, which are distinctly shorter and narrower 

than the outer posterolateral tooth; in P. macrochelos 

these anterolateral teeth are much larger, 

reaching almost the size of the outer posterolat- 

FIGURE 85.—Parthmope notialis, new species (from Monod, 1956, fig. 859).


FIGURE 86.—Parthenope notialis, new species, paratype, male, 

cl 17.5 mm, Pillsbury Sta 260: a, front; b, abdomen. Parthenope 

macrochelos (Herbst), male, cl 32.8 mm, Naples: c, front; d, 

abdomen. 

eral. The general shape and tuberculation of the 

carapace of P. notialis is very similar to that of P. 

macrochelos. 

The epistome, oral field, and third maxilliped 

are like in P. macrochelos, only the spines are 

sharper and more distinct. The subhepatic region 

just lateral of the oral cavity is smooth in P. 

notialis, uniformly tuberculated in P. macrochelos. 

In P. notialis the chelipeds of the adult males 

are relatively distinctly longer than in P. macrochelos. 

The second pereiopod in P. notialis does not 

attain the end of the merus of the chela, while in 

P. macrochelos it reaches beyond. The fingers of the 

chelipeds in P. notialis are more laterally compressed 

than in P. macrochelos, where they are more 

cylindrical. The teeth on the palm in P. notialis 

also are more compressed and have the margins 

with small serrations, sometimes small tubercles 

may be seen on the lateral surfaces of these teeth. 

In P. macrochelos the teeth on the palm are more 

** 

conical and have spinules on all sides. The upper 

surface of the palm of the chelipeds in P. notialis 

is almost smooth, only a few indistinct tubercles 

are seen; in P. macrochelos this surface is very rough 

with many spines and tubercles. The lower outer 

surface of the palm is slightly convex with a few 

rows of small tubercles, of which the median is 

the most conspicuous. In P. macrochelos this surface 

is more convex and the tubercles are larger. In P. 

notialis the merus of the cheliped has the slightly 

convex upper surface almost smooth, except for 

a median row of spinules; in P. macrochelos the 

surface is more uniformly spinuliferous. The 

spines on the margins of the upper surface of the 

merus are more flattened and less numerous in P. 

notialis than in P. macrochelos. 

The following pereiopods are very similar in P. 

notialis and P. macrochelos. The dactylus is covered 

with a very short felt-like pubescence; it is practically 

as long as the propodus in the second 

pereiopod, becoming gradually relatively longer 

in the following legs. The carpus also is about as 

long as the propodus in pereiopod 2, but it becomes 

gradually shorter in the following legs. The 

merus is the longest segment and is as long as 

propodus and carpus combined. A row of spinules 

is present on the upper and the lower margins of 

the merus, and on the upper margin of the carpus 

and propodus of all legs. The spinules are distinct 

on the merus, faint to very faint on the carpus 

and propodus; in the posterior legs they are more 

distinct than in the anterior. 

The shape and armament of the male thoracic 

sternum and abdomen is about the same in the 

two species. The second abdominal segment 

shows one median and two lateral teeth with 

tubercles in between; in P. notialis (Figure 86A) 

these teeth are large and bluntly lobiform, flattened 

and higher than wide, the tubercles are 

few, small and inconspicuous; in P. macrochelos 

(Figure 86W) the teeth are placed wider apart, are 

less flattened, lower and more triangular, while 

the tubercles are distinct and rather large. The 

general shape of the abdomen is the same in the 

two species, but the last somite is narrower in P. 

notialis. The third to fifth somites of the abdomen

334 

are fused: sutures rather than articulations indicate 

the lines between the somites. 

Figures: Figures of the whole animal were published 

by Doflein (1904, pi. 32: fig. 5), Capart 

(1951, fig. 34), and Monod (1956, fig. 859). 

Male Pleopod: Figures of the male pleopods are 

provided by Capart (1951, pi. 2: figs. 5, 6) (Zaire) 

and Monod (1956, figs. 860-861) (Senegal). They 

do not essentially differ from those of P. macrochelos. 

MEASUREMENTS.—The carapace length of the 

specimens seen by us varies between 5 and 21 

mm, that of the ovigerous females between 14 

and 18 mm. Odhner's (1923) specimen had the 

carapace width between 11 and 19 mm. Both 

Capart's (1951) largest male and largest female 

had the carapace 18.5 mm long and 21 mm wide. 

Monod (1956:585, 586) gave the following measurements 

for six of his males: cl 12 to 18 mm, cb 

13 to 23 mm; and for three ovigerous females: cl 

10 to 14 mm, cb 11 to 16 mm. The single specimen 

reported upon by Guinot and Ribeiro (1962:80) 

was a male with the carapace length and width 

both 5 mm. It is clear that the present species is 

a small one compared to P. macrochelos, which 

may attain a carapace length of 39 mm. The eggs 

of P. notialis are numerous and small, they are 

spherical and are 0.3 to 0.35 mm in diameter. 

Zariquiey Alvarez (1968:441), in citing ovigerous 

females of P. macrochelos with cl 10 mm, based his 

information on Monod's (1956) data of the present 

new species. 

REMARKS.—Parthenope notialis very strongly resembles 

P. macrochelos, and it is not surprising that 

the two species have always been confused. They 

may finally prove to be only subspecifically distinct. 

Our extensive material of the present species, 

however, differs consistently from the specimens 

of P. macrochelos, with which we could compare 

it. 

So far as we could ascertain all previous records 

of/*, macrochelos from tropical West Africa pertain 

to the present species. 

Studer (1883), in discussing his Liberian specimens 

(males and females), described the upper 

surface of the palm of the cheliped (which he 


inadvertently named "Carpus") as "fast glatt, 

nur mit wenig Hockern besetzt" and also described 

the outer lower surface of the palm with 

its median row of tubercles, and remarked that 

in these features his specimens differed from Mediterranean 

P. macrochelos. This clearly proves the 

identity of his material with the present species. 

Doflein (1904:87) mentioned two lots of"Lambrus 

macrocheles" from West Africa. One of these 

consisted of a single damaged male from Seine 

Seamount, NE of Madeira at 33°43.8'N, 14° 

20'W, ca. 150 m deep. This male might well 

belong to Parthenope massena (Roux). Doflein remarked 

that it had "auffallend kiirzere Scheren 

mit dunkel-rotbraunen Fingerenden," which 

would fit P. massena quite well, as we examined 

numerous specimens of that species which, after 

long preservation, still showed distinct reddish 

brown finger tips. Bouvier (1922:76, pi. 2: fig. 3), 

under the name Lambrus Miersi, described and 

figured (in color) P. massena, remarking that "le 

bout des doigts des pinces [sont] d'un brun noiratre," 

which color also is clearly shown on the 

figure. Doflein's specimen had the rostrum broken 

("dessen Rostrum abgebrochen ist"), and thus 

lacked a character that undoubtedly would have 

made Doflein recognize the true identity of the 

species. 

Doflein's second lot came from off the mouth 

of the Congo River and the male figured by him 

(1904, pi. 32: fig. 5) clearly belongs to the present 

species. 

Odhner (1923) considered his specimens to be 

juveniles of P. macrochelos as none had a carapace 

width of more than 19 mm; it is more probable 

that his specimens are P. notialis. 

Capart's (1951:102, fig. 34) illustration of a 

female from off Angola identified as "Lambrus 

macrocheles" clearly shows that it belongs to P. 

notialis as probably does also Capart's other material, 

since he remarked that the specimens in his 

collection "ne montrent pas entre eux de variations 

notables." As shown above the measurements 

of Capart's specimens fall entirely within 

the range that we found for the present species. 

Monod (1956:585, figs. 859-861) gave excellent


figures of the present species, while the measurements 

given by him check well with those of P. 

notialis (p. 334). Of one of his lots (from S of Cap 

Vert) Monod stated specially "petite forme, face 

superieure des chelipedes plus ou moins lisses 

entre les cretes." There is thus no indication that 

any of his specimens do not belong to P. notialis. 

Longhurst's (1958) and Gauld's (1960) specimens 

were identified by Monod. 

We examined a specimen from Guinea-Bissau 

reported upon by Forest and Guinot (1966) as P. 

macrochelos and found that it belongs to P. notialis. 

The characters used in various keys (Bouvier, 

1940:309, 310; Monod, 1956:572; Zariquiey Alvarez, 

1968:437, 438) to distinguish between P. 

macrochelos and P. miersii (A. Milne Edwards and 

Bouvier), also serve to distinguish P. macrochelos 

from P. notialis. This notwithstanding, we believe 

that P. notialis is closer to P. macrochelos than to P. 

miersii. Parthenope miersii has the carapace narrower 

than in either P. macrochelos or P. notialis 

and the chelipeds are shorter. Parthenope verrucosa 

(Studer, 1883) from Ascension has the rostrum 

similar to that of P. macrochelos and P. notialis, but 

its chelipeds are shorter than in these two species, 

and the tuberculation of the carapace seems to 

be different. Additional information on both P. 

miersii and P. verrucosa is badly needed; material 

of neither species was available to us. 

TYPE-LOCALITY.—The male holotype was collected 

at Pillsbury Sta 260, off Cameroon, 03°45'N, 

09°05'E to 03°43'N, 09° 10'E, depth 46 m. 


D. 31545) is deposited in the collection of the 

Rijksmuseum van Natuurlijke Historie, Leiden. 

Some of the paratypes are in the Leiden Museum 

and some in the National Museum of Natural 


DC. 


Latin notialis (southern). 


depths between 18 and 162 m (one of Maurin's 

uncertain records is from 300 to 500 m); however, 

more than 80% of the records are from depths 

between 30 and 110 m. The species usually is 

found on bottoms of mud, sandy mud or sand, 

almost always mixed with broken shells, bryozoans, 

branched or foliate Foraminifera, corals 

or rocks. The records in the literature are the 

following: shelly mud (Longhurst, 1958; Forest 

and Guinot, 1966); sand, mud, coral and rock; 

sand, brown mud and coral; sand, mud and rock 

(Capart, 1951); sand, mud and shells; mud, shells 

and Cidaris; mud, stones, calcareous algae, sand 

and Foraminifera; gravel, shells and Foraminifera 

(Forest and Guinot, 1966); sandy mud with shells; 

mud with dense Jullienella (Voss, 1966); bottom 

with Jullienella foetida Schlumberger (Monod, 

1956); sand and broken shells; sand, clayish sand 

and broken shells (Odhner, 1923); rock and shells 

(Forest and Guinot, 1966); sand and mud or 

sandy mud (Capart, 1951; Guinot and Ribeiro, 

1962; Forest and Guinot, 1966); and mud (Capart, 

1951; Forest and Guinot, 1966). The present 

material has been taken on similar types of bottom. 

Ovigerous females were observed in February, 

May, July, and September (Monod, 1956), 

March (Crosnier, 1970), May (Forest and Guinot, 

1966), May and June (present paper), November 

(Capart, 1951). 

DISTRIBUTION.—Most of the records of P. macrochelos 

from tropical West Africa are based on the 

present species. Whether Maurin's (1968a, 1968b) 

material from off Spanish Sahara and Mauritania 

belongs here, to P. macrochelos, or to P. miersii is 

not clear; this can only be decided after reexamination 

of his material. The range of the present 

species extends at least from Senegal to Angola, 

but it might go north as far as the Spanish 

Sahara. If we include Maurin's material in the 

present species, the published records for it are 


Spanish Sahara: Medano de Aaiiin and W of Cabo 

Bojador, 300-500 m (Maurin, 1968b). 

Mauritania: Bane d'Arguin, 40-60 m (Maurin, 1968b), 

and 90-100 m (Maurin, 1968a,b). Tamzak, 70-75 m 

(Maurin, 1968a, 1968b). 

Senegal: Off Saint-Louis, 100-300 m; off Kayar; S of 

Cap Vert, 97-98 m; SE of lie de la Madeleine, 48 m; off 

Dakar, 140 m; near Goree, 40-41 m, 50 m, % m, 132 m (all

336 

Monod, 1956). 13°01'N, 17°24'W, 51-55 m; 12°55.5'N, 





1966). 


1958). 

Liberia: 04°40'N, 09°40.6'W, 90 m (Studer, 1882, 1883). 

04°34.5'N, 08°31'W, 64 m (Forest and Guinot, 1966). 

Ivory Coast: O5°O7'N, 04°32'W to O5°O7'N, 04°36'W, 

38-42 m, and 04°35'N, 06°40'W to 04°35'N, 06°41'W, 64 

m (Voss, 1966). 

Ghana: 04°40'N, 02°08'W to 04°39'N, 02°05'W, 50 m; 

04°36.5'N, 01°31'W, 50 m; 04°37'N, 00°50'W, 90-100 m 

(Forest and Guinot, 1966). Accra, 43 m, 44 m, 65 m (Monod, 

1956; Gauld, 1960). 

Cameroon: No specific locality, depth more than 50 m 

(Crosnier, 1964). 

Gabon: 00°25'N, 09°00'E, 73 m (Forest and Guinot, 

1966). Off Pointe Banda, 03°57.5'S, 10°36.5'E, 85 m (Capart, 

1951). 

Cabinda: West of Landana, 50-65 m (Rossignol, 1962). 

Angola: Off Moita Seca, 06° 16'S, 12°O7'E, 50 m; 06°21'S, 

H°53'12"E, 100 m (Capart, 1951). Off the mouth of the 

Congo River, 06°18.7'S, 12°02.1'E, 44 m (Donein, 1904). 

09°47'S, 13° 1 l'E, 30-35 m; 09°40'S, 13°02'E, 80 m (Capart, 

1951). Baia Farta, Benguela, 22-28 m (Guinot and Ribeiro, 

1962). Baia dos Elefantes, 13°05'S, 12°46'E, 100-110 m; 

13°05'S, 12°45'E, 100-110 m; Baia de Salinas, 14°05'S, 

12°17'E, 110 m (Capart, 1951). Off Porto Alexandre, 72 m, 

108 m (Odhner, 1923). 16°37'S, 11°22'E, 126 m; 16°41'S, 

11°21'E, 162 m (Crosnier, 1970). 


(L). Sta 49, 73-77 m, 26, 5? (4 ov) (L). Sta 50, 128-192 m, 

26, 1? (W). Sta 59, 55-64 m, mud with dense branched 

Foraminifera, Id (L). Sta 60, 79-82 m, coral or rock, lcJ (L). 

Nigeria: Sta 237, 101 m, 49 (1 ov) (W). 

Undaunted Material: Angola: Sta 96, 162 m, 19 ov (L). 

Other Material: Congo: Off Pointe-Noire, 04°56'S, 11°- 

31'E, 95 m, muddy sand, 3 Sep 1965, A. Crosnier, Id (W). 



593, figs. 868-870. 


(Zaire). 

MEASUREMENTS.—The ovigerous females in the 

present collection had the carapace length 6 to 9 

mm. Capart (1951) mentioned ovigerous females 

with cl 7 and 10 mm, cb 7 and 11 mm; Monod's 

(1956) ovigerous females had cl 7 to 9 mm and 

cb 8 to 10 mm. The carapace length in the rest of 

our material varies between 6 and 12 mm. Capart's 

(1951) largest specimen had cl 11 and cb 

11.5 mm; Monod's (1956) smallest and largest 

male had cl 5 and 11.5 and cb 4.5 and 12 mm, 

respectively, while Crosnier's (1967) males had cl 

6.6 to 8.7 mm and cb 7.0 to 9.1 mm. 

BIOLOGY.—The species is known from depths 

between 64 and 215 m; the shallowest record is 

55-64 m (present material), the deepest 215-220 

m (Capart, 1951; Monod, 1956). Of the records 

90% are from between 70 and 140 m. It has been 

reported from mud (Capart, 1951; Crosnier, 

Genus Solenolambrus Stimpson, 1871 1967), sandy mud (Capart, 1951; Crosnier, 1967), 

Solcnolambrus Stimpson, 1871a: 132 [type-species: Solenolambrus and shelly mud (Longhurst, 1958). 

typicus Stimpson, 1871, by use of typicus; gender: mascu- Ovigerous females have been found in the 

line]. 

months of February, March, May, July, and No- 

Pisolambrus A. Milne Edwards, 1878, in 1873-1881:157 [typevember (Capart, 1951; Monod, 1956; Crosnier, 

species: Pisolambrus nitidus A. Milne Edwards, 1878, by 

1970; present paper). 


DISTRIBUTION.—Tropical West Africa, from localities 

between Senegal and Angola. It has not 

* Solenolambrus noordendei (Capart, 1951) 

been recorded previously from Liberia or Nigeria, 

Heterocrypta noordendei Capart, 1951:108, fig. 37, pi. 2: figs. 14, but these records are well within its known range. 

15. 

Records since Monod's (1956) include the follow- 

Solenolambrus noordendei.—Monod, 1956:593, figs. 868-870. ing: 

Longhurst, 1958:89.—Gauld, 1960:72.—Crosnier, 1967: 

340; 1970:1215 [listed], 1219. 

Sierra Leone: No specific locality, 72 m (Longhurst, 1958). 

Ivory Coast: No specific locality (Crosnier, 1967). 

MATERIAL EXAMINED.—Pillsbury Material: Liberia: Ghana: Off Accra, 80 m (Gauld, 1960). 


RV Pillsbury COLLECTIONS 

Collections made in the Gulf of Guinea in 1964 

and 1965 (data from Bayer, 1966; OT = otter 

trawl). 

5 

Appendix I: Station Data 

1. Nigeria. Lagos harbor, 06°28'N, 03°23'E, shore collecting, 

23 May 1964: Callinectes marginatus, Cyclograpsus 

integer, Geograpsus lividus, Goniopsis pelii, Metagrapsus curvatus, 

Pachygrapsus gracilis, Panopeus africanus, Sesarma 

(Chiromantes) buettikoferi, Uca tangeri 

2. Nigeria. Lagos harbor, 06°28'N, O3°23'E, dipnet at 

surface, outgoing tide, 23 May 1964: Callinectes amnicola, 

Callinectes pallidus 

16. Ghana. 05°40'N, 00°30'E to 05°40'N, 00° 17'E, 46 m, 

mud with Foraminifera, shells, 6' OT, 26 May 1964: 

Capartiella longipes, Eurynome parvirostris, Ilia spinosa, Inachus 

angolensis, Inachus nanus, Machaerus oxyacantha, Macropodia 

gilsoni, Macropodia spinulosa, Medorippe lanata 

17. Ghana. 05°35'N, 00°10'E to 05°36'N, 00°11.5'E, 48 

m, fine sand and green mud, 40' OT, 26 May 1964: 

Achaeus buderes, Achaeus foresti, Calappa pelii, Calyps- 

v 3Q 

A : 

N H 

pis 

/' /• /' ^3 

%\) 

1*64 CRUISE * 

achaeus calypso, Ebalia affinis, Inachus biceps, Macropodia 

spinulosa, Parthenope notialis, Pisa carinimana 

18. Ghana. [05°04'N, 00°12'E] to 05°01'N, 00°12'E, 

3047-3129 m, soft dark gray clay, Blake Trawl, 26 

May 1964: Ethusina beninia 

22. Ghana. 05°25'N, 00°01'W to 05°22'N, 00°02'W, 51 

m, rough bottom, 6' OT, 27 May 1964: Achaeus buderes, 

Calappa pelii, Calypsachaeus calypso, Ebalia tuberculata, 

Ethusa vossi, Herbstia condyliata, Ilia spinosa, Inachus angolensis, 

Inachus nanus, Macropodia hesperiae, Nanocassiope 

melanodactyla, Parthenope massena, Parthenope notialis, 

Stenorhynchus lanceolatus, Stemodromia spinirostris 

23. Ghana. 05°10'N, 00°25'W to 05°08'N, 00°28'W, 42 

m, foliate brown to orange bryozoans, 6' OT, 28 May 

1964: Achaeus buderes, Calappa pelii, Capartiella longipes, 

Ebalia affinis, Ebalia tuberculata, Ethusa vossi, Eurynome 

parvirostris, Heterocrypta maltzami, Ilia spinosa, Inachus 

biceps, Inachus nanus, Macropipus rugosus, Macropodia gils&ni, 

Macropodia spinulosa, Medorippe lanata, Nanocassiope 

melanodactyla, Parthenope massena, Parthenope notialis, Pilumnus 

perrieri, Pisa carinimana, Stenorhynchus lanceolatus 

24. Ghana. 04°56'N, 00°47.5'W to 04°56'N, 00°50 / W, 

35-37 m, dark red bryozoans, 6' OT, 28 May 1964: 

GULF 

t 

/ ^ 

OF GUINEA . IMS C«UIS£ tf 

Z 300 • / 

\% / 

A? 98 (J Sae Tom 

FIGURE 87.—Cruise track of the RA^ Pillsbury in the Gulf of Guinea, 1964 and 1965 

(from Voss, 1966, fig. 2). 

337 

\ 

* 1

338 


Achaeus buderes, Calappa rubroguttata, Capartiella longipes, gilsoni, Monodaeus rouxi, Pseudomyra mbizi, SoUnolambrus 

Ebalia qffinis, Ebalia tuberculata, Ethusa vossi, Heterocrypta noordendei 

maltzami, Ilia spinosa, Inachus biceps, Machaerus atlanticus, 46. Ivory Coast. O5°O7'N, 04°32'W to O5°O7'N, 04°36'W, 

Macropipus rugosus, Macropodia spinulosa, Medorippe lan- 38-42 m, mud with dense Jullienella, 6' OT, 30 May 

ata, Nanocassiope melanodactyla, Palicus caronii, Parthenope 1964: Calappa pelii, Capartiella longipes, Cronius ruber, 

massena, Parthenope notialis, Pilumnus stebbingi, Pisa cari- Ethusa vossi, Heterocrypta maltzami, Ilia spinosa, Leopoldius 

nimana, Portunus inaequalis, Sttnorhynchus lanceolatus, Ster- pisifer, Machaerus atlanticus, Machaerus oxyacantha, Macronodromia 

spinirostris, Typhlocarcinodes integrifrons 

podia spinulosa, Medorippe lanata, Nanocassiope melanodac- 

26. Ghana. 04°57'N, 01°16'W to 04°59'N, 01°16.5'W, 27 tyla, Parthenope notialis, Phyllodorippe armata, Pisa carini- 

m, shell bottom (scallops), 6' OT, 28 May 1964: mana, Stenorhynchus lanceolatus, Stemodromia spinirostris 

Dromia monodi, Ethusa vossi, Ilia spinosa, Inachus biceps, 47. Ivory Coast. 05°04.5'N, 04°51.5'W, 37 m, bottom 

Parthenope massena, Portunus inaequalis, Stenorhynchus lan- with Jullienella, 6' OT, 31 May 1964: Calappa pelii, 

ceolatus 

Calappa rubroguttata, Ebalia affinis, Ethusa vossi, Leopoldius 

27. Ghana. 04°48'N, 01°42'W to 04°49'N, 01°47'W, 33 pisifer, Machaerus oxyacantha, Macropodia gilsoni, Macro- 

m, 6' OT, 28 May 1964: Ebalia tuberculata, Pilumnus podia spinulosa, Medorippe lanata, Nanocassiope melanodac- 

perrieri, Pisa carinimana, Portunus inaequalis 

tyla, Neodorippe armata, Philyra laevidorsalis, Phyllodorippe 

28. Ghana. 04°40'N, 02°00'W to [04°39'N, 02'02'W], 49- armata, Pisa carinimana, Portunus inaequalis, Stemodromia 

53 m, 6' OT, 28 May 1964: Capartiella longipes, Inachus spinirostris 

angolensis, Machaerus oxyacantha, Macropodia gilsoni, Ma- 48. Ivory Coast. [05°05'N, 04°59.5'W], 22 m, 6' OT, 31 

cropodia spinulosa, Medorippe lanata, Pseudomyra mbizi, May 1964: Cronius ruber, Ilia spinosa, Machaerus oxyacan- 

Raninoides bouvieri, Stenorhynchus lanceolatus, Stemodromia tha, Macropodia spinulosa, Phyllodorippe armata, Stenorhyn- 

spinirostris 

chus lanceolatus 

29. Ghana. [04°38'N, 02°02'W to 04°36'N, 02°00'W], 49. Ivory Coast. 05°00'N, 05°00'W to 04°59'N, 05°00'W, 

58-60 m, 40' OT, 28 May 1964: Inachus nanus, Pseudo- 73-77 m, 6' OT, 31 May 1964: Calappa pelii, Ethusa 

myra mbizi. 

rugulosa, Inachus angolensis, Inachus nanus, Monodaeus rouxi, 

30. Ghana. 04°46'N, O2°3O'W to 04°45'N, 02°33'W, 61- Phyllodorippe armata, Pseudomyra mbizi, SoUnolambrus noor- 

64 m, coral, 40' OT, 28 May 1964: Macropodia gilsoni, dendei 

Medorippe lanata, Stenorhynchus lanceolatus 

50. Ivory Coast. 04°58'N, 05°00'W to 04°57'N, 05°01'W, 

32. Ghana. 04°37'N, O2°32'W to 04°38'N, 02°35'W, 110 

128-192 m, 6' OT, 3 May 1964: Ethusa rugulosa, Inachus 

m, 40' OT, 28 May 1964: Macropodia gilsoni, Pseudomyra angolensis, Macropodia gilsoni, Monodaeus rouxi, Parthenope 

mbizi 

notialis, Pseudcmyra mbizi, SoUnolambrus noordendei 

34. Ghana. 03°53'N, 02°33'W to [03°47'N, 02°33'W], 51. Ivory Coast. 04°56'N, 05°01'W to 04°56.6'N, 05°- 

1948-1984 m, mud, Blake Trawl, 29 May 1964: Ethu- 03'W, 329-494 m, 6' OT, 31 May 1964: Acanthocarpus 

sina beninia 

brevispinis, Bathynectes piperitus, Carcinoplax bamardi, Ger- 

41. Ivory Coast. 04°47'N, O3°33'W to 04°47'N, 03°35'W, yon maritae 

641-842 m, 6' OT, 30 May 1964: Ethusa rosacea, Geryon 

maritae 

42. Ivory Coast. 05°02.5'N, 03°49.5'W to 05°05'N, 03°- 

59. Ivory Coast. 04°57.5'N, 05°- 22W to 04°57'N, 05°- 

30'W, 55-64 m, mud with dense branched Foraminifera, 

6' OT, 1 Jun 1964: Macropodia gilsoni, Parthenope 

52^, 62-75 m, mud with brown, branched Forami- notialis, Pseudomyra mbizi, SoUnolambrus noordendei 

nifera, 6' OT, 30 May 1964: Atlantotlos rhombifer, Ca- 60. Ivory Coast. 04°55'N, 05°34.5'W to 04°- 54'N, 05°lappa 

pelii, Capartiella longipes, Ilia spinosa, Inachus ango- 37^, 79-82 m, coral or rock, 6' OT, 1 Jun 1964: 

lensis, Inachus nanus, Macropipus rugosus, Macropodia gil- Capartiella longipes, Ethusa rugulosa, Inachus nanus, Macrosoni, 

Macropodia straeleni, Medorippe lanata, Nanocassiope podia hesperiae, Macropodia straeleni, Medorippe lanata, 

melanodactyla, Parthenope notialis, Pisa carinimana, Pseudo- Parthenope notialis, Pisa carinimana, Pseudomyra mbizi, Somyra 

mbizi, SoUnolambrus noordendei, Stenorhynchus lanceo- Unolambrus noordendei 

latus 

62. Ivory Coast. 04°45'N, 06°13.5'W to 04°44'N, 

43. Ivory Coast. Surface tow during Sta. 42, 0.5 m net, 30 06° 16^, 46 m, brown, branched and foliate Fora- 

May 1964: Callinectes pallidus 

minifera, 6' OT, 1 Jun 1964: Calappa pelii, Ilia spinosa, 

44. Ivory Coast. 05°05'N, 04°00'W to 05°04'N, 04°02'W, Inachus angolensis, Machaerus atlanticus, Macropodia gilsoni, 

403-586 m, hard dark gray mud, 6' OT, 30 May Medorippe lanata, Monodaeus rouxi, Parthenope notialis, Pisa 

1964: Balhynectes piperitus, Carcinoplax bamardi, Ethusa carinimana, Pseudomyra mbizi, Raninoides bouvieri, Stemo- 

rosacea, Geryon maritae 

dromia spinirostris 

45. Ivory Coast. 05°05'N, 04°04.5'W to 05°06'N, 04°- 

06'W, 73-97 m, 40' OT, 30 May 1964: Macropodia 

63. Ivory Coast. 04°35'N, 06°40'W to 04°35'N, 06°4l'W, 

64 m, sandy mud with shells, 6' OT, 2 Jun 1964:


Atlantotlos rhombifer, Ilia spinosa, Inachus angolensis, Ma230. 

Nigeria. 06° 1 l'N, 03°36'E to 06° 10'N, 03°38'E, 82cropodia 

gilsoni, Parthenope notialis, Pseudomyra mbizi, Ster- 97 m, hard ground, with gorgonians, coral, rock, 40' 

nodromia spinirostris 

OT, 11 May 1965: Macropodia hesperiae, Pisa armata 

64. Ivory Coast. 04°23'N, 07°06.5'W to 04°22'N, 232. Nigeria. 05°56'N, 04°27'E to 05°54'N, 04°27'E, 101- 

07°08.5'W, 68 m, 6' OT, 2 Jun 1964: Calappa pelii, 

132 m, green mud, 40' OT, 11 May 1965: Pseudomyra 

Inachus angolensis, Inachus nanus, Macropipus rugosus, Ma- mbizi 

cropodia gilsoni, Medorippe lanata, Parthenope notialis, Pisa236. 

Nigeria. 05°20'N, 04°45'E to 05° 19'N, 04°48'E, 101carinimana, 

Pseudomedaeus africanus, Raninoides bouvieri 128 m, coral ground, rough, 40' OT, 12 May 1965: 

65. Ivory Coast. 04°15'N, 07°32'W to 04°12'N, Pseudomyra mbizi 

O7°35.5'W, 46-49 m, 40' OT, 2 Jun 1964: Macropipus 237. Nigeria. 05°19'N, 04°48'E to O5°O7'N, 04°55'E, 101 

rugosus, Macropodia hesperiae, Macropodia spinulosa, Ma- m, 10' OT, 12 May 1965: Calappa pelii, Macropodia 

cropodia straeleni, Medorippe lanata, Stenorhynchus lanceola- gilsoni, Monodaeus rouxi, Pseudomyra mbizi, Solenolambrus 

tus 

noordendei 

68. Liberia. 04°23'N, 08°05.5'W to 04°24'N, 08°07.5'W, 239. Nigeria. 04°56'N, 05°00'E to 04°54'N, 05°05'E, 73 m, 

70 m, broken shell, 6' OT, 3 Jun 1964: Achaeus foresti, 10' OT, 13 May 1965: Atlantotlos rhombifer, Capar- 

Atlantotlos rhombifer, Calappa pelii, Calypsachaeus calypso, tiella longipes, Inachus angolensis, Inachus nanus, Macropodia 

Capartiella longipes, Ebalia affinis, Ethusa vossi, Eurynome gilsoni, Macropodia straeleni, Monodaeus rouxi, Pseudomyra 

aspera, Heterocrypta maltzami, Homola barbata, Ilia spinosa, mbizi 

Inachus nanus, Macropipus rugosus, Macropodia gilsoni, 240. Nigeria. 04°44'N,05°17'E to04°41'N,05°19'E, 37 m, 

Macropodia straeleni, Medorippe lanata, Monodaeus rouxi, 10' OT, 13 May 1965: Capartiella longipes 

Parthenope notialis, Pseudomyra mbizi, Raninoides bouvieri, 241. Nigeria. 04°35'N, 05°18'E to 04°34'N, 05°19'E, 59- 

Solenolambrus noordendei, Stemodromia spinirostris 

63 m, mud and shell, 10' OT, 13 May 1965: Atlantotlos 

69. Liberia. 04°29.5'N, 08°06'W to 04°29.5'N, rhombifer, Calappa pelii, Capartiella longipes, Homola bar- 

08°07.5'W, 29 m, coral or rock, 6' OT, 3 Jun 1964: bata, Inachus angolensis, Machaerus oxyacantha, Macropipus 

Inachus nanus, Pisa carinimana, Pseudomyra mbizi 

rugosus, Macropodia gilsoni, Medorippe lanata, Portunus 

70. Liberia. 04°30'N, 08°09'W to 04°29.5'N, 08°09'W, validus, Pseudomyra mbizi, Raninoides bouvieri 

33 m, branched Foraminifera, 6' OT, 3 Jun 1964: 245. Nigeria. 04°32'N, 05°07'E to °31'N, 05°13'E, 64- 

Apiomithrax violaceus, Ethusa vossi, Nanocassiope melano- 119 m, mud, 40' OT, 13 May 1965: Pseudomyra mbizi 

dactyla, Paractaea margaritaria, Parthenope massena, Pisa246. 

Nigeria. 04° 13'N, 05°30'E to 04° 10'N, 05°33'E, 37 m, 

carinimana, Stenorhynchus lanceolatus 

40' OT, 13 May 1965; Ilia spinosa, Portunus inaequalis, 

73. Liberia. ?04°38'N, ?09°20'W to 04°40'N, 09°20'W, Pseudomyra mbizi, Stenorhynchus lanceolatus 

311-366 m, 40' OT, 4 Jun 1964: Bathynectes piperitus 248. Nigeria. 04°03'N, 05°4l'E to 04°07'N, O5°4O'E, 33 m, 

74. Liberia. 04°20'N, 09°26'W to 04°30'N, 09°22'W, 

10' OT, 13 May 1965: Achaeus turbator, Atlantotlos 

641-733 m, 40' OT, 4 Jun 1964: Ethusa rosacea, Geryon rhombifer, Calappa rubroguttata, Calypsachaeus calypso, Ca- 

maritae 

partiella longipes, Dromia monodi, Ebalia affinis, Ebalia 

82. Liberia. 04°57'N, 09°30'W to 04°58'N, 09°32'W, tuberculata, Ethusa vossi, Heterocrypta maltzami, Ilia spinosa, 

146-150 m, 40' OT, 5 Jun 1964: Macropodia macrocheles Inachus biceps, Leopoldius pisifer, Macropodia spinulosa, 

83. Liberia. 04°59'N, 09°37'W to 04°57.5'N, 09°33'W, Medorippe lanata, Parthenope massena, Phyllodorippe armata, 

156-220 m, 40' OT, 5 Jun 1964: Macropipus rugosus 

Pilumnus perrieri, Pisa carinimana, Portunus inaequalis, 

224. Nigeria. Lagos, 06°28'N, 03°23'E, shore, sand beach, Stenorhynchus lanceolatus 

ichthyocide, 9 May 1965: Ocypode cursor, Pilumnopeus 250. Nigeria. 04°06'N, 05°58'E to 04°02'N, 06°04'E, 24 m, 

africanus 

brackish water, mud, 10' OT, 14 May 1965: Callinectes 

225. Nigeria. Lagos harbor, 06°28'N, 03°23'E, shore at pallidus, Dromia monodi, Machaerus oxyacantha, Nanocas- 

dock, 9 May 1965: Plagusia depressa 

siope melanodactyla, Phyllodorippe armata, Parthenope mas- 

226. Nigeria. Lagos harbor, 06°28'N, O3°23'E, surface at 

sena, Portunus inaequalis 

dock, night light, 9 May 1965: Callinectes pallidus 251. Nigeria. O4°O3'N, 06°03'E to 04°04'N, 06°04'E, 27 m, 

227. Nigeria. Lagos harbor, 06°28'N, O3°23'E, shore, on 

mud, 10' OT, 14 May 1965: CallinecUs pallidus, Ma- 

rocks and seawall, by hand, 10 May 1965: Goniopsis chaerus oxyacantha, Phyllodorippe armata 

pelii, Pachygrapsus gracilis 

252. Nigeria. 04°04'N, 06° 18'E to 04°05'N, 06°22'E, 30 m, 

228. Nigeria. Lagos harbor, 06°28'N, 03°23'E, surface at 

mud, 10' OT, 14 May 1965: Callinectes pallidus, Ilia 

dock, 10 May 1965: Callinectes pallidus 

spinosa, Machaerus oxyacantha, Phyllodorippe armata, Por- 

229. Nigeria. Lagos harbor, 06°28'N, O3°23'E, surface at tunus inaequalis, Portunus validus, Stenorhynchus lanceolatus 

dock, dip net, 10 May 1965: Callinectes amnicola, Calli- 253. Nigeria. 04°04'N, 06°35'E to 04°03'N, 06°38'E, 33nectes 

pallidus 

40 m, mud, 10' OT, 14 May 1965: Calappa pelii,

340 

Leopoldius pisifer, Parthenope massena, Phyllodorippe armata, 

Stenorhynchus lanceolatus 

254. Nigeria. 03°50'N, 07°08'E to 03°51'N, 07° 12'E, 148- 

174 m, 40' OT, 14 May 1965: Pseudonym mbizi 

255. Nigeria. 03°49'N, 07°38'E to 03°48'N, 07°42'E, 264- 

269 m, 40' OT, 14 May 1965: Acanthocarpus brevispinis, 

Carcinoplax bamardi, Inachus grallator, Macropodia macrocheles 

256. Nigeria. 03°45'N, 08°03'E to 03°45'N, 08°02'E, 409- 

485 m, 40' OT, 14 May 1965: Acanthocarpus brevispinis, 

Carcinoplax bamardi 

257. Fernando Poo. 03°45'N, 08°48'E, shore, ichthyocide, 

15 May 1965: Callinectes marginatus, Callinectes pallidus, 


Eurypanopeus blanchardi, Pachygrapsus transversus, Xanthod- Pilumnus stebbingi, Sakaila africana 

ius inaequalis inaequalis 

316. Nigeria. Lagos, 06°28'N, 03°23'E, shore, sand beach, 

258. Fernando Poo. 03°45'N, 08°48'E, shore, ichthyocide, 

ichthyocide, 28 May 1965: Ocypode cursor, Pachygrapsus 

15 May 1965: Callinectes pallidus, Epixanthus hellerii, 

Eurypanopeus blanchardi, Grapsus grapsus, Pachygrapsus 

transversus, Percnon gibbesi, Xanthodius denticulatus, Xan- 

transversus 

thodius inaequalis inaequalis 

Geronimo COLLECTIONS 

259. Cameroon. 03°53'N, 08°53'E to 03°51'N, 08°54'E, 

59 m, mud and broken shell, 10' OT, 16 May 1965: 

Calappa pelii, Inachus angolensis, Macropipus rugosus, Macropodia 

gilsoni, Pseudomyra mbizi, Raninoides bouvieri 

260. Cameroon. 03°45'N, 09°05'E to 03°43'N, 09°10'E, 

46 m, 10' OT, 16 May 1965: Atlantotlos rhombifer, 

Calappa pelii, Calypsachaeus calypso, Ilia spinosa, Inachus 

minus, Macropipus rugosus, Macropodia gilsoni, Medorippe 

lanata, Parthenope notialis, Raninoides bouvieri 

271. Annobon. 01°25'S, 05°38'E, NE coast between Punta 

Yoyo and Punta Pedrinha, shore, ichthyocide, 19 May 

1965: Acanthonyx minor, Grapsus grapsus, Nanocassiope 

melanodactyla, Pachygrapsus transversus, Percnon gibbesi, 

Plagusia depressa, Sirpus gordonae, Xanthodius denticulatus, 

Xanthodius inaequalis inaequalis 

273. Annobon. 01°24'S, O5°37'E, N coast, near Islote Piramide, 

shore, ichthyocide, 19 May 1965: Acanthonyx 

minor, Domecia acanthophora africana, Grapsus grapsus, Ocypode 

cursor, Pachygrapsus transversus, Plagusia depressa 

275. Annobon. 01°24'S, O5°37'E to 01°24'S, 05°38'E, 9- 

69 m, rubble of coralline algae, dredge, 20 May 1965: 

Cataleptodius Jloridanus, Cronius ruber, Dynomene filholi, 

Euryozius pagalu, Globopilumnus stridulans, Herbstia nitida, 

Nanocassiope melanodactyla, Nanopilumnus boletifer, Paractaea 

margaritaria, Paractaea rufopunctata africana, Pisa calva, 

Ranilia constricta, Stenorhynchus lanceolatus, Xanthodius denticulatus 

278. Annobon. 01°24'S, 05°37'E, shore [collection made in 

transit to Crater Lake], 20 May 1965: Ocypode cursor 

281. Annobon. 01°24'S, O5°37'E, shore, ichthyocide, 20 

May 1965: Callinectes marginatus, Grapsus grapsus, Ocy- 

pode cursor 

282. Annobon. 01°28'S, 05°36'-37'E to 01°29'S, 05°36'E, 

18-37 m, nodular coralline algae, dredge, 21 May 

1965: Domecia acanthophora africana, Dynomene filholi, 

Euryozius pagalu, Globopilumnus stridulans, Glyptoxanthus 

angolensis, Herbstia nitida, Microcassiope rufopunctata, Nanocassiope 

melanodactyla, Paractaea margaritaria, Paractaea 

rufopunctata africana, Pisa calva 

283. Annobon. 01°29'S, 05°35'E, 51-55 m, nodular coral- 

line algae, dredge, 21 May 1965: Dynomene filholi, 

Ebalia tuberculata, Euryozius pagalu, Herbstia nitida, Homola 

barbata, Nanocassiope melanodactyla, Paractaea margaritaria, 

Parthenope expansa, Parthenope massena, Pilumnus 

stebbingi, Pisa calva 

284. Annobon. 01°30'S, 05°36'E, 73 m, black basaltic 

rocks, dredge, 21 May 1965: Nanocassiope melanodactyla, 

Collections made by National Marine Fisheries 

Service vessel Geronimo off Gabon in 1963. 

179. Off Gabon River, OO°O2.2'S, 08°50.2'E, 161 fm (293 

m), 31 Aug 1963: Acanthocarpus brevispinis, Bathynectes 

piperitus, Carcinoplax bamardi 

185. 00°32'S, 08°42'E, 110 fm (200 m), 1 Sep 1963: 

Pseudomyra mbizi 

187. OO°32'S, 08°40'E, 165 fm (300 m), 1 Sep 1963: 


191. 01°28'S, 08°24.5'E, 165 fm (300 m), 3 Sep 1963: 

Bathynectes piperitus, Geryon maritae 

197. 01°30'S, 08°27.5'E, 110 fm (200 m), Sep 1963: Macropodia 

macrocheles 

198. 01°28'S, 08°24.5'E, 165 fm (300 m), 3 Sep 1963: 

Acanthocarpus brevispinis, Bathynectes piperitus, Carcinoplax 

bamardi, Geryon maritae 

199. 01°26.4'S, 08°24'E, 220 fm (400 m), 3 Sep 1963: 

Bathynectes piperitus 

202. 02°00'S, 08°55'E, 55 fm (100 m), 4 Sep 1963: Acanthocarpus 

brevispinis, Macropipus rugosus 

203. 02°01'S, 08°50.5'E, 110 fm (200 m), 4 Sep 1963: 

Acanthocarpus brevispinis, Bathynectes piperitus, Carcinoplax 

bamardi, Geryon maritae 

206. 02°00'S, 08°46.5'E, 250-335 fm (455-610 m), 4 Sep 

1963: Bathynectes piperitus 

211. 02°32'S, 09°05'E, 55 fm (100 m), 5 Sep 1963: Atelecyclus 

rotundatus, Euchirograpsus liguricus, Macropipus rugosus, 

Pseudomedaeus africanus 

212. 02°30'S, 08°58'E, 110 fm (200 m), 5 Sep 1963: Calappa 

pelii 

213. 02°31'S, 08°51'E, 165 fm (300 m), 5 Sep 1963: Bathynectes 

piperitus, Macropipus rugosus


214. 02°30'S, 08°52'E, 300 fm (546 m), 5 Sep 1963: Bathynecles 

piperitus 

220. 03°02'S, 09°21'E, 165 fm (300 m), 6 Sep 1963: Bathynectes 

piperitus 

228. 03°31'S, 09°53'E, 165 fm (300 m), 7 Sep 1963: Macropipus 

rugosus 

235. 04°01'S, 10°35'E, 55 fm (100 m), 8 Sep 1963: Calappa 

pelii, Macropodia gilsoni, Parthenope notialis, Sakaila africana 

247. 04°38.4'S, ll°01.2'E, 220 fm (400 m), 9 Sep 1963: 

Bathynectes piperitus 

Undaunted COLLECTIONS 

Collections made by National Marine Fisheries 

Service vessel Undaunted off Angola and South- 

West Africa (below 17°15'S) in 1968. 

94. 16°27'S, 11°35'E, 90 m, 18 Mar 1968: Goneplax rhomboides, 

Inachus angolensis, Macropipus australis 

95. 16°37'S, 11°22'E, 126 m, 18 Mar 1968: Calappa pelii, 

Goneplax rhomboides, Macropipus australis, Macropodia gilsoni, 

Macropodia spinulosa, Parthenope notialis 

96. 16°41'S, 11°21'E, 162 m, 18 Mar 1968: Calappa pelii, 

Eurynome aspera, Inachus angolensis, Inachus grallator, Macropipus 

australis, Parthenope notialis, Pisa armata, Solenolambrus 

noordendei 

102. 17°O2'S, ll°40'E, 54 m, 24 Mar 1968: Calappa pelii, 

Medorippe lanata 

103. 17°O6'S, 11°35'E, 90 m, 24 Mar 1968: Calappa pelii, 

Inachus angolensis, Macropipus australis, Medorippe lanata, 


104. 17°09'S, ll°30'E, 126 m, 24 Mar 1968: Macropipus 

australis 

105. 17°13'S, 11 O 27'E, 155 m, 24 Mar 1968: Macropipus 

australis 

106. 17° 18'S, 11 °24'E, 225 m, 24 March 1968: Euchirograpsus 

liguricus, Macropipus australis, Monodaeus couchii 

107. 17°23'S, 11 °20'E, 359 m, 24 Mar 1968: Euchirograpsus 

liguricus, Geryon maritae 

111. 10°36'S, 13°12'E, ca 366 m, 12 Apr 1968: Bathynectes 

piperitus

Localities from the literature or from specimenassociated 

data cited in the species accounts are 

listed here, alphabetically by country, along with 

coordinates. Spellings and coordinates are from 

gazetteers of the United States Board on Geographic 

Names. Alternate spellings and coordinates 

from sources other than those gazetteers are 

given in brackets. Localities that we have not 

been able to locate are identified in the text by a 

question-mark in brackets, [?], and are cross-referenced 

in this gazetteer by species name. Specific 

localities identified in the text as "near" another 

locality (e.g., Boulbinet, near Conakry) or as in 

another locality, (e.g., Samba, Luanda) are not 

listed separately here if they could not be located 

in the Board on Geographic Names gazetteers. 

Localities in the literature or from specimen labels 

originally accompanied by coordinates are not 

repeated here; in the list, coordinates added by 

us are set off in brackets. We have anglicized 

country names but not names for localities within 

countries. We have used older names for two of 

the offshore islands of the Gulf of Guinea, now 

part of Equatorial Guinea: Annobon, instead of 

Pagalu, and Fernando Poo, instead of Macias 

Nguema Biyogo. 

Other sources, which have been helpful in identifying 

West African localities either by listing 

coordinates or by giving maps, include: Forest 

and Games, 1960 (Morocco); Guinot and Ribeiro, 

1962 (Cape Verde Islands, Angola); 

Longhurst, 1958 (Sierra Leone and Guinea 

Shelf); Rathbun, 1921 (a list of West African 

localities, mostly in the Congo and Zaire); Rossignol, 

1962 (localities between Nigeria and 

southern Angola); Sourie, 1954a (Senegal, especially 

around Dakar); and Tiirkay, 1976b (Madeira). 

Appendix II: Gazetteer 

342 

Angola 

Baia do Ambriz 

Baia de Ambrizete 

Baia de Benguela 

Baia da Caota, Benguela 

Baia dos Elefantes 

Baia Farta, Benguela 

Baia do Lobito 

Baia de Luanda 

Baia de Mocamedes 

Baia de Porto Amboim 

Baia de Santa Marta 

Baia dos Tigres 

Baia das Vacas [Ponta das 

Vacas] 

Benguela 

Cacuaco 

Chiloango 

Egito Praia 

Ilha de Luanda 

Luanda [St. Paul do 

Loanda] 

Lucira 

Mocamedes 

Morro da Cruz 

Musserra [Mussera, 

Massera] 

Mussulo Grande 

Ponta da Caruita 

Ponta da Moita Seca 

Ponta do Morro [Cap 

Morro?] 

Ponta de Sao Jose 

Ponta do Sombreiro 

Porto Alexandre 

Porto Amboim 

Praia Amelia, Mocamedes 

Praia das Conchas, Mocamedes 

Quicembo [Kinsembo] 

Rio Chiloango 

Rio Cuanza 

Rio Cunene [Kunene River] 

Santo Antonio do Zaire [San 

Antonio, Saint-Antoine] 

07° 50'S, 13 

07° 

12° 

12° 

13° 

12° 

12° 

08° 

15° 

10° 

13° 

16° 

12° 

C 

13'S, 12 C 

35'S, 13 C 

36'S, 13 C 

13'S, 12° 

36'S, 13' 

20'S, 13 C 

47'S, 13 C 

10'S, 12' 

42'S, 13' 

51'S, 12 C 

'06'E 

'51'E 

'21'E 

'16'E 

'44'E 

'13'E 

'34'E 

'16'E 

"08'E 

'45'E 

'28'E 

( 

38'S, 11' 46'E 

( 

37'S, 13' 14'E 

12° 35'S, 13 

08° 

05° 

11° 

08° 

08° 

C( 

25'E 

47'S, 13'>22'E 

12'S, 12''08'E 

59'S, 13''46'E 

48'S, 13''13'E 

48'S, 13''14'E 

13° 51'S, 12''31'E 

15° 10'S, 12'WE 

08° 57'S, 13''04'E 

07° 29'S, 12''58'E 

08° ll'S, 13' 

D 

12° '35'S, 13' 16'E 

06° 07'S, 12''16'E 

10° 45'S, 13'>43'E 

12 C 

12c 

15 C 

10 c 

15 C 

15 C 

07 c 

05 c 

09 c 

17 C 

06 c 

5 17'E 

3 12'E 

36'S, 13' 

3 

35'S, 13' 18'E 

3 

49'S, 11' 53'E 

'44'S, 13'°44'E 

"12'S, 12'°06'E 

'07'S, 12°07'E 

'44'S, 13°03'E 

'12'S, 12°07'E 

'19'S, 13°08'E 

'20'S, 11 °50'E 

'07'S, 12°18'E

NUMBER 306 

Annobon [Pagalu, Equatorial 

Guinea] 

Isla Tortuga 

Islote Piramide 

Punta Pedrinha 

Punta Yoyo 

San Antonio 

Santa Cruz 

Ascension Island 

Azores 

Ilha do Corvo 

Ilha do Faial [Fayal] 

Almoxarife. See Acanthonyx 

brevifrons 

Caldeirinhas [Caldeira Inferno] 

Feteira 

Horta 

Pasteleiro 

Ilha das Flores 

Ilha da Muda. See Dromia 

marmorea 

Ilha do Pico 

Madalena 

Ilha de Sao Miguel 

Caloura 

Ponta Delgada 

Ponta da Galera 

Ilha Terceira 

Ponta Sao Diego. See 

Dromia marmorea 

Ilheu da Praia 

Rochas dos Capelinhos [Volcano 

Capelinhos] 

Cabinda 

Cabinda 

Chinchoxo [Tschintschotscho, 

Quinchoxo] 

Landana 

Quila [River] 

Cameroon 

Batanga 

Bibundi 

Douala 

Kribi 

Souellaba [Souelaba] 

Wouri River 

Yaounde 

Canary Islands 

Estrecho de la Bocaina 

Isla de Fuertaventura 

Puerto Cabras 

Punta Morro Jable [Ponta 

da Matorra, Punta del 

Mattorral] 

01°25'S, 05°36'E 

01°24'S, 05°38'E 

01°24'S, O5°37'E 

[01°24'35"S, O5°37'25"E] 

[01°24'45"S, 05°37'35"E] 

O1°27'S, 05°37'E 

O1°27'S, 05°37'E 

O7°57'S, 14°22'W 

38°30'N, 28°0O'W 

39°42'N, 31°06'W 

38°34'N, 28°42'W 

38°30'N, 28°37'W 

38°31'N, 28°41'W 

38°32'N, 28°38'W 

38°31'N, 28°38'W 

39°26'N, < i\°\3'^M 

38°28'N, 28°20'W 

38°32'N, 28°32'W 

37°47'N, 25°30'W 

37°42'N, 25°30'W 

37°44'N, 25°40'W 

37°42'N, 25°31'W 

38°43'N, 27°13'W 

39°03'N, 27°57'W 

38°36'N, 28°50'W 

05°33'S, 12°12'E 

[05°09.24'S, 12°03.75'E] 

05°13'S, 12°08'E 

05°58'S, 14°47'E 

[ca 02°54'N, 09°55'E] 

04°13'N, 08°59'E 

04°03'N, 09°42'E 

02°57'N, 09°55'E 

03°49'N, 09°33'E 

04°06'N, 09°43'E 

03°52'N, 11°31'E 

28°00'N, 15°3O'W 

28°47'N, 13°50'W 

28°20'N, 14°00'W 

28°29'N, 13°51'W 

28°02'N, 14°20'W 

Isla de la Gomera 

Isla de Gran Canaria 

Isla de Lanzarote 

Arrecife 

Playa Quemada 

Isla de La Palma 

Santa Cruz de La Palma 

Isla de Tenerife 

Ensenada de Crist ianos 

Playa de los Abrigos 

Puerto de la Cruz 

Puerto de Orotava 

Cape Verde Islands 

Baixo Joao Leitao 

Boa Vista, Ilha da 

Ilheu de Sal Rei 

Ponta Rodrigo 

Porto de Sal Rei 

Ribeira do Rabil 

Branco, Ilheu 

Brava, Ilha 

Baia de Faja di Agua 

[Porto da Faja] 

Ponta Garbeiro. See Dorrucia 

acanthophora africana 

Porto da Furna 

Fogo, Ilha do 

Ponta da Areia 

Ilheu Luis Carneiro 

Maio, Ilha de 

Porto Ingles 

Sal, Ilha do 

Baia da Murdeira [Mordeira] 

Pedra Lume 

Santo Antao, Ilha de 

Baia do Monte Trigo 

Ponta do Esbarradeiro 

Ponta da Praia Formosa 

Ponta do Sol 

Porto dos Carvoeiros [Baio 

do Porto Novo] 

Santa Luzia, Ilha de 

Sao Nicolau, Ilha de 

Tarrafal 

Sao Tiago, Ilha de [Santiago] 

Baia de Santa Clara 

Porto da Praia [La Praya, 

Praia] 

Porto de Sao Francisco 

Tarrafal 

Sao Vicente, Ilha de 

28°06'N, 17°08'W 

28°00'N, 15°36'W 

29°00'N, 13°40'W 

28°57'N, 13°32'W 

28°54'N,13°43'W 

28°40'N, 17°52'W 

28°41'N, 17°45'W 

28°19'N, 16°34'W 

28°02'N, 16°42'W 

28°01'N, 16°35'W 

28°24'N, 16°33'W 

28°46'N, 17°45'W 

16°00'N, 24°00'W 

15°48'N, 23°!!^ 

16°05'N, 22°50'W 

16°10'N,22°56'W 

16°12'N, 22°43'W 

16°10'N, 22°56'W 

16°09'N, 22°59'W 

16°39'N, 24°41'W 

14°52'N, 24°43'W 

14°52'N, 24°45'W 

14°53'N, 24°41'W 

14°55'N, 24°25'W 

14°53'N, 24°31'W 

14°58'N, 24°40'W 

15°15'N,23 o 10'W 

15°08'N,23°13'W 

16°45'N, 22°55'W 

16°41'N, 22°57'W 

16°46'N, 22°54'W 

WOb'W, 25°10'W 

17°OrN,25°2O'W 

16°55'N, 25°14'W 

16°55'N, 25°13'W 

17°12'N, 25°06'W 

17°O2'N, 25°04'W 

16°46'N, 24°45'W 

16°35'N, 24°15'W 

16°34'N, 24°22'W 

15°05'N, 23°40'W 

15°01'N, 23°45'W 

14°54'N,23°31'W 

14°58'N, 23°28'W 

15°17'N, 23°46'W 

16°50'N, 25°00'W 

343

344 

Baia da Fateixa 

Baia das Gat as 

Ilheu Raso 

Ponta da Calheta [Baia de 

Calheta] 

Porto Grande 

Praia da Matiota [Baia da 

Matiota] 

Conception Bank 

Congo 

Bahua, See Cardisoma armatum 

Baie de Loango 

Baie de Pointe-Noire 

Banga [Banda] 

Djeno 

Loango 

Loya River 

Pointe Indienne 

Pointe Kounda 

Pointe-Noire 

Songololo River [Songolo 

River] 

Dahomey 

Cotonou 

Grand-Popo 

Lac Nokoue 

Zogbo, NW of Cotonou on 

Lac Nokoue 

Fernando Poo [Macias Nguema 

Biyogo, Equatorial Guinea] 

Mongola 

San Carlos 

Gabon 

Baie du Cap Lopez 

Barre des Portugais 

Cap Lopez 

Cap Santa Cfara [Pointe 

Santa Clara] 

Crique Banjia [Banya] 

Ivindo River 

Libreville 

Mayumba [Mayoumba] 

Nyanga 

Ogooue [stream] 

Owendo 

Pointe Banda 

Pointe Claire 

Pointe Gombe 

Pointe Panga 

Port-Gentil 

Sette Cama 

Gambia 

Bathurst 

Gambia River 

16°52'N, 25°04'W 

16°12'N, 22°43'W 

16°37'N, 24°36"W 

16°47'N, 24°58'W 

16°53'N, 25°O1'W 

16°53'N, 25°00'W 

29°30'N, 12°45'W 

04°36'S, 11°44'E 

04°47'S, 11°51'E 

04°34'S, 12°23'E 

04°55'S, 11°57'E 

04°39'S, 11°48'E 

04°49'S, 12°52'E 

04°40'S, 11°47'E 

04°ll'S, 11°23'E 

04°48'S, 11°51'E 

04°45'S, 11°51'E 

06°21'N, 02°26'E 

06°17'N, 01°50'E 

06°26'N, 02°27'E 

[06°24'N, 02°25'E] 

O3°3O'N, 08°42'E 

03°46'N, 08°44'E 

03°27'N, 08°33'E 

00°40'S, 08°50'E 

01°16'S, 09°00'E 

00°37'S, 08°43'E 

00°30'N, 09°19'E 

00°14'N, 09°40'E 

00°09'S) 12°09'E 

OO°23'N, 09°27'E 

03°25'S, 10°39'E 

02°59'S, 10°17'E 

00°49'S, 09°00'E 

00°17'N, 09°30'E 

03°46'S, ll°00'E 

01°08'S)09°26'E 

00°18'N, 09°18'E 

03°15'S,.10°32'E 

00°43'S, 08°47'E 

02°32'S, 09°45'E 

13°27'N, lG'^'W 

13°28'N, 16°34'W 


Gunjur 

Ghana 

Accra 

Ada 

Angaw Lagoon 

Ankobra River [Ancobra] 

Apam 

Axim 

Baya River 

Butre [Boutry, Butry] 

Chorkor, near Accra 

Christiansborg 

Densu 

Denu 

Dixcove 

Elmina [St. George 

d'Elmina] 

Hwini River 

Komenda 

Kumasi 

Labadi 

Ningo 

Prampram 

Sakumo Lagoon 

Sekondi 

Shama Bay 

Takoradi 

Tema harbor 

Tenkpobo[Tenpobo] 

Teshi [Teshie] 

Volta River delta 

Winneba 

Guinea 

Conakry 

Fotoba 

lie Blanche 

lie de Corail 

lie Kassa [Cassa] 

lies de Los 

lie Marara 

lie Roume 

lie Tamara 

Matakong 

Rio Pongo 

Tanene 

Guinea-Bissau [Portuguese 

Guinea] 

Arquipelago dos Bijagos 

[Bissagos Islands] 

Bissau 

Ilha de Bissau 

Ilha de Bubaque 

Ilha Caravela 

Ilha de Rubane [Rouban] 

l'N, 16°46'W 

05°33'N, 00°13'W 

05°46'N, OO°37'E 

05°48'N, OO°47'E 

04°53'N, O2°17'W 

05°17'N, 00°44'W 

04°52'N, 02°14'W 

[05°05'N ,01 

04°49'N, 

[05°33'N 

O5°33'N, 

05°31'N, 

06°05'N, 

04°48'N, 

05°05'N, 

o 22'W] 

01°55'W 

, 00°15'W] 

00°ll'W 

00°19'W 

01°08'E 

O1°57'W 

01°21'W 

04°53'N, 

05°03'N, 

06°41'N, 

05°33'N, 

06°03'N, 

05°42'N, 

05° 3 l'N, 

04°56'N, 

05°00'N, 

04°53'N, 

05°38'N, 

05°43'N, 

05°35'N, 

05°55'N, 

05°20'N, 

09° 3 l'N, 

09°30'N, 

09°26'N, 

09°26'N, 

09°29'N, 

09°30'N, 

10°02'N, 

09°28'N, 

09°29'N, 

09° 16'N, 

10°03'N, 

10°03'N, 

01°47"W 

01°29'W 

O1°37'W 

00°09'W 

00°ll'W 

00°07'E 

00°19'W 

01°42'W 

01°36'W 

01°45'W 

00°01'E 

00°08'W 

00°06'W 

01°00'E 

OO°37'W 

13°43'W 

13°48'W 

13°46'W 

13°45'W 

13 "48^ 

14°02'W 

13°48'W 

13°49'W 

13°25'W 

14°04'W 

13°54'W 

, 16°05'W 

11°51'N, 1 

11°52'N, 15°46'W 

11°15'N, 15°52'W 

11°32'N, 16°20'W 

ll°20'N, 15°49'W

NUMBER 306 

Uha de Soga [Soja] 

Ivory Coast 

Fresco 

Grand-Bassam 

Grand-Lahou 

Jacqueville 

Lagune Ebrie 

Port-Bouet 

San-Pedro 

Sassandra 

Tabou 

Vridi 

Josephine Seamount 

La Chapelle Bank [Bane de la 

Chapelle] 

Liberia 

Bromley. See Sesarma (Chiromantes) 

buettikoferi 

Bushrod Island 

Cape Mesurado 

Farmington River 

Grand Cape Mount, near 

Robertsport 

Harbel [Habel] 

Junk River 

Lake Piso [Fisherman's 

Lake] 

Mesurado River 

Monrovia 

Robertsport 

St. Paul River 

Upper Buchanan 

Madeira Islands 

Agua de Pena 

Canical 

Enseada da Abra 

Funchal 

Ilha de Porto Santo 

Ilhas Desertas 

Deserta Grande 

Ilheu do Bugio 

Ilheu do Gorgulho 

Machico Bay 

Ponta da Garajau 

Ponta da Oliveira 

Ponta dos Reis Magos 

Ponta de Sao Lourenco 

Prainha. See Portunus 

hastatus 

Ribeira Brava 

Mauritania 

Baie de Cansado 

Baie de l'Ouest [on Cap 

Blanc peninsula] 

H°21'N, 15°52'W 

05°05'N, 05°34'W 

O5°12'N, 03°44'W 

05°08'N, 05°01'W 

05°12'N, 04°25'W 

05°14'N, 04°26'W 

05 o 15'N,03 o 58'W 

04°44'N, 06°37'W 

04°57'N, 06°05'W 

04°24'N, 07°21'W 

05°15'N, 04°02'W 

36°52'N, 14°20'W 

47°37'N, 07°18'W 

06°21'N, 10°47'W 

06°19'N, 10°49'W 

06°08'N, 10°22'W 

[06°44'N, 11°24'W] 

06°16'N, 10°21"W 

06°09'N, 10°23'W 

06°45'N, 11°16'W 

06°20'N, 10°48'W 

06°19'N, 10°48'W 

06°45'N, 11°22'W 

06°23'N, 10°48'W 

05°56'N, 10°04'W 

32°44'N,17°00'W 

32°42'N, 16°46'W 

32°44'N, 16°44'W 

32°44'N, 16°42'W 

32°38'N, 16°54'W 

33°04'N, 16°20'W 

32°30'N, l6 o 3ffYf 

32°30'N, lG^CW 

32°26'N, 16°29'W 

32°37'N, le^^ 

32°42'N, 16°46'W 

32°38'N, 16°51'W 

32°38'N, ^"SCW 

32°38'N, \6°A:9rW 

32°44'N, 16°41'W 

32°39'N) 17°O4'W 

20°53'N, 17°01'W 

[ca. 20°47'N, 19°24 

Baie du Repos 

Baie de Saint-Jean 

Bane d'Arguin 

Bilaouakh. See Eurypanopeus 

blanchardi 

Cansado 

Cap Blanc 

Cap Timiris 

Nouakchott 

Pointe de Cansado 

Portendick [Portendic] 

Port-Etienne 

Tamzak [Tamxat] 

Morocco 

Agadir 

Asilah [Arzila] 

Bane de Spartel [off Cap 

Spartel] 

Cap Draa [Cap Noun] 

Cap Juby 

Cap Rhir [Cap Ghir] 

Cap Spartel 

David 

El Hank. See Pachygrapsus 

transversus 

Essaouira 

Fosse de Rabat 

Foum Agouitir [Puerto-Cansado] 

Foum Assaka 

Larache 

Lucien Saint. See Percnon 

gibbesi 

Mannesman 

Oued Bou Regrcg 

Oued Cherrat 

Plage David 

Pointe d'El Hank 

Rabat 

Sidi Bou Knadel [Bouknadel] 

Sidi Moussa 

Skhirat 

Tangier 

Temara 

Temara—plage 

Tillet [PTillelt]. See Acanthonyx 

lunulatus 

Nigeria 

Adu River 

Ajudaibo 

Bonny River 

Brass 

Forcados 

20°55'N, 17°O2'W 

19°27'N, 16°22'W 

20°10'N, 16°45'W 

20°51'N, 17°O2'W 

20°46'N, 17°03'W 

19°23'N, 16°32'W 

18°06'N, 15°57'W 

20°51'N, 17°02'W 

18°35'N, 16°O7'W 

20°54'N, 17°04'W 

17°26'N, 16°02'W 

30°24'N, 09°36'W 

35°28'N, 06°02'W 

[ca. 35°54'N, 06°14'W] 

28°44'N, ll'WW 

27°57'N, 12°55'W 

30°38'N, 09°54'W 

35°48'N, 05°56'W 

[33°50'N, O7°O7'W] 

31°31'N, 09°46'W 

[ca. 34°N, 07°W] 

28°04'N, 12° 14^ 

29°08'N, 10°24'W 

35°12'N, 06°09'W 

[33°37'N, O7°33'W] 

34°03'N, Oe'SCW 

33°50'N, O7°O7'W 

[33°50'N, O7°O7'W] 

33°37'N, 07°40'W 

34°02'N, 06°50'W 

34°08'N, 06°44'W 

32°53'N, 08°51'W 

33°52'N, O7°O3'W 

35°48'N, 05°48'W 

33°55'N, Wbb'VJ 

33°56'N, 06°58'W 

345 

07°38'N, 10°05'E 

[05°34.5'N, 05°11.75'E] 

04°23'N, 07°06'E 

04°19'N, 06°14'E 

05°22'N, 05°26'E

346 

Lagos 

Ogudu River [creek] 

Port Harcourt 

Tarkwa Bay 

Victoria Beach 

Yaba 

Princesse Alice Bank 

Principe, Ilha do 

Baia das Agulhas 

Baia de Santo Antonio 

Ilheu Caroco 

Ilheu Santana [Sta. Ana] 

Hheus dos Mosteiros 

Morro de Praia Grande 

Ponta Capitao 

Ponta da Mina 

Ponta Novo Destino 

Ponta do Pico Negro 

Ponta da Praia Salgada 

Praia Pequena 

Rio Papagaio 

Santana 

Santo Antonio 

Tinhosa Grande [Hermano 

Grande] 

Rio Muni [Equatorial Guinea] 

Cabo San Juan 

Corisco Bay 

Isla de Corisco 

Islas de Elobey 

Rio Muni mouth 

Saint Helena 

James Bay 

Ruperts Bay 

Sao Tome, Ilha de 

Angra de Sao Joao dos Angolares 

Baia do Almoxarife 

Baia de Ana de Chaves 

Baia de Iogoiogo 

Baia de Sao Miguel 

Binda 

Ilheu das Cabras 

Ilheu Gago Coutinho [Ilheu 

das Rolas] 

Ilheu Macaco [ilot dos 

Cocos] 

Ilheu de Sant'Ana 

Morro Peixe 

Oquedelrei 

Ponta Diogo Nunes 

Ponta Diogo Vaz 

Ponta Furada 

Ponta de Sao Sebastiao 

06°27'N, 03°23'E 

06°34'N, 03°24'E 

04°46'N, 07°01'E 

06°24'N, 03°24'E 

06°25'N, 03°24'E 

06°32'N, 03°23'E 

38°00'N, 29°15'W 

O1°37'N, 07°25'E 

O1°37'N, 07°21'E 

01°39'N, O7°27'E 

01 o 31'N,07°26'E 

01°40'N, O7°27'E 

01°41'N,07°29'E 

01°41'N, O7°27'E 

01°40'N, 07°28'E 

01°39'N, 07°26'E 

Ol^N, 07°26'E 

01°32'N,07°24'E 

01°39'N, O7°27'E 

01°38'N, O7°27'E 

01°39'N, 07°25'E 

01°40'N, 07°26'E 

01°39'N, 07°26'E 

01°21'N, 07°18'E 

0ri0'N,09°21'E 

00°55'N, 09°25'E 

00°55'N, 09°19'E 

00°59'N, 09°30'E 

01°01'N, 09°36'E 

15°57'S, 05°42'W 

15°55'S, 05°43'W 

15°55'S, 05°43'W 

00°12'N, 06°39'E 

00°07'N, 06°39'E 

00°17'N, 06°45'E 

00°21'N, 06°44'E 

00°02'N, 06°33'E 

00°08'N, 06°30'E 

00°13'N, 06°28'E 

00°24'N, 06°43'E 

00°01'S, 06°32'E 

00 o 06'N,06°3rE 

00°15'N, 06°46'E 

00°24'N, 06°39'E 

OO°22'N, 06°44'E 

00°23'N, 06°43'E 

00°19'N, 06°30'E 

00°15'N, 06°28'E 

00°21'N, 06°44'E 


Praia das Conchas 

Praia de Fernao Dias 

Praia Lagarto 

Praia Melao 

Praia Pantufo 

Praia de Santa Catarina 

Ribeira Peixe 

Sant'Ana 

Seine Seamount [Bane de la 

Seine] 

Senegal 

Anse Bernard 

Anse de Hann 

Baie de Dakar 

Baie de Goree 

Bakoy [?=Bako or 

Bakor River] 

Baling [?=Bafing] 

Bane du Seminole 

Bargny Gouddou 

Bel-Air 

Camberene 

Cap Manuel 

Cap Vert 

Casamance River 

Dakar 

Faleme 

Faleme River. See Callinectes 

amnicola 

Fosse de Kayar [Cayar] 

Goree 

lie de Goree 

lie de la Madeleine 

Joal 

Lac de Nguer [N'Guer] 

Lagoba. See Dromia monodi 

Leybar 

Mbao 

Mboro 

Mbour 

Ngaparou 

Ngazobil 

Ngor 

Pointe des Almadies [?Les 

Almadies] 

Pointe de Sangomar 

Pointe Thionk 

Presqu'ile du Cap Vert 

Rosso 

Rtombo. See Cardisoma 

armatum 

Rufisque 

Saint-Louis 

Saloum 

00°24'N, 06°37'E 

OO°25'N, 06°41'E 

0O°22'N, 06°43'E 

00°19'N, 06°45'E 

00°19'N,06°45'E 

00°16'N, 06°29'E 

00°05'N, 06°37'E 

00°16'N, 06°45'E 

33°45'N, 14°25'W 

14°40'N, 17°26'W 

14°43'N, 17°25'W 

14°41'N, 17°25'W 

14°40'N, 17°2O'W 

14°00'N, 15°16'W 

12°55'N, 14°44'W 

14°23'N, 16°57'W 

14°36'N, 17°28'W 

14°42'N, 17°14'W 

14°42'N, 17°25"W 

14°46'N, 17°26'W 

14°39'N, 17°26'W 

14°43'N, 17°28'W 

12°33'N, 16°46'W 

14°40'N, 17°26'W 

14°46'N, 12°14'W 

14°57'N, 17°14'W 

14°40'N, 17°24'W 

14°40'N, 17°24'W 

14°39'N, 17°28'W 

14°10'N, 16°51'W 

14°52'N, 14°53'W 

15°58'N, lG^CW 

14°44'N, 17°19'W 

15°09'N, 16°54'W 

14°24'N, 16°58'W 

14°28'N, 17°04'W 

14°12'N, 16°52'W 

14°45'N, 17°31'W 

14°45'N, 17°32'W 

13°50'N, 16°46'W 

12°40'N, 16°40'W 

14°43'N, 17°28'W 

16°30'N, 15°49'W 

14°43'N, 17°17'W 

16°02'N, 16°30'W 

13°50'N, 16°45'W

NUMBER 306 

Senegal River 

Soumb-Dioun 

Thiaroye-sur-Mer [Tiaroye] 

Sierra Leone 

Aberdeen 

Bullom Shore 

Cline Bay [Kline] 

Freetown 

Great Scarcies River 

Kissy 

Murray Town 

Robene Point 

Rokupr 

Sierra Leone River 

Spanish Sahara 

Angra da Cintra (Bay) 

Cabo Barbas 

Cabo Blanco 

Cabo Bojador 

Cabo Corbeiro [Corveiro] 

Islote Virginia 

Medano de Aaiun 

15° 48'N, 16°32'W 

3 

[14' 41'N, 17°28'W] 

14° 44'N, 17°23'W 

08° 30'N, 13°17'W 

[08' >34'N, 13°12'W] 

08° 29'N, 13°12'W 

08° 30'N, 13°15'W 

08° 55'N, 13°08'W 

08° 28'N, 13°12'W 

08° 30'N, 13°16'W 

a 

[08' 29'N, 13°06'W] 

08° 40'N, 12°23'W 

08° 30'N, ^"lS'W 

23° 00'N, 16°15'W 

22° 18'N, 16°41'W 

20° 46'N, 17°O3'W 

26° 08'N, 14°30'W 

21° 48'N, 16°58'W 

22° 09'N, 16°50'W 

27° 07'N, 13°25'W 

Morro Garnet [Cabo 

Garnet] 

Pulpito Bay [?Buen Jardin] 

Villa Cisneros 

Togo 

Lome 

Zaire 

Banana 

Boma 

Congo River 

lie de Mateba 

Katala 

Kunga 

Malela 

Moanda 

Shiloango River [Loango R. 

(Congo), Chiloango R. 

(Angola)] 

Tumuna. See Sesarma (Chinmantes) 

angolense 

Vista 

Zambi 

24°45'N, 14°55'W 

[24°41'N, 14°51'W] 

23°42'N, 15°56'W 

06°08'N, 01°13'E 

06°01'S, 12°24'E 

05°51'S, 13°03'E 

06°04'S, 12°24'E 

05°54'S, 12°50'E 

05°58'S, 12°44'E 

[ca. 05°55'S, 12°35'E] 

05°59'S, 12°37'E 

[05°55'S, 12°25'E] 

05°12'S, 12°O7'E 

347 

05°52'S, 12°17'E 

[ca. 05°52'S, 12°50'E]

Addendum 

After this manuscript went to the printer, Mr. C. B. Powell forwarded to us 

representatives of what proved to be a fourth new genus and species of 

camptandriine crab from West Africa; it is described here. This species raises 

the number of East Atlantic Ocypodidae to six genera and seven species; six 

of the species occur off tropical West Africa. 

Genus Lillyanella, new genus 

TYPE-SPECIES.—Lillyanella plumipes, new species. 

ETYMOLOGY.—The name Lillyanella is chosen in 

recognition of the important contribution to the 

present study made by Lilly King Manning, to 

whom we are indebted for preparation of the 

illustrations. Gender is feminine. 


slightly convex, subhexagonal, broader 

than long, widest in the middle. Regions poorly 

indicated, surface even, with sparse, scattered, 

very short, stiff, dark pubescence; in posterolateral 

part some small granules. Front wide, anterior 

margin broadly truncate, slightly sinuous. 

Epigastric lobes barely visible. H-shaped groove 

distinct. 

Anterolateral margins with two large and one 

or more small to very small teeth far behind outer 

orbital angle. Posterior margins of the orbits 

slightly curved, almost in one line, outer half 

somewhat granular. 

Cornea well developed, slightly narrower than 

eyestalk. Antennules obliquely folded. Antennae 

short, flagellum consisting of a single segment 

with a long apical seta. Lower orbital margin 

distinct, with several distinct denticles and granules. 

Suborbital ridge and pterygostomian region 

with granules. 

Epistome deeply concave with raised anterior 

and posterior margins. Third maxilliped not fill- 

348 

ing oral field. Merus slightly longer than ischium 

and partly covering exopod; anteromedian part 

of merus with quadrangular excision. Anterior 

margin of ischium oblique. 

Chelipeds equal. Those of adult male much 

larger and more robust than those of female, and 

with molariform tooth on dactylus. Walking legs 

short and robust. Merus with dorsal row of teeth, 

of which distal and subdistal large, the following 

smaller but still distinct; lower surface of merus 

with teeth or tubercles. The other segments without 

teeth or denticles, some with a few granules. 

All legs covered with dense short dark pubescence, 

especially dorsally. Propodus of third leg of 

adult male with conspicuous tuft of long soft 

hairs. 

Male abdomen elongate triangular, only 

slightly constricted at fifth somite, but entirely 

covering the pleopods; second to fourth somites 

of male abdomen fused. First somite not wider 

than second and failing to reach the coxae of the 

fifth pereiopods. Male gonopods strongly recurved, 

slightly swollen apically with some distinct 

recurved spines. 

REMARKS.—Lillyanella differs from all other 

genera of Camptandriinae in the shape and armament 

of the ambulatory legs. In this respect it 

resembles the genus Leipocten most closely, in 

which the rneri likewise have spines on the lower 

surface. However, in Leipocten the large distal and 

subdistal dorsal teeth of the merus are absent, 

and the other segments of the legs are different.


Lillyanella plumipes, new species boundary of freshwater and oligohaline area, un- 

FIGURE 88 

der bark of large sticks, 5 February 1980, C. B. 

Powell, \6 holotype (L), 86, 99 ov, 4 non-oviger- 

MATERIAL EXAMINED.—Pillsbury Material: None. ous 9, 1 juv, paratypes (L); 76, 169 (8 ov), 3 juv, 

Other Material: Nigeria: New Calabar River at paratypes(W). 

Choba bridge, 04° 54'N, 06°54'E, intertidal zone, DESCRIPTION.—Carapace subhexagonal to al- 

9 

FIGURE 88.—Lillyanella plumipes, new genus, new species. Ovigerous female: a, carapace, dorsal 

view; b, pterygostomian region, orbit, and third maxilliped, ventral view; c, second leg, d, third 

leg; e, fourth leg; /, fifth leg. Male: g, carapace, dorsal view; k, abdomen; i, first pleopod. 

Smaller male: j, first leg. Male holotype: *, first leg, /, propodus and dactylus of third leg. Nonovigerous 

female: m, first leg; n, chela. 

f

350 

most quadrangular, somewhat (about 1.2 times) 

broader than long. Surface slightly convex but 

even, without conspicuous elevations. H-shaped 

groove distinct, but not very deep. Epigastric 

lobes low and inconspicuous. Posterior half of 

anterolateral margin with two large teeth, between 

which 1 to 3 very small. Outer orbital 

angle sharp, about as large as the larger anterolateral 

teeth. Posterolateral margins with several 

small granules. Posterior margin almost straight, 

about as wide as the base of front. Front wide 

and not very prominent, anterior margin broad 

and somewhat sinuous: slightly concave in the 

middle, slightly convex laterally; anterolateral 

angle widely rounded; lateral margin evenly 

merging, under a broad curve, with the posterior 

orbital margin. Posterior orbital margins almost 

straight, the two lying in a practically single line; 

no incisions on this margin, but granules are 

present in its greater part. Dorsal surface of carapace 

with a sparse pubescence of short, stiff, 

curved, black hairs. The posterolateral portions 

of carapace sparsely granulated. 

Eyes almost filling the orbits, with the cornea 

well developed, rounded, slightly narrower than 

the eyestalk. Eyestalk with short, curved, stiff, 

dark hairs on anterior surface. Lower margin of 

orbit with distinct denticles and granules over its 

full length, and with a denticulated process near 

the base of the antenna. A short suborbital row 

of granules below outer half of orbit. Pterygostomian 

region granulate. 

Antennules covered by front, folding obliquely. 

Antennae short, entering orbit, the latter widely 

open at base. Antennal flagellum consisting of a 

single segment with a very long apical seta. 

Epistome wide, deeply concave, with anterior 

and posterior margins elevated. 

Third maxillipeds not filling oral cavity. Merus 

rectangularly excavated anteromedially; outer 

anterior angle broadly rounded. Ischium longer 

than merus, inner anterior angle somewhat produced 

forward. Exopod at base about 1/3 as wide 

as ischium; distal part, including multi-articulated 

flagellum, hidden behind merus. 

First pereiopods equal left and right, in adult 

males much larger and more robust than in fe- 


males. Fingers of adult male cheliped somewhat 

shorter than palm, curved and gaping, tips ending 

in horn-colored hoofs; cutting edges with 

small denticles, in addition to which a large 

molar-like tooth in the dactylus. Palm in adult 

males almost as high as long. Surface of palm and 

fingers smooth, except for some minute granules 

on lower surface of palm. Carpus short, cupshaped, 

upper anterior margin with tubercles. 

Merus about twice as long as carpus, but still 

short and robust; lower surface with granulae, 

both inner and outer lower margin with a longitudinal 

row of teeth; also dorsal margin with a 

row of teeth; upper part of anterior margin granulate. 

Ischium short, lower margin with teeth. 

First leg of female more slender than that of male, 

palm more than twice as long as high; fingers as 

long as palm, slightly gaping with denticles along 

cutting edge, but no molariform tooth, tips of 

fingers hooved; rest of leg as in male, but more 

slender. 

First three walking legs of about the same size, 

the third leg (P3) slightly longer than the other 

two; the fifth leg shortest of all legs. Dactylus of 

walking legs simple with sharp, curved tip; about 

3/4 of length of propodus. Propodus two to three 

times as long as high (relatively longest in third 

leg), unarmed. Carpus short, about as long as 

dactylus, narrowing proximally, upper anterior 

margin sometimes with granules. Merus with dorsal 

margin ending in a large distal tooth and a 

subdistal tooth of about same size. Behind subdistal 

tooth a longitudinal row of up to 10 rather 

small but quite distinct teeth. Lower surface of 

merus with teeth or tubercles. Sometimes tubercles 

on outer surface near upper or lower margin. 

In adult males propodus of third pereiopod with 

a conspicuous large brush of very long soft hairs. 

In all specimens a distinct pubescence of short, 

curved, stiff, dark hairs on all segments of the 

legs, especially dorsally; here and there a few 

longer and softer hairs may be present. 

Female abdomen wide, reaching the coxae of 

the pereiopods; all segments free; distal segment 

widely triangular with broadly rounded slightly 

protruding top. 

Male abdomen elongate triangular. First som-


ite about as wide as second, failing to reach coxae 

of fifth pereiopods. Second, third, and fourth 

somites fused; indentation in lateral margin 

marking separation between second and third 

somites. Second somite very short, only slightly 

longer than first. Third and fourth somites of the 

same length, about twice as long as second and 

as wide; fourth segment narrowing slightly distally. 

Fifth and sixth somites free. Fifth longer 

than fourth, constricted in the basal half, but not 

strongly enough to expose the gonopods. Sixth 

somite about as long as fifth, narrowing distally. 

Seventh somite as long as sixth, but much narrower, 

broadly rounded at apex. 

Male gonopod with the recurved shape characteristic 

of Camptandriinae, narrowing distally, 

but widening slightly just before the tip. Apex 

broadly rounded, provided with three large 

spines; inner part of widened subapical portion 

with a group of recurved spines; further numerous 

very small spinules in distal part of gonopod. 

Second pleopod of male very small, L-shaped. 

MEASUREMENTS.—In the adult male holotype 

cl is 4.2, cb 5.2 mm. The carapace width in the 

ovigerous females is 4.3 to 6.3 mm, in the nonovigerous 

females 4.0 to 6.5 mm, in the males 3.7 

to 6.5 mm, and in the juveniles 1.3 to 4.5 mm. 

The eggs are about 0.2 mm in diameter. 

REMARKS.—Lillyanella plumipes differs from all 

other West African Camptandriinae by the presence 

of a row of teeth on the dorsal margin of the 

merus of the walking legs, and by having teeth or 

tubercles on the lower surface of that segment. 

Also the peculiar brush of long setae on the 

propodus of the third leg of the adult males is 

quite characteristic. In the presence of anterolateral 

teeth on the carapace, Lillyanella differs from 

Ecphantor and resembles the two other West African 

genera. The wide and broadly truncated 

front it has in common with Telmatothrix; in Calabarium 

the front is produced in two broadly 

triangular teeth separated by a V-shaped incision. 

The gonopods also are different from those of the 

other genera. 

TYPE-LOCALITY.—New Calabar River, slightly 

NW of Port Harcourt at Choba bridge, Nigeria, 

4°54'N 6°54'E. 

DISPOSITION OF TYPES.—The holotype male 

(Crust. D 32752) is placed in the collection of the 

Rijksmuseum van Natuurlijke Historie, Leiden. 

The paratypes are in the same museum and in 

the National Museum of Natural History, Smithsonian 


ETYMOLOGY.—The name plumipes is inspired by 

the presence of a brush of long hairs on the 

propodus of the third leg of adult males. 

BIOLOGY.—The site at which the species was 

collected was described by Mr. Powell (in litt., 5 

February 1980) as follows: 

Its proper habitat is under the bark of dead tree branches 

etc., in the intertidal zone. The same site has Telmatothrix 

(common or abundant), Calabarium, Sesarma buettikoferi (abundant) 

and juveniles of S. angolense and S. alberti. The main 

vegetation is Pandanus and freshwater swamp trees, with the 

aquatic Crinum (subtidal) and Nymphaea (intertidal); a few 

specimens ofRhizophora occur, but these are small individuals 

which probably will not flower—there is no mature Rhizophora 

within sight of the bridge. The two other invertebrates 

occurring under the bark of the same branches are Potamalpheops 

monodi (Sollaud) (Alpheidae) and the large burrowing 

and filter-feeding mayfly nymph Povilla. The site is slightly 

beyond the upstream limit of isopods, amphipods and snails 

and bivalves. The New Calabar River is a soft-water river, 

lacking a freshwater mollusc fauna. Once I did find a single 

specimen of the oligohaline (?) snail Potamopyrgus ciliatus 

(Gould) at the collecting site, but have not been able to find 

any more. Occasionally the small and very thin specimen of 

the polychaete Namalycastis is found; also a red nemertean. 

The shrimps present are Potamalpheops monodi (Sollaud), an 

undescribed species of Potamalpheops, juveniles of Macrobrachium 

felicinum Holthuis (uncommon), M. vollenhovenii (Herklots), 

M. macrobrachion (Herklots) (most common) and Caridina, 

but no Palaemonetes. 

Mr. Powell remarked that the species "has the 

same distinctive odor as does Telmatothrix, when 

preserved in formalin. Also it sheds its legs much 

more readily than any other crab I've met." 

Ovigerous females were collected in February. 

DISTRIBUTION.—Known only from the type locality.


ADDENDUM TO KEY 

Our key to the genera of Camptandriinae (p. 193) can be altered to include 

this additional new genus by substituting the following two couplets for 

couplet 8 and changing couplets 9-11 to 10-12. 

8. Merus of pereiopods with blunt spines or teeth on lower (flexor) 

surface 9 

Merus of pereiopods without ventral spines or teeth 10 

9. Upper margin of merus of pereiopods granular, without large distal or 

subdistal teeth; lower margin of propodus of these legs ending in a 

tooth. Carapace granular all over, without large anterolateral teeth 

Leipocten 

Upper margin of merus of pereiopods ending in a large distal tooth and 

a subdistal tooth of the same size; lower margin of propodus without a 

distal tooth. Carapace smooth for the larger part, only granular in the 

posterolateral area; anterolateral margin with 2 large and some small 

teeth Lillyanella, new genus

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Premiere Partie. Expeditions Scientifiques du 

Travailleur et Talisman pendant les Annies 1880, 

1881, 1882, 1883. 396 pages, 32 plates. Paris. 

1902. Les Dromiaces et Oxystomes. In Reports on the 

Results of Dredging, under the Supervision of 

Alexander Agassiz, in the Gulf of Mexico (1877- 

78), in the Caribbean Sea (1878-79), and along 

the Atlantic Coast of the United States (1880), by 

the U.S. Coast Survey Steamer "Blake," Lieut.- 

Com. C. D. Sigsbee, U.S.N., and Commander J. 

R. Bartlett, U.S.N., Commanding, XXXIX. Memoirs 

of the Museum of Comparative Zoology at Harvard 

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1923. Les Porcellanides et des Brachyures. In Reports on 

the Results of Dredging, under the Supervision of 

Alexander Agassiz, in the Gulf of Mexico (1877- 

78), in the Caribbean Sea (1878-79), and along 

the Atlantic Coast of the United States (1880), by 

the U.S. Coast Survey Steamer "Blake," Lieut.- 

Com. C. D. Sigsbee, U.S.N., and Commander J. 

R. Bartlett, U.S.N., Commanding. XLVII. Memoirs 

of the Museum of Comparative Zoology at Harvard 

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Milne Edwards, H. 

1834. Histoire naturelle des Crustaces, comprenant Vanatomic, la 

physiologie et la classification de ces animaux. Volume 

1: xxxv + 468 pages. Paris. 

1836-1844. Les Crustaces. In G. Cuvier, Le regne animal, 

distribue dapres son organisation, pour servir de base a 

rhistoire naturelle des animaux, et a"introduction a 

Canaiomie comparee, 278 pages. Atlas, plates 1-80. 

Paris. [According to Cowan (1976:60), the text 

and plates of this work were published in 23 parts 

(livraisons) between September 1836 and March 

1844. These parts are summarized here as follows: 

main part number (equivalent Crustacea part 

number): pages, plates, (date): 8(1): 1-8, pis. 1, 7, 

22, 26, September 1836. 20(2):9-16, pis. 2, 17, 24, 

41, March 1837. 33(3): 17-24, pis. 25, 42, 47, 48, 

October 1837. 38(4):25-32, pis. 37, 38, 44, 50, 

January 1838. 51(5):33-4O, pis. 8, 20, 29, 33, July 

1838. 60(6) :41-48, pis. 9, 21, 27, 40, November 

1838. 70(7):49-56, pis. 23, 30, 31, 45, April 1839. 

73(8):57-64, pis. 43, 46, 58, 61, June 1839. 87(9): 

65-72, pis. 28, 49, 65, 66, January 1840. 90(10): 

73-80, pis. 3, 36, 71, IXbis, February 1840. 95(11): 

81-88, pis. 19, 32, 34bis, 67, May 1840. 99(12) :89- 

96, pis. 10, llbis, 34, 51, July 1840. 107(13):97- 

104, pis. 4, 12, 76, November 1840. 110(14): 105- 

112, pis. 52, 53, 55, 59, December 1840. 118(15): 

113-120, pis. 5, 18, 54, bbbis, April 1841. 120(16): 

121-128, pis. 16, 39, 56, 77, May 1841. 149(17): 

129-136, pis. 6, 15, 35, 60, July 1842. 157(18): 

137-144, pis. 11, 14, 64, 69, November 1842. 

176(19): 145-176, pis. 13, 68, 70, July 1843. 

180(20): 177-184, pis. 57, 62, 62bis, 63, August 

1843. 184(21): 185-216, pis. 54bis, 74, 75, October 

1843. 188(22):217-248, pis. 70bis, 72, 78, December 

1843. 194(23):249-278 + 5, pis. 73, 79, 80, 

March 1844.] 

1837. Histoire naturelle des Crustaces, comprenant Panatomie, la 

physiologie et la classification de ces animaux. Volume 

2, 532 pages. Atlas [1834, 1837, 1840]: 32 pages, 

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1852. De la famille des ocypodides (Ocypodidae). Second 

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1854. Notes sur quelques Crustaces nouveaux ou peu 

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naturelle (Paris), 7:145-192, plates 9-16. [Although 

volume 7 of the Archives is dated 1854-1855, the 

reprint of this article carries the date 1854, and 

the paper must have been published in that year, 

even though it is cited by page in H. Milne 

Edwards (1853). In Neave's Nomenclator Zoologicus 

the present paper is dated 1853, and the generic

370 

names proposed in it are given precedence over 

those published in the previous paper (H. Milne 

Edwards, 1853). Apart from the fact that H. Milne 

Edwards in his 1853 paper cited the pages of his 

1854 paper, we cannot find any evidence that the 

latter was published earlier than indicated both 

on the volume and on the reprint. H. Milne 

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Milne Edwards, H., and H. Lucas 

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