LIFE SCIENCES CONTRIBUTIONS 148 Shallow-Water Hydroids of Bermuda The Athecatae Dale R. Calder ROM ROYAL ONTARIO MUSEUM ROYAL ONTARIO MUSEUM PUBLICATIONS IN LIFE SCIENCES The Royal Ontario Museum publishes three series in the Life Sciences. CONTRIBUTIONS: a numbered series of original scientific publications. OCCASIONAL PAPERS: a numbered series of original scientific publications, primarily short and of taxonomic significance. MISCELLANEOUS PUBLICATIONS: an unnumbered series on a variety of subjects. All manuscripts considered for publication are subject to the scrutiny and editorial policies of the Life Sciences Editorial Board, and to independent refereeing by two or more persons, other than Museum staff, who are authorities in the particular field involved. LIFE SCIENCES EDITORIAL BOARD Senior editor: J. L. Eger Editor: D. C. Darling Editor: R. W. Murphy External editor: C. S. Churcher Manuscript editor: J. C. Barlow Production editor: J. E. Hawken Dale R. Calder is associate curator in charge in the Department of Invertebrate Zoology, Royal Ontario Museum, and associate professor in the Department of Zoology, University of Toronto. Canadian Cataloguing in Publication Data Calder, Dale R. Shallow-water hydroids of Bermuda : the Athecatae (Life sciences contributions, ISSN 0384-8159 ; 148) Bibliography: p. Includes index. ISBN 0-88854-339-5 1. Hydroida. 2. Coelenterata — Bermuda. I. Royal Ontario Museum. II. Title. III. Series. QL377.H9C35 1988 593.7' 1'097299 C88-093898-6 Publication date: 15 June 1988 ISBN 0-88854-339-5 ISSN 0384-8159 © Royal Ontario Museum, 1988 100 Queen's Park, Toronto, Canada M5S 2C6 PRINTED AND BOUND IN CANADA AT UNIVERSITY OF TORONTO PRESS Contents Abstract 1 Introduction 1 Materials and Methods 2 Systematic Account 5 Family Clavidae McCrady, 1859a 5 Subfamily Corydendriinae, subf. nov. 5 Genus Corydendrium van Beneden, 1844a 5 Corydendrium parasiticum (Linnaeus, 1767) Genus Turritopsis McCrady, 1859b 7 Turritopsis nutricula McCrady, 1859b 8 Genus Rhizodendrium, gen. nov. 10 Rhizodendrium sterreri, sp. nov. 10 Family Bougainvilliidae Liitken, 1850 12 Subfamily Pachycordylinae Cockerell, 1911 13 6 Genus Millardiana Wedler and Larson, 1986 13 Millardiana longitentacidata Wedler and Larson, 1986 Genus Pachycordyle Weismann, 1883 14 Pachycordyle napolitana Weismann, 1883 15 Genus Silhouetta Millard and Bouillon, 1973 16 Silhouetta uvacarpa Millard and Bouillon, 1973 Subfamily Rhizorhagiinae, subf. nov. 18 Genus Parawrightia Warren, 1907 18 19 Parawrightia robusta Warren, 1907 Subfamily Bimeriinae Allman, 1872 21 Genus Bimeria Wright, 1859 13 17 21 Bimeria vestita Wright, 1859 21 Subfamily Bougainvilliinae Liitken, 1850 23 Genus Bougainvillia Lesson, 1830 23 Bougainvillia nmscus (Allman, 1863) 24 Family Cytaeididae L. Agassiz, 1862 28 Genus Cytaeis Eschscholtz, 1829 29 Cytaeis sp. 29 Family Hydractiniidae L. Agassiz, 1862 31 Genus Stylactaria Stechow, 1921a 32 Stylactaria arge (Clarke, 1882), comb. nov. 33 Superfamily Pandeoidea Haeckel, 1879 35 Genus Pelagiana Borstad and Brinckmann-Voss, 1979 36 Pelagiana trichodesmiae Borstad and Brinckmann-Voss, 1979 Family Eudendriidae L. Agassiz, 1862 38 Genus Eudendrium Ehrenberg, 1834 38 Eudendrium bermudense, sp. nov. 39 Eudendrium capillare Alder, 1856a 41 Eudendrium carneum Clarke, 1882 Genus Myrionema Pictet, 1893 43 46 Myrionema amboinense Pictet, 1893 Family Corymorphidae Allman, 1872 Genus Zyzzyzus Stechow, 1921a Zyzzyzus warreni, nom. nov. 36 46 48 49 49 in Family Tubulariidae Fleming, 1828 52 Genus Ectopleura L. Agassiz, 1862 52 Ectopleura pacifica Thornely, 1900 53 Family Halocordylidae Stechow, 1921a 55 Genus Halocordyle Allman, 1872 55 Halocordyle disticha (Goldfuss, 1820) 56 Family Sphaerocorynidae Prevot, 1959 60 Genus Sphaerocoryne Pictet, 1893 60 Sphaerocoryne bedoti Pictet, 1893 61 Family Corynidae Johnston, 1836 63 Genus Coryne Gaertner, 1774 63 Coryne sargassicola, sp. nov. 64 Family Cladonematidae Gegenbaur, 1857 66 Genus Cladonema Dujardin, 1843a 66 Cladonema radiatum Dujardin, 1843a 67 Family Zancleidae Russell, 1953 68 Genus Zanclea Gegenbaur, 1857 69 Zanclea alba (Meyen, 1834), comb. nov. 69 Family Milleporidae Fleming, 1828 72 Genus Millepora Linnaeus, 1758 73 Millepora alcicornis Linnaeus, 1758 73 Family Porpitidae Goldfuss, 1818 76 Genus Porpita Lamarck, 1801 77 Porpita porpita (Linnaeus, 1758) 77 Genus Velella Lamarck, 1801 80 Velella velella (Linnaeus, 1758) Acknowledgements Literature Cited Index IV 103 84 85 81 Shallow-Water Hydroids of Bermuda The Athecatae Abstract The shallow-water athecate hydroids known from Bermuda and vicinity, comprising 26 species in 24 genera, are surveyed. Synonymies are updated for each family-, genus-, and species-group taxon represented. Six taxa are described as new to science: Corydendriinae, subf. nov., Rhizorhagiinae, subf. nov., Rhizodendrium, gen. nov., Rhizo dendrium sterreri, sp. nov., Eudendrium bermudense, sp. nov., and Coryne sargassicola, sp. nov. Zyzzyzus warreni is proposed as a replacement name for the junior primary homonym Tubularia solitaria Warren, 1906b (not Tubularia solitaria Rapp, 1829). The tribe Pachycordylini Cockerell, 1911, is elevated to the rank of subfamily. Pachycordyle Weismann, 1883, Parawrightia Warren, 1907, and Stylactaria Stechow, 1921a, are re established as the valid names of genera. Tubularia muscoides Linnaeus, 1761, is desig nated as type species of the nominal genus Fistulana O. F. Muller, 1776a. A lectotype is designated for the conglomerate Podocoryne alderi Hodge, 1861, a nominal species based on hydroid and medusa stages referable to different genera. The invalid name Bougainvillia ramosa (van Beneden, 1844a) is replaced with the name Bougainvillia muscus (Allman, 1863). Descriptions and illustrations are provided for each species studied, and data on ne- matocyst complement and size are given for all but one of them. Of the 26 species discussed, 23 occur elsewhere in the western Atlantic and 9 are reportedly circumglobal in warm waters. Eleven of the 23 previously known species are reported from Bermuda for the first time. Introduction Hydroids of the oceanic island of Bermuda have been the subject of several previous investigations. The most com prehensive taxonomic accounts to date have been by Congdon (1907) and Bennitt (1922). Hydroids collected by H.M.S. Challenger on Challenger Bank in the vicinity of Bermuda were discussed by Allman (1888) and Ritchie (1909). Reports by Verrill (1900, 1907), Smallwood (1910), Stechow (1912), Jaderholm (1920), and Moore (1969) dis cussed one or more hydroid species from the Bermuda platform. Fraser (1944) included most of the species known from the area, based on literature records. Calder (1986) listed common and otherwise noteworthy hydrozoan spe cies. Hydroids on pelagic Sargassum were studied by Burkenroad {in Parr, 1939), by Morris and Mogelberg (1973), and by Ryland (1974). Morphological, developmental, and physiological investigations have been undertaken on Ber muda material by Congdon (1906), Cowden (1965a, 1965b), Bermudian waters have been published by Fewkes (1883), Verrill (1900), Bigelow (1918), Totton (1936), and Calder (1986). Millepora alcicornis Linnaeus, 1758, a conspic uous member of the Bermudian coral reef community, has been mentioned in papers such as those of Nelson and Duncan (1876), Moseley (1876, 1879,1880), Rice (1878), Quelch (1886), Verrill (1900, 1902a, 1902b, 1907), Moore (1969), and Calder (1986). (1986). Accounts of the neustonic species Porpita porpita The purpose of this report was to provide a taxonomic account of the athecate hydroids currently known from Bermuda and vicinity, to a depth of 100 m, based largely on collections made by the author since 1977. The athecate fauna of the study area is rather depauperate, accounting for only about one-quarter of the total number of hydroid species known from Bermuda (Calder, unpublished data). A decision was made early in the study to include as complete a synonymy of each family-, genus-, and speciesgroup taxon as possible. Original spellings of the names of taxa were verified, authorship and dates of these names (Linnaeus, 1758) and Velella velella (Linnaeus, 1758) in were rechecked, and matters of nomenclature were con- Summers (1972a), Lesh-Laurie (1976), and Clark and Cook 1 sidered according to provisions of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature [iczn], 1985). Some of the nomenclatural problems encountered have been resolved here, while others will require submissions to the commission. Questions regarding the identity, synonymy, and sys tematic positions of taxa arose repeatedly during the course of this study. One quandary in particular involved the extent to which nominal species should be combined or divided. The hydrozoan literature is replete with extremes of taxonomic "lumping" and "splitting," and the con fusion resulting from both. For example, Duchassaing and Michelotti (1864) viewed practically every morphological form of the hydrocoral Millepora Linnaeus, 1758, from the Caribbean as a distinct species, whereas Hickson (1898a, 1898b) recognized only one species in the genus world wide. Most authors now follow Boschma (1948) in rec ognizing three species in the Caribbean, and about a dozen worldwide. Nevertheless, determining how far to go in combining or splitting nominal species is largely a matter of personal opinion. Most recent hydrozoan systematists have tended to be "taxonomic lumpers," and generally broad taxa have been recognized here. Reasons why rel atively few species of hydroids are believed to exist world wide were briefly stated by Cornelius (1981). Related to the question of lumping or splitting of taxa is the interpretation of hydroid species distribution. Ac cording to literature records, many species of hydroids are virtually cosmopolitan. Admittedly, certain hydroids are well adapted for long-range dispersal, and their rate of speciation seems to be rather slow (Cornelius, 1981). Yet the question arises whether some species are as widely distributed as records indicate, or whether their reported range is partly an artifact of the hydrozoan taxonomist's inability to discriminate distinct but closely related species. Hydrozoan classification is complicated by many fac tors, including the following: (1) the existence of separate hydroid and medusa generations in many species; (2) the legacy of separate classifications for hydroids and medu sae; (3) the production of free medusae and fixed gonophores in closely related species; (4) the differential reduction of male and female gonophores in certain species; (5) the production in some taxa of morphologically dissimilar me dusae by virtually indistinguishable hydroids, and vice versa; (6) the morphological variation sometimes displayed within a given taxon; (7) the scarcity of reliable taxonomic characters in various taxa; (8) the general lack of knowl edge concerning the biology of these animals, including life cycles of many species. Classification of the order Athecatae Hincks, 1868, in particular is currently in a state of flux. Most authors over the past 30 years have regarded the Capitata Kiihn, 1913, as the most primitive suborder of the Hydrozoa, largely following Rees (1957). Athecate classification has been extensively modified recently by Petersen (1979), Werner (1984), and Bouillon (1985). Fol lowing the last two authors, families of the suborder Filifera Kiihn, 1913, are discussed first here. Yet it is unlikely that a stable classification, accurately reflecting relation ships within the Athecatae, has been achieved by the tra ditional approaches used in studies to date. A re-examination of relationships within athecate hydroids and their me dusae, especially using methods of phylogenetic systematics (Wiley, 1981), is greatly needed. The only such study within the Hydrozoa up to now is that by Cairns (1984) for stylasterids. The arrangement of families adopted here, somewhat modified from Werner (1984), is recognized as unsatisfactory. Unfortunately, no demonstrably superior classification exists at present. The known range given here is based on reported oc currences at Bermuda, and elsewhere in the Atlantic, Pa cific, and Indian oceans. In most cases, only one significant record has been cited to document occurrences outside Bermuda. Materials and Methods Hydroids were collected during six field trips to the Ber muda Islands, on the following dates: 1-22 September 1977, 26 February-10 March 1982, 17 July-6 August 1982, 15 June-13 July 1983, 20 September-11 October 1984, and 24 September-8 October 1986. Specimens were also obtained on 23-24 May 1979 during a two-day va cation cruise to the islands. Most collections were made by snorkelling, although scuba gear was used on occa sional dives. Collecting efforts were concentrated in shallowwater areas of the northeastern half of Bermuda, especially in Flatts Inlet, Castle Harbour, Harrington Sound, and Whalebone Bay (Fig. 1), and only those hydroids taken in depths of 0-100 m are included in this report. Intensive sampling was undertaken around ledges, bridges, pilings, floats, and moorings; in grass beds; and in ponds and caves. Considerable time was spent searching beaches for stranded specimens of the neustonic hydroids Porpita and Velella. Large quantities of pelagic Sargassum were col lected and examined for attached hydroids during each field trip. Collections at depths of 60-100 m in offshore waters south of Castle Harbour were made by dredging from the M/V Northstar (3 September 1977), R/V Culver (1 July 1983), and R/V Weatherbird (27 September 1984). Dredg- 64°|45' 64° 50' 64° 40' ^&^ V ^ Whalebone Bay^^^^^^^^PK^U^^^Bm^^^^H .,»~^^^K CASTLE ^^^^^V^K ^^^^^^ b Nonsuch!^ HARBOUR :** • ^^^^^^ ^^* Castle b CN CN CN CO flf ,A<>^ Flatts InletJEJ^ ^ ^^ Natural Arches CN CO Beach ^^^^^^^^^^H^r ^ SOUND #|V^^^^^^B^| ATLANTIC ^fr r^^^^^^^^Warwick Long Bay OCEAN »o m CN CO cn CO 64°|45' 64°|50' 64°|40' Fig. 1. The Bermuda Islands. ing in 75 m of water on Challenger Bank, southwest of Bermuda (Fig. 2), was undertaken on 3 October 1984 Dr W. Sterrer on 18 August 1974; (7) Castle Harbour, collected by H. E. Lehman in June 1966; (8) surveys of aboard R/V BBS II; on the return cruise from Challenger Bermudian caves, collected by Dr T. Iliffe during the Bank, a 10-minute plankton tow was made 6 km offshore from Gibbs Hill lighthouse with a plankton net 1 m in summer of 1982. diameter. Hydroids in reference collections at the Bermuda Bio logical Station were examined. Among these were spec imens from (1) buoy chains, collected by Dr John Markham and colleagues between 5 October and 23 December 1976; (2) waters south of Castle Harbour in 60-100 m, collected by Dr Markham using a dredge aboard M/V Northstar on 6 August and 27 August 1977; (3) the Castle Harbour area, collected by Prof. H. Mergner on 22 and 24 June 1967; (4) the Castle Harbour area, collected by Dr H. Thiel on 3-4 August 1974; (5) Argus Tower on Plantagenet Bank (Fig. 2), collected by Dr W. Sterrer on 23 April 1976; (6) the wreck Pelinaion off St David's Island, collected by Specimens collected during this investigationhave been deposited in the Department of Invertebrate Zoology, Royal Ontario Museum (romiz). All descriptions and illustra tions herein are from Bermuda material, as indicated, ex cept for a paratype of Coryne sargassicola from the Gulf Stream off New York City. As complete a synonymy as possible has been given for each taxon, although many of the lists may be less than exhaustive. Virtually all of the listed synonyms have been verified by examination of cited references. Nematocysts were examined in preserved material by compressing pieces of tissue, or entire individuals of mi nute specimens, between a slide and coverslip. Occasion- tion. Length and width measurements were made on undischarged and horizontally oriented nematocysts using an ocular micrometer. At least 10 nematocysts of each type in each species were measured in determining size ally, materials were treated with a 5 per cent solution of sodium hypochlorite for 15 to 45 seconds and rinsed in fresh water prior to slide preparation. All observations were made by brightfield microscopy. Nematocyst cate gories were identified based on Weill's (1934) classifica ranges. I Bermuda ATLANTIC C.h a 11 e n g e r OCEAN Bank Plant agenet Bank 65°|io' 650|oo' I 648|50, Fig. 2. Bermuda, Challenger Bank, and Plantagenet Bank. 640|40, Systematic Account Family Clavidae McCrady, 1859a Clavidae McCrady, 1859a: 123. Turridae Allman, 1872:259. Cordylophorinae von Lendenfeld, 1885a:221. DIAGNOSIS Hydroids solitary or colonial. Colonies stolonal or erect, arising from a creeping hydrorhiza; growth in erect col onies monopodial with terminal hydranths. Perisarc soft or firm, investing hydrorhiza only or covering both hy drorhiza and hydrocaulus, usually terminating at base of hydranth. Hydranths elongate, clavate to fusiform, with scattered filiform tentacles. Hypostome conical. Nematophores present or absent. Gonophores fixed sporosacs or free medusae, arising from hydrorhiza, hydrocaulus, branches, pedicels, or en tire or reduced hydranths. Medusa bell-shaped with short manubrium; mouth of medusa surrounded by four lips, margins of lips with clusters of nematocysts. Radial canals four. Marginal tentacles solitary, numerous in adult. Ocelli present. Gonads on interradial walls of manubrium. REMARKS The family name Turridae was constituted by Allman (1872) for clavid-like hydroids having medusiform gonophores. Taxa referred by Allman to the nominal family are now generally included in the Clavidae McCrady, 1859a. The identity of Turris Lesson, 1843, type genus of the Turri dae, is discussed below under Turritopsis nutricula McCrady, 1859b. The family Clavidae has been used as a catch-all group for a seemingly rather disparate assemblage of hydroids having scattered filiform tentacles on the hydranth. Dif ferences among taxa within the Clavidae are sufficiently large to warrant recognition of several subfamilies at least. In addition to the nominotypical subfamily Clavinae McCrady, 1859a, the Corydendriinae, subf. nov., is rec ognized below. Earlier, von Lendenfeld (1885a) founded the subfamily Cordylophorinae, including in it the genera Cordylophora Allman, 1844, and Merona Norman, 1865. Of these two, only Cordylophora should be retained in the Cordylophorinae, in my opinion. One or more addi tional new subfamilies should be established for clavid genera not represented in Bermuda, but doing so is beyond the scope of this report. Bouillon (1985) included 11 genera of hydroids and hydromedusae in this family. Subfamily Corydendriinae, subf. nov. DIAGNOSIS Clavid hydroids with stolonal or erect colonies; branches of erect colonies adnate to hydrocaulus for a varying dis tance basally. Hydranths elongate, more or less cylindri cal, not polymorphic; tentacles filiform, scattered over much of hydranth. Nematophores absent. Gonophores fixed sporosacs or free medusae, arising from hydrorhiza, hydrocaulus, branches, or pedicels, but not from hydranths. the genera Corydendrium van Beneden, 1844a, Turritopsis McCrady, 1859b, and Rhizodendrium, gen. nov. It is pos sible that the poorly known genus Tubiclava Allman, 1863, if it is valid, belongs in this group. Merona Norman, 1865, having polymorphic hydranths and nematophores, is ex cluded; without doubt it should be referred to another new subfamily within the Clavidae. Genus Corydendrium van Beneden, 1844a REMARKS Hydroids of the Corydendriinae, subf. nov., differ from the Clavinae McCrady, 1859a, in having cylindrical in stead of club-shaped hydranths, tentacles scattered over much of the hydranth rather than restricted to a bulbous region distally, and gonophores borne on hydrorhiza, stem, or branches instead of on the hydranth. Unlike on the Cordylophorinae von Lendenfeld, 1885a, the branches are adnate for some distance at their origin rather than becom ing immediately free, and the hydranths are elongate and tubular rather than spindle-shaped to vasiform. The subfamily Corydendriinae, as defined here, encompasses Corydendrium van Beneden, 1844a:313. Soleniopsis Ritchie, 1908:494. DIAGNOSIS Clavid hydroids with erect, irregularly branched colonies; hydrocaulus polysiphonic. Branches adnate to hydro caulus, or to other branches, over part or all of their lengths. Perisarc firm, terminating near hydranth base. Hydranths elongate, tubular; tentacles filiform, scattered over much of hydranth. Gonophores fixed sporosacs, arising as blind, elongate sacs of coenosarc below hydranths and within perisarcal tubes of branchlets. TYPE SPECIES Sertularia parasitica Linnaeus, 1767, by monotypy. REMARKS L. Agassiz (1862) believed that Corydendrium van Beneden, 1844a, and Cordylophora Allman, 1844, were congeneric, and he referred both to the nominal genus Syncoryna Ehrenberg, 1834. Allman (1872) showed why neither should be referred to Syncoryna, identical with Coryne Gaertner, 1774, and argued that they represented two distinct genera. Colony form, hydranth shape, and characteristics of the gonophores are sufficiently distinctive in Corydendrium to warrant its separation from Cordylophora. Ritchie (1908) recognized the similarities between Cor ydendrium and his nominal genus Soleniopsis, but he mis takenly believed that the former produced free medusae. Stechow (1911) recognized this error and referred Solen iopsis to Corydendrium. Kramp (1935) regarded Turritopsis McCrady, 1859b, as congeneric with Corydendrium, suggesting that hy droids of the two differed only in the type of gonophore produced. Petersen (1979) seems to have adopted this view, but both genera are recognized as valid here and in most other recent publications. Kramp himself, in later reports (e.g., Kramp, 1959, 1961, 1965, 1968), employed the name Turritopsis for the medusa. Corydendrium parasiticum (Linnaeus, 1767) Figs. 3, 4 Sertularia parasitica Linnaeus, 1767:1315. Sertolara parassita—Cavolini, 1785:181; pi. 6, figs. 8-13 [incorrect subsequent spelling]. Pennaria parasitica—Goldfuss, 1820:89. Sertulariam parasiticam—Ehrenberg, 1834:71 [incorrect subsequent spelling]. Syncoryna parasitica—Ehrenberg, 1834:71. Corydendrium parasiticum—van Beneden, 1844b:313. Sertularia (Syncoryne) parasitica—Frey and Leuckart, Fig. 3. Corydendrium parasiticum, part of hydrocaulus with hydranths, romiz B136. Scale equals 1 mm. Corydendrium flabellatum Fraser, 1938:11; pi. 1, figs. a,b. Corydendrium parasticum—Wedler and Larson, 1986:71 [incorrect subsequent spelling]. TYPE LOCALITY "Habitat in Oceano, saepe in Corallina rubente" (Lin naeus, 1767). 1847:30. Syncoryne parasitica—Allman, 1864a:352. Clava parasiticum—Bonnevie, 1899a:9. Clava {Corydendrium) parasiticum—Bonnevie, 1899a: 39. Soleniopsis dendriformis Ritchie, 1908:495; figs. 142, 143; pi. 26, fig. 1. Corydendrium sessile Ritchie, 1910a:802; pi. 76, figs. 1,2. Corydendrium dendriformis—Ritchie, 1910a:803. Corydendrium dendriforme—Gravely, 1927:7; pi. 2, fig. 2. MATERIAL EXAMINED Flatts Inlet, on underside of large, flat rock, - 3 m, 2 August 1982, one colony, 2.2 cm high, without gono phores, romiz B136. Atlantic Ocean, 2 km southeast of Castle Roads, on calcareous rubble, —60 to —90 m, 3 September 1977, one colony, 4.5 cm high, without hy dranths and gonophores, romiz B158. Harrington Sound, near Flatts Inlet bridge, on ledge near shore, —1.5 m, 5 March 1982, two colonies, 4 cm high, without gono phores, romiz B173. Fig. 4. Corydendrium parasiticum, nematocysts of hydranth, romiz B158. Scales equal 10 \xm. a, Desmoneme. b, Heterotrichous microbasic eurytele. DESCRIPTION Colonies erect, straggly, up to 4.5 cm high, arising from a creeping hydrorhiza. Hydrocaulus polysiphonic in all but very young colonies, individual tubes relatively stout, 1910a, as conspecific with C. dendriforme, suggesting that the former was based on a young, immature colony of the latter. Its name, too, is referred to C. parasiticum here. Fraser's (1938) account of Corydendrium flabellatum 0.45 mm in diameter. Branching irregular, in one or more from the Pacific coasts of Mexico and Panama corresponds planes, branches adnate to hydrocaulus basally, gradually curving outwards and becoming free distally; secondary branches arising in like manner from primary branches; ultimate branchlets typically alternate. Perisarc moderately with C. parasiticum, and the former is regarded here as conspecific with the latter. The status of C. fruticosum thick over most of colony, becoming thin at hydranth base and terminating below tentacles, smooth or with occa sional wrinkles but not annulated, clear to straw-coloured, often encrusted with detritus and silt. Hydranths cylindri Fraser, 1914, from the Vancouver Island region is unclear. The colony form of this hydroid appears to have resembled that of C. parasiticum and other nominal species of the genus, but the species was referred to Corydendrium van Beneden, 1844a, with some doubt by Fraser (1914, 1937, 1946) because its gonophores had not been observed. Ac cording to Fraser (1914), hydranths of C. fruticosum have cal through clavate to fusiform, constricted basally below orifice of perisarc tube, up to 2.8 mm long from basal constriction to tip of hypostome, about 0.3-0.5 mm wide. ent in C. parasiticum. Corydendrium fruticosum merits Tentacles filiform, often as many as 40 or more, scattered further study to determine its affinities. 12 to 15 tentacles, far fewer than the number usually pres over distal three-quarters of hydranth, proximal tentacles shorter and more slender than distal ones. Hypostome elon gate, conical. Gonophores not seen. Nematocysts— Hydroids: desmonemes 5.3-5.7 \xm x 3.7-3.8 p,m; KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Wedler, 1975); eastern At lantic (Ritchie, 1908); Indian Ocean (Millard, 1975); west ern Pacific (Leloup, 1937); eastern Pacific (Fraser, 1938). heterotrichous microbasic euryteles 8.2-8.4 p,m x 3.84.3 p-m. Genus Turritopsis McCrady, 1859b REMARKS Ritchie (1910a) believed that Corydendrium dendriforme (Ritchie, 1908) was distinct from C. parasiticum (Lin naeus, 1767) in having (1) thicker, more definite stems; (2) branchlets arranged in a pseudopinnate fashion; and (3) the free portion of the pedicels much less elongate. Rees and Thursfield (1965) also considered C. dendriforme to be valid. However, the characters used to distinguish them seem variable, and Vervoort (1941) and Millard (1959a, 1975) have been followed in regarding the name C. dendriforme as a junior synonym of C. parasiticum. Leloup (1937) regarded Corydendrium sessile Ritchie, Clavula Wright, 1859:106. Turritopsis McCrady, 1859b:58. Dendroclava Weismann, 1883:26. Turrutopsis Wedler and Larson, 1986:71 [incorrect sub sequent spelling]. DIAGNOSIS Clavid hydroids with stolonal or erect and irregularly branched colonies; hydrocaulus monosiphonic or polysi phonic. Branches adnate to hydrocaulus, or to other branches, over part of their length. Perisarc firm, termi- nating near hydranth base. Hydranths elongate, tubular; tentacles filiform, scattered over much of hydranth calyx. Gonophores free medusae, developing on pedicels or branches below hydranths. Medusa with eight or more simple, solitary marginal tentacles. Radial canals sur rounded by mass of vacuolated cells at apex of stomach. Ocelli present. TYPE SPECIES Turritopsis nutricula McCrady, 1859b, by monotypy. REMARKS The familiar and widely used name Turritopsis McCrady, 1859b, is predated by the nearly forgotten ClavulaWright, 1859, and the two are considered synonyms here, as dis cussed below. Although McCrady"s (1859b) paper pro posing the generic name Turritopsis was presented orally before a meeting of the Elliott Society of Natural History Fig. 5. Turritopsis nutricula, two hydranths arising from sponge substrate, romiz B172. Scale equals 0.5 mm. of Charleston, South Carolina, on 1 December 1856, the proceedings of the meeting were not published until 1859. Only the year of publication is indicated on the cover of these proceedings. In the absence of other information, the date must be taken as the last day of the year [Art. 21c (ii)]. Wright's (1859) account of Clavula, in the July 1859 issue of the Edinburgh New Philosophical Journal, Modeeria multitentacula Fewkes, 1881:149; pi. 3, figs. 7-10 [medusa]. Modeeria nutricula—Fewkes, 1882:294; pi. 10, fig. 8 [medusa]. Modeeria {Turritopsis) nutricula—Fewkes, 1883:80 must be interpreted as having been published first. Ap plicationwill be made to the iczn to use its plenarypowers [Art. 79] to suppress the virtually unused name Clavula Wright, 1859, in favour of the well-known Turritopsis. Russell (1953) included Turris neglecta Lesson, 1843, as questionably conspecific with Turritopsis nutricula, the Modeeria multitentaculata—Brooks, 1883a: 144[medusa] [incorrect subsequent spelling]. Oceania nutricula—Brooks, 1883b:465 [medusa]. Modeeria nutricola—Brooks, 1886:388 [medusa] [incor type species of Turritopsis. However, he noted that Les Turritopsis polycirrha—Hartlaub, 1897:480; pi. 16c, fig. son's medusa was inadequately described and its identity uncertain. The genus name Turris Lesson, 1843, is re garded here as a nomen dubium. Mueller (1766) had earlier applied the name Turris to a genus of the Mollusca, but Mueller's publication has been placed on the Official Index [medusa]. rect subsequent spelling]. 2 [medusa]. Turritopsis nutricola var. pacifica Maas, 1911:14; pi. 1, figs. 6-8; pi. 2, fig. 9 [medusa] [incorrect subsequent spelling]. Corydendrium nutricula—Kramp, 1935:11. Turrutopsis nutricula—Wedler and Larson, 1986:71 [in of Rejected and Invalid Works in Zoological Nomenclature by the iczn (Opinion 701). correct subsequent spelling]. Turritopsis nutricula McCrady, 1859b TYPE LOCALITY Figs. 5, 6 Charleston Harbour, South Carolina, United States. ITurris neglecta—Forbes, 1848:23; pi. 3, figs. 2a-i [me dusa] [Turris neglecta Lesson, 1843, a nomen dubium]. Clavula gossii Wright, 1859:106; pi. 8, fig. 1. Oceania {Turritopsis) nutricula McCrady, 1859b:56; pi. 4,figs. l-10,12-15,28a;pl.5,figs. 11,16-18,28b[medusa]. Turritopsis nutricula McCrady, 1859b:58 [medusa]. Turritopsis nutricola—L. Agassiz, 1862:347 [medusa] [incorrect subsequent spelling]. Oceania polycirrha Keferstein, 1863:26; pi. 2, figs. 1113 [medusa]. Turritopsispolynema Haeckel, 1879:66 [medusa]. 8 MATERIAL EXAMINED Whalebone Bay, on sponge, - 1 m, 7 September 1977, two colonies, 5 mm high, without gonophores, romiz B162. Flatts Inlet, on sponge, -2 m, 5 March 1982, one colony, with hydranths extending 1.5 mm above sponge substrate, without gonophores, romiz B172. DESCRIPTION Colonies erect, with hydrorhiza and much of hydrocaulus embedded in sponge, sparingly and irregularly branched, reaching 5 mm high. Hydrocaulus monosiphonic, branches Fig. 6. Turritopsis nutricula, nematocysts of hydranth, romiz B172. Scales equal 10 am. a, Desmoneme. b, Heterotrichous microbasic eurytele. adnate to hydrocaulus at their origin, curved outwards and becoming free distally; hydrocaulus and branches slender basally, 0.12 mm wide, gradually expanding in diameter distally, reaching 0.23 mm wide at base of hydranth. Perisarc moderately thick, clear to straw-coloured, with fine longitudinal creases and occasional wrinkles but no annulations, terminating below tentacles on hydranth base. Hydranths fusiform, reaching 0.9 mm long from base to tip, 0.2 mm wide. Tentacles filiform, 12 to 20 in number; about 4 in an irregular whorl distally, remainder scattered over distal two-thirds to three-quarters of hydranth, those at proximal end shorter and more slender than those at distal end. Hypostome elongate, conical. Gonophores not seen. Nematocysts— Hydroids: desmoncmes 4.6-4.8 am x 2.8-3.1 am; heterotrichous microbasic euryteles (small) 6.5-6.8 am X 3.1-3.3 am. REMARKS Both hydroid and medusa stages of this species have long been known as Turritopsis nutricula McCrady, 1859b, but this binomen is actually predated by the virtually forgotten name Clavula gossii Wright, 1859. This nomenclatural problem arises in part from Wright's application of a sep arate name to the hydroid of a medusa he believed was conspecific with Turris neglecta Lesson, 1843. His con temporaries (e.g., Allman, 1864a, 1872; Hincks, 1868) realized that this contravened nomenclatural principles, and the name C. gossii was included in the synonymy of T. neglecta in their monographs. However, T. neglecta is now generally considered to be a nomen dubium. Ac cording to Russell (1953), Lesson's (1843) description of the medusa was inadequate for positive identification. Rus sell (1953) believed that the medusa described by Wright (1859) was identical with T. nutricula, and he included the name C. gossii as its junior synonym. This interpre tation of relative priority was based on the widespread but mistaken belief (e.g., see A. Agassiz, 1865; Mayer, 1910; Fraser, 1944; Russell, 1953; Kramp, 1961; Vervoort, 1968; Millard, 1975) that McCrady's (1859b) original descrip tion of T. nutricula was published in 1856, as discussed earlier (see p. 8). Clavula gossii has not been used as a senior synonym since it was instituted by Wright (1859), to my knowledge. Upon completion of this study, appli cation will be made to the iczn to use its plenary powers [Art. 79] to suppress the name Clavula gossii Wright, 1859, in favour of Turritopsis nutricula McCrady, 1859b. I have followed Mayer (1910) and Russell (1953) in regarding Oceania polycirrha Keferstein, 1863, Turritop sis polynema Haeckel, 1879, and Modeeria multitentacula Fewkes, 1881, as conspecific with this species. Most authors have adopted the view of Stechow (1923a) that Turritopsis dohrnii (Weismann, 1883) is conspecific with 7'. nutricula. Hydroids of the two appear to differ in both colony form and habitat. Unlike colonies of T. nu tricula, which are small with a monosiphonic hydrocaulus and usually found in shallow water, hydroids of T. dohrnii are larger with a polysiphonic hydrocaulus and known from deeper waters. Numerous specimens of T. nutricula have been observed from shallow-water habitats of Vir ginia (Calder, 1971) and South Carolina, including the type locality of Charleston Harbour (Calder and Hester. 1978). None of these hydroids were more than a few mil limetres in height, and none had polysiphonic stems like T. dohrnii. Specimens of T. nutricula from nearshore waters of Bermuda resembled those examined from the American east coast in colony form. Both species are recognized as valid here because conclusive evidence that differences in colony form in the two may be environmentally induced is lacking. Young medusae have been described from both species, but a critical comparison of the two has not been made. The adult medusa of T. dohrnii is apparently un known. Specimens identified as T. dohrnii (romiz B139) were found on a brachyuran crab collected at a depth of 256 m off Castle Roads, Bermuda, during this study. However, the species has been excluded from this report, which includes hydroids from the upper 100 m only. Turritopsisfascicularis Fraser, 1943b, collected from a depth of 118 fathoms (216 m) off Florida, appears to be identical in most respects with descriptions of T. dohrnii, and is regarded here as conspecific with the latter rather than with T. nutricula. McCrady (1859b) mistook the parasitic actinula larva of the narcomedusa Cunina octonaria McCrady, 1859a, a frequent parasite on the medusa stage of Turritopsis nutricula, for the hydroid of this species. Excellent de scriptions and illustrations of the hydroids and young me dusae of T. nutricula were given by Brooks (1886). KNOWN RANGE Bermuda: Castle Harbour, medusa stage (Fewkes, 1883); in shallow inshore waters and on buoy chains (Calder, 1986). Elsewhere: western Atlantic (Fraser, 1944); eastern At lantic (Russell, 1953); Indian Ocean (Millard, 1975); west ern Pacific (Ralph, 1953); eastern Pacific (Fraser, 1948). Genus Rhizodendrium, gen. nov. DIAGNOSIS Clavid hydroids with creeping hydrorhiza and sessile, elongate hydranths. Hydranths invested with thin perisarc proximally, occasionally forming a basal collar; tentacles filiform, as many as 20 or more, arranged in a more or less regular whorl around mouth, scattered or in several irregular whorls proximally, those of distal end longer and stouter than those proximally. Hypostome short, domeshaped. Gonophores fixed sporosacs, borne on hydrorhiza, spherical where known. siformis (mcz 52), confirmed Agassiz's original descrip tion of Rhizogeton in most respects, although the specimens were in rather poor condition. Nevertheless, it was ap parent from this examination that Rhizodendrium can be held distinct from Rhizogeton, on the following characters. The hypostome is short and dome-shaped, instead of very elongate and conical. Tentacles number as many as 20 or more, instead of a maximum of 10. The tentacles are arranged in an oral whorl distally, and sometimes occur in more or less regular whorls elsewhere on the hydranth, instead of being decidedly scattered. The shape of the hydranth in Rhizodendrium is much like that of Turritopsis McCrady, 1859b, and somewhat less like Corydendrium van Beneden, 1844a. The colony form is strictly stolonal, unlike that of Turritopsis and Corydendrium, and the gonophores arise from the stolon rather than from branchlets or pedicels. In Tubiclava Allman, 1863, gonophores were believed to arise in dense clusters from the hydranth, as in Clava (Allman, 1863, 1872), and not from the hydrorhiza, as in Rhizodendrium. Two nominal species referred to Rhizogeton—R. nudus Broch, 1909, from Spitzbergen and R. ezoense Yamada, 1964, from Japan—are transferred here to Rhizodendrium. The only species thus remaining in Rhizogeton is R. fu siformis. Rhizogeton nematophorus Antsulevich and Polteva, 1986, from the USSR is a polymorphic species with nematophores, and seems sufficiently distinct to be rec ognized as a new genus. Gonophores have not been observed in the type species of Rhizodendrium. However, those of R. nudum and R. ezoense are spherical rather than fusiform, as in Rhizogeton. Rhizodendrium sterreri, sp. nov. Figs. 7, 8 TYPE SPECIES Rhizodendrium sterreri, sp. nov., designated herein. ETYMOLOGY The name is a combination of parts of the names Rhizogeton and Corydendrium, and is derived from the Greek words rhiza (root) and dendron (tree). The gender of the MATERIAL EXAMINED Holotype: Whalebone Bay, on pelagic Sargassum, 2 Sep tember 1977, one colony, 2 mm high, without gono phores, romiz B150. Paratype: Whalebone Bay, on pelagic Sargassum, 2 September 1977, one colony, 2 mm high, without gonophores, romiz B305. name is neuter. DESCRIPTION REMARKS Rhizodendrium, gen. nov., resemblesRhizogeton L. Agassiz, 1862, a genus established to accommodate the hydroid R. fusiformis L. Agassiz, 1862, from tide pools on the Massachusetts coast. L. Agassiz (1862) observed that Rhi zogeton was similar to Clava Gmelin, 1790, but differed in having a thin covering of perisarc over the proximal part of the hydranth, a different hydranth shape, and gonophores on the hydrorhiza instead of the hydranth. Examination of type material of the type species, R. fu 10 Colonies stolonal, with a creeping reticular hydrorhiza bearing sessile hydranths. Perisarc smooth or wrinkled, moderately thin on hydrorhiza, extending up 1 mm or more over base of hydranth or restricted to little more than a short collar between hydranth and hydrorhiza. Hydranths elongate, cylindrical, up to 2.0 mm long, 0.3 mm wide, with as many as 20 or more tentacles. Tentacles filiform, 4 to 6 in an irregular whorl distally, remainder scattered over distal two-thirds to three-quarters of hydranth, those at distal end longer and stouter than those proximally. Fig. 7. Rhizodendriumsterreri, sp. nov., hydranth from holo type colony, romiz B150. Scale equals 0.5 mm. a * (.^ I > , ,::: •' » • ':":•. tpfe, J • 1 > % m*.^ !!• * * #"# b \ ••• 'W • :>:* * Fig. 8. Rhizodendrium sterreri, sp. nov., nematocysts of hydranth of holotype colony, romiz B150. Scales equal 10 ^m. a, Desmonemes. b, Heterotrichous microbasic eurytele. tacles. They also differ from those of R.fusiformis in being much smaller, in possessing more tentacles, and in having longer and stouter tentacles at the distal than at the prox Hypostome dome-shaped. Gonophores lacking. Nematocysts— Hydroids: desmonemes 4.6-5.2 |xm x 2.8-3.1 heterotrichous microbasic euryteles 6.6-7.6 (xm x 2.9 |xm. • m uum; 2.6- REMARKS Hydranths of Rhizodendrium sterreri, sp. nov., are distinct from L. Agassiz's (1862) description and illustrations of Rhizogeton fusiformis in having an irregular oral whorl of four to six tentacles instead of widely scattered distal ten imal end. Rhizodendrium sterreri is very similar to descriptions of its two congeners, R. nudum (Broch, 1909) and R. ezoense (Yamada, 1964), in hydranth shape and size, as well as in tentacle number, size, and arrangement. How ever, in R. sterreri the base of the hydranth is sheathed in perisarc, as is obvious when the soft tissues are dissolved in sodium hypochlorite, whereas perisarc is reportedly ab sent from the base in/?, nudum (Broch, 1909; Dons, 1913). 11 Hydranths of R. sterreri appear to be smaller than those of/?, ezoense. Nematocyst data exist only from R. sterreri, but cnidome and nematocyst size are unlikely to differ much in these hydroids. Rhizodendrium nudum has been reported several times from warm-water localities, but the records seem ques tionable on zoogeographic grounds. Ritchie (1910b) iden tified a hydroid from the Christmas Islands in the Indian Ocean as "Rhizogeton nudum Broch (?)," and indicated that his specimens lacked perisarc at the base of the hy dranth. Rees and Thursfield (1965) expressed some doubt that Ritchie's specimens were identical with R. nudum, and suggested that they might represent a juvenile colony of another species. Mammen (1963) referred specimens from south India to R. nudum, but noted that they possibly belonged to another species. Millard and Bouillon (1974) reported R. nudum from Mozambique, and stated that their record confirmed the existence of the species in the Indian Ocean. Their hydroids, unlike those of/?, nudum described by Broch (1909), had a collar of perisarc at the base of the hydranth. Some, if not all, of these records may be based on specimens of R. sterreri. In a study of the hydroids on pelagic Sargassum in the western Atlantic, Burkenroad {in Parr, 1939) mentioned finding a hydroid resembling the genus Clava Gmelin, 1790. It seems highly probable that it was R. sterreri. ETYMOLOGY The species is named after Dr Wolfgang Sterrer, former director of the Bermuda Biological Station, who provided the initial stimulus for this study. KNOWN RANGE Known only from the type locality. Family Bougainvilliidae Lutken, 1850 Bougainvilleae Lutken, 1850:29 (emended to Bougain villiidae by Allman, 1876). Hippocrenidae McCrady, 1859a: 158. Nemopsidae L. Agassiz, 1862:345. Dicorynidae Allman, 1864a:366. Atractylidae Hincks, 1868:87. Bimeridae Allman, 1872:294 (emended to Bimeriidae by Torrey, 1902). Margelidae Haeckel, 1879:68. Lizusidae Haeckel, 1879:80. Thamnostomidae Haeckel, 1879:84. Pachycordylini Cockerell, 1911:77. Lizziinae Russell, 1953:144. Clavopsellidae Thiel, 1962:249. DIAGNOSIS Hydroid colonies stolonal or erect, arising from a creeping hydrorhiza; growth monopodial with terminal hydranths. Perisarc on hydrorhiza and hydrocaulus of varied thick ness, terminating at base of hydranth or extending over hydranth as a thin, filmy pseudohydrotheca. Hydranths cylindrical through fusiform to vasiform, with one or more whorls of filiform tentacles beneath conical to nipple-shaped hypostome. Gonophores fixed sporosacs or free medusae, borne on hydrorhiza, hydrocaulus, branches, and pedicels, or on entire or reduced hydranths. Medusae bell-shaped with short manubrium; mouth circular; oral tentacles simple or dichotomously branched, inserted above mouth. Radial canals four. Marginal tentacles either solitary or in clus ters, borne on 4, 8, or 16 tentacle bulbs. Ocelli present or absent. Gonads on manubrium, either forming a con tinuous ring or on interradial, adradial, or perradial axes. 12 REMARKS Characters of hydroids belonging to the Bougainvilliidae Lutken, 1850, have recently been reviewed by Millard (1975). She noted (1975:71) that difficulties may be en countered when drawing dividing lines between hydroids of the Clavidae McCrady, 1859a, Hydractiniidae L. Agas siz, 1862, Eudendriidae L. Agassiz, 1862, and Bougain villiidae. Medusae of the Bougainvilliidae have a number of characteristics, including the presence of oral tentacles, in common with those of the Cytaeididae L. Agassiz, 1862, and Russelliidae Kramp, 1957. Because of these similarities, Petersen (1979) placed the three families to gether in the superfamily Bougainvillioidea Lutken, 1850. The family Bougainvilliidae, as currently classified, in cludes a seemingly disparate assemblage of hydroids and medusae. Russell (1953) recognized three subfamilies, the Bougainvilliinae Lutken, 1850, Lizziinae Russell, 1953, and Thamnostominae Haeckel, 1879, within the group. Four subfamilies are distinguished here among the bougainvilliids of Bermuda. In addition to the Bougainvillii nae, Bimeriinae Allman, 1872, and Pachycordylinae Cockerell, 1911, a new subfamily is recognized and de fined, the Rhizorhagiinae. The subfamily Bimeriinae, as used here, is roughly equivalent in scope to the Tham nostominae as defined by Russell (1953). Neither hydroids nor medusae of the Lizziinae have been reported from Bermuda. According to Haeckel (1879), the family-group names Margelidae, Lizusidae, and Thamnostomidae were first used in his 1877 manuscript "Prodromus Syst. Medusen." However, this was an unpublished document (Kramp, 1961:400), and the three names were not published [Art. 8] until the appearance of the later work (Haeckel, 1879). Subfamily Pachycordylinae Cockerell, 1911 DIAGNOSIS Bougainvilliid hydroids with perisarc terminating at base of hydranth. Hydranths club-shaped through spindle-shaped to amphora-shaped; hypostome dome-shaped; tentacles in Genus Millardiana Wedler and Larson, 1986 Millardiana Wedler and Larson, 1986:90. two or more close whorls. DIAGNOSIS Gonophores, where known, fixed sporosacs or free but sometimes degenerate medusae. Hydroid colonies mostly stolonal, with perisarc terminat ing at hydranth base. Hydranth thick, clavate; tentacles filiform, scattered around distal end of hydranth. Gonophores sporosacs, borne on gonozooids beneath REMARKS Whereas hydroids of the subfamily Bougainvilliinae Lut ken, 1850, have tentacles arranged more or less in a single whorl on the hydranth, representatives of the tribe Pachycordylini Cockerell, 1911, herein elevated to the rank of subfamily, have tentacles in two or more whorls. Often these whorls are rather indistinct, and the tentacles may be essentially scattered. Nevertheless, the tentacles are restricted to a relatively narrow band on the hydranth and are thereby distinguishable from hydroids of the family Clavidae McCrady, 1859a. Thiel (1962) established the nominal family Clavopsellidae for Clavopsella Stechow, 1919, and Balella Stechow, 1919, but Stechow (1922) had earlier constituted the family name Balellidae for the latter genus. Nutting (1905) applied the name Tubidendridae to the Balellidae, but this name is not available because it was not based on whorl of four to five tentacles. TYPE SPECIES Millardiana longitentaculata Wedler and Larson, 1986, by monotypy. REMARKS Millardiana was established by Wedler and Larson (1986) for a hydroid resembling Pachycordyle Weismann, 1883, but differing from that genus in having gonophores on gonozooids with four to five tentacles instead of zooids with the normal complement of tentacles. They referred the genus to the family Bougainvilliidae Lutken, 1850. It is tentatively referred here to the Pachycordylinae Cock erell, 1911, although the presence of gonozooids is anom alous. Millardiana is a monotypic genus. a name then valid for a contained genus [Art. 1If (i)(l)]. Little is known about the medusa stages of either of these two genera. All of the species heretofore included in Cla vopsella possess fixed gonophores or degenerate medusae. Millardiana longitentaculata Wedler and Larson, 1986 Figs. 9, 10 Jaderholm (1919) observed medusa buds in Balella mir- abilis (Nutting, 1905), but was unable to provide infor mation beyondsize and presence of short marginal tentacles. Balella is regarded here as closer to the Clavidae than to the Bougainvilliidae because tentacles are present on the proximal as well as on the distal part of the hydranth. Accordingly, the family Balellidae is recognized here as a valid taxon. The nominal genus Clavopsella is regarded as a junior synonym of Pachycordyle Weismann, 1883, in this report, and the name Pachycordylinae has priority over Clavopsellinae. The affinities of the genus SilhouettaMillard and Bouil lon, 1973, are uncertain, but it has been included here in the Pachycordylinae because of the scattered arrangement of the tentacles and the peculiar domelike shape of the hypostome. Representatives of the Pachycordylinae appear inter mediate between bougainvilliids and clavids, but they are retained in the Bougainvilliidae here based on character istics of the medusa of Silhouetta. Medusa buds of Sil houetta uvacarpa Millard and Bouillon, 1973, were described as being "without doubt'' bougainvilliid in char acter by Millard and Bouillon (1973, 1975). Millardiana longitentaculata Wedler and Larson, 1986:90; figs. 7Ba,b;pl. 1, fig. 8. TYPE LOCALITY La Parguera, Puerto Rico. MATERIAL EXAMINED Green Bay, Harrington Sound, on shells of Cerithium lit teratum from Cladophora bed, -2.5 m, 21 September 1984, colonies without gonophores on four shells, romiz B371. DESCRIPTION Colonies stolonal, with hydrorhiza adhering to gastropod shells. Pedicels very short, each bearing a terminal hy dranth. Perisarc thin, smooth or slightly wrinkled, ter minating at base of hydranth; pseudohydrotheca absent. Hydranths clavate to ovate to nearly fusiform, reaching 1.2 mm long, 0.34 mm wide at widest point, distal end with about 10 to 25 long, filiform tentacles in two to three close whorls, those of one whorl more or less alternating with those of adjacent whorls, proximal tentacles often 13 Fig. 9. Millardiana longitentaculata, hydranth, romiz B371. Scale equals 0.5 mm. Fig. 10. Millardiana longitentaculata, nematocysts of hydranth, romiz B371. Scales equal 10 am. a, Desmoneme. b, Heterotrichous microbasic eurytele. smaller than distal ones. Hypostome prominent, extensi ble, and proboscis-like. Colour of hydranth deep red in Bermuda: first record. live material. Elsewhere: western Atlantic (Wedler and Larson, 1986). Gonophores not seen. Nematocysts— Hydroids: desmonemes 5.6-6.4 am x 3.4-3.8 am; heterotrichous microbasic euryteles 7.6-10.4 am x 2.83.8 p.m. REMARKS Although superficially similar in colony form to Pachy cordyle napolitana Weismann, 1883, Millardiana longi tentaculata Wedler and Larson, 1986, is clearly a distinct species. Characteristics of the latter distinguishing it from the former include the intense red pigmentation of the hydranths and the extremely long tentacles and hypostome. According to Wedler and Larson (1986), gonophores are sporosacs borne on polyps having only four to five ten tacles. From their description and illustration, these re markable polyps somewhat resemble the gonozooids of the family Hydractiniidae L. Agassiz, 1862, and are atyp ical of hydroids of the Bougainvilliidae Lutken, 1850. 14 KNOWN RANGE Genus Pachycordyle Weismann, 1883 Pachycordyle Weismann, 1883:87. IParvanemus Mayer, 1904:6. Pachycordile Lo Bianco, 1909:544 incorrect subsequent spelling]. Clavopsella Stechow, 1919:21. DIAGNOSIS Bougainvilliid hydroids with the characters of the subfamily. Gonophores fixed sporosacs or free but degenerate me dusae. Medusa ephemeral, pyriform, without mouth, ra dial canals, marginal tentacles, oral tentacles, or ocelli. TYPE SPECIES Pachycordyle napolitana Weismann, 1883, by monotypy. REMARKS Pachycordyle weismanni Hargitt, 1904a:553; pi. 21, figs. Pachycordyle napolitana Weismann, 1883, and P. weis manni Hargitt, 1904a, type species of the nominal genera Pachycordyle Weismann, 1883, and Clavopsella Stechow, 1919, respectively, are regarded here as conspecific fol lowing Picard (1958) and Morri (1981). The name Cla vopsella can therefore be regarded as a junior subjective synonym of the name Pachycordyle, resurrected here. Picard (1958) and Morri (1981) included P. napolitana in the genus Cordylophora Allman, 1844. Pachycordyle differs from Cordylophora in having tentacles arranged in 1-8. two or more close whorls rather than scattered over much of the hydranth. Stechow (1919) proposed the generic names Clavopsella and Balella for species of Filifera having more than one whorl of tentacles on the hydranth. Unlike hydroids of Balella, which have two widely separated whorls of ten tacles, hydroids of Clavopsella (= Pachycordyle) have Cordylophora annulata Motz-Kossowska, 1905:66; fig. 5. Pachycordyle neapolitana—Motz-Kossowska, 1905:70 [incorrect subsequent spelling]. Perigonimus neapolitanus—Motz-Kossowska, 1905:75; fig. 8 [incorrect subsequent spelling] [not Perigonimus napolitanus Hargitt, 1904a]. Pachycordile weismanni—Lo Bianco, 1909:544 [incorrect subsequent spelling]. Tubiclava annulata—Stechow, 1912:343; pi. 13, fig. 8. Clavopsella weismanni—Stechow, 1919:22. Clavopsella annulata—Stechow, 1921a:250. Rhizorhagium {Pachycordyle) napolitanum—Stechow, 1923a:56. Cordylophora neapolitana—Picard, 1958:189 [incorrect subsequent spelling]. from two to four close whorls of tentacles. This charac teristic is shared with Silhouetta Millard and Bouillon, 1973, a genus with well-developed medusa buds instead of fixed sporosacs or free but degenerate medusae. The nominal genus Clavopsella was initially referred to the family Bougainvilliidae Lutken, 1850, by Stechow (1919), but was later transferred to the Clavidae McCrady, 1859a, by Stechow (1923a). Thiel (1962) placed Clavop sella with Balella in a new family, the Clavopsellidae, a family he considered intermediate between the Clavidae and the Bougainvilliidae. As noted above, Balella is re garded as closer to the Clavidae than the Bougainvilliidae and is returned here to the family Balellidae. Clavopsella (i.e., Pachycordyle) is classified among the Bougainvil liidaein this report, following Millard (1975) and Bouillon (1985). The definition of Pachycordyle adopted above differs from that used by Thiel (1962) and Millard (1975) for Clavopsella. They included Clavopsella quadranularia Thiel, 1962, and Rhizorhagium navis Millard, 1959b, nominal species likely referable to Aselomaris Berrill, 1948, in Clavopsella. Aselomaris, in my opinion, belongs in the subfamily Bougainvilliinae rather than in the Pachycordylinae. Parvanemus Mayer, 1904, established for a species {P. degeneratus Mayer, 1904)whosemedusalackedtentacles, radial canals, marginal sense organs, and presumably a ring canal, was included by Mayer (1910) in the synonymy of Pachycordyle. Pachycordyle napolitana Weismann, 1883 Figs. 11, 12 Pachycordyle napolitana Weismann, 1883:87; pi. 6, fig. 6. Fig. 11. Pachycordyle napolitana, hydranth, romiz B154. Scale equals 0.5 mm. 15 Fig. 12. Pachycordyle napolitana, nematocysts of hydranth, romiz B154. Scales equal 10 |jum. a, Des monemes. b, Heterotrichous microbasic eurytele. TYPE LOCALITY Naples, Italy. MATERIAL EXAMINED Green Bay, Harrington Sound, on gastropod from Thalassia bed, —2 m, 8 March 1982, one colony, 2 mm high, with an incipient gonophore, romiz B154. Branching of the hydrocaulus is not regarded here as par ticularly reliable in separation of the two. Radial canals were not reported in the degenerate and ephemeral me dusae of either nominal species in earlier descriptions (Har gitt, 1904a; Motz-Kossowska, 1905; Stechow, 1919, 1923a). A vestigial ring canal, observed in the medusa of Pachy cordyle weismanni by Hargitt (1904a), was not reported in P. annulata by Motz-Kossowska (1905). Hydroids and DESCRIPTION degenerate medusae of both nominal species appear similar Colony stolonal, with reticular hydrorhiza growing over a gastropod shell. Pedicels of varied length but usually less than 1 mm long, slender basally, widening distally, bearing a terminal hydranth. Perisarc moderately thin, wrinkled throughout, terminating at base of hydranth; pseudohydrotheca absent. Hydranths club-shaped to spindle-shaped, reaching 1.0 mm long, 0.45 mm wide at widest point, distal end of fully developed hydranth with based on existing descriptions, and the two have been regarded conspecific by Picard (1958), Morri (1981), and others. I have followed Picard and Morri in regarding both of these as conspecific with P. napolitana Weismann, 1883. Picard (1958) included Mediterranean records of Tubiclava fruticosa Allman, 1871, under this species as well. Material from Bermuda discussed here closely resem bles specimens (romiz B695) and published descriptions about 16 to 20 filiform tentacles in three or four close of Pachycordyle napolitana from the Mediterranean, and whorls, tentacles of one whorl alternating with those of adjacent whorls, proximal tentacles often smaller than dis tal ones. Hypostome dome-shaped. Incipient gonophore(?) arising singly from hydranth pedicel on short, wrinkled stalk, completely invested with perisarc. Sex indeterminable. Nematocysts— Hydroids: desmonemes 5.6-6.0 p,m x 3.0-3.6 |xm; heterotrichous microbasic euryteles 9.1-9.6 u,m x 3.8- has been identified as such. However, specimens with well-developed gonophores are needed for more definitive identification. KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Wedler and Larson, 986); Mediterranean Sea (Morri. 1981). 4.5 p,m. Genus Silhouetta Millard and Bouillon, 1973 REMARKS Thiel (1962) distinguished Clavopsella weismanni (Har gitt, 1904a) and C. annulata (Motz-Kossowska, 1905) on the basis of hydroid colony shape and presence or absence of radial canals in the medusa. According to his key, the hydrocaulus of C. weismanni is slightly branched and its medusa has radial canals, whereas in C. annulata the hy drocaulus is unbranched and radial canals are lacking. 16 Silhouetta Millard and Bouillon, 1973:25. DIAGNOSIS Hydroid colonies stolonal or erect, with firm perisarc ter minating at hydranth base. Hydranths large, amphorashaped; tentacles filiform, in two or more close whorls, those of one whorl alternating with those of adjacent whorls. Gonophores free medusae, arising in clusters from stem or branches. Medusae at liberation with four simple or dichotomously branched oral tentacles arising above mouth. Tentacle bulbs four; marginal tentacles four, solitary. Ocelli present. distally, terminating at hydranth base. Hydranth amphorashaped, with conical hypostome, wrinkled basally, reach ing 1.5 mm long from base to tip of hypostome, up to 0.9 mm wide at widest point, bearing filiform tentacles distally. Young hydranths with about 8 tentacles in two whorls; older hydranths with up to 24 tentacles in four TYPE SPECIES ratherclose whorls, tentacles of one whorlalternating with Silhouetta uvacarpa Millard and Bouillon, 1973, by those of adjacent whorls, proximal tentacles often shorter monotypy. and more slender than distal ones. REMARKS The genus Silhouetta was founded by Millard and Bouillon (1973) to accommodate the hydroid S. uvacarpa Millard and Bouillon, 1973, from the Seychelles. Although they recognized the similarity of their hydroid to specimens previously referred to Clavopsella Stechow, 1919, a new genus was established because well-developed medusa buds were present instead of fixed sporosacs or degenerate Gonophores lacking. Nematocysts— Hydroids: desmonemes 6.7-7.6 |xm x 3.7-3.9 |xm; heterotrichous microbasic euryteles 9.6-12.7 |xm x 4.45.6 (Jim. medusae. Silhouetta appears to be distinct from Clavopsella, and its senior synonym Pachycordyle Weismann, 1883, even if gonophore type is not regarded as a valid generic char acter. Although hydranth shape is typically variable in bougainvilliid hydroids, all the hydranths of 5. uvacarpa from Bermuda, like those illustrated by Millard and Bouil lon (1973) from the Seychelles, had a characteristic am phora shape. Silhouetta uvacarpa Millard and Bouillon, 1973 Figs. 13, 14 Silhouetta uvacarpa Millard and Bouillon, 1973:25; figs. 3A-D; pis. 2, 3. Silhouetta puertoricensis Wedler and Larson, 1986:91; figs. 9Aa,b. TYPE LOCALITY Silhouette, Seychelles. MATERIAL EXAMINED Sailor's Choice Cave, near Walsingham Pond, Hamilton Parish, - 1.0 m, 6 July 1982, 16 colonies, 1.0-2.3 cm high, without gonophores, coll. T. Iliffe, romiz B138. DESCRIPTION Colonies initially stolonal, later erect, reaching 2.3 cm high, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic in young colonies, polysiphonic in older ones, irregularly branched; primary branches unbranched or irregularly branched; hydrocaulus and branches curved and twisted, imparting a straggly appearance to colony. Perisarc irregularly wrinkled and creased but with annulations absent, moderately thick and golden in colour basally, becoming progressively thinner and more colourless Fig. 13. Silhouetta uvacarpa, hydranth, romiz B138. Scale equals 0.5 mm. 17 Fig. 14. Silhouetta uvacarpa, nematocysts of hydranth, romiz B138. Scales equal 10 (xm. a, Desmoneme. b, Heterotrichous microbasic eurytele. 1.0 mm wide, with four dichotomously branched oral REMARKS These hydroids lacked gonophores, but resembled the orig inal description of Silhouetta uvacarpa Millard and Bouil lon, 1973, in all other major respects, including nematocyst complement and size. Fully developed medusa buds and newly liberated me dusae of Silhouetta uvacarpa were described by Millard and Bouillon (1973, 1975). Large medusa buds bore four marginal tentacle bulbs, each with a single tentacle and black ocellus, and simple oral tentacles inserted above the mouth. Newly liberated medusae were 0.9 mm high and tentacles. Silhouetta puertoricensis, described by Wedler and Lar son (1986) from Puerto Rico, is regarded here as conspe cific with S. uvacarpa. Wedler and Larson established a new species for their material because oral tentacles could not be seen in the medusa buds. KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Wedler and Larson. 1986); Indian Ocean (Millard and Bouillon, 1973). Subfamily Rhizorhagiinae, subf. nov. DIAGNOSIS Bougainvilliid hydroids with perisarc extending as a pseu dohydrotheca around hydranth. Hydranths vasiform; hy postome nipple-shaped; tentacles in two or more close whorls. Gonophores, where known, fixed sporosacs. Bimeriinae, in which perisarc extends as a sheath over the bases of the tentacles and the hypostome. Included in the subfamily are the genera Rhizorhagium M. Sars, 1874, and Parawrightia Warren, 1907. Although only Parawrightia is represented in Bermuda, Rhizorha gium was chosen as the type genus of the taxon because it is better known [Recommendation 64A], REMARKS The hypostome of the Rhizorhagiinae, subf. nov., is elon gate and nipple-shaped instead of dome-shaped as in the Bougainvilliinae Lutken, 1850, Pachycordylinae Cocker Genus Parawrightia Warren, 1907 ell, 1911, and Bimeriinae Allman, 1872. Moreover, the hydranths are decidedly vasiform, resembling those of the Eudendriidae L. Agassiz, 1862, as much or more than those of other subfamilies of the Bougainvilliidae. This is not to imply that the subfamily is especially close to the Eudendriidae. Unlike in eudendriids, in the Rhizorhagiinae the hypostome is not flared or knob-shaped and gonophores do not arise from the hydranths. Like the Pachycordylinae, this subfamily has hydroids with tentacles arranged in two or more close whorls on the hydranth. A pseudohydrotheca is present, but it is not as extensively developed as in the 18 Parawrightia Warren, 1907:187. DIAGNOSIS Bougainvilliidae with branched or unbranched hydrocau lus; perisarc extending as a distinct pseudohydrotheca over base of hydranth nearly to tentacles. Hydranth vasiform; manubrium nipple-shaped; tentacles in several close, al ternating whorls. Gonophores fixed sporosacs, enveloped in perisarc, borne on hydrocaulus and branches. TYPE SPECIES are distinct. In this report, Clavopsella is regarded as iden Parawrightia robusta Warren, 1907, by monotypy. tical with Pachycordyle. Parawrightia shows some remarkable similarities to REMARKS Cordylophora Allman, 1844, a genus usually included in the family Clavidae McCrady, 1859a. In bothgenera, col onies are typically branched, gonophores are fixed spo rosacs occurring on hydrocaulus andbranches, andhydranths Parawrightia Warren, 1907, is recognized as a valid name here, although it has not been widely adopted even for its type species. It bears some resemblance to Wrightia Allman, 1872, but the name of the latter is an invalid junior homonym of Wrightia L. Agassiz, 1862, a name given to a genus of thecate hydrozoans. Berrill (1948) instituted the new genus Aselomaris to accommodate Atractylis arenosa Alder, 1862, type species of Wrightia Allman, 1872, by monotypy, and for a new species, A. michaeli. Neither of the two species originally included was des ignated by Berrill (1948) as type of his new nominal genus. Atractylis arenosa is herein designated as type species of Aselomaris. Wrightia Allman, 1872 (not Wrightia L. Agassiz, 1862) thus becomes an objective synonym of Aselomaris, having the same type species. Aselomaris dif fers from Parawrightia in having fusiform to clavate hy dranths with a dome-shaped hypostome, like hydranths of Bougainvillia Lesson, 1830, instead of vasiform hydranths with a nipple-shaped hypostome. Hydranths of many bou gainvilliid hydroids are admittedly variable in shape, de pending upon degree of expansion, but those of Parawrightia are quite consistent in form and clearly distinct from those are similar in shape. Nevertheless, tentacles are somewhat less scattered and a pseudohydrotheca is present in Parawrightia, and this genus is considered distinct from Cordylophora here. Finally, Parawrightia differs from Silhouetta Millard and Bouillon, 1973. Unlike in Silhouetta, gonophores of Parawrightia are solitary, fixedsporosacsinsteadof grape like clusters of medusa buds that become free medusae. In addition, hydranths are vasiform in Parawrightia in stead of spindle-shaped. Parawrightia was founded by Warren (1907) for a single new nominal species, Parawrightia robusta, and is still monotypic. It seems debatable whether the genus should be referred to the Bougainvilliidae Lutken, 1850, although it is discussed under that family here. Parawrightia robusta Warren, 1907 Figs. 15, 16 of Aselomaris. Stechow (1923a), Millard (1975), and Bouillon (1985) included Parawrightia as a synonym of Rhizorhagium M. Sars, 1874. It is improbable that/?, roseum M. Sars, 1874, and P. robusta Warren, 1907, type species of Rhizorha gium and Parawrightia respectively, are congeneric. Rhizorhagium differs in having simple unbranched stems, tentacles arranged in a single whorl on the hydranth, and gonophores on the hydrorhiza or rhizocaulome rather than on the hydrocaulus (Rees, 1938). Therefore, both genera are recognized as valid here. The name Rhizorhagium has on occasion (e.g., Rees, 1938; Millard, 1975) been at tributed to M. Sars (1877), but it was actually made avail able earlier by M. Sars in G. O. Sars (1874). Millard (1975) and Bouillon (1985) mistakenly included Wrightia Allman, 1872, as a synonym of Rhizorhagium; instead, it is an objective synonym of Aselomaris, as noted above. Stechow (1923a) regarded Parawrightia and Pachycor dyle Weismann, 1883, as synonyms. The genus Pachy cordyle is poorly understood at present, but it seems highly unlikely that it is congeneric with Parawrightia. Pachy cordyle weismanni Hargitt, 1904a, type species of the nominal genus Clavopsella Stechow, 1919, liberates free but degenerate medusae (Hargitt, 1904a; Brinckmann-Voss, pers. comm., 1986); its hydranths are claviform to fusi form, and its hypostome is subconical. Based on such differences, I conclude that Parawrightia and Clavopsella Parawrightia robusta Warren, 1907:187; figs. 1, 2B, 3, 4; pis. 33, 34. Rhizorhagium robustum—Millard, 1966:452. Garveia robusta—Wedler and Larson, 1986:90; figs. 8Ba-d; pl. 1, fig. 7. TYPE LOCALITY Natal, South Africa. MATERIAL EXAMINED Flatts Inlet, on rocks and stems of Eudendrium carneum, - 1 m, 9 July 1983, several colonies, up to 1.5 cm high, with male gonophores, romiz B357. Flatts Inlet, on Thyroscyphus marginatus, —2 m, 4 July 1983, one colony, up to 1 cm high, with male gonophores, romiz B358. DESCRIPTION Colonies stolonal or erect, reaching 1.5 cm high, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic, slender, unbranched or irregularly branched; hydrocaulus and branches usually somewhat crooked; colonies tangled and straggly. Perisarc fairly thick, smooth or wrinkled, extending as a thin pseudohydrotheca over hydranth base nearly to tentacles. Hydranths vasiform, about 1.1 mm long from base to tip of hypostome, 0.5 mm wide at widest point; hypostome conical, elongate. Tentacles filiform, ta pering gradually from base to tip, up to about 26 in num19 ber, arranged in two or three close whorls, those of one whorl more or less alternating with those of adjacent whorls, proximal tentacles smaller than distal ones. Gonophores fixed sporosacs, completely enveloped in thin perisarc, arising singly on short, smooth pedicels from hydrocaulus, or from hydrocaulus and branches. Nematocysts— Hydroids: desmonemes 4.8-5.3 am X 2.7-2.9 am; heterotrichous microbasic euryteles 7.3-7.5 am x 3.73.8 am. REMARKS Warren (1907) described this hydroid as a new genus and species, naming it Parawrightia robusta. Millard (1966, 1975), without comment, placed it in the genus Rhi zorhagium M. Sars, 1877. According to Rees (1938), Rhizorhagium has simple, unbranched stems and tentacles arranged in a single whorl. Warren's hydroid, with typi cally branched hydrocauli and tentacles in several whorls, is recombined here with Parawrightia. In Bermuda, the hydroid was found on ceilings of cavi ties in the rocky shoreline near the bridge at Flatts Inlet. KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Florez Gonzalez, dian Ocean (Millard, 1975). 983); In- Fig. 15. Parawrightia robusta, hydranth and gonophore, romiz B357. Scale equals 1 mm. Fig. 16. Parawrightia robusta, nematocysts of hydranth, romiz B357. Scales equal 10 am. a, Desmo nemes. b, Heterotrichous microbasic eurytele. 20 Subfamily Bimeriinae Allman, 1872 DIAGNOSIS Bougainvilliid hydroids with perisarc enveloping hydranth and extending as a sheath over proximal ends of tentacles. Hydranths ovoid to vasiform; hypostome dome-shaped; tentacles in a single whorl or in two close whorls. Gonophores fixed sporosacs or free medusae; medusae with characteristics of the family. Marginal tentacles soli tary or in eight groups. Oral tentacles dichotomously 1938, are similar to those of Bimeria vestita, type species of Bimeria, although B. vestita produces fixed gonophores instead of free medusae. Petersen and Vannucci (1960) referred P. cidaritis and T. russelli, as well as material identified as Thamnostoma sp. by Browne (1905), to Koellikerina fasciculata (Peron and Lesueur, 1807). Petersen and Vannucci noted that hydroids of Thamnostoma Haeckel, 1879, are unknown. Rees (1938), Vervoort (1964), and others have pointed to the relationship between hydroids of Garveia Wright, branched. 1859, and Bimeria. The two have been regarded as con REMARKS Allman (1872) originally included in the family Bimeriidae those genera of bougainvilliids whose hydroids produced fixed sporosacs. This unnatural grouping of taxa was recombined with the Atractylidae Hincks, 1868, by Torrey (1902). Russell (1953) tentatively referred a hydrozoan to the genus Thamnostoma Haeckel, 1879, under the subfamily generic by some authors (e.g., Torrey, 1902; Browne, 1907; Stechow, 1919, 1923a; Bouillon, 1985). Bimeria differs from Garveia in having a perisarcal sheath over the base of the tentacles, and I agree with authors such as Rees (1938), Vervoort (1964), and Millard (1975) that the two should be regarded as distinct. In fact, as reported by Mammen (1963), Garveia appears to resemble Bougain- Thamnostominae Haeckel, 1879. The hydrozoan that Rus sell studied has since been referred to KoeHikerina Kramp, villia Lesson, 1830, more than it does Bimeria. 1939, by Petersen and Vannucci (1960). This genus is included in a subfamily with Bimeria Wright, 1859, here, species, B. vestita, were published in two differentjournals by Wright (1859, 1863a). Also duplicated in these two papers were accounts of the nominal species Coryne implexa (Alder, 1856b), Coryne margarica Wright, 1859, under the name Bimeriinae Allman, 1872. Identical accounts of the genus Bimeria and its type and Garveia nutans Wright, 1859. Genus Bimeria Wright, 1859 Bimeria Wright, 1859:109. Manicella Allman, 1859a:51. DIAGNOSIS Hydroids with the characters of the subfamily. Gonophores fixed sporosacs. TYPE SPECIES Bimeria vestita Wright, 1859, by monotypy. Bimeria vestita Wright, 1859 Figs. 17, 18 Bimeria vestita Wright, 1859:109; pi. 8, fig. 4. Manicella fusca Allman, 1859a:51. Bimeria humilis Allman, 1877:8; pi. 5, figs. 3,4. Perigonimus vestitus—Motz-Kossowska, 1905:74. not Bimeria vestita—Annandale, 1907:141; fig. 3 [ = Garveia franciscana (Torrey, 1902)]. Bimeria vestita f. nana Leloup, 1932:142; fig. 14. The generic names Bimeria Wright, 1859, and Manicella Leuckartiara vestita f. nana—Vervoort, 1946a:294. Leuckartiara vestita—Vervoort, 1946a:295. Allman, 1859a, are simultaneous synonyms, having both Perigonimus vestita—Mammen, 1963:42 [incorrect sub been published in July 1859 for the same species. Allman sequent spelling]. REMARKS (1864a) seemed uncertain thai Manicella and Bimeria were Garveia humilis—Vervoort, 1968:7. congeneric. Later, he conceded (Allman, 1872) that the type species of the two genera, Bimeria vestita Wright, Bimeria {Garveia) umilis—Wedler and Larson, 1986:71 [incorrect subsequent spelling]. 1859, and Manicella fusca Allman, 1859a, were "almost certainly the same species" and recognized Bimeriaas the TYPE LOCALITY valid name. Hincks (1868), acting as first reviser [Art. 24], had earlier chosen Bimeria as the name having precedence. Firth of Forth, Scotland. Life-cycle studies by Bruckner (1914) and Rees (1938) showed that hydroids of the medusae of Perigonimus ci- Flatts Inlet, on undersides of flat rocks, - 3 m, 2 August 1982, two colonies, 4 and 5 mm high, without gono daritis Weismann, 1883, and Thamnostoma russelli Rees, phores, romiz B137. Green Bay Cave, Harrington Sound, MATERIAL EXAMINED 21 on Eudendrium carneum, 8 July 1982, one colony, 7 mm high, with male gonophores, romiz B146. Harrington Sound, at Flatts Bridge, on algae, 21 September 1977. one colony, 1.5 mm high, with female gonophores, romiz B160. DESCRIPTION Colonies minute, stolonal or erect, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic, slender basally, gradually expanding distally, either unbranched or spar ingly and more or less alternately branched. Perisarc of moderate thickness, annulated or wrinkled at base of hy drocaulus and branches, becoming encrusted with silt and detritus in older colonies, extending as a filmy covering over hydranth and around base of hypostome, forming tubular sheaths around bases of tentacles. Hydranth vasi form, merging almost imperceptibly with pedicel; hypo stome conical, tentacles 9 to 16 in number, filiform, in two close whorls. Nematocysts— Hydroids: desmonemes 3.8—4.7 |xm X 2.6-2.9 (xm; heterotrichous microbasic euryteles 6.6-7.3 u.m X 3.74.5 u,m. Gonophores fixed sporosacs lacking radial canals and tentacular rudiments, completely enveloped in perisarc, arising singly on short, annulated or wrinkled pedicels from hydrocaulus and branches, less frequently from hy drorhiza. Female gonophores pear-shaped, each bearing a single egg or embryo. Male gonophores pear-shaped to elongate-oval. REMARKS Fig. 17. Bimeria vestita, part of hydrocaulus with two hy dranths and gonophore, romiz B160. Scale equals 0.25 mm. Bimeria vestita Wright, 1859, and Manicella fusca Allman, 1859a, are simultaneous synonyms, as noted above Fig. 18. Bimeria vestita, nematocysts of hydranth. Scales equal 10 (j-m. a, Desmoneme, romiz B160. b, Heterotrichous microbasic eurytele, romiz B137. 22 under discussion of the genus Bimeria Wright, 1859. Hincks (1868), the first reviser [Art. 24], chose B. vestita as hav ing precedence over M. fusca. as B. humilis. Congdon observed dense growths of this hydroid on species of Eudendrium Ehrenberg, 1834, and Halocordyle Allman, 1872, as well as on sponges. Allman (1877) introduced the name Bimeria humilis for certain hydroids dredged in shallow water off the Tortugas, Florida. He indicated that these specimens differed from B. vestita in having "massive hydranths" and a somewhat less branched hydrocaulus. The two nominal species are nevertheless regarded as conspecific here. Bimeria vestita has been reported from Bermuda before by Congdon (1907), Bennitt (1922), and Calder (1986), KNOWN RANGE Bermuda: on hydroids and sponges. (Congdon, 1907); listed, with no comment (Bennitt, 1922); in shallow inshore waters, and on wrecks and buoy chains (Calder, 1986). Elsewhere: western Atlantic (Vervoort, 1968); eastern At lantic (Picard, 1958); Indian Ocean (Millard, 1975); west ern Pacific (Leloup, 1937); eastern Pacific (Fraser, 1948). Subfamily Bougainvilliinae Lutken, 1850 DIAGNOSIS Bougainvilliid hydroids with perisarc terminating at base of hydranth, or extending upwards as a pseudohydrotheca. Hydranth fusiform to clavate; hypostome dome-shaped; tentacles more or less in a single whorl. Gonophores fixed or motile sporosacs, or free medusae; medusae with characteristics of the family. Marginal ten tacles in four perradial groups. Oral tentacles almost al ways dichotomously branched. REMARKS The subfamily Bougainvilliinae Lutken, 1850, as defined here, encompasses hydroids and medusae of the genera Bougainvillia Lesson, 1830, and Nemopsis L. Agassiz, 1849, as well as hydroids of the genera Dicoryne Allman, 1859b, Garveia Wright, 1859, and Aselomaris Berrill, 1948. Of these, only Bougainvillia is known at present from Bermuda. Bourgainvillia Allman, 1872:433 [incorrect subsequent spelling]. Atractilis Allman, 1872:433 [incorrect subsequent spelling]. Lizusa Haeckel, 1879:80. Bouganvilleia Brooks, 1883b:468 [incorrect subsequent spelling]. Perigonemus Fewkes, 1891:29 [incorrect subsequent spelling]. Perigonismus Mobius, 1893:89 [incorrect subsequent spelling]. Lizuza Delage and Herouard, 1901:56 [incorrect subse quent spelling]. Bougainvilla Hargitt, 1902:13 [incorrect subsequent spelling]. Perigominus Hartlaub, 1905:533 [incorrect subsequent spelling]. Hyppocrene Bedot, 1912:259 [incorrect subsequent spelling]. DIAGNOSIS Genus Bougainvillia Lesson, 1830 Bougainvillia Lesson, 1830:118. Hippocrene Brandt, 1835:29 [invalid junior homonym of Hippocrene Oken, 1817 (Mollusca)]. Perigonimus M. Sars, 1846:8. Bougainvillea Forbes, 1848:61 [incorrect subsequent spelling]. Perigonymus Forbes, 1848:81 [incorrect subsequent spelling]. Margelis Steenstrup, 1850:35. Atractylis Wright, 1858a:447. Bourgainvillea Wright, 1858a:449 [incorrect subsequent spelling]. Perigommus Allman, 1871:24 [incorrect subsequent spelling]. Parigonimus Allman, 1872:325 [incorrect subsequent spelling]. Bougainvilliidae with colonies stolonal or erect, with mon osiphonic or polysiphonic hydrocaulus. Perisarc soft or firm, terminating at hydranth base or forming a pseudo hydrotheca. Hydranth with a single distal whorl of filiform tentacles; hypostome conical. Gonophores free medusae, arising singly or in clusters from hydrocaulus, branches, or hydrorhiza. Medusae with short manubrium; oral tentacles perradial, usually branched dichotomously. Radial canals four; ring canal present. Marginal tentacles all alike in structure, arising in clusters from four tentacle bulbs. Ocelli usually present. Gonads on manubrium; medusa buds occasionally produced. TYPE SPECIES Bougainvillia macloviana Lesson, 1830, by monotypy. REMARKS The name Bougainvillia has been attributed by many au23 thors to Lesson (1836), although it was actually founded in an earlier paper by the same author (Lesson, 1830). not Tubularia {Sertularia) ramosa—Dalyell, 1847:64; pi. Likewise, there has been confusion over the name of the Goodsir, 1851)]. type species of the genus because Lesson (1830) first re ferred to it as Cyanaea bougainvillii. Below this name, but on the same page, Lesson commented that his medusa was not at all a "cyanee" but the type of a new genus named Bougainvillia. He then applied the name B. macloviana to the species, making C. bougainvillii and B. macloviana simultaneous synonyms. Lesson (1836), act ing as first reviser [Art. 24], chose the name/?, macloviana for the species. Bougainvillia macloviana is the type spe cies of the genus, not B. ramosa (van Beneden, 1844b) as designated by Allman (1872). Atractylis ramosa—Wright, 1858a:449 [not Eudendrium Rees (1938) demonstrated that Perigonimus M. Sars, 1846, is a junior subjective synonym of Bougainvillia be cause the hydroid of its type species, P. muscoides M. Sars, 1846, gives rise to medusae referable to Bougain villia. In a revision of the genus Perigonimus, Rees (1956a) noted that nearly 40 other nominal species had been re ferred to Perigonimus at one time or another. These were shown to belong not only to a number of genera, but to at least six families. Totton (1930) designated Eudendrium ramosum sensu van Beneden, 1844b (not Sertularia ramosa Linnaeus, 1758), referred to Bougainvillia, as type species of Atrac tylis Wright, 1858a. As noted by Totton (1930) and Rees (1938), Atractylis is, therefore, a subjective synonym of Bougainvillia. Margelis principis Steenstrup, 1850, which is type spe cies of Margelis Steenstrup, 1850, and the two originally included nominal species of Lizusa Haeckel, 1879, are now all included in Bougainvillia. The names Margelis and Lizusa are, therefore, synonyms of Bougainvillia. Vannucci and Rees (1961) noted that hydroids of the various species of Bougainvillia are difficult to separate; they differ little from each other, yet are greatly influenced by environmental factors. Vannucci and Rees concluded from their review of the genus that many of the nominal species are of questionable validity. The same opinion was expressed many years earlier by Mayer (1910). Of the numerous incorrect subsequent spellings [Art. 33c] of the name Bougainvillia and its synonyms, the most commonly encountered is Bougainvillea, firstused by Forbes (1848). The list given here should not be considered com plete; I was unable to trace and verify several others listed by Neave (1939, 1940a, 1940b). Bougainvillia muscus (Allman, 1863) Figs. 19, 20 11, figs. 1-8 [= Wougainvillia pyramidata (Forbes and ramosum (Linnaeus, 1758)]. Bourgainvillea britannica—Wright, 1858a:449 [medusa] [incorrect subsequent spelling] [not Bougainvillia britan nica (Forbes, 1841)]. Podocoryne alderi Hodge, 1861:82; pi. 2, figs. 11-15 [medusa, not hydroid]. Margelis ramosa—L. Agassiz, 1862:344 [hydroid and medusa] [not Eudendrium ramosum (Linnaeus, 1758)]. Perigonymus muscus Allman, 1863:12 [incorrect subse quent spelling]. Perigonymus ramosus—Allman, 1863:12 [incorrect sub sequent spelling] [not Eudendrium ramosum (Linnaeus, 1758)]. Atractylis {Eudendrium) ramosa—Wright, 1863b:35 [not Eudendrium ramosum (Linnaeus, 1758)]. Corynopsis alderi—Allman, 1864a:354 [medusa, not hydroid]. Bougainvillia ramosa—Allman, 1864a:366 [not Euden drium ramosum (Linnaeus, 1758)]. Bougainvillia muscus—Allman, 1864a:366. Bougainvillia fruticosa Allman, 1864a:366 [nomen nudum]. Bougainvillia fruticosa Allman, 1864b:58. Lizusa octocilia—Haeckel, 1879:80 [part] [not Medusa octocilia Dalyell, 1847 = Wougainvillia pyramidata (Forbes and Goodsir, 1851)]. Bougainvillea ramosa—Pictet, 1893:11 [incorrect subse quent spelling] [not Eudendrium ramosum (Linnaeus, 1758)]. Bougainvillea muscus—Pictet, 1893:11 [incorrect subse quent spelling]. Bougainvillea {Margelis) ramosa—Garstang, 1894:214 [medusa] [not Eudendrium ramosum (Linnaeus, 1758)]. Bougainvillia flavida Hartlaub, 1897:456; pi. 14, fig. 5 [female medusa only]. Bougainvillia autumnalis Hartlaub, 1897:465; pi. 15, figs. 11-13 [medusa]. Lizusa octociliata—Aurivillius, 1898:114 [medusa] [in correct subsequent spelling]. Lizusa 8-ciliata—Aurivillius, 1898:424 [medusa] [incor rect subsequent spelling]. Wougainvillia v. benedenii Bonnevie, 1898:468. Wougainvillia benedenii Bonnevie, 1898:484; pi. 26, figs. 34,35 [hydroid and medusa]. Wougainvillia vanbenedeni—Bonnevie, 1899a:43 [incor rect subsequent spelling]. Bougainvillia gibbsi Mayer, 1900a:5; pi. 4, figs. 14,15 [medusa]. Eudendrium ramosum—van Beneden, 1844b:56; pi. 4, figs. 1-13 [hydroid and medusa] [not Eudendrium ramo sum (Linnaeus, 1758)]. 24 Margelis autumnalis—Browne, 1900:708 [medusa]. Wougainvillia van benedeni—Jaderholm, 1909:46; pi. 3, fig. 5 [incorrect subsequent spelling]. Wougainvillia van benedenii—Broch, 1909:198. Bougainvillia ramosa var. nana Hartlaub, 1911:189 [medusa]. Bougainvillia triestina Hartlaub, 1911:154; fig. 138 [medusa]. Bougainvillia ramosa var. minima Kramp and Damas, 1925:254 [medusa]. Bougainvillia autumnalis var. magna Babnik, 1948:290; fig. 2 [medusa]. Bougainvillia ramosa f. musca—Millard, 1975:99. Bougainvillia ramosa f. fruticosa—Millard, 1975:99. Bougainvillia ramosa f. ramosa—Millard, 1975:99. Bougainvillia ramosa f. vanbenedenii—Millard, 1975:99. MATERIAL EXAMINED Green Bay Cave, Harrington Sound, on survey line 40 m from entrance, —6 m, 3 March 1982, three colonies, 614 mm high, with medusa buds, coll. T. Iliffe, romiz B152. Flatts Inlet, on Cliona sp., -0.5 m, 27 February 1982, one colony, 4 mm high, without medusa buds, romiz B153. St George's Island, north shore, on oyster on moor ing chain, - 2 m, 5 October 1976, one colony, 10 mm high, without medusa buds, coll. J. Markham, L. Coen, romiz B163. Flatts Inlet, on sponge, - 1.5 m, 5 March 1982, one colony, 7 mm high, without medusa buds, romiz B169. Flatts Inlet, on rocks and Eudendrium carneum, - 2 m, 4 July 1983, three colonies, up to 13 mm high, with medusa buds, newly liberated medusae, and laboratoryreared adult medusae, romiz B328. TYPE LOCALITY Torquay (Torbay), Devon, Great Britain. m m 0$ Mtfr Fig. 19. Bougainvillia muscus. Scales for a and b equal 0.5 mm; scale for c equals 1 mm. a, Hydroid colony, romiz B152. b, Newlyliberated medusa, romiz B328. c, Three-day-old adult female medusa, romiz B328. 25 DESCRIPTION Hydroid colony initially stolonal, later erect, reaching 14 mm high, growing from a creeping hydrorhiza. Hydrocaulus slender, monosiphonic, profusely and more or less alter nately branched; primary branches unbranched or some what regularly branched; terminal branchlets slender basally, gradually increasing in diameter distally; hydrocaulus and branches curved and twisted, occasionally exhibiting sto lonal growth. Perisarc of moderate thickness, smooth or irregularly wrinkled (especially at bases of branches) but not annulated, heavily encrusted with particles of silt and detritus, and extending over base of hydranth as a pseudohydrotheca; pseudohydrotheca cup-shaped in retracted hydranths; perisarc not investing tentacles or hypostome. Hydranth cylindrical when extended, fusiform when con tracted, with conical hypostome, bearing tentacles distally. Tentacles slender, filiform, in two close whorls, 10 to 16 in number. Nematocysts— Hydroids: desmonemes 3.9^1.6 (xm x 2.7-3.0 |xm; heterotrichous microbasic euryteles 5.7-6.5 p.m x 2.83.4 (xm. Medusa buds globular, arising singly on stalks of mod erate length from pedicels below hydranths, completely invested with perisarc. Newly liberated medusae thimbleshaped, 0.8 mm high, 0.7 mm wide; mesoglea of moderate thickness; umbilical canal present; peduncle lacking. Exumbrella with four distinct interradial longitudinal furrows and four somewhat less distinct perradial longitudinal fur rows; exumbrellar nematocysts lacking. Manubrium small, tubular, with simple quadrate mouth; oral tentacles four, unbranched, inserting just above mouth, each oral tentacle terminating with small cluster of nematocysts. Radial ca nals four, joining a narrow ring canal. Marginal bulbs conical, rounded basally, each with two tentacles and two conspicuous black ocelli, each ocellus occurring at base Fig. 20. Bougainvillia muscus, nematocysts, romiz B328. Scales for a-d equal 10 p.m; scale for e equals 100 p.m. a, Desmoneme of hydroid. b, Heterotrichous microbasic eurytele of hydroid. c, Desmoneme of medusa, d, Heterotrichous microbasic eurytele of medusa, e, Eggs, each with outer envelope of heterotrichous microbasic euryteles. 26 of a tentacle. Velum broad. Endoderm of manubrium and tentacle bulbs cream-coloured. Nematocysts— Medusae: desmonemes 4.5-5.1 |xm x 3.0-3.5 (xm; heterotrichous microbasic euryteles 5.3-6.5 |xm x 2.93.4 (xm. One day after liberation, medusae 1.2 mm high, 1.1 mm wide; mesoglea thicker, especially at apex. One medusa with an incipient third tentacle on one of four marginal bulbs, incipient tentacle lacking an ocellus basally. Two days after liberation, medusae about 1.5 mm high and wide, oral tentacles distally bifurcated, and gonads present interradially on manubrium. Each tentacle bulb with a developing third tentacle, varying in length from about half that of original pair to a mere stub. Developing ten tacles without ocelli initially, later with ocelli. Three days after liberation, medusae about 2.0 mm high and wide, oral tentacles divided once, marginal tentacles 12 in num ber, 3 on each tentacle bulb, and ocelli 10 to 12 in number. Gonads large, mature eggs present, ova with an outer envelope bearing numerous heterotrichous microbasic euryteles (6.5 x 3.6 |xm). REMARKS The invalid name Bougainvillia ramosa (van Beneden, 1844b) has been extensively applied to this species in the literature of both hydroids and hydromedusae (e.g., see Hincks, 1868; Allman, 1872; Stechow, 1919; Vervoort, 1946b; Russell, 1953; Kramp, 1961; Vannucci and Rees, 1961; Millard, 1975; and numerous others). Van Beneden (1844b) applied the name Eudendrium ramosum to a bougainvilliid hydrozoan in the mistaken belief that it was conspecific with Tubularia ramosa Linnaeus, 1758, a spe cies now referred to the genus Eudendrium Ehrenberg, 1834. Even though the hydrozoan studied by van Beneden has since been referred to a different genus from the true Eudendrium ramosum (Linnaeus, 1758), the name B. ra mosa cannot be retained for the species [Art. 49]. The name B. ramosa is replaced here with its oldest available synonym, Bougainvillia muscus (Allman, 1863). Russell (1953) noted that the synonymy of this species is in doubt because several different hydroid growth forms apparently produce similar medusae. It is still unresolved whether these different hydroid growth forms belong to a single, variable species, or to more than one species. The list of synonyms given here, taken largely and selectively from Bedot (1905, 1910, 1912, 1916, 1918), Russell (1953), Kramp (1961), and Vannucci and Rees (1961), is provi sional and not intended to be exhaustive. A thorough taxonomic reassessment of the species is needed. Records of the hydroid Tubularia {Sertularia) ramosa and the hydromedusa Medusa ocilia {M. octocilia, M. sexdecilia, M. duodecilia) by Dalyell (1847) have been excluded from the synonymy of this species based on the conclusions of Edwards (1966a), who suggested that they were likely referable to Bougainvillia pyramidata (Forbes and Goodsir, 1851). Edwards noted that Dalyell's illus tration of the hydroid, found on a sea pen, closely resem bled specimens of B. pyramidata from the same substrate examined by Edwards (1964a). Wright (1858a) thought that the medusa of this hydroid was identical with B. britannica (Forbes, 1841), and Mayer (1910) among others regarded the two as conspecific. However, these are now considered to be different species (e.g., see Russell, 1953; Kramp, 1961; Vannucci and Rees, 1961; Edwards, 1964b, 1966a). The nominal species Podocoryne alderi Hodge, 1861, was almost certainly based on a hydroid belonging to Podocoryna M. Sars, 1846, and a medusa belonging to Bou gainvillia muscus (Vannucci and Rees, 1961; Edwards, 1966a, 1972). Allman (1864a) proposed the generic name Corynopsis for the conglomerate P. alderi. Hodge (1861) reported sending "two or three depauperated specimens," along with some sketches, to J. Alder for identification. This material, labelled "Podocoryne alderi Seaham H" (Seaham Harbour, northeast England), and relabelled "Corynopsis alderi Hodge," is represented in the Alder Collection at the Hancock Museum, Newcastle upon Tyne. The hydroid colony consists of a branching and anasto mosing stolon network, with occasional short spines and a few hydractiniid hydranths, growing over serpulid polychaete tubes. Since the more widely used name B. mus cus is predated by Hodge's (1861) nominal species, the name P. alderi is restricted here to the hydroid only; the above-cited material from the Hancock Museum is des ignated as the lectotype [Art. 74]. The name P. alderi thus is restricted to Podocoryna, and no longer threatens the name B. muscus. Moreover, the generic name Corynopsis Allman, 1864a, becomes a junior subjective synonym of Podocoryna, instead of Podocoryna in part and Bougain villia in part. Edwards (1972) included P. alderi as a questionable synonym of P. borealis (Mayer, 1900a), which in turn was viewed as a questionable synonym of P. tubulariae M. Sars, 1857. Bougainvillia flavida Hartlaub, 1897, has been regarded as a synonym, or a synonym in part, of this species (e.g., see Hartlaub, 1911; Kramp, 1937; Vannucci and Rees, 1961). Edwards (1964a, 1964b) concluded that Hartlaub's (1897) hydroid ofB. flavida was referable toB. britannica, his male medusae to B. pyramidata, and his female medusa to "B. ramosa." Bougainvillia autumnalis Hartlaub, 1897, is also a synonym, at least in part, of this species (e.g., see Russell, 1953; Kramp, 1961; Vannucci and Rees, 1961). Bougainvillia vanbenedenii Bonnevie, 1898, has been in cluded as a questionable synonym here following Van nucci and Rees (1961), although these authors left open the possibility that Bonnevie's species might be identical with B. superciliaris (L. Agassiz, 1849). However, the 27 hydroid of B. superciliaris is now known to be stolonal (Werner, 1961; Edwards, 1966a). Hydroids examined here from Bermuda bear consider able resemblance to the description of Bougainvillia longicirra Stechow, 1914, from the Caribbean. Like B. muscus, Stechow's (1914) colonies were small but often exten sively branched, the hydrocaulus was monosiphonic, the perisarc was wrinkled and encrusted with particles of silt and detritus, a pseudohydrotheca was present, and the tentacles numbered about 16. Medusa buds were present on Stechow's hydroids, but the newly liberated medusa was not observed. Fraser (1944) was almost certain that B. longicirra was conspecific with B. superciliaris, but Vannucci and Rees (1961) correctly disputed this, noting that the latter is a boreal species occurring well to the north of the locality where B. longicirra was found. As noted above, the hydroid of B. superciliaris is now known to be stolonal. Stechow (1914) indicated that B. longicirra might represent the hydroid of the medusa B. niobeMayer, 1894. Vannucci and Rees (1961) stated that this was based on surmise, and noted that the type locality of Charlotte Amalie in the West Indies is within range of B.frondosa Mayer, 1900b, as well as that of B. niobe. I have not regarded B. longicirra as conspecific with B. muscus because of a lack of knowledge about its medusa stage. Medusae described here were isolated and reared in the laboratory following liberation from the hydroid. Speci mens were maintained at 28-29° C in covered fingerbowls containing natural seawater, and fed pieces of newly hatched nauplii of Artemia several times daily. Observations were made on both living and preserved material. Sexual ma turity was attained about three days after liberation, when eggs were observed being shed from the gonads of female medusae; none of the medusae reared to maturity were male. The eggs were surrounded by an envelope containing nematocysts, as reported previously for this species (Rus sell, 1953). Of several dozen medusae isolated initially, none survived longer than 3.5 days in the laboratory. A short life span in medusae of this species might partly explain Russell's (1953) observation that they are scarce in the plankton around the British Isles, considering the abundance of the hydroid there. Neither hydroid nor medusa of Bougainvillia muscus has been reported previously from Bermuda, but two other species of medusae belonging to this genus have been reported from the area. Bigelow (1918, 1938) identified B. niobe in collections of hydromedusae from Bermuda, and Moore (1949) noted that the species was an abundant winter form there. Bougainvillia platygaster (Haeckel, 1879) has been recorded from a number of areas in the western North Atlantic, including the Sargasso Sea near Bermuda (Kramp, 1959). KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Mayer, 1910, as B. autum nalis); eastern Atlantic (Russell, 1953); Indian Ocean (Mil lard, 1975); western Pacific (Yamada, 1959). Family Cytaeididae L. Agassiz, 1862 CytaeidaeL. Agassiz, 1862:341 [emended to Cytaeididae free medusae were included in Cytaeis Eschscholtz, 1829. by Kramp, 1961]. Species with fixed gonophores were placed in Perarella Stechow, 1922, except for the poorly known Stylactis ver- DIAGNOSIS Hydroid colonies stolonal; hydranths arising from a creep ing hydrorhiza. Perisarc covering hydrorhiza, terminating at base of hydranths, often in form of a collar. Hydranths columnar; tentacles filiform, in one more or less regular oral whorl; hypostome conical. Gonophores free medusae or fixed sporosacs, arising from hydrocaulus on stalks. Medusa bell-shaped; manu brium bulbous, with simple, circular mouth and four or more unbranched oral tentacles inserting above mouth. Radial canals four. Marginal tentacles solid, four or ex ceptionally eight, each arising from a tentacle bulb. Ocelli absent. Gonads on manubrium. REMARKS Rees (1962) recognized three genera of hydroids in the family Cytaeididae L. Agassiz, 1862. All species having 28 micola Allman, 1888, which was retained in Stylactella Haeckel, 1889. The diagnosis of the family Cytaeididae given here en compasses the recently described Paracytaeis Bouillon, 1978a, but excludes Cnidostoma Vanhoffen, 1911, fol lowing Picard (in Kramp, 1961:444) and Rees (1962). Haeckel (1889) believed that Stylactella, Stylactis Allman, 1864a, and Hydranthea Hincks, 1868, were related genera, and stated, "Perhaps Stylactella and the allied genera may represent together a distinct family, the Stylactidae." Although the poorly known genus Stylactella is apparently a cytaeid, the nominal family Stylactidae is not a synonym of the Cytaeididae, as implied by Rees (1962). The name Stylactidae is derived from the nominal genus Stylactis rather than Stylactella, and the former is therefore type genus of the family [Art. 63]. The nominal genus Stylactis, and Haeckel's Stylactidae, are included here in the Hydractiniidae L. Agassiz, 1862. Hydranthea is now included in the family Haleciidae Hincks, 1868 (e.g., see Cornelius, 1975; Bouillon, 1985). Genus Cytaeis Eschscholtz, 1829 Cytaeis Eschscholtz, 1829:104. Cytaeis de Blainville, 1834:284 [incorrect subsequent spelling]. Cyteis van Beneden, 1867:18 [incorrect subsequent spelling]. Cytheis van Beneden, 1867:18 [incorrect subsequent spelling]. Nigritina Haeckel, 1879:73. Cytaeidium Haeckel, 1879:75. Cytaesis Bouillon, 1978a: 129 [incorrect subsequent spelling]. DIAGNOSIS Hydroids with characters of the family. Gonophores free medusae, arising on stalks from hy drorhiza. Medusa with characters of the family, with only four marginal tentacles. TYPE SPECIES Cytaeis tetrastyla Eschscholtz, 1829, by monotypy. REMARKS Haeckel (1879) established two subgenera within the genus Cytaeis Eschscholtz, 1829, Nigritina for those species lacking a gastric peduncle andCytaeidium forthose having a peduncle. He attributed Nigritina to J. Steenstrup, but apparently the name was derived from an unpublished manuscript by that author. Neither nominal subgenus is recognized in the recentliterature on Cytaeis (e.g., Kramp, 1959, 1961, 1965; Rees, 1962; Uchida, 1964; Vervoort, 1967; Millard, 1975; Hirohito, 1977; Bouillon, 1978a, 1980, 1985). Kramp (1961) regarded Cytaeis pusilla Ge- genbaur, 1857, type species of Cytaeidium, as doubtfully conspecific with Cytaeis tetrastyla Eschscholtz, 1829. Mayer (1910) included the nominal genus Cubogaster Haeckel, 1879 (original spelling Cybogaster Haeckel, 1864), Komai (1931) was the first to link hydroid and medusa stages of Cytaeis, working with what he thought was C. japonica Uchida, 1927. Rees (1962) regarded Komai's hydrozoan as a distinct species and named it C. uchidae. Worldwide, three nominal species of Cytaeis medusae were listed by Kramp (1961). Six species, including the hydroids of five of these, were included in the genus by Rees (1962). Rees suggested that more than one species may have been combined under C. tetrastyla by Kramp (1961). Cytaeis sp. Figs. 21, 22 MATERIAL EXAMINED Green Bay, Harrington Sound, on shell of Cerithium lit teratum from Cladophora bed, -2.5 m, 21 September 1984, one colony, with medusa buds, two medusae lib erated in laboratory from hydroid, romiz B353. DESCRIPTION Colony stolonal, with hydrorhizal network adhering to gas tropod shell. Perisarc smooth, thin, terminating at baseof hydranth, not dilated in form of cup-shaped collar. Spines absent. Hydranths all gastrozooids, clavate to nearly col umnar, up to 0.6 mm long, 0.2 mm wide, with four to five tentacles in each of two closely placed oral whorls, those of one whorl alternating with those of adjacent whorl. Tentacles filiform, but with nematocyst batteries arranged in more or less distinct rings medially and distally. Hy postome dome-shaped. Nematocysts— Hydroids: desmonemes 5.0-5.8 |xm x 3.3-3.8 (xm; heterotrichous microbasic euryteles 7.4-8.3 |xm x 3.53.8 |xm. Medusa buds pear-shaped to globular, arising singlyon relatively longstalksfromhydrorhiza, completely invested in perisarc. Newly liberated medusae bell-shaped, about 0.4 mm high and wide; mesoglea thin. Exumbrella with four distinct perradial longitudinal furrows and four less distinct interradial furrows in preserved and contracted in the synonymy of Cytaeis. However, Cybogaster gemmascens Haeckel, 1864, type species of Cybogaster by monotypy, was regarded by Russell (1953) as conspecific material; scattered exumbrellar nematocysts present. Manubrium tubular to somewhat fusiform, extending about with Lizzia blondina Forbes, 1848, type species of the four, unbranched, inserted just above mouth, each with a small terminal cluster of nematocysts. Radial canals four; bougainvilliid genus Lizzia Forbes, 1846. Thus, neither Cybogaster nor its junior objective synonym Cubogaster [Art. 33a (ii)] can be regarded as synonymous withCytaeis. Stylactella Haeckel, 1889, and Perarella Stechow, 1922, aredistinguished from Cytaeis largelyby the type of gonophore produced (Rees, 1962), a practice criticized by Pe tersen (1979). However, the relationships of these nominal genera are unclearand they have not been combined here. halfway to velar opening; mouth simple; oral tentacles narrow ring canal present. Tentacle bulbs four, perradial, subspherical, each with a single contracted filiform ten tacle. Ocelli absent. Gonads undeveloped. REMARKS This hydrozoan cannot be assigned at present, with any degree of confidence, to any of the nominal species cur29 Fig. 21. Cytaeis sp., romiz B353. Scales equal 0.25 mm. a, Hydranth and gonophore. b, Newly liberated, living medusa, c, Newly liberated, preserved medusa. Fig. 22. Cytaeis sp., nematocysts of hydranth, romiz B353. Scales equal 10 u.m. a, Desmoneme. b, Heterotrichous microbasic eurytele. 30 rently referred to Cytaeis Eschscholtz, 1829. While bear ing considerable resemblance to C. nassa (Millard, 1959a), Bermuda specimens have not been referred to that species for several reasons. First, hydranths were smaller than those of C. nassa described by Vervoort (1967) and Mil lard and Bouillon (1973), and much smaller than those measured by Millard (1959a) and Rees (1962). Admit tedly, hydranthmeasurements in the literatureon C. nassa are quite varied, and differences noted here may be taxonomically insignificant. Secondly, Millard (1959a) re ported that young hydranths of C. nassa had 8 tentacles in a single verticil, while older ones had 16 tentacles in two alternating verticils. The smaller hydranths of Cytaeis sp. studied here had 4 to 5 tentacles in each of two closely placed whorls. This seeming difference may also be taxonomically unimportant, because Vervoort's (1967) ma similar to descriptions of C. imperialis Uchida, 1964, from Japan, but has not been identified as that species largely on zoogeographic grounds. ROM specimens may well be long to Cytaeis tetrastyla Eschscholtz, 1829, an epipelagic medusa known to occur near Bermuda (Bigelow, 1918). However, such an identification cannot be established at present because of lack of information on the life cycle of C. tetrastyla. Rearing of the medusa of Bermudian ma terial will likely be necessary to ascertain whether it be longs to any of the previously named species of Cytaeis, or to an undescribed species. The status of the various nominal species of Cytaeis is in need of clarification. Confusion exists in particular over C. japonica Uchida, 1927. It seems uncertain whether C. japonica is identical with C. uchidae (alternative spelling C. uchidai, as emended by Kramp, 1965). Rees (1962) terial of C. nassa had 4 tentacles per whorl on the majority believed that the two were distinct, while Uchida (1964) of hydranths. Finally, the manubriumof the young medusa in present collections was proportionately smaller than that regarded them as conspecific. If they are conspecific, the specific name japonica has priority [Art. 23] and cannot described or illustrated in C. nassa by Millard (1959a, 1975), Rees (1962), and Millard and Bouillon (1973). As in C. niotha (Pennycuik, 1959), an inadequately known by Uchida (1964). The name C. japonica originally en compassed at least two and possibly three different species nominal species from Australia, hydranths of C. nassa appear to have been substantially larger than those of Bermudian specimens. Gonophores of C. niotha, although arising from the hydrorhiza, were unlike my material in being clustered around the bases of the hydranths. Stylactis indica Stechow, 1920, referred to Cytaeis by Rees (1962), is likewise poorly known; questions remain as to the nature of its gonophores and to its generic identity. Bermuda specimensdiffer from C. uchidae Rees, 1962, and C. nuda Rees, 1962, in having undeveloped instead of developed gonads in the newly liberated medusa. My material is be abandoned in favour of the name uchidae, as proposed of medusae. Uchida (1930) realized that what he originally thought (Uchida, 1927) was the young medusa with me dusa buds of C. japonica was identical with Podocoryna simplex Kramp, 1928, instead. In addition, he later indi cated (Uchida, 1964) that the name C. japonica encom passed C. imperialis as well. Kramp (1961,1965) regarded C. japonica as a synonym of C. tetrastyla. Further research is needed to clarify the relationships of the Japanese species of Cytaeis. Hydroids of the genus Cytaeis have not been reported before from Bermuda. Family Hydractiniidae L. Agassiz, 1862 Hydractinidae L. Agassiz, 1862:339 [emended to Hy dractiniidae by Hincks, 1868:18]. Podocorynidae Allman, 1864a:353. Stylactidae Haeckel, 1889:79. Janariidae Stechow, 1921b:29. DIAGNOSIS Hydroid colonies stolonal. Hydrorhiza consisting of tubes covered with chitinous perisarc, or an encrustation of na ked coenosarc with or without a calcareous skeleton, fre quentlywith spines, less frequently with calcareous branches. Hydranths sessile, naked, polymorphic, as gastrozooids, gonozooids, and occasionally dactylozooids. Gastrozooids of one or more types, usually with one or more close whorls of filiform tentacles encircling a conical to club- shaped hypostome; gonozooids with or without filiform tentacles; dactylozooids elongate, lacking tentacles. Gonophores fixed sporosacs or free medusae, usually borne on gonozooids. Medusa well developed to degen erate, more or less bell-shaped. Manubrium tubular to sacshaped, with or without a peduncle; rim of manubrium tubular or with four branched or unbranched lips, having terminal batteries of nematocysts; mouth present or absent. Radial canals four. Marginal tentacles solid, four, eight, or more. Ocelli present or absent. Gonads on manubrium, sometimes extending along proximal portions of radial canals. REMARKS Rees (1962) provided the first clear distinction between hydroids of the closely related families Hydractiniidae L. Agassiz, 1862, and Cytaeididae L. Agassiz, 1862. Ac31 cording to his diagnosis, followed here, cytaeid hydroids differ from hydractiniids in completely lacking spines on the hydrorhiza, and in having gonophores on the hydro rhiza instead of on gonozooids. The status of the nominal family Stylactidae Haeckel, 1889, referred to the Hydrac tiniidae here, is discussed under the family Cytaeididae elsewhere in this report (see pp. 28-29). Bouillon (1978b) briefly discussed the Hydractiniidae, including it together with the families Stylasteridae Gray, 1847, Ptilocodiidae Coward, 1909, and Rhysiidae Brinckmann, 1965, in a newly recognized superfamily Hydractinioidea L. Agassiz, 1862 [Art. 36]. Generic limits within the Hydractiniidae are problematic covered with naked coenosarc, nor forming a calcareous skeleton. Hydrorhizal spines present or absent. Hydranths typical of the family. Gonophores fixed sporosacs or free but degenerate me dusae, borne on gonozooids beneath whorl of oral tenta cles; newly liberated medusa sac-shaped. Manubrium simple, tubular, without mouth, oral lips, or oral tentacles. Tentacles 8 to 10 in number, rudimentary. Radial canals four. Ocelli absent. Gonad surrounding manubrium. TYPE SPECIES Stylactis inermis Allman, 1872, by monotypy. (Motz-Kossowska, 1905; Goette, 1916; Stechow, 1923a; REMARKS Kramp, 1932; Iwasa, 1934; Rees, 1962; Bouillon, 1971, 1985; Millard, 1975). Stechow (1923a), for example, rec ognized 14 recent (nonfossil) genera in his classification of the subfamily Hydractiniinae. In a re-examination of Stechow's classification, Kramp (1932) included no more than five of these in the group: he combined seven within Hydractinia (Halerella Stechow, 1922, Stylactis Allman, 1864a, Cionistes Wright, 1861, Podocoryna M. Sars, 1846, Hydronema Stechow, 1921a, Hydractinia van Beneden, 1841, and Hydrissa Stechow, 1921a), excluded four from discussion (Clavactinia Thornely, 1904, Rhizohydra Cope, 1884, Hydrocorella Stechow, 1921b, and Hydractomma Stechow, 1921a), and dismissed the remaining three as pandeids (Perigonella Stechow, 1921c, Podocorella Ste chow, 1921c, and apparently Clavopsis Graeffe, 1883). Kramp recognized two subgenera, Hydractinia and Sty lactis, within the genus Hydractinia. More recently, Bouil lon (1985) included the nominal genera Clavactinia, Hansiella Bouillon, 1980, Hydractinia, Hydrocorella, Janaria Stechow, 1921b, Kinetocodium Kramp, 1921, Po docoryna (as Podocoryne; but Podocoryne is an incorrect subsequent spelling of Podocoryna, first used by Lutken, 1850), Stylactis, and Tregoubovia Picard, 1958, in the Hydractiniidae. Kinetocodium possesses characters of both the Hydractiniidae and the Cytaeididae, and its systematic position is unclear at present. Stylactis is a synonym of Hydractinia, as noted by Stechow (1923a) and others, and Stylactis auct., to which Bermuda material belongs, is replaced here by Stylactaria Stechow, 1921a. Much con fusion remains at the generic level within this family, and taxonomic revision is badly needed. The nominal genus Stylactis Allman, 1864a, was consti tuted to accommodate two species, Podocoryne sarsii Steenstrup, 1850, and Podocorynafucicola M. Sars, 1857. A retiform hydrorhiza, consisting of anastomosing tubes covered with perisarc, was regarded as diagnostic of the genus (Allman, 1864a). Bonnevie (1898) discovered that the hydrorhiza in type material of P. sarsii was encrusting and covered with naked coenosarc. She did not, as sug gested by Iwasa (1934) and others, mention examining material of P. fucicola as well. However, the hydrorhiza of P. fucicola also appears to be encrusting with naked coenosarc (see Castric-Fey, 1970). I concur with Goette Genus Stylactaria Stechow, 1921a Stylactaria Stechow, 1921a:250. DIAGNOSIS Hydractiniidae with reticular hydrorhiza; hydrorhizal sto lons covered with perisarc; hydrorhiza not encrusting, nor 32 (1916), Stechow (1923a), Iwasa (1934), and others that these two species, with their encrusting hydrorhizae, are best referred to the genus Hydractinia van Beneden, 1841. The type species of Stylactis must be one of these two originally included species [Art. 69], not one subsequently referred to the genus such as Stylactis inermis Allman, 1872 (see Millard, 1975). Mayer (1910) designated P. fucicola (misspelled as Stylactis fuciola) type species of Stylactis. Podocoryna fucicola is referred to Hydractinia (e.g., see Stechow, 1923a; Iwasa, 1934; Castric-Fey, 1970), and the name Stylactis is a junior subjective synonym of Hydractinia, as noted earlier by Stechow (1923a). Stechow (1923a) suggested, incorrectly, that the name Stylactella Haeckel, 1889, be used for species of Stylactis auct. Three nominal species were originally included in Stylactella by Haeckel (1889), S. abyssicola Haeckel, 1889, S. spongicola Haeckel, 1889, and S. vermicola Allman, 1888. Gonophores in all three reportedly arise from the hydrorhiza and not from gonozooids (Allman, 1888; Hae ckel, 1889; Iwasa, 1934; Rees, 1962); they should be included in the family Cytaeididae rather than in the Hy dractiniidae (Rees, 1962). Thus, the name Stylactella can not replace Stylactisauct., species of which have gonophores on gonozooids. Rees (1962) regarded Stylactella as an insufficiently described genus of cytaeid, and included S. vermicola as its type and only known species. He trans ferred both S. spongicola and S. abyssicola to Perarella Stechow, 1922, also included in the family Cytaeididae. Stechow (1921a) proposed that the new name Stylac taria be applied to those species of Stylactis auct. with gonophores on gonozooids, if such hydroids were to be recognized as a distinct genus, and designated Stylactis inermis Allman, 1872, as type species of the genus. Sty lactaria is recognized here as the valid name of the genus. Stylactaria inermis reportedly produces fixed sporosacs (Bouillon, 1971), while some species of Stylactis sensu Mayer (1910) may liberate degenerate medusae. Given the variation of gonophore development among species of the group, a separate genus for those liberating a degenerate Stylactis hooperii Sigerfoos, 1899:802; figs. 1-5. Stylactis hooperi—Hargitt, 1901a:311 [incorrect subse quent spelling]. Stylactis hoopei—Komai, 1932:451 [incorrect subsequent spelling]. Stylactis sp. Crowell, 1947:206. Hydractinia arge—Calder, 1971:31; pi. 2, fig. B; pi. 7, fig. A. Styllactis hooperi var. minor Wedler and Larson, 1986:94; fig. 10c [incorrect subsequent spelling]. medusa seems unjustified. Stechow (1923a) included the nominal genus Clavopsis Graeffe, 1883, characterized by the presence of free but TYPE LOCALITY degenerate medusae, in the Hydractiniidae L. Agassiz, 1862. He believed that Stylactis sensu Mayer (1910) cor responded with that nominal genus. However, Graeffe's account of the type species of Clavopsis, C. adriatica Graeffe, 1883, was of a hydroid that was more likely a pandeoid or possibly a bougainvillioid than a hydractiniid. The hydranth pedicel of C. adriatica was enveloped in thin perisarc, and there was no clear evidence given of polymorphism in the species. Clavopsis is certainly not a synonym of Stylactis sensu Mayer (1910), and hence not of Stylactaria either. Stylactaria differs from Hydractinia, Podocoryna M. Sars, 1846, Clavactinia Thornely, 1904, Hydractomma Stechow, 1921a, and Hydrissa Stechow, 1921a, in having a reticular, nonencrusting hydrorhiza devoid of naked coenosarc. The hydrorhiza does not form a calcareous skeleton, as in Hydrocorella Stechow, 1921b, and Janaria Stechow, 1921b. Unlike Podocoryna, which liberates a well-developed medusa, Stylactaria has fixed gonophores or a degenerate, short-lived medusa. Stylactaria differs from Kinetocodium Kramp, 1921, in having well-developed instead of reduced oval tentacles, and gonophores arising from gonozooids rather than from the hydrorhiza. Species assigned to Stylactaria here, in addition to S. inermis, are S. arge (Clarke, 1882), S. arctica (Jaderholm, 1902), S. ingolfi (Kramp, 1932), S. pisicola (Komai, 1932), S. yerii (Iwasa, 1934), S. carcinicola (Hiro, 1939), and S. claviformis (Bouillon, 1971). I have not included Hy dractinia pruvoti Motz-Kossowska, 1905, in the genus because of its hydrorhiza, which is encrusting rather than reticular. Moreover, its medusa is campanulate instead of sac-shaped, and four tentacles are present instead of eight. It was included by Stechow (1921a) as the only species in his new genus Hydractomma. Crisfield, Maryland, on Chesapeake Bay, United States. MATERIAL EXAMINED Green Bay, Harrington Sound, on shell of Cerithium lit teratum from Cladophora bed, —2 m, 11 July 1983, 1 colony, with gonozooids and male gonophores, two me dusae liberated in laboratory from hydroid, romiz B354. Green Bay, Harrington Sound, on shells of C. litteratum from Cladophora bed, -3 m, 11 July 1983, 2 colonies, one male and one female, with gonozooids and gono phores, romiz B355. Green Bay, Harrington Sound, on shells of C. litteratum from Cladophora bed, - 1.5 m, 27 June 1983, 13 colonies, with gonozooids and gono phores, romiz B356. Green Bay, Harrington Sound, on shells of C. litteratum from Cladophora bed, - 3 m, 4 October 1986, 2 colonies, with gonozooids and gono phores, romiz B367. DESCRIPTION Colony stolonal, with branching and anastomosing hy drorhizal stolons adhering to gastropod shell. Perisarc thin, smooth or with irregular wrinkles, terminating at base of hydranth, not dilated as cup-shaped collar. Spines chitinous, simple, inconspicuous in unstained material, up to 0.3 mm high, arising from hydrorhizal stolons. Polyps of two types, gastrozooids and gonozooids. Gastrozooids contractile, variable in shape but generally clavate, up to 2 mm high, 0.3 mm wide, somewhat bulbous and rugose beneath tentacular whorl, slightly to distinctly constricted at insertion with hydrorhiza. Tentacles filifbimf b\o 16 in number on fully developed gastrozooids, in two closely placed whorls, those of upper whorl held somewhat more erect than those of lower whorl in life. Hypostome domeshaped to clavate to knob-shaped. Gonozooids columnar, contractile, up to 1.7 mm high, 0.2 mm wide; region be neath tentacular whorl smooth and slender. Tentacles fili form, 5 to 10 in number, in one whorl. Medusa buds Stylactaria arge (Clarke, 1882), comb. nov. Figs. 23, 24 Stylactis arge Clarke, 1882:135; pi. 8, figs. 18-20. Sytlactis arge Clarke, 1882:138 [lapsus]. arising proximal to tentacular whorl, usually with two buds on opposite sides of gonozooid. Hypostome bulbous to dome-shaped. Sexes separate. Nematocysts— Gastrozooids: desmonemes 5.0-5.7 jxm x 2.8-3.3 jxm; 33 Fig. 23. Stylactaria arge. Scales equal 0.5 mm. a, Gastrozooid, and gonozooid with female medusa buds, romiz B355. b, Gastrozooid, and gonozooid with male medusa buds, romiz B354. c, Male medusa, one to two hours old, romiz B354. 0% -I' .:' I -' a b Q Fig. 24. Stylactaria arge, nematocysts of gastrozooid and gonozooid. Scales equal 10 p,m. a, Desmonemes of gastrozooid, romiz B354. b, Heterotrichous microbasic eurytele of gastrozooid, romiz B354. c, Haploneme of gonozooid, romiz B367. 34 heterotrichous microbasic euryteles 7.3-7.8 yon x 2.62.8 |xm. Gonozooids: desmonemes 5.5-6.5 (xm x 3.0-3.8 |xm; haplonemes 4.9-6.6 |xm x 2.4-2.7 |xm; heterotrichous microbasic euryteles 8.3-10.2 jxm x 2.9-3.7 |xm. Medusa sac-shaped, degenerate, about 1.1 mm high and 1.2 mm wide in formalin-preserved specimens; mesoglea thin. Manubrium tubular, extending nearly to velar open ing; mouth, oral arms, and oral tentacles lacking. Radial canals four; narrow ring canal present. Tentacles eight in number, rudimentary. Ocelli absent. Gonads encircling manubrium, fully developed prior to liberation of medusa. Nematocysts— Medusae: desmonemes 4.8-5.6 |xm x 2.6-3.0 |xm; heterotrichous microbasic euryteles 5.9-6.9 |xm x 2.42.9 [im. REMARKS Medusae of Stylactaria arge (Clarke, 1882) and S. hooperii (Sigerfoos, 1899) are inseparable from existing de scriptions (Clarke, 1882; Sigerfoos, 1899). Accounts of their hydroids differ only in minor respects (Fraser, 1944; Crowell, 1947; Calder, 1971, 1975). From the original descriptions (Clarke, 1882; Sigerfoos, 1899), S. hooperii seems to differ from S. arge in having (1) somewhat smaller gastrozooids, (2) gastrozooids that do not reproduce asexually by autotomy, (3) gastrozooid tentacles in a single row rather than in two closely placed whorls, and (4) spines on the hydrorhiza. I have been unable to locate type ma terial of either nominal species for comparison. Apparent differences such as those noted above are interpreted here as nothing more than variation that might be expected within a single species, and S. hooperii is regarded here as conspecific with S. arge. Both nominal species were originally described from eelgrass beds on the middle At lantic coast of the United States. Calder (1971, 1975) earlier referred Crowell's (1947) report of Stylactis sp. to this species. Specimens referred here to Stylactaria arge were less robust than the hydroids described by Clarke (1882) and Sigerfoos (1899), as also were colonies described earlier (romiz B666) from Chesapeake Bay (Calder, 1971). There seems little likelihood, from their overall similarity to 5. arge, that the specimens from Bermuda represent a distinct species. As in material from the United States, medusae were degenerate, short-lived, sexually mature at release, and liberated from the hydroids only at dusk. KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Calder, 1975). Superfamily Pandeoidea Haeckel, 1879 ?Trichydridae Hincks, 1868:215. Tiaridae Haeckel, 1879:40 [invalid name, type genus a junior homonym]. Pandaeidae Haeckel, 1879:46 [emended to Pandeidae by Bigelow, 1913]. Amphinemidae Haeckel, 1879:46. Protiaridae Haeckel, 1879:46. Bythotiaridae Maas, 1905:434. Calycopsidi Mayer, 1910:104. Stomotocini Cockerell, 1911:79. Hydrichthyinae Stechow, 1922:142. ?Timoididae Kramp, 1961:138. Niobiidae Petersen, 1979:133. Halimedusidae Arai and Brinckmann-Voss, 1980:62. a conical hypostome; those of parasitic forms elongate, degenerate, lacking tentacles. Gonophores free medusae, arising from hydrorhiza, hy drocaulus, branches, pedicels, or hydranths. Medusae bellshaped, with or without an apical projection; manubrium quadrate, with or without a peduncle; oral tentacles absent; mouth surrounded by four, or infrequently eight, lips with or without marginal nematocyst batteries. Radial canals four, or infrequently eight; centripetal canals usually ab sent. Marginal tentacles hollow, two, four, or more, with or without conical basal bulbs. Ocelli present or absent. Gonads on manubrium, extending outwards along radial canals in some species. REMARKS DIAGNOSIS Pelagiana trichodesmiae Borstad and Brinckmann-Voss, Hydroid colonies stolonal or erect, with a creeping hy drorhiza; some taxa ectoparasitic on fishes and ichthyoparasitic copepods, arising from a basal plate embedded in tissues of host. Perisarc variably developed; pseudohydrotheca present or absent. Hydranths of free-living forms with a single, oral whorl of filiform tentacles surrounding 1979, is evidently referable to the superfamily Pandeoidea Haeckel, 1879, but the family to which it should be re ferred is uncertain at present. Accordingly, only the su perfamily for this species is given here. Classification of the Pandeoidea is based largely on the medusa stage because hydroids of most genera within the 35 superfamily are unknown. A thorough revision of the Pandeidae by Hartlaub (1914) improved taxonomic under standing of the family, though he retained the name Tiaridae Haeckel, 1879, for the group. The name Tiaridae Haeckel, 1879, is invalid (Bigelow, 1913) because Tiara Lesson, 1843, its! nominal type genus, is a junior homonym of Tiara Swainson, 1832, a mollusc [Art. 39]. Bigelow's (1913) use of the name Pandeidae for the family has been widely followed in the subsequent literature (e.g., Bige Genus Pelagiana Borstad and Brinckmann-Voss, 1979 Pelagiana Borstad and Brinckmann-Voss, 1979:1233. DIAGNOSIS Medusa taxonomists have recently recognized that the family Pandeidae, as envisaged earlier this century, en compassed a mixed assemblage of genera. Some authors (e.g., Russell, 1953; Arai and Brinckmann-Voss, 1980) Hydroids on planktonic blue-green "algae" (Trichodesmium thiebautii); body globular with conical hypostome. Tentacles filiform, in an oral whorl; tentacular nematocysts arranged in two spiral bands. Gonophores free medusae, arising from hydranth prox imal to tentacular whorl. Young medusa thimble-shaped, with four radial canals. Mouth quadrate, without(?) clus ters of nematocysts or oral tentacles. Tentacle bulbs four, triangular, with one opposite pair larger than the other; marginal tentacles two, solitary, filiform, arising from the larger pair of tentacle bulbs. Ocelli absent. Adult medusa have divided the Pandeidae into a number of subfamilies. unknown. Petersen (1979) recognized a group of families, including the Calycopsidae Mayer, 1910, Protiaridae Haeckel, 1879, Pandeidae, and Niobiidae Petersen, 1979, within a superfamily, the Pandeoidea. The Trichydridae Hincks, 1868, Pelagiana trichodesmiae Borstad and Brinckmann-Voss, 1979, by original designation. low, 1918; Russell, 1953; Rees, 1956a; Yamada, 1959; Kramp, 1961; Goy, 1972; Millard, 1975; Petersen, 1979; Arai and Brinckmann-Voss, 1980; Bouillon, 1980, 1985). TYPE SPECIES and Halimedusidae Arai and Brinckmann-Voss, 1980, were also included in this superfamily by Bouillon (1985). How ever, if the Trichydridae is included in this taxon, the superfamily name Trichydroidea Hincks, 1868, would pre date the name Pandeoidea Haeckel, 1879 [Arts. 23a, 36]. It seems likely that further refinements to the classification of the group will become necessary as knowledge of these hydrozoans advances. Stechow (1922) established the subfamily Hydrichthyinae based on the parasitic hydroid genus Hydrichthys Fewkes, 1887. Fraser (1944) recognized the group as a distinct family, the Hydrichthyidae. Millard (1975) in cluded Hydrichthys in the Pandeidae based on the mor phology of its medusa stage. Through life-cycle studies, Larson (1982) identified the medusa of a Hydrichthys hy droid as Stomotoca pterophylla Haeckel, 1879, a pandeid. The family-group name Hydrichthyinae is, therefore, in cluded here within the synonymy of the Pandeoidea. Lar son (1982) concluded that the genus name Hydrichthys was a synonym of Stomotoca L. Agassiz, 1862, but Arai (in press) disagreed. She noted that medusae of the genus Stomotoca possess two tentacles, whereas immature me dusae of Hydrichthys mirus Fewkes, 1887, type species of Hydrichthys, have four tentacles (Fewkes, 1887). Bouillon (1980) observed that the gonads of Timoides agassizii Bigelow, 1904, arose from the manubrium of the medusa rather than from the radial canals as believed ear lier, and placed the family name Timoididae Kramp, 1961, in synonymy with the Pandeidae. Hydroids of several pandeoid medusa genera have been placed in the nominal genus Perigonimus M. Sars, 1846. However, Perigonimus is a junior subjective synonym of Bougainvillia Lesson, 1830 (see p. 24). 36 REMARKS Borstad and Brinckmann-Voss (1979) tentatively referred their nominal genus Pelagiana to the family Pandeidae Haeckel, 1879. They noted that the young medusa of P. trichodesmiae Borstad and Brinckmann-Voss, 1979, type species of the genus, resembles the Pandeidae in having large, triangular tentacle bulbs, filiform tentacles, four ra dial canals, and a four-cornered mouth. The hydroid also has a number of characters found in pandeids: it is monomorphic, and it has a single whorl of filiform tentacles and a conical hypostome. Nevertheless, the systematic position of this species is uncertain because its medusa has yet to be reared to maturity, and in the absence of more detailed information it is referred here only to the superfamily Pandeoidea. Borstad and Brinckmann-Voss (1979) observed that pandeid medusae undergo considerable morphological change during development, and suggested that life-cycle studies may demonstrate that this hydrozoan belongs to a species that is already known. Pelagiana trichodesmiae Borstad and Brinckmann-Voss, 1979 Figs. 25, 26 Pelagiana trichodesmiae Borstad and Brinckmann-Voss, 1979:1233; figs. 1-3 [hydroid and young medusa]. TYPE LOCALITY Barbados, West Indies. Fig. 25. Pelagiana trichodesmiae, hydroid on clump of Trichodesmium thiebautii, romiz B352. Scale equals 0.25 mm. Fig. 26. Pelagiana trichodesmiae, nematocysts of hydranth, romiz B352. Scales equal 10 p,m. a, moneme. b, Heterotrichous microbasic eurytele. Des- 37 Borstad and Brinckmann-Voss (1979) described the me MATERIAL EXAMINED Atlantic Ocean, 6 km southwest of Gibbs Hill lighthouse, dusa of Pelagiana trichodesmiae from a single specimen on Trichodesmium thiebautii, —1 m, 3 October 1984, two newly liberated from its hydroid, and did not find this colonies, 0.3 mm high, without gonophores, romiz B352. stage in the plankton. Clarification of the possible syn DESCRIPTION a more advanced stage. Although the hydroid of Pelagiana trichodesmiae is in onymy of this species awaits the rearing of its medusa to Hydroids minuscule, inconspicuous, solitary or forming stolonal colonies of a few hydranths; hydrorhiza with thin perisarc, embedded in filamentous tufts of planktonic Tri chodesmium thiebautii. Hydranths sac-shaped to pearshaped, up to 345 |xm high, 195 fjim wide, constricted at juncture with hydrorhiza. Tentacles four to five in number, filiform, in an oral whorl. Hypostome relatively large, conspicuous and poorly known, it appears to be relatively common. Geiselman (1977) found it in about 40 per cent of her near-surface collections of Trichodesmium thiebautii from the subtropical North Atlantic. Borstad and Brinckmann-Voss (1979) found it everymonth of the year at Barbados between August 1974 and June 1976. It was dome-shaped, with terminal mouth. Gonophores not seen. Nematocysts— Hydroids: desmonemes 4.6-5.2 (xm x 3.4-3.8 (xm; heterotrichous microbasic euryteles 6.8-7.9 |xm x 3.04.0 |xm. pothesized that the species may have a wide distribution. Hydroids of this species were collected by Geiselman (1977) during a cruise "from Spain to Bermuda," but the REMARKS closest of her records to Bermuda was in mid-North At lantic near 30° N, 44° W. immediately locatedin a plankton sampletakenoff Bermuda during this study. Given the occurrence of T. thiebautii in all tropical oceans, Borstad and Brinckmann-Voss hy Geiselman (1977) collected hydroids of this species in plankton samples from the subtropical Atlantic, but did not name them. Borstad and Brinckmann-Voss (1979) de scribed and named Pelagiana trichodesmiae from material collected at Barbados. Its hydroid has been reported only on Trichodesmium thiebautii, an oceanic blue-green "alga." KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Borstad and BrinckmannVoss, 1979); eastern Atlantic (Geiselman, 1977). Family Eudendriidae L. Agassiz, 1862 Eudendroidae L. Agassiz, 1862:342 [emended to Euden driidae by Hincks, 1868]. Myrionemidae Pictet, 1893:18. DIAGNOSIS Hydroid colonies usually erect with branched hydrocauli, arising from a creeping hydrorhiza; growth monopodial with terminal hydranths. Perisarc firm, reaching to groove at base of hydranth. Hydranth often large, urn-shaped to elongate; hypostome typically flared, flexible. Tentacles filiform, in a single whorl or in two or more close whorls. Gonophores fixed sporosacs, originating on hydranth beneath tentacles, often arranged in a whorl, reproductive hydranth often reduced to a blastostyle. Male gonophore and usually flaring distally. The classification of the Eu dendriidae has been spared the dual nomenclature found in many other families of athecate hydroids because all known species reproduce sexually by fixed sporosacs. The sexes are usually on separate colonies, although Euden drium motzkossowskae Picard, 1951, is reportedly her maphroditic (Picard, 1951). The family comprises two genera. Eudendrium Ehrenberg, 1834, a well-known genus with a worldwide distri bution, includes many nominal species. Myrionema Pictet, 1893, has been reported infrequently, is apparently re stricted to shallow-water habitats in the tropics and subtropics, and includes only three nominal species. Both genera are found in Bermuda. with one or more bulbous chambers, successive chambers in a linear series. Female gonophore initially with curved spadix, each spadix supporting a single egg. REMARKS Species of the family Eudendriidae L. Agassiz, 1862, are immediately distinguishable from other athecate hydroids by the shape of the hypostome, which is large, flexible, 38 Genus Eudendrium Ehrenberg, 1834 Calamella Oken, 1815:55 [invalid name, published in work rejected for nomenclatural purposes by the iczn, Opinion 417]. Eudendrium Ehrenberg, 1834:72. Corymbogonium Allman, 1861:171. Edendrium Allman, 1872:295 [incorrect subsequent spelling]. Erudendium Thompson, 1899:583 [incorrect subsequent spelling]. / Watson (1985) emphasized the value of the cnidome in identification of species of Eudendrium from Australia. Interspecific differences that were notedby Watson in the complement of nematocyst categories present, and in the length-width ratios of nematocyst capsules of a given category, could be used as a taxonomic character. DIAGNOSIS Eudendriidae with calyx of hydranth moderately short, urn-shaped. Tentacles in one whorl, of varied number but Eudendrium bermudense, sp. nov. usually fewer than 35. Figs. 27-29 TYPE SPECIES MATERIAL EXAMINED Tubularia ramosa Linnaeus, 1758, by subsequent desig nation by Allman (1872). Holotype: Sailor's Choice Cave, Hamilton Parish, on ledge at entrance, - 1.5 m, 30 June 1983, one colony, 2.0 cm high, with female gonophores, romiz B333. Paratypes: REMARKS Sailor's Choice Cave, Hamilton Parish, on ledges and Cornelius (1976) noted that the widely used generic name Eudendrium Ehrenberg, 1834, was threatened by the in colonies, 3.2 and 3.8 cm high, with male gonophores; one survey line at entrance, - 1 to - 2 m, 30 June 1983, two frequently used name Thoa Lamouroux, 1816. Two spe cies had originally been included in Thoa by Lamouroux (1816), T. savignii Lamouroux, 1816, and Sertularia hal colony, 2.0 cm high, with female gonophores; romiz B334. ecina Linnaeus, 1758. The former is a junior objective high, with female gonophores, romiz B335. Castle Har bour, under causeway near halfway point, on rocks and synonym of Tubularia ramea Pallas, 1766, a species now Sailor's Choice Cave, Hamilton Parish, on ledges at en trance, - 1 m, 27 June 1983, three colonies, 2.0-3.5 cm referred to Eudendrium. Sertularia halecina had com shells, - 1 m, 1 October 1986, five colonies, up to 6.6 cm monly been referred to Halecium Oken, 1815, a generic name invalidated (Opinion 417) because it was originally published in a work that was not consistently binominal. high, with female gonophores; four colonies, up to 4.6 cm high, with male gonophores; one sterile colony, 3.5 cm high; romiz B360. As a solution to these nomenclatural problems, Lemche (1976) recommended that the iczn (1) validate, under the DESCRIPTION plenary powers, the generic name Halecium Oken, 1815, as requested by Cornelius (1976), and (2) designate Ser Colonies slender, straggly, up to 3.8 cm high, arising from a creeping hydrorhiza. Hydrocaulus upright, monosi- tularia halecina Linnaeus, 1758, as the type species of phonic or slightly polysiphonic basally, more or less al ternately branched in one plane, branches similarly Thoa. This proposal was adopted (Opinion 1220), with Thoa becoming a junior objective synonym of the valid name Halecium. Another threat to Eudendrium has been removed else where in this report (p. 64) by designating Tubularia muscoides Linnaeus, 1761, as the type species of Fistulana O. F. Muller, 1776a. In so doing, Fistulana becomes a rebranched or with alternate pedicels. Perisarc of moderate thickness, horn-coloured to dark brown basally, becoming progressively thinner and more colourless towards extrem ities, terminating at groove around base of hydranth, annulated at bases of hydrocaulus, branches, and pedicels, with occasional irregularly placed annulations elsewhere junior subjective synonym of Coryne Gaertner, 1774. The genus Eudendrium comprises a common and dis tinctive group of hydroids. However, there are many nom inal species in the genus that are by no means easily distinguished, and a large number of these are of ques tionable validity. Fraser (1944) noted that descriptions of many of the 22 nominal species of Eudendrium reported from the western North Atlantic are meagre and based on incomplete specimens. The taxonomy of the genus has been further complicated because species have often been # r described on the basis of taxonomically unreliable char acters. In addition, there has been little evidence, partic ularly in the older literature, of an appreciation of the considerable range of colony form that can occur within a species. Fig. 27. Eudendrium bermudense, sp. nov., colony form. Natural size, a, Holotype, romiz B333. b, Paratype, romiz B334. 39 Fig. 28. Eudendrium bermudense, sp. nov., hydranthsand gonophores. Scales equal 0.5 mm. a, Hydranth, holotype colony, romiz B333. b, Hydranth with female gonophores, holotype colony, romiz B333. c, Hydranth with male gonophores, paratype colony, romiz B334. but mostly smooth. Hydranths urn-shaped, 447-638 |xm long from proximal end to base of hypostome when fully developed, 298-532 |xm wide, with a shallow but distinct perisarc groove basally; hypostome large, knob-shaped to flared. Tentacles 21 to 26 in number, solid, filiform, in one whorl. Gonophores fixed sporosacs, originating distal to peri sarc groove on hydranth. Female gonophores borne in a whorl on entire hydranths, spadix unbranched, curving over egg. Male gonophores with one or two chambers, borne on entire hydranths. Nematocysts— Hydroids: macrobasic euryteles (on hypostome, hy dranth base, coenosarc) 29.4-34.5 |xm X 12.3-14.0 |xm; heterotrichous microbasic euryteles (on tentacles, hypo stome, hydranth, coenosarc) 8.0-8.4 |xm x 3.4-3.7 (xm. REMARKS Eudendrium bermudense, sp. nov., differs from its Bermudian congeners in having macrobasic euryteles in ad dition to microbasic euryteles. In possessing nematocysts of the former category, E. bermudense resembles E. infundibuliforme Kirkpatrick, 1890, E. glomeratum Picard, 1951, and E. motzkossowskae Picard, 1951, as well as 40 Myrionema amboinense Pictet, 1893. Unlike E. infundibuliforme, hydranth pedicels of E. bermudense are not expanded distally. In contrast with E. glomeratum, ma crobasic euryteles of E. bermudense are scattered rather than being aggregated in dome-shaped batteries on the hydranth column. There is no evidence that the gonophores of E. bermudense are hermaphroditic, as has been reported in E. motzkossowskae (e.g., see Motz-Kossowska, 1905; Picard, 1951; Millard, 1975; Boero, 1981). Eudendrium bermudense is immediately distinguishable from M. am boinense in having more regularly branched colonies, smaller hydranths, and fewer tentacles, and in lacking algal symbionts. The hydroids resemble descriptions ofE. angustum Warren, 1908, which has large nematocysts of uncertain identity (Millard, 1975), but the hypostome of the latter is reportedly "blocked" by a solid plug of endodermal cells (Warren, 1908; Millard, 1975). No such plug was apparent in specimens of E. bermudense. Finally, large nematocysts, believed by Watson (1985) to be macrobasic euryteles, occur in E. aylingae Watson, 1985, and E. currumbense Watson, 1985. However, E. aylingae is ap parently a much smaller species than E. bermudense, and the macrobasic euryteles(?) of E. currumbense are con siderably smaller than those of E. bermudense. Fig. 29. Eudendrium bermudense, sp. nov., nematocysts of hydranth, romiz B333. Scales for a and c equal 10 p.m; scales for b and d equal 100 p-m. a, Macrobasic eurytele. b, Hypostome with macrobasic euryteles. c, Heterotrichous microbasic eurytele. d, Hydranth base with macrobasic euryteles. ETYMOLOGY TYPE LOCALITY The specific name refers to the occurrence of the species Embleton Bay, Northumberland, Great Britain. in Bermuda. MATERIAL EXAMINED KNOWN RANGE Known only from the type locality. Eudendrium capillare Alder, 1856a Figs. 30-32 Eudendrium capillare Alder, 1856a:355; pi. 12, figs. 9-12. Corymbogonium capillare—Allman, 1861:171. Dicoryne capillare—Alder, 1862:230. Eudendrium tenue A. Agassiz, 1865:160; fig. 250. Eudendrium capillare var. mediterranea Neppi, 1917:30. Castle Island, Castle Harbour, on underside of flat rocks, —2 m, 30 July 1982, one colony, 13 mm high, with male gonophores; two colonies, 6-13 mm high, without gono phores; romiz B142. Hungry Bay, on underside of flat rocks, —1.5 m, 6 September 1977, two colonies, 1117 mm high, with female gonophores, romiz B161. DESCRIPTION Colonies small, slender, straggly, up to 17 mm high, aris not Eudendrium Icapillare—Millard, 1966:454 [= Euden ing from a creeping hydrorhiza. Hydrocaulus upright, monosiphonic, 100-125 |xm wide, irregularly to more or less alternately branched; branches in turn alternately to irregularly branched; pedicels often long and bent. Perisarc relatively thick and golden-coloured at base of colony, becoming progressively thinner and colourless towards ex drium ramosum (Linnaeus, 1758)]. tremities, terminating at groove around base of hydranth, Eudendriumparvum Warren, 1908:272; fig. 1; pi. 45, figs. 1-4. 41 Fig. 30. Eudendrium capillare, colony form, romiz B142. Natural size. Fig. 31. Eudendrium capillare, hydranths and gonophores. Scales equal 0.25 mm. a, Hydranth, romiz B142. b, Reduced hydranth with female gonophores, romiz B161. c, Reduced hydranth with male gono phores, romiz B142. annulated at bases of hydrocaulus, branches, and pedicels, and with irregularly placed annulations frequent else where. Hydranths urn-shaped, 295-385 (xm long from proximal end to base of hypostome, 250-340 (xm wide, with a shallow but distinct perisarc groove basally. Hy postome large, flared. Tentacles 15 to 20 in number, solid, filiform, in one whorl. Gonophores fixed sporosacs, originating distal to peri sarc groove on hydranth. Female gonophores borne in a whorl on hydranths with partially atrophied tentacles; spa dix unbranched, curving over egg. Male gonophores with up to three chambers each, borne on atrophied hydranths; terminal chamber with an apical tubercle. Nematocysts— Hydroids: heterotrichous microbasic euryteles (on ten tacles, hydranth, and elsewhere) 7.1-8.0 |xm x 3.03.2 fim. REMARKS Records suggest that Eudendrium capillare Alder, 1856a, is widely distributed (Vervoort, 1959), but Millard (1975) 42 cautioned that most records provide insufficient informa tion to verify identification. Although the species has been frequently recorded from warm waters (e.g., Fraser, 1912, 1948; Mammen, 1963; Millard and Bouillon, 1974; Cooke, 1975; Millard, 1975), hydroids from Bermuda were never theless referred to E. capillare with some reservation given the northerly type locality of this species. Unfortunately, type material of E. capillare could not be located (Cor nelius and Garfath, 1980), but specimens from Bermuda corresponded with Alder's (1856a) figures and brief de scription of the species. Bermuda specimens are also iden tical in all major respects, including the complement and arrangement of nematocysts, with more detailed accounts of the species given by Millard and Bouillon (1974) from East Africa and by Millard (1975) from South Africa. A small hydroid somewhat resembling Eudendrium capT illare in colony form was described and named E. tenellum by Allman (1877) from material collected at a depth of 471 fathoms (861 m) off Florida. Hydranths and gono phores, both of which are now generally regarded as es sential for diagnosis of any species of the genus Eudendrium therefore, not regarded it as conspecific with E. capillare. Hincks (1868) regarded Eudendrium tenue A. Agassiz, 1865, as possibly conspecific with E. capillare. Vervoort (1946b) thought likewise, and I concur. Eudendrium parvum Warren, 1908, also seems conspecific with E. capil lare, for reasons given by Millard (1966). KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Fraser, 1944); eastern At lantic (Hincks, 1868); Indian Ocean (Millard, 1975); west ern Pacific (Yamada, 1959); eastern Pacific (Fraser, 1937). Eudendrium carneum Clarke, 1882 Figs. 33-35 Eudendrium ramosum—McCrady, 1859a: 166. —A. Fig. 32. Eudendrium capillare, heterotrichous microbasic eurytele from hydranth, romiz B161. Scale equals 10 |j.m. Agassiz, 1865:160. —Congdon, 1906:27; figs. 1-4; 1907:464. —Fraser, 1912:349; figs. 8A-C; 1943:87. — Bennitt, 1922:245 Inot Eudendrium ramosum (Linnaeus, 1758)]. Ehrenberg, 1834, were lacking in Allman's material. In fact, Allman was not certain that his specimen belonged to Eudendrium. He stated (1877:8): "Its reference to this genus is probably correct, but as neither hydranths nor gonophores were present in the specimen, it may possibly have its true place in some other." Nonetheless, many authors have identified small hydroids of this genus as E. tenellum, and the species has been reported from Atlantic, Pacific, Arctic, and Indian oceans (e.g., Stechow, 1923a; Fraser, 1937, 1944, 1948; Kramp, 1943; Yamada, 1959; Calder, 1972; Hirohito, 1977). It is unfortunate that the name E. tenellum, based on unrecognizable material and regarded here as a nomen dubium, should subsequently have been applied to what is seemingly a recognizable species. It is not possible to ascertain whether any of the later records of E. tenellum are actually conspecific with the hydroid upon which the name is based. These records are founded on one or more species of Eudendrium, ap parently differing from E. capillare in having gonophores on entire rather than on atrophied hydranths. Naumov (1960) regarded E. tenellum auct. as a synonym of E. capillare, and Christiansen (1972) concurred with this view. Calder (1972), Hirohito (1977), and others have regarded the two as distinct based on differences in colony form and degree of reduction of hydranths bearing gonophores. Naumov (1960) also regarded E. hyalinum Bonnevie, 1899b, as conspecific with E. capillare. Bonnevie's (1899b) material was sterile and her description was so general that the identity of her hydroid is uncertain, although it could be identical with the E. tenellum of some authors. I have, Eudendrium carneum Clarke, 1882:137; pi. 7, figs. 10-17. Eudendrium cunninghami Kirkpatrick, 1910:127; pi. 7, figs. 1-3. TYPE LOCALITY Fort Wool, Hampton Roads, Virginia, United States. MATERIAL EXAMINED Hamilton Harbour, on mooring chain, —2.5 m, 12 No vember 1976, two colonies, 6.4 cm high, with female gonophores, coll. J. Markham, L. Coen, G. Rupp, romiz B134. Flatts Inlet, on concrete pier, - 2 m, 24 May 1979, five sterile colonies, up to 6 cm high; seven colonies, up to 6 cm high, with male gonophores; five colonies, up to 10 cm high, with female gonophores; romiz B135. Castle Grotto, Castle Harbour, on cave wall 25-50 m from en trance, - 1 m, 20 July 1982, one sterile colony, 5.8 cm high, romiz B148. Flatts Inlet, on rocks, -0.5 m, 27 February 1982, one colony, 9.3 cm high, with male gono phores, romiz B166. Ferry Reach, St George's Island, on rope, -0.5 to —2 m, 2 September 1977, one male col ony, 3.6 cm high; one sterile colony, 10.5 cm high; romiz B176. Somerset Bridge, on concrete wall, —2 m, 15 Sep tember 1977, one male colony, 10 cm high, romiz B178. DESCRIPTION Colony extensively branched and bushy, up to 10.5 cm high, arising from a creeping mass of hydrorhizal stolons. Hydrocaulus upright, polysiphonic, more or less alter nately branched; primary branches also polysiphonic and 43 alternately or somewhat irregularly branched; secondary branches often polysiphonic basally and branched in like manner. Perisarc thick and brownish-coloured in older parts of colony, thinner and paler towards extremities, annulated or wrinkled at bases of branches and hydranth pedicels, with occasional annulations elsewhere but mostly smooth, terminating almost imperceptibly at groove around hy dranth base. Hydranths urn-shaped, about 0.8 mm long from proximal end to base of hypostome, 0.65 mm wide, with a shallow perisarc groove and a ring of anisorhiza nematocysts basally; hypostome very large, flared to knobshaped. Tentacles about 27 to 32 in number, solid, fili form, in one whorl. Gonophores fixed sporosacs, developing on hydranth distal to perisarc groove. Female gonophores on reduced hydranths with partially atrophied tentacles; spadix bifid, curving over egg. During development, spadices shed, embryos borne in perisarc-covered capsules arranged ir regularly along pedicel, perisarc of gonophore pedicel ex tensively wrinkled, terminal hydranth eventually lost. Male gonophores with up to five chambers each, borne on atro phied hydranths; distal end of gonophore with scattered anisorhiza nematocysts. Nematocysts— Hydroids: heterotrichous anisorhizas (on hydranth base, hypostome, and tips of male gonophores) 20.3-23.0 (xm x 9.4-11,3 |xm; heterotrichous microbasic euryteles (on tentacles, hydranth, and elsewhere) 8.3-9.4 (xm x 3.64.0 |xm. Fig. 33. Eudendrium carneum, colony form, romiz B135. Natural size. Eudendrium, but all Allman's species appear to have been different from E. carneum. So does E. distichum, de scribed by Clarke (1879) from material obtained southwest of Key West, Florida, in 339 fathoms (620 m) of water. Although E. distichum appears to have resembled E. car neum in colony size and general shape, it seems to have had only 16 to 20 tentacles on the hydranth. Some of Allman's (1877) species were evidently distinguished on the basis of relatively minor differences, and their status REMARKS needs to be re-evaluated. Allman (1877) described and named eight new nominal In Bermuda, Eudendrium carneum was abundant during warm months in areas swept by strong tidal currents, such as Flatts Inlet and the Somerset Bridge area. Specimens with active hydranths and gonophores were also collected in Flatts Inlet at 18° C during the winter of 1982 (romiz species of Eudendrium Ehrenberg, 1834, from the south eastern United States, where E. carneum Clarke, 1882, is now known to be frequent (Fraser, 1944; Calder and Hes ter, 1978). One of these, E. tenellum, was discussed ear lier. Of the remaining seven, all differ from E. carneum in one or more respects. The hydrocaulus was described as monosiphonic, rather than polysiphonic, in E. attenu- B166). Eudendrium carneum has been reported previously from Bermuda as E. ramosum (Linnaeus, 1758) by Congdon atum, E. laxum, and E. cochleatum. The number of ten (1906, 1907) and Bennitt (1922). Records of E. ramosum tacles borne by the hydranth was reported to be only about 20, instead of 25 or more, in E. eximium, E. exiguum, and E. fruticosum. Annulations at the bases of branches and pedicels, well marked in E. carneum, were faint or even absent in E. eximium, E. fruticosum, and E. gracile. Reproductive hydranths, reduced or aborted in E. car neum, were little if at all aborted in E. eximium, E. fru ticosum, and E. laxum. Unfortunately, gonophores were lacking in Allman's material of E. exiguum and E. gracile, and both gonophores and hydranths were lacking in his specimens of E. attenuatum. Characters such as tentacle number, degree of atrophy of reproductive hydranths, ex tent of perisarc annulation, and number of tubes compris ing the hydrocaulus are known to vary intraspecifically in from the southeastern United States by McCrady (1859a), A. Agassiz (1865), and Fraser (1912, 1943b) were also likely based on material of E. carneum. Ultrastructural studies on spermatozoa of E. carneum were conducted in Bermuda by Summers (1972a), again on hydroids mis- 44 identified as E. ramosum. KNOWN RANGE Bermuda: no specific locality given (Congdon, 1906, 1907); Hamilton Harbour (Bennitt, 1922); Flatts Inlet (Summers, 1972a); shallow inshore waters (Calder, 1986). Elsewhere: western Atlantic (Fraser, 1944); eastern At lantic (Kirkpatrick, 1910); Indian Ocean (Millard, 1975); eastern Pacific (Fraser, 1948). Fig. 34. Eudendrium carneum, hydranths and gonophores, romiz BOS. Scales equal 0.5 mm. a, Hy dranth. b, Reduced hydranth with female gonophores. c, Pedicel with capsules containing embryos, d, Reduced hydranth with male gonophores. Fig. 35. Eudendrium carneum, nematocysts of hydranth, romiz B135. Scales equal 10 p.m. a, Hetero trichous anisorhiza. b, Heterotrichous microbasic eurytele. 45 Genus Myrionema Pictet, 1893 Myrionema Pictet, 1893:18. DIAGNOSIS Eudendriidae with calyx of hydranth elongate, often col DESCRIPTION Colonies straggly, growing in clumps up to 5.6 cm high; hydrorhiza creeping. Hydrocaulus monosiphonic, 0.160.25 mm in diameter, sparingly and irregularly branched; primary branches unbranched or irregularly branched, often directed upwards and resembling hydrocaulus in appear umnar below tentacles. Tentacles in two or more close ance and size. Perisarc thin, flexible, straw-coloured to whorls, number varied but commonly 40 or more. virtually colourless, usually annulated or faintly wrinkled at bases of branches, mostly smooth elsewhere, terminat ing at groove around hydranth base. Hydranth reaching about 1.7 mm long from proximal end to base of hypo stome, urn-shaped to club-shaped with a long, cylindrical calyx, widest at tentacle-bearing region, with a shallow perisarc groove and a ring of macrobasic eurytele nema tocysts basally. Hypostome large, flared to knobbed. Ten tacles up to 2 mm long, about 35 to 60 in number, solid, filiform, in two or more close whorls. Hydranth and ten tacles bearing large numbers of zooxanthellae. TYPE SPECIES Myrionema amboinense Pictet, 1893, by monotypy. REMARKS This genus was instituted by Pictet (1893) for a new spe cies, Myrionema amboinense, from the western Pacific. The greater number of tentacles, which occur in several close whorls, and the elongate calyx of the hydranth dis tinguish hydroids of this genus from the more familiar Eudendrium Ehrenberg, 1834. Hydroids of this genus con tain symbiotic zooxanthellae in their tissues and are ap parently restricted to shallow water. Gonophores fixed sporosacs, originating on hydranth proximal to tentacles. Male gonophores with one to four chambers, borne on entire hydranths. Female gonophores borne in a whorl of as many as eight or more on entire Myrionema amboinense Pictet, 1893 Figs. 36, 37 hydranths; spadix unbranched, curving over egg. During development, spadices shed, embryos borne in perisarccovered capsules arranged irregularly along pedicel. Nematocysts— Myrionema amboinensis Pictet, 1893:19, 62; pi. 1, figs. 12,13; pi. 3, figs. 55,56. Eudendrium hargitti Congdon, 1906:27; figs. 5-11. "Eudendrium" hargettii—Wallace, 1909:137 [incorrect subsequent spelling]. Eudendrium griffini Light, 1913:333; figs. 1-5; pis. 1, 2. Eudendrium amboinensis—Leloup, 1932:143; fig. 15; pi. Hydroids: macrobasic euryteles (on hydranth base, hy postome) 21.8-23.4 |xm X 9.7-11.3 jim; heterotrichous microbasic euryteles (on tentacles, hydranth, and else 16, fig. 1. Morphological differences between Myrionema hargitti (Congdon, 1906) from the tropical western Atlantic and Myrionema amboinense—Briggs and Gardner, 1931:184; pi. 1, figs. 1-3. Myrionema hargitti—Spracklin, 1982:240; fig. 114b. Myrionema griffini—Watson, 1985:180. TYPE LOCALITY Baton-Mera, Ambon, Moluccas, Indonesia. MATERIAL EXAMINED Tucker's Town Bay, Castle Harbour, on pontoon anchor chain, - 1 m, 23 July 1982, one colony, 3.5 cm high, with male gonophores, romiz B141. Tucker's Town Bay, Castle Harbour, on pontoon anchor chain, -0.5 m, 5 March 1982, three colonies, 3.5-5.6 cm high, without gonophores, romiz B171. Walsingham Pond, on rocky cliff, - 2 m, 5 September 1977, one colony, 3.4 cm high, without gonophores, romiz B179. Whalebone Bay, on where) 8.5-9.4 |xm x 3.5-3.8 p.m. REMARKS M. amboinense Pictet, 1893, from the Indo-west Pacific appear to be negligible. Hydroids of both are shallowwater inhabitants harbouring large numbers of algal symbionts, and are brownish in colour when alive. Specimens of M. amboinense illustrated by Millard and Bouillon (1973) have longer calyces than hydroids referred to M. hargitti, but calyx length is highly variable in this genus. So too is tentacle number, although counts have usually been higher in M. amboinense. In M. hargitti, tentacle number has been reported as 35 to 45 (Congdon, 1907) and 35 to 60 (Bennitt, 1922). Tentacle number in M. amboinense, with which M. griffini (Light, 1913) is conspecific, has been reported as 80 to 120 (Pictet, 1893), 50 to 70 (Light, 1913), 40 to 50 (Hargitt, 1924), and 70 to 90 (Millard and colonies, 1.3-4.0 cm high, with female gonophores, romiz Bouillon, 1973). Nematocyst types and sizes appear to be similar in the two. In proposing M. griffini, Light (1913) noted that the tentacles were heavily armed with nema tocysts. Congdon (1906, 1907) reported that nematocysts B329. were few in number on the tentacles of M. hargitti. An rocks in Thalassia bed, -0.5 m, 24 June 1983, seven 46 a Fig. 36. Myrionema amboinense. Scale for a equals 3 mm; scale for b equals 1 mm; scale for c equals 0.5 mm. a, Hydrocaulus and hydranths, with a male gonophore, romiz B141. b, Hydranth, with female gonophores, romiz B329. c, Hydranth, romiz B141. Fig. 37. Myrionema amboinense, romiz B171. Scale for a equals 40 p-m; scales for b and c equal 10 |xm. a, Part of a tentacle, with nematocyst battery (arrow) and algal symbionts. b, Heterotrichous microbasic eurytele. c, Macrobasic eurytele. 47 examination of the tentacles of Bermuda specimens of M. amboinense during this study revealed that nematocysts were present in moderate numbers (Fig. 37a). Although hydroids of the two nominal species are isolated geo graphically, I regard M. hargitti as conspecific with M. 1907) and later seen by Smallwood (1910). Bennitt also located it just below low tide on buoys, timbers, ledges, and turtle grass throughout Hamilton Harbour and Great Sound. amboinense. KNOWN RANGE Myrionema amboinense was observed in Bermuda dur ing this study only in shallow waters of relatively sheltered areas such as Tucker's Town Bay. It was also one of the few hydroid species collected in the quiet waters of Walsingham Pond. Bennitt (1922) reported this hydroid to be extremely abundant in Hungry Bay on the south shore of Bermuda, where it was first observed by Congdon (1906, Bermuda: no specific locality given (Congdon, 1906); inlet on south shore (Congdon, 1907); Hungry Bay (Smallwood, 1910); Hungry Bay, Hamilton Harbour, Great Sound (Bennitt, 1922); quiet inshore waters (Calder, 1986). Elsewhere: western Atlantic (Fraser, 1944); eastern At lantic (Picard, 1958); Indian Ocean (Millard and Bouillon, 1973); western Pacific (Pennycuik, 1959). Family Corymorphidae Allman, 1872 Corymorphidae Allman, 1872:386. Monocaulidae Allman, 1872:395. Amalthaeidae Haeckel, 1879:37. Steenstrupiini Cockerell, 1911:78. Branchiocerianthidae Broch, 1916:21. Paragotoeidae Ralph, 1959:176. DIAGNOSIS Hydroids solitary. Hydrocaulus upright, more or less cy lindrical, invested with thin, flexible perisarc; usually with papillae and anchoring filaments basally. Hydranths ter minal, flask-shaped, with oral and aboral bands of ten tacles; oral tentacles capitate or filiform, in one or more close whorls; aboral tentacles filiform, in one whorl. Gonophores fixed sporosacs or free medusae, generally borne on blastostyles arising from hydranth just distal to aboral tentacles. Medusa, when present, bell-shaped; exumbrella without tracks of nematocysts; manubrium short, with simple, circular mouth. Radial canals four. Marginal tentacles one to four. Ocelli absent. Gonads surrounding manubrium. REMARKS Allman (1872) established the family Corymorphidae to accommodate Corymorpha M. Sars, 1835, and several other nominal genera now considered identical with it. In the same publication, he founded the Monocaulidae to include Monocaulus Allman, 1864a. For much of the pres ent century, the genera Monocaulus and Corymorpha, and the two families based on them, have been regarded as identical. The name Monocaulidae has fallen into disuse in favour of the name Corymorphidae, but to my knowl edge criteria of the First Reviser Principle in nomenclature [Art. 24] have never been met in this case. The two are considered synonyms here, and the name Corymorphidae 48 is assigned precedence over Monocaulidae in the interests of nomenclatural stability. Rees (1957) recognized four subfamilies within the Corymorphidae. "Lower corymorphines," such as Euphysa Forbes, 1848, Hypolytus Murbach, 1899, and pos sibly Gymnogonos Bonnevie, 1898, were grouped in the Euphysinae Haeckel, 1879. "Higher corymorphines," including Corymorpha, were placed in the Corymorphinae. Boreohydra Westblad, 1937, considered an aber rant corymorphine by Rees, was retained in the Boreohydrinae Westblad, 1947. Branchiocerianthus Mark, 1898, viewed as a corymorphine with secondarily acquired bilateral symmetry, was placed in the Branchiocerianthinae Broch, 1916. Although there has been little outright criticism of this classification, Millard (1975) cautioned that subdivision of the family is still not settled. For ex ample, Calder (1974) suggested that Boreohydra may be better referred to the family Myriothelidae Hincks, 1868 (= Candelabridae Stechow, 1921a) rather than to the Cor ymorphidae, and Petersen (1979) elevated the Euphysinae to the rank of family. Bouillon (1985) recognized the Euphysidae and Boreohydridae as distinct families. The family-group names Amalthaeidae Haeckel, 1879, and Steenstrupiini Cockerell, 1911, are synonyms of the Corymorphidae because their type genera, Amalthaea Schmidt, 1852, and Steenstrupia Forbes, 1846, are now generally considered congeneric with Corymorpha (e.g., see Broch, 1916; Brinckmann-Voss, 1970; Millard, 1975; but also see Rees and Thursfield, 1965; Bouillon, 1985). Brinckmann-Voss (1970) has been followed in regarding the Paragotoeidae Ralph, 1959, as identical with the Cory morphidae. The family Euphysidae, including the Trichorhizini Cockerell, 1911, and Hypolytidae Fraser, 1943a, was considered valid by Petersen (1979) and Bouillon (1985). Hydroids of the families Corymorphidae and Tubulariidae Goldfuss, 1818, are obviously related, and several authors (e.g., Broch, 1916; Stechow, 1923a; Russell, 1953; Naumov, 1960; Kramp, 1961) have united the two. Dif ferences between them are usually more pronounced in the hydroid stage than in the medusa. The family Cory morphidae is recognized as a distinct taxon in this report, a position held previously by authors such as Kramp (1949), Rees (1957), Brinckmann-Voss (1970), Vervoort (1972), Calder (1975), Millard (1975), Petersen (1979), and Bouil lon (1985). Genus Zyzzyzus Stechow, 1921a the Corymorphidae in my opinion. For example, the hy droid is strictly solitary; the hydrocaulus is thick and par enchymatous, with well-developed endodermal canals; the perisarc is thin, soft, and flexible; and the internal anatomy of the hydranth is more like that of Corymorpha M. Sars, 1835 (Allman, 1872; Rees, 1957) than that of Tubularia or the "lower corymorphines" (Gronberg, 1897; Rees, 1957). The gastrovascular cavity of the hydranth in Zyzzy zus is separated into oral and aboral chambers by a welldeveloped diaphragm (Fig. 38). For these reasons, the genus is included here in the Corymorphidae. The absence of papillae on the hydrocaulus, the presence of one or more anchoring stolons basally, and the existence of an actinula larva in its life cycle distinguish Zyzzyzus from Corymorpha. Zyzzyzus Stechow, 1921a:249. Zyzzygus Neave, 1940a:712 [incorrect subsequent spelling]. Zyzzyzuz Bouillon, 1985:243 [incorrect subsequent spelling]. Zyzzyzus warreni, nom. nov. Figs. 38-40 DIAGNOSIS Tubularia solitaria Warren, 1906b:83; pis. 10, 11 [invalid junior primary homonym of Tubularia solitaria Rapp, 1829 Corymorphid hydroids epizoic on sponges. Hydrocaulus parenchymatous, with endodermal canals, with rooting processes present basally, covered with thin perisarc. Hy dranths radially symmetrical, with two whorls of tentacles; aboral tentacles long, filiform, in one whorl; oral tentacles relatively short, capitate in young hydroids, otherwise fili form, forming a band around hypostome. Gonophores fixed sporosacs, arising from blastostyles distal to aboral tentacles, forming actinulae; hydroids monoecious. TYPE SPECIES (not a hydroid)]. Zyzzyzus solitarius—Stechow, 192la:249. Corymorpha solitaria—Kramp, 1933:12. TYPE LOCALITY Natal, South Africa. MATERIAL EXAMINED Flatts Inlet, on sponge, - 1 to - 2 m, 13 September 1977, several small hydroids, up to 5 mm high, some with de veloping blastostyles, romiz B133. Flatts Inlet, on sponges Tubularia solitaria Warren, 1906b (not Tubularia solitaria and Eudendrium carneum on underside of flat rock, —3 m, Rapp, 1829), by monotypy. 2 August 1982, several hydroids, up to 10 mm high, with developing blastostyles, romiz B147. Castle Grotto, Cas REMARKS Stechow (1921a) founded Zyzzyzus to accommodate Tu bularia solitaria Warren, 1906b (not Tubularia solitaria Rapp, 1829), a hydroid having characteristics of both the Corymorphidae Allman, 1872, and Tubulariidae Goldfuss, 1818. The genus was referred to the subfamily Corymorphinae, within the family Tubulariidae, by Stechow (1923a). Kramp (1933) initially regarded Zyzzyzus as congeneric with Corymorpha Allman, 1872, but concluded later (Kramp, 1949) that the genus was based on a species of Tubularia Linnaeus, 1758, which was adapted for life as an epizoite on sponges. Millard (1975), Watson (1978), Petersen (1979), and Bouillon (1985) recognizedZyzzyzus as valid, but referred it to the Tubulariidae rather than the Corymorphidae. Zyzzyzus resembles Tubularia and theTubulariidae mainly in having an actinulalarva in its life cycle (Warren, 1906b), but in most other respects it resembles Corymorpha and tle Harbour, about 25 m inside cave entrance, on sponge, - 1 m, 20 July 1982, several hydroids, up to 11 mm high, with gonophores, romiz B165. Stream PassageCave, Har rington Sound, 3 m inside cave entrance, on sponge, - 1 m, 20 June 1983, several hydroids, up to 17 mm high, with gonophores, romiz B370. DESCRIPTION Hydroids solitary, up to 17 mm high, usually embedded in sponge tissue basally. Base bulbous, lacking papillae but with one or more stout anchoring stolons of varying length. Hydrocaulus parenchymatous, with branching and anastomosing endodermal canals, reaching 1.5 mm wide basally, tapering distally. Perisarc thin, flexible, termi nating just below hydranth in a distinct circular perisarc groove. Hydranths up to 1.7 mm high, 1.5 mm wide, dis tinctly demarcated from hydrocaulus, vasiform with two whorls of tentacles. Aboral tentacles filiform, reaching 49 3 mm long; oral tentacles much shorter, capitate in young polyps, otherwise filiform; tentacle number varying with hydranth size, larger hydranths with 22 to 25 aboral and If 15 to 20 oral tentacles. Blastostyles short, arising just distal to aboral tentacles, bearing clusters of fixed, cryptomedusoid gonophores. In examined specimens, gonophores incompletely devel oped, lacking actinulae. Nematocysts— Hydroids: desmonemes 3.7-4.2 |xm x 2.8-2.9 \±m; isorhizas (ovate) 6.6-7.5 p.m x 2.8-3.7 p.m; ?isorhizas (reniform) 6.4—7.3 p.m x 1.9-2.1 p,m; ?mastigophores 6.88.2 |xm x 4.3-5.4 pun; microbasic euryteles 9.5-10.8 p.m x 4.9-6.1 p,m; stenoteles (small) 5.5-5.7 p,m x 4.54.7 (i.m; stenoteles (medium) 6.7-7.3 fjun X 5.7-6.4 p,m; stenoteles (large) 9.6-12.2 (ira X 8.7-11.2 |xm. REMARKS The name Tubularia solitaria Warren, 1906b, is an invalid junior primary homonym of Tubularia solitaria Rapp, 1829, and must be replaced |Art. 52]. Zyzzyzus warreni is pro posed here as a new replacement name for Warren's taxon. Zyzzyzus warreni resembles descriptions and illustra tions of Tubularia spongicola von Lendenfeld, 1885b, by von Lendenfeld (1885b) and Watson (1978). According to Watson, Z. warreni differs from Z. spongicolus in hav ing male and female gonophores on separate blastostyles instead of on the same ones. From existing descriptions, the two seem otherwise remarkably similar and may prove conspecific. According to Watson (1978), von Lendenfeld's (1885b) description and figure of Z. spongicolus do not correspond well with the type material, which she re o oc VH 'j^^** mm.f I 0 ^'*'*•*'*' *w» % * ~>m Fig. 38. Zyzzyzus warreni, photomicrograph of cross-section of hydranth, showing oral (oc) and aboral (ac) chambers sepa rated by a well-developed diaphragm, romiz B147. Scale equals 250 pm. examined. In Bermuda, this hydroid is usually, but not exclusively, epizoic on sponges. A few specimens were found attached to the hydrocaulus of the hydroid Eudendrium carneum Clarke, 1882, during this study (romiz B147). Zyzzyzus warreni is evidently dormant during winter in Bermuda. No specimens were observed on a collecting trip in February and March 1982, although it was specif ically looked for in areas where it was known to occur earlier and where it was collected on subsequent summer trips. ETYMOLOGY The replacement name for this species honours Ernest Warren, who first discovered and named this species, and who contributed significantly to knowledge of the hydroids of Natal. KNOWN RANGE Bermuda: first record. Elsewhere: western Atlantic (Millard, 1975); eastern At lantic (Ritchie, 1908); Indian Ocean (Millard, 1975). 50 Fig. 39. Zyzzyzus warreni, hydroid, romiz B147. Scale equals 2 mm. a b f ; Fig. 40. Zyzzyzus warreni, nematocysts of hydranth, romiz B165. Scales equal 10 p,m. a, Desmoneme. b, Ovate isorhiza. c, Reniform isorhiza. d, ?Mastigophore. e, Microbasic eurytele. /, Small stenotele. g. Medium stenotele. h, Large stenotele. 51 Family Tubulariidae Fleming, 1828 Tubulariadae Fleming, 1828:552 [corrected to Tubularii dae by Hincks, 1868]. Hybocodonidae Allman, 1872:421. Hybdocoridae Pennington, 1885:71 [incorrect subsequent spelling of Hybocodonidae Allman, 1872]. DIAGNOSIS Hydroids solitary or colonial. Hydrocaulus upright, cylin drical, invested with thick, rigid perisarc. Hydranths ter minal, flask-shaped, with oral and aboral whorls of tentacles. Oral tentacles relatively small, usually filiform in mature hydranths, capitate in immature ones; aboral tentacles large, filiform, in one whorl. Gonophores fixed sporosacs or free medusae, borne on blastostyles arising from hydranth just distal to aboral ten tacles; developmental stages including an actinula larva. Medusa, when present, bell-shaped; exumbrella with or without tracks of nematocysts; manubrium short, with sim ple, circular mouth. Radialcanals four. Marginal tentacles one to four. Ocelli absent. Gonads surrounding manubrium. REMARKS Goldfuss (1818) appears to have been the first to establish a family-group taxon under the name "Tubulariae." Four genera were included in the group, Clava Gmelin, 1790, Coryne Gaertner, 1774, Calamella Oken, 1815, and Ser tularia Linnaeus, 1758. The name, based on a genus ex cluded from the group, does not meet the criteria of availability [Art. 1If (i)(l)]. Goldfuss recognized the ge nus Tubularia Linnaeus, 1758, as valid, but it was included with a heterogeneous assemblage of invertebrate genera in another family, named the Polypi. Fischer von Waldheim (1823) also recognized a family "Tubulariae," but did not include Tubularia in the group. The family-group vast majority of species in the present state of our knowledge.'' The matter is of relevance in delimitation of several genera within the Tubulariidae. In most contemporary classifications, Ectopleura L. Agassiz, 1862, and Hybo codon L. Agassiz, 1862, are distinguished from Tubularia Linnaeus, 1758, chiefly in having free medusae instead of fixed gonophores. Ectopleura and Hybocodon in turn are distinguished on the basis of differences in the morphology of their medusae. These three genera were redefined by Petersen (1979) in a step towards a single classification system for the athecate hydroids and their medusae. Gono phores were considered in his scheme, but the presence or absence of a free medusa was not viewed as a diagnostic character at the generic level. He included in Tubularia those solitary species of tubularian hydroids producing either asymmetrical fixed gonophores or asymmetrical me dusae. Under such a redefinition of the genus, Hybocodon becomes congeneric with Tubularia. Primitively colonial tubularian hydroids, having either symmetrical fixed gono phores or symmetrical medusae, were referred to the genus Ectopleura. Petersen's proposal eliminates vestiges of the illogical dual classification for hydroids and medusae in the Tubulariidae, and makes it possible to refer hydroids and medusae, independently of each other, to the appro priate genus. Further evaluation is needed to test whether it is tenable for all species of the family, and whether it constitutes a natural classification. Significant advances have been made over the past three decades towards achieving a single classification of the Hydrozoa. Nevertheless, there is at present little alterna tive but to retain separate genera for many hydroids and hydromedusae until the identity and affinities of the two generations are resolved through life-cycle and systematic name Tubulariidae was first made available by Fleming (1828), as the Tubulariadae. studies. Even when life cycles are known, serious diffi culties may arise in formulating a single natural classifi Allman (1872) founded the family Hybocodonidae to accommodate Hybocodon L. Agassiz, 1862, but the genus cation (Rees, 1957). is now generally referred to the Tubulariidae (e.g., Millard, of the Tubulariidae known from Bermuda, is the conven 1975; Petersen, 1979; Bouillon, 1985). Pennington(1885) incorrectly spelled this nominal family 4tHybdocoridae." tional one followed by authors such as Brinckmann-Voss (1970), Millard (1975), and Bouillon (1985). There has been disagreement among hydrozoan systematists as to whether hydroids with fixed gonophores should Genus Ectopleura L. Agassiz, 1862 The definition adopted for Ectopleura, the only genus be recognized as different genera from those producing free medusae, when other criteria supporting generic sep aration are lacking. The view that hydroids should not be referred to different genera if they differ only in gonophore type hasbeenadvocated by authors such as Levinsen (1893), Broch (1916), Kramp (1949), and Petersen (1979). Con versely, Rees (1957:498) reviewed various points of view in the debate and concluded that "the use of separate genera is justifiable and the only suitable course for the 52 Ectopleura L. Agassiz, 1862:342. Acharadria Wright, 1863c:378. Acharadrium Allman, 1872:376 [incorrect subsequent spelling]. Acharadia Brinckmann-Voss, 1970:25 [incorrect subse quent spelling]. Archaradia Bouillon, 1985:112 [incorrect subsequent spelling]. DIAGNOSIS Primitively colonial hydroids, with the characters of the family. Gonophores free medusae, with the characters of the family. Medusa radially symmetrical; exumbrella with eight longitudinal tracks of nematocysts; marginal tentacles two or four. TYPE SPECIES Tubularia dumortierii van Beneden, 1844b, by subsequent designation by Mayer (1910). REMARKS The genus Ectopleura was established by L. Agassiz (1862) to accommodate Tubularia dumortierii van Beneden, 1844b, and four other nominal species, including corynids as well as tubulariids. Agassiz did not designate a type species for the genus, but E. dumortierii was later so designated by Mayer (1910). hydranth. Perisarc straw-coloured, moderately thick ba sally, tapering to thin distally, terminating just below hy dranth base. Hydranth vasiform, up to 1.8 mm high, 1.0 mm wide, bearing two whorls of tentacles. Aboral tentacles filiform, numbering 17 to 22 in mature hydranths, reaching 2 mm long; oral tentacles capitate, numbering 15 to 20 in mature hydranths, 0.3 mm long. Gonophores free medusae, arising in clusters from short, slender blastostyles on hydranth just distal to aboral ten tacles. Well-developed medusa buds dome-shaped, 2.8 mm high, 2.5 mm wide, with an apical projection; exumbrella with eight meridional tracks of nematocysts; mesoglea thin. Manubrium simple, tubular, reaching two-thirds distance to velar opening. Radial canals four. Tentacle bulbs four; two of these, opposite each other, bearing well-developed, capitate marginal tentacles. Nematocysts— Hydroids: desmonemes (on hydranths, medusa buds) Brinckmann-Voss (1970) obtained medusae referable to Ectopleura from the hydroid of Acharadria larynx Wright, 1863c, type species of Acharadria. The name Achara dria Wright, 1863c, is, therefore, a junior synonym of Ectopleura. Ectopleura pacifica Thornely, 1900 Figs. 41, 42 Ectopleura sp. Fewkes, 1883:85; pi. 1, fig. 11 [medusa]. EctopleurapacificaThorncly, 1900:452;pi. 44, figs. 1,1a. Ectopleura minerva Mayer, 1900b:31; pi. 16, fig. 38; pi. 37, fig. 125 [medusa]. ITubularia pacifica—Borradaile, 1905:838. TYPE LOCALITY Blanche Bay, New Britain, Papua New Guinea. MATERIAL EXAMINED Stream Passage Cave, Harrington Sound, on limestone wall, - 1.5 m, 27 July 1982, one hydroid, 2.3 cm high, with blastostyles, romiz B140. St George's Island, north shore, on mooring chain, —9 m, 5 October 1976, one colony, 2.5 cm high, with well-developed medusa buds, a coll. J. Markham, L. Coen, romiz B164. Stream Passage Cave, Harrington Sound, on rock, - 1 m, 20 June 1983, one hydroid, 3.0 cm high, with medusa buds, romiz B369. DESCRIPTION Hydroids with creeping, branched hydrorhiza invested with thick, straw-coloured, smooth or occasionally wrinkled perisarc, giving rise to upright hydrocauli. Hydrocauli up to 2.5 cm high, 0.7 mm wide, invested with perisarc, widely separated or in small clumps, each with a single terminal Fig. 41. Ectopleurapacifica, romiz B164. Scale for a equals 1 mm; scale for b equals 0.5 mm. a, Hydranth with medusa buds, b, Medusa bud. 53 Fig. 42. Ectopleura pacifica, nematocysts of hydranth and medusa bud. romiz B369. Scales equal 10 pm. a, Desmoneme of hydranth. b, ?Basitrichous isorhiza of hydranth. c, Heterotrichous anisorhiza of medusa bud. d, ?Microbasic mastigophore of medusa bud. e, Small stenotele of hydranth. /, Medium stenotele of hydranth. 4.6-A.9 p.m x 3.3-3.6 p,m; ?basitrichous isorhizas (on hydranths) 8.3-9.1 p,m x 3.7-4.0 p.m; heterotrichous anisorhizas (on medusa buds) 7.3-7.8 p.m x 6.1-6.7 p,m; ?microbasic mastigophores (on medusa buds) 6.8-7.5 p.m X 3.4-3.8 p.m; stenoteles (small) (on hydranths, medusa buds) 6.5-7.3 p,m X 4.9-6.6 p,m; stenoteles (medium) (on hydranths, medusa buds) 9.3-11.9 p.m x 8.0-9.8 p.m. REMARKS Thornely (1900) described Ectopleura pacifica from a hy droid collected off New Britain in the western Pacific. Hydroids from Bermuda correspond in all respects to Thornely's original description of this species. While re cognizing that major zoogeographic barriers separate these Bermudian hydroids from the type locality of E. pacifica (see Ekman, 1953; Briggs, 1974), I nevertheless regard them as conspecific based on morphological evidence. Mayer (1900b) applied the name Ectopleura minerva to medusae collected at the Tortugas, Florida. The same spe cies had earlier been reported from Bermuda by Fewkes 54 (1883), as Ectopleura sp. Neither Mayer nor Fewkes ob served the hydroid of E. minerva, but their descriptions of the medusa are applicable to medusa buds in present material (romiz B164) and to Thornely's (1900) descrip tion of E. pacifica, and the two are regarded here as con specific. The name E. pacifica has priority over E. minerva because Thornely's (1900) account of the former appeared in May, while Mayer's (1900b) report of the latter ap peared in July. Free medusae ascribed to Ectopleura pacifica have been reported in the literature only by Mammen (1963) and Calder (1986), the latter based on Fewkes's (1883) record of Ectopleura sp. from Bermuda. Mammen's identification is believed to be wrong because his medusa bore four equally developed tentacles instead of two, and the oral tentacles of his hydroid were filiform instead of capitate. Notably, however, medusae identified as E. minerva have been reported from several locations in the Indian Ocean and the western Pacific (Nair, 1951; Kramp, 1965, 1968; Sugiura, 1977; Bouillon, 1978a, 1978c). Hydroids of Ectopleura pacifica resemble descriptions of E. larynx (Wright, 1863c). In both species, hydroids are small and "primitively" colonial with capitate oral Brinckmann-Voss (1970) believed that E. larynx and E. minerva were different species based on the shape of the medusa. tentacles, and the medusa buds have two tentacles instead of four. In spite of these similarities, the two are regarded as separate species here because of apparent differences in their morphology. In hydroids of E. larynx, only 2 KNOWN RANGE Bermuda: Castle Harbour, medusa stage (Fewkes, 1883); inshore on floats and offshore on buoy chains (Calder, to 8 oral tentacles and 4 to 16 aboral tentacles have 1986). been observed on the hydranth (Wright, 1863a, 1863c; Brinckmann-Voss, 1970), far fewer than in E. pacifica. Elsewhere: western Atlantic (Mayer, 1900b); Indian Ocean (Jarvis, 1922); western Pacific (Sugiura, 1977). Family Halocordylidae Stechow, 1921a Halocordylidae Stechow, 1921a:249. DIAGNOSIS Colonial, capitate hydroids with creeping hydrorhiza and upright, pinnately branched hydrocaulus bearing branches on upper side only. Perisarc tubular, thick, and firm. Hy dranths terminal, clavate to pear-shaped, each with an aboral whorl of long, filiform or slightly capitate tentacles, an oral whorl of short capitate tentacles, and one or more distinct or indistinct whorls of short capitate tentacles between. Gonophores borne just distal to aboral tentacles, either liberated as short-lived medusae or remaining attached to hydranth as eumedusoids. Medusa thimble-shaped with thin mesoglea; manubrium short; mouth lacking. Radial canals four. Tentacle bulbs four; tentacles rudimentary or absent. Ocelli present or absent. Gonads surrounding manubrium. Pennaria Oken, 1815, is invalid on grounds in addition to those noted by Stechow (1922, 1923a). Oken's (1815) publication has been rejected for nomenclatural purposes by the iczn (Opinion 417) because it did not consistently adhere to the Principle of Binominal Nomenclature [Art. 5a]. Halocordyle Allman, 1872, type genus of the family Halocordylidae, is regarded as congeneric with Eucoryne Leidy, 1855. For reasons noted below (see p. 56), usage of the junior synonym Halocordyle as the name of the genus is maintained in this report. The name Halocordy lidae is retained as the valid name of the family. Ten genera were included in the family Halocordylidae by Stechow (1923a). With the exception of the type genus, all have since been transferred to other families (Mammen, 1963). Hydroids of this family are unusual among the Athecatae in their regularly pinnate colony form (Brinckmann-Voss, 1970; Millard, 1975). Genus Halocordyle Allman, 1872 REMARKS Stechow (1921a) proposed that the name Pennariidae McCrady, 1859a, as applied to this taxon, be replaced for reasons elaborated upon in subsequent publications (Ste chow, 1922, 1923a). In these last two papers, he noted that the type genus Pennaria Oken, 1815, originally in cluded eight species, five of which were thecates and the other three of which were of indeterminable identity. Ste chow noted that the name Pennaria should not have been applied later by Goldfuss (1820) to two species of athecate hydroids, including the familiar Pennaria disticha. He rec ognized Halocordyle Allman, 1872, as the valid name of the genus, and established the name Halocordylidae for the family. This family name has now gained widespread use (e.g., Rees, 1957; Pennycuik, 1959; Vervoort, 1959; Mammen, 1963; Calder, 1971; Cooke, 1975; Millard, 1975; Pennaria auct. [not Pennaria Oken, 1815:93 (invalid name, published in a work rejected for nomenclatural purposes by the iczn, Opinion 417)]. Globiceps Ayres, 1854:193 [invalid junior homonym of Globiceps Le Peletier de Saint-Fargeau and Serville, 1825 (Hemiptera)]. Eucoryne Leidy, 1855:136. Eucoryna van Beneden, 1867:17 [incorrect subsequent spelling]. Halocordyle Allman, 1872:368. Halocordile Wedler and Larson, 1986:69 [incorrect sub sequent spelling]. DIAGNOSIS Hirohito, 1977; Petersen, 1979; Bouillon, 1985; Garcia- With the characters of the family. Corrales and Aguirre, 1985). The family names Penna riidae and Halocordylidae cannot be regarded as synonyms because the nominal genera Pennaria and Halocordyle are TYPE SPECIES not even in the same order. subjective synonym of Pennaria disticha Goldfuss, 1820. Globiceps tiarella Ayres, 1854, by monotypy, a junior 55 REMARKS The invalid name Pennaria Oken, 1815, came into wide spread use for this genus after Goldfuss (1820) referred his familiar species P. disticha to it. As noted above, Pennaria originally included five thecate species and three other species of indeterminable identity (Stechow, 1922, 1923a). The binomen "Sertolara pennara" had earlier been ap plied by Cavolini (1785) to the hydroid referred to by Goldfuss (1820) as Pennaria disticha. Cavolini obviously believed his material was conspecific with Sertularia pen naria Linnaeus, 1758, but the latter is now known to be a species of thecate hydroid (see p. 57). Sertolara as used by Cavolini is an incorrect subsequent spelling of Sertu laria Linnaeus, 1758. As such, it is an unavailable name [Art. 33c] and cannot be used as the name of this genus. Ayres (1854) founded the name Globiceps tiarella for an American hydroid now generally regarded as conspe cific with Pennaria disticha. The name Globiceps Ayres, 1854, cannot replace Pennariaauct. because it is an invalid junior homonym of GlobicepsLe Peletier de Saint-Fargeau and Serville, 1825, a name applied to a hemipteran (L. Agassiz, 1862; Allman, 1872). One year after the publication of the name Globiceps tiarella, Leidy (1855) described the same species under the name Eucoryneelegans. L. Agassiz (1862) and Allman (1872) believed that the name Eucoryne Leidy, 1855, was preoccupied by Eucorynus Schoenherr, 1823, a coleopteran. Allman (1872) proposed Halocordyle as a substitute name for Globiceps Ayres, 1854, and Eucoryne Leidy, 1855. However, Eucoryne Leidy, 1855, and Eucorynus Schoenherr, 1823, are not homonyms [Art. 56b]. Euco ryne, seldom used as a valid name since it was published (e.g., seeBedot, 1910, 1912, 1916, 1918, 1925), is none theless available as a senior synonym of Halocordyle, now in widespread use (e.g., see the 11 works cited above to demonstrate widespread use of the family name Halocor dylidae). In the interests of nomenclatural stability, a sub mission will be made to the iczn requesting suppression of the genus name Eucoryne Leidy, 1855, and placement of the name Halocordyle Allman, 1872, on the Official List of Generic Names in Zoology. Halocordyle disticha (Goldfuss, 1820) Figs. 43-45 Sertolara pennara—Cavolini, 1785:134; pi. 5, figs. 1-6 [incorrect subsequent spelling] [not Sertularia pennaria Linnaeus, 1758]. Sertularia pennaria—Gmelin, 1790:3856 [not Sertularia pennaria Linnaeus, 1758]. Aglatophenia pinnaria—Costa, 1839:185 [incorrect sub sequent spelling]. Pennaria caulini delle Chiaje, 1841:145. Anisocalyx pinnarium—Costa, 1842:18. Globiceps tiarella Ayres, 1854:193. Eucoryne elegans Leidy, 1855:136; pi. 10, figs. 1-5. lEuphysaglobator Leuckart, 1856:28; pi. 2, fig. 4 [medusa]. Pennaria tiarella—McCrady, 1859a: 153. Pennaria gibbosa L. Agassiz, 1860, pi. 15, figs. 1,2. Pennaria distycha—L. Agassiz, 1862:344 [incorrect sub sequent spelling]. Eucoryna elegans—van Beneden, 1867:17 [incorrect sub sequent spelling]. Pennaria cavolini—van Beneden, 1867:50 [incorrect sub sequent spelling]. Halocordyle tiarella—Allman, 1872:369. Pennaria symmetrica Clarke, 1879:240; pi. 1, figs. 2,3. Globiceps globator—Haeckel, 1879:40 [medusa]. Pennaria inornata Brooks, 1883a: 144. Pennaria australis Bale, 1884:45. Pennaria rosea von Lendenfeld, 1885b:594; pi. 24, figs. 40^12. Pennaria adamsia von Lendenfeld, 1885b:595; pi. 25, figs. 45-48; pi. 26, fig. 49. Pennaria pennaria—Marktanner-Turneretscher, 1890:201. Halocordyle australis Bale, 1894:94. Pennaria cavolina—Spencer, 1892:13 [incorrect subse quent spelling]. Halocordyle cooperi Warren, 1906a:73; pi. 9. Pennaria pacifica Clarke, 1907:6; pi. 1, figs. 1-6. Pennaria australis var. cooperi—Warren, 1908:282. Pennaria australis—Warren, 1908:283. Pennaria disticha var. australis—Ritchie, 1910a:806. Pennaria wilsoni Bale, 1913:116. Halocordyle disticha—Stechow, 1923a:48. Halocordyle australis—Stechow, 1923a:48. Halocordyle wilsoni—Stechow, 1923a:48. Corydendrium splendidum Boone, 1938:33; pi. 4. Halocordyle disticha var. australis—Vervoort, 1941:192. Halocordyle fragilis Vannucci, 1951:76; pi. 1, figs. 2,3. Halocordyle pennaria var. australis—Mammen, 1963:54; figs. 22-24. Pennaria "americana" Garcia-Corrales and Aguirre, 1985:86 [nomen nudum]. Pennaria {Halocordyle) tiarella—Garcia-Corrales and Aguirre, 1985:86. Pennaria "europea" Garcia-Corrales and Aguirre, 1985:86 [nomen nudum]. Pennaria {Halocordyle) disticha—Garcia-Corrales and Aguirre, 1985:86. Pennaria disticha Goldfuss, 1820:89. Pennaria symetrica—Garcia-Corrales and Aguirre, 1985:86 [incorrect subsequent spelling]. Plumularia pennaria—de Blainville, 1830:442. Halocordile disticha—Wedler and Larson, 1986:69 [in Pennaria cavolinii Ehrenberg, 1834:297. correct subsequent spelling]. 56 TYPE LOCALITY Gulf of Naples, Italy. MATERIAL EXAMINED Castle Harbour near Tucker's Town, on patch reef, -7 m, 3 August 1982, two colonies, 6.7 and 10.2 cm high, with well-developed medusa buds, romiz B131. Harts Inlet, on underside of flat rocks, —3 m, 2 August 1982, one colony, 2.2 cm high, without gonophores, romiz B143. Whalebone Bay, on ledges at entrance, — 1 m, 4 March 1982, two colonies, 2.6 and 3.1 cm high, without gono phores, romiz B167. DESCRIPTION Colonies erect, arising from a creeping, branching hy drorhiza; growth monopodial with terminal hydranths. Hydrocaulus monosiphonic, reaching about 0.4 mm wide, zigzag to nearly straight, annulated basally, divided at more or less regular intervals by one or more well-developed annulations; internodes 0.6-4.0 mm long, each typically supporting a branch distally. Perisarc thick, black through brown to deep horn-coloured basally, becoming progres sively thinner and lighter coloured distally, terminating abruptly below hydranths of hydrocaulus, branches, and ramuli. Branches up to 27 mm long, annulated basally, given off alternately from opposite sides of hydrocaulus, reduced. Ocelli lacking. Gonads surrounding manubrium, filling subumbrellar cavity. Nematocysts— Medusa buds: heterotrichous microbasic euryteles 7.58.3 |xm x 3.1-4.0 [im. REMARKS There has been some question as to whether Sertularia pennaria Linnaeus, 1758, may belong to the same species as this hydroid (e.g., see Cavolini, 1785; L. Agassiz, 1862; Marktanner-Turneretscher, 1890; Bedot, 1901; Mayer, 1910). Bedot (1912) reported that the Linnaean species is an Aglaophenia Lamouroux, 1812, rather than an athecate hydroid. Examination of the Linnaean type has confirmed Bedot's (1912) conclusion (P. F. S. Cornelius, pers. comm.) that it is an aglaopheniid. The oldest available specific name for this well-known species is that of Goldfuss (1820), who applied the name Pennaria disticha to Cavolini's (1785) "Sertolara pennara." Cavolini's hydroid was identical with what is widely referred to today as Halocordyle disticha (Goldfuss, 1820) and Pennaria tiarella (Ayres, 1854), rather than Lin naeus ' s (1758) Sertularia pennaria. curved outwards, divided into internodes; these internodes 1.5-4.0 mm long, marked by distinct to rather faint an nulations proximally and distally; each internode giving rise to a ramulus from both its upper surface and its distal end. Ramuli unbranched, annulated basally or throughout entire length, each terminating in a hydranth. Hydranths clavate to pear-shaped, up to 1.7 mm long, 0.3 mm wide; with a whorl of about 10 to 16 long, filiform or faintly knobbed tentacles aborally; a varied number of short, cap itate tentacles in one or more regular or irregular verticils medially; and a whorl of about four to six short, capitate tentacles orally. Hypostome dome-shaped. Nematocysts— Hydroids: desmonemes 4.5-5.3 jxm x 3.3-3.8 (xm; basitrichous haplonemes 5.7-9.4 |xm x 2.5-3.6 |xm; het erotrichous microbasic euryteles 10.8-13.6 (xm X 6.16.8 |xm; stenoteles (very small) 5.7-6.6 |xm x 4.3-4.6 jim; stenoteles (small) 7.4-7.8 |xm x 5.5-5.7 jxm; stenoteles (medium) 14.2-17.7 |xm x 10.0-12.0 jxm; stenoteles >*^Uuz^ *< (large) 28.1-39.8 \x,m x 16.3-20.2 jxm. Gonophores free but degenerate medusae, those of a given colony either all male or all female, arising on short pedicels just distal to aboraltentacles. Well-developed me dusa buds elongate-oval, about 0.9 mm high, 0.6 mm wide, covered with an ectodermal sheath; mesoglea thin; manu brium simple, tubular, reaching nearly to velar opening. Radial canals four. Tentacle bulbs four, reduced; tentacles Fig. 43. Halocordyle disticha, colony form. Natural size, a, Colony from exposed area, romiz B167. b, Colony from shel tered area, romiz B131. 57 Fig. 44. Halocordyle disticha, romiz B131. Scale for a equals 1 mm; scale for b equals 0.5 mm. a, Hydrocaulus and hydranths. b, Medusa bud. The synonymy list provided above has been taken largely from Bedot (1901, 1905, 1910, 1912, 1916, 1918, 1925), Mayer (1910), Millard (1975), Cooke (1977), Hirohito (1977), and Garcia-Corrales and Aguirre (1985). The last two papersincluded extensive discussion of the synonymy of Halocordyle disticha, and the matter will not be repeated here. In neither paper, however, was it noted that Bale (1894) had described Halocordyle australis as a different species from Pennaria australisBale, 1884. Both nominal species are included here in the synonymy of Halocordyle disticha. Colonies of Halocordyle disticha displayed considerable variation in form from one location to another in Ber- mudian waters (Fig. 35). Specimens from sheltered waters near the base of a patch reef in Castle Harbour (romiz B131) were large and gracile. Internodes of both hydro caulus and branches were long and slender, and the branches and ramuli were elongate. Specimens from wave-swept 58 ledges at the entrance of Whalebone Bay (romiz B167), as well as those subjected to strong tidal currents near the bridge at Flatts Inlet (romiz B143), were small and com pact. Internodes of the hydrocaulus and hydrocladia in these colonies were thicker and much shorter, and the branches and ramuli were relatively stunted. The degree of annulation on stem and branches, a variable character in this species (Millard, 1975), did not appear to differ significantly from one colony form to another in Bermudian material. Hydroids of Halocordyle disticha are inactive during colder months in temperate areas (Hargitt, 1900; McDougall, 1943; Brinckmann-Voss, 1970; Calder, 1971). In Bermuda, a few colonies with active hydranths were found in winter even at water temperatures as low as 17°C, but the species was observed in far greater abundance during the warmer seasons. Medusae of this species are short-lived and are liberated / i Fig. 45. Halocordyle disticha, nematocysts of hydranth and medusa bud. Scales equal 10 pm. a, Des moneme of hydranth, romiz B143. b, Basitrichous haploneme of hydranth, romiz B143. c, Heterotrichous microbasic eurytele of hydranth, romiz B143. d, Heterotrichous microbasic eurytele of medusa bud, romiz B131. e, Very small stenotele of hydranth, romiz B143. /, Small stenotele of hydranth, romiz B143. g, Medium stenotele of hydranth, romiz B167. h, Large stenotele of hydranth, romiz B167. 59 in the eveningin Bermuda (Weill, 1937a), as noted in spe cimens from ChesapeakeBay (Calder, 1971).BrinckmannVoss (1970) and others noted that medusa buds of Halo cordyle disticha often shed their gametes while attached to the hydroid and are sometimes not released. The phe nomenon of egg release by attached medusa buds in this species was described by Cavolini (1785), as noted by times previously (Verrill, 1900; Congdon, 1907; Bennitt, 1922; Calder, 1986). It has also been utilized in mor phological and developmental studies conducted at the Bermuda Biological Station (Weill, 1937a;Cowden, 1964, 1965a, 1965b; Summers and Haynes, 1969; Summers, 1970, 1972b; Lesh-Laurie, 1976; Clark and Cook, 1986). Cornelius (1977). According to Weill (1934), the cnidome of hydroids of this species included desmonemes, stenoteles, microbasic mastigophores, and heteronemes of an undetermined cat egory. Bouillon (1985) has been followed here inregarding KNOWN RANGE the last two of these as microbasic euryteles and basitri Bermuda: no specific locality given (Verrill, 1900; Cong don, 1907; Weill, 1937a); Hamilton Harbour and Great Sound (Bennitt, 1922); near the Bermuda Biological Sta tion (Cowden, 1964, 1965a, 1965b); Flatts Inlet (Summers chous haplonemes. The nematocyst complement of me and Haynes, 1969; Summers, 1970, 1972b; Lesh-Laurie, dusa buds in Bermudian material typically consisted of 1976); inshore in shallow water and offshore on buoy microbasic euryteles, although a few stenoteles and hap chains (Calder, 1986). Elsewhere: western Atlantic (Fraser, 1944); eastern At lonemes were observed in some specimens. Euryteles of medusabuds almost alwayslackedthe spherical inclusions found in those of the hydroid. This species has been reported from Bermuda several lantic (Brinckmann-Voss, 1970); Indian Ocean (Millard, 1975); western Pacific (Yamada, 1959); eastern Pacific (Fraser, 1948). Family Sphaerocorynidae Prevot, 1959 Sphaerocorynidae Prevot, 1959:108. nemes. Petersen believed that this family was more closely related to the Moerisiidae Poche, 1914, than to the Cory- DIAGNOSIS Hydroids colonial, with creeping stolons and upright, un branched or sparingly branched hydrocauli with terminal hydranths. Perisarc moderately thin, reaching to hydranth base. Hydranths pyriform, with bulbous base and conical hypostome. Tentacles capitate, scattered in an irregular nidae as suggested by Prevot (1959). Genus Sphaerocoryne Pictet, 1893 Sphaerocoryne Pictet, 1893:9. band about bulbous hydranth base. Gonophores free medusae, arising from hydranth on short pedicels among or distal to tentacles. Medusae bellshaped, with or without exumbrellar nematocyst tracks; manubrium tubularin juveniles,cruciform in cross-section in adults, not extending beyondvelaropening; mouth sim ple. Radial canals four. Marginal tentacles two or four, capitate. Ocelli present. Gonads on perradii ofmanubrium. REMARKS Sphaerocoryne Pictet, 1893, and Linvillea Mayer, 1910, DIAGNOSIS Hydroids with the characters of the family. Medusa buds arising in clusters on hydranth just distal to tentacles. Medusae with scattered exumbrellar nema tocysts; manubrium of adult somewhat cruciform in crosssection. Marginal tentacles four, capitate or moniliform, equally developed at liberation. TYPE SPECIES Sphaerocoryne bedoti Pictet, 1893, by monotypy. the latter a genus formerly included in the family Corynidae Johnston, 1836, were grouped by Petersen (1979) inthe Sphaerocorynidae Prevot, 1959. Petersen noted that these genera differ inseveral respects from typical corynid hydrozoans. Tentacles in the hydroid stage are restricted to the expanded basal region of the hydranth, and medusa buds arise among or distal to the tentacles. Gonads in the medusa do not surround the manubrium but are located perradially on it. Finally, the nematocyst complement of both hydroid and medusa includes stenoteles and desmo 60 REMARKS Thehydroids ofSphaerocoryne Pictet, 1893, and Linvillea Mayer, 1910, resemble one another morphologically. Ya mada and Konno (1973) reported finding only minor dif ferences in a comparison of Sphaerocoryne multitentaculata (Warren, 1908) (= S. bedoti Pictet, 1893) from Japan and Linvillea agassizii (McCrady, 1859a) from Virginia, United States. Medusa buds in L. agassizii arise among the ten tacles on the hydranth, while they occur distal to the ten- tacles in S. bedoti. Differences are more pronounced in medusae of the two species. In L. agassizii, medusae have eight exumbrellar nematocyst tracks, the manubrium is distinctly cruciform in cross-section in the adult, and two of the four marginal tentacles are undeveloped in young medusae. In S. bedoti, medusae have scattered exum brellar nematocysts, the manubrium is less distinctly cru ciform in cross-section than in L. agassizii, and the four marginal tentacles are equally developed at all stages of growth. Adult medusae of Sphaerocoryne peterseni Bouillon, 1984a, have adaxial batteries of nematocysts along each of four marginal tentacles, and an abaxial ocellus is present on each tentacle bulb. The hydroid stage of this species Gonophores developing in small clusters, borne on hy dranth just distal to tentacles. Well-developed medusa buds and newly liberated medusae thimble-shaped, 450 jxm high, 350 |xm wide in alcohol-preserved specimens; exumbrella with scattered nematocysts; mesoglea thin; manubrium simple, conical, reaching about halfway to velar opening. Radial canals four. Tentacle bulbs four, equally developed; marginal tentacles scarcely if at all developed. Ocelli and gonads undeveloped. Nematocysts— Medusae: desmonemes 8.3-9.5 |mm x 3.8^1.5 [xm; basitrichous haplonemes 9.2-10.8 |xm X 7.9-9.4 |xm; stenoteles (small) 9.1-9.7 (xm x 6.5-7.5 jxm; stenoteles (large) 12.4-13.6 [Jim x 8.8-10.5 |xm. is unknown. REMARKS Sphaerocoryne bedoti Pictet, 1893 Figs. 46, 47 Sphaerocoryne bedoti Pictet, 1893:10; pi. 1, figs. 5,6. Clavatella multitentaculata Warren, 1908:278; pi. 45, figs. 7-9. Three nominal species of the genus Sphaerocoryne Pictet, 1893, S. bedoti Pictet, 1893, S. multitentaculata (Warren, 1908), and S. peterseni Bouillon, 1984a, have been de scribed in the literature. Only the medusa stage of S. pe terseni is known. Hydroids of the other two, found on sponges, are virtually inseparable based on currently avail able descriptions. Yamada and Konno (1973) preferred to Sphaerocoryne multitentaculata—Stechow, 192la:248. Eleutheria multitentaculata—Bedot, 1925:179. Sphaerocoryne sp.—Gravely, 1927:8; pi. 2, fig. 3. Coryne (?) multitentaculata—Pennycuik, 1959:158. TYPE LOCALITY Ambon, Moluccas, Indonesia. MATERIAL EXAMINED Major's Bay, Harrington Sound, on sponge, -2 m, 13 September 1977, several hydranths with medusa buds, and several newly liberated medusae, romiz B132. Major's Bay, Harrington Sound, on sponge, —2 m, 13 September 1977, several hydranths with medusa buds, romiz B177. Walsingham Pond area, cave entrance, on underside of rock, - 1 m, 29 June 1983, two hydranths, 1 cm high, without gonophores, romiz B331. DESCRIPTION Hydrorhiza embedded in sponge. Hydrocaulus monosiphonic, unbranched, about 1 cm high, 120 |xm wide, sup porting a terminal hydranth. Perisarc rather thin, hyaline, smooth or with a few wrinkles; distinct annulations not apparent. Hydranths pyriform, 0.8 mm high, 0.5 mm wide, with an elongate hypostome. Tentacles capitate, of varying length, about 30 in number, scattered in a narrow band around bulbous hydranth base. Nematocysts— Hydroids: desmonemes 12.4-13.2 |mm x 5.6-5.9 |xm; stenoteles (small) 11.2-12.2 |xm x 7.7-9.2 jxm; steno teles (large) 24.4-26.3 |xm x 16.8-18.0 |xm. Fig. 46. Sphaerocoryne bedoti. Scales equal 0.5 mm. a, Part of hydrocaulus, and hydranth, romiz B177. b, Newly liberated medusa, romiz B132. 61 1 L ,1, IP' g ! Fig. 47. Sphaerocoryne bedoti, nematocysts of hydranth and medusa bud. Scales equal 10 |xm. a, Des moneme of hydranth, romiz B331. b, Desmoneme of medusa bud, romiz B132. c, Basitrichous haploneme of medusa bud, romiz B132. d. Small stenotele of hydranth, romiz B331. e. Small stenotele of medusa bud, romiz B132. /, Large stenotele of hydranth, romiz B331. g, Large stenotele of medusa bud, romiz B132. recognize S. multitentaculata as a valid species because of a general lack of information on living material from areas other than Japan, but Mammen (1963) and Millard (1975) have been followed here in regarding it as conspe cific with S. bedoti. Yamada and Konno (1973) described the hydroid of this species and outlined the development of the medusa from liberation to maturity. They found that the four marginal tentacles developed concurrently in the medusa, and that an abaxial ocellus appeared on each tentacle bulb about 8 to 10 days after liberation. Medusae raised on Artemia in the laboratory were mature two weeks after release from the hydroid. Male medusae were ovoid, 2.5-3.5 mm high, 2.0-3.0 mm wide, and possessed a manubrium two-thirds to four-fifths the length of the subumbrellar cavity. Female medusae were conical, 3.0^1.5 mm high, 2.0-3.0 mm wide, and bore a manubrium about two-thirds the length of the subumbrellar cavity. These laboratory-reared spec imens have not been identified with any previously named species of medusa. Bermudian hydroids of Sphaerocoryne bedoti tended to 62 be slightly smaller than those described from Indonesia (Pictet, 1893), South Africa (Warren, 1908), India (Mam men, 1963), and Japan (Yamada and Konno, 1973). How ever, the range of hydranth size and shape in this species is reflected in both living and preserved material examined by Mammen (1963). Tentacle number is likely to vary somewhat with hydranth size, and tentacle arrangement appears to differ depending upon the degree of expansion or contraction of the hydranth. Newly liberated medusae of this species from Bermuda, measured after five years in preservative, were decidedly smaller than those de scribed by Yamada and Konno (1973). Part of this dif ference may be attributable to contraction of my specimens on preservation, and their subsequent shrinkage in the al cohol preservative. KNOWN RANGE Bermuda: inshore, on sponges (Calder, 1986). Elsewhere: western Atlantic (Mergner and Wedler, 1977); Indian Ocean (Millard and Bouillon, 1974); western Pa cific (Yamada and Konno, 1973). Family Corynidae Johnston, 1836 Corynidae Johnston, 1836:107. Codonium Haeckel, 1879, is congeneric with Sarsia (Mayer, Sarsiadae Forbes, 1848:54. 1910). Syncorynidae Allman, 1872:274. The corynid genera Coryne, Sarsia, and Dipurena were recently redefined by Petersen (1979). In his classification, tentacle arrangement and gonophore location on the hy dranth of the hydroid, as well as manubrium length, extent Codonidae Haeckel, 1879:9. DIAGNOSIS Hydroid colonies stolonal or erect, with creeping hydro rhiza and upright, unbranched or irregularly branched hy drocaulus. Perisarc firm. Hydranths variable in shape but often clavate, with an oral whorl of capitate tentacles, frequently with scattered or verticillate capitate tentacles extending proximally over hydranth, and with or without an aboral whorl of reduced filiform tentacles. Hypostome dome-shaped to conical. Gonophores fixed sporosacs or free medusae, borne on hydranth proximal to or among capitate tentacles. Me dusae, when present, bell-shaped, without exumbrellar nematocyst tracks; manubrium tubular, short and enclosed within subumbrellar cavity or long and extending beyond velar opening; mouth simple, circular. Radial canals four. Marginal tentacles four. Ocelli present. Gonads surround ing manubrium, arranged in one or more rings. of the manubrium functioning as a "stomach," and position of the gonad on the manubrium in the medusa, were used to distinguish genera. It is possible, following Petersen's diagnoses, to refer corynid hydroids to genus in the absence of information on type of gonophore pro duced. It remains to be determined whether this classifi cation is more "natural" than the one adopted by most other current authors. The filiform tentacles of corynid hydroids, which may be present or absent even within a given species, were regarded as evolutionary vestiges by Rees (1957). Edwards and Harvey (1983) disputed this, suggesting instead that they are highly specialized sense organs used for detection of prey as indicated by Tardent and Stossel (1971), Stossel and Tardent (1971), and Tardent and Schmid (1972). Genus Coryne Gaertner, 1774 REMARKS The scope of the family Corynidae Johnston, 1836, adopted here encompasses only the genera Coryne Gaertner, 1774, Sarsia Lesson, 1843, and Dipurena McCrady, 1859a. The poorly known genus Dicodonium Haeckel, 1879, has gen erally been regarded as a corynid, but apparently is not (Brinckmann-Voss, 1970). Accordingly, the name of the tribe Dicodoniini, established for this genus by Cockerell (1911), is excluded from the synonymy of the Corynidae given above. Petersen (1979) has been followed in refer ring Sphaerocoryne Pictet, 1893, and Linvillea Mayer, 1910, to the Sphaerocorynidae Prevot, 1959, as well as Dicyclocoryne Annandale, 1915, and Bicorona Millard, 1966, to the Dicyclocorynidae Petersen, 1979. Earlier, Rees (1957) removed Hydrocoryne Stechow, 1907, to a new family, the Hydrocorynidae. Among other nominal genera frequently encountered in discussions of corynids, Syncoryna Ehrenberg, 1834, and Staurocoryne Rotch, 1872, are regarded as congeneric with Coryne (e.g., see Millard, 1975), and Stauridiosarsia Mayer, 1910, is considered to be congeneric with Sarsia (e.g., see Brinckmann-Voss, Coryne Gaertner, 1774:40. Fistularia O. F. Muller, 1776a:254 [invalid junior hom onym of Fistularia Linnaeus, 1758 (Pisces)]. Fistulana O. F. Muller, 1776a:282. Capsularia Modeer, 1793:256 [nomen nudum]. Capsularia Cuvier, 1798:665. Corine Cuvier, 1798:656 [incorrect subsequent spelling]. Coryna Bosc, 1802:238 [incorrect subsequent spelling]. Corina Schweigger, 1820:409 [incorrect subsequent spelling]. Stipula M. Sars, 1829:4. Syncoryna Ehrenberg, 1834:294. Hermia Johnston, 1838:111. Syncoryne Steenstrup, 1842:10 [incorrect subsequent spelling]. Halybotrys de Filippi, 1866:383. lActinogonium Allman, 1871:95 [invalidjunior homonym of Actinogonium Schomburgk, 1847 (?Protozoa)]. Syncorine Spagnolini, 1871:211 [incorrect subsequent 1970; Millard, 1975; Arai and Brinckmann-Voss, 1980; spelling]. Staurocoryne Rotch, 1872:126. Bouillon, 1985). The systematic affinities of Sarsiella Hartlaub, 1907, and Bibrachium Stechow, 1919, are un Halobotrys Cams, 1885:2 [incorrect subsequentspelling]. Eucoryne Broch, 1909:138 [invalid junior homonym of clear, although both were included in the Corynidae by Bouillon (1985). The family-group name Codonidae Hae ckel, 1879 (correctly spelled Codoniidae) is included as a synonym of the name Corynidae because its type genus, Eucoryne Leidy, 1855 (Hydrozoa)]. Halybothrys Bedot, 1910:310 [incorrect subsequent spelling]. lActigia Stechow, 1921a:248. 63 DIAGNOSIS Corynid hydroids with stolonal or erect and branching colonies. Hydranths with several more or less distinct whorls of capitate tentacles distally, those of one whorl often alternating with those of adjacent whorls; hydranths oc casionally with a whorl of reduced filiform tentacles proximally. Gonophores fixed sporosacs, arising on hydranth either in axils of capitate tentacles or just proximal to capitate tentacles. TYPE SPECIES Coryne pusilla Gaertner, 1774, by monotypy. Gaertner, 1774, a hydroid that was subsequently shown by Hincks (1868) and Allman (1872) to be a different species. Hincks (1868) proposed the name Coryne vanbenedenii for this hydroid. Allman (1871, 1872) removed it to a new nominal genus, Actinogonium, chiefly because an actinula larva was reported in its life cycle, and he renamed the species A. pusillum. Stechow (1921a) noted that Actinogonium Allman, 1871, is a junior homonym of Actinogonium Schomburgk, 1847, a protozoan, and pro posed Actigia as a replacement name. Brinckmann-Voss (1970) listed the species from the Mediterranean, based on a questionable record by Schneider (1898a), under the genus Coryne. cles either present or absent; gonophores arising either in the axils of the tentacles or in a whorl replacing the lowest verticil of capitate tentacles; and medusae, when present, with a short manubrium completely or almost completely surrounded by the gonad. Although not followed in this report, Petersen's ideas concerning the scope of the genus Stechow (1923a) has been followed in regarding Stipula M. Sars, 1829, Syncoryna Ehrenberg, 1834, Hermia John ston, 1838, and Halybotrys de Filippi, 1866, along with Fistularia O. F. Muller, 1776a, Fistulana, Capsularia Cuvier, 1798, and Eucoryne Broch, 1909, as congeneric with Coryne. Stechow also regarded Acrochordium Meyen, 1834, as a synonym of Coryne, but it belongs in the Zancleidae Russell, 1953 (see p. 69). Staurocoryne Rotch, 1872, has been regarded as congeneric with Coryne, fol lowing Millard (1975), Edwards and Harvey (1983), and Bouillon (1985). Under the definition of this genus pro posed by Petersen (1979), Stauridia Wright, 1858b [not Stauridia Forbes, 1848], Stauridiosarsia Mayer, 1910, and Perinema Stechow, 1921c, would be congeneric with Cor yne. They are regarded here as congeneric with Sarsia merit further evaluation. Lesson, 1843. REMARKS The diagnosis of Coryne Gaertner, 1774, given above is similar to that adopted by Brinckmann-Voss (1970), Mil lard (1975), Bouillon (1985), and others. It differs from the definition of the genus proposed by Petersen (1979), who included in Coryne those species of corynids having a stolonal or erect and branching colony form; capitate tentacles alternating in successive whorls; filiform tenta The name of this genus was originally spelled Coryne by Gaertner (1774); Corine Cuvier, 1798, Coryna Bosc, 1802, and Corina Schweigger, 1820, are incorrect sub sequent spellings. Broch (1909) founded the subgenus Eucoryne within the genus, but that name is an invalid junior homonym of Eucoryne Leidy, 1855. Although the name Fistularia had been applied to a genus of fishes by Linnaeus (1758), O. F. Muller (1776a) applied the same name to a genus of hydroids. Later in the same publication, Muller employed the name Fistulana as a replacement name for Fistularia O. F. Muller, 1776a. One of the hydroids included in Fistulana by Muller was Tubularia ramosa Linnaeus, 1758, the type species of Eudendrium Ehrenberg, 1834. Had this species been des ignated type species of Muller's nominal genus, the vir tually forgotten name Fistulana would be a senior objective synonym of the widely used name Eudendrium. To my knowledge, no type species has ever been designated for Fistulana. Accordingly, Tubularia muscoides Linnaeus, 1761, another species included by Muller (1776a) in his genus but now included in Coryne, is hereby designated as type species of Fistulana. The name Fistulana thus becomes a junior subjective synonym of Coryne, and its potential nomenclatural threat to Eudendrium is removed. Van Beneden (1844b) identified as Syncoryna pusilla 64 Coryne sargassicola, sp. nov. Figs. 48, 49 Syncoryne mirabilis—Fraser, 1912:347; fig. 3 [not Sarsia mirabilis L. Agassiz, 1849 (= Sarsia tubulosa (M. Sars, 1835))]. Syncoryne sp. Burkenroad in Parr, 1939:23. Syncoryne {Sarsia) mirabilis—Morris and Mogelberg, 1973:10; fig. 2. MATERIAL EXAMINED Holotype: Natural Arches Beach, washed ashore on pe lagic Sargassum, 8 March 1982, one colony, 2.7 mm high, without gonophores, romiz B159. Paratypes: Whalebone Bay, on pelagic Sargassum, 6 September 1977, one col ony, 2.7 mm high, without gonophores, romiz B156; Nat ural Arches Beach, washed ashore on pelagic Sargassum, 8 March 1982, one colony, 2.5 mm high, without gono phores, romiz B302; Gulf Stream, east-southeast New York City, 40°02' N, 70°50' W, on pelagic Sargassum, 1 October 1969, one colony, 4.5 mm high, with gono phores, coll. M. Fine on R/V Eastward, romiz B490. Fig. 48. Coryne sargassicola, sp. nov., part of colony with hydrorhiza, hydrocaulus, and hydranth. romiz B302. Scale equals 0.5 mm. Fig. 49. Coryne sargassicola, sp. nov., nematocysts of hydranth, romiz B302. Scale equals 10 um. a, Small stenotele. b. Large stenotele. DESCRIPTION Hydroid colonies typically stolonal, occasionally with an irregular branch; pedicels arising from a creeping hydro rhiza. Pedicels up to 3 mm long, 0.12 mm wide, sup porting terminal hydranth. Perisarc of moderate thickness basally, thinning out distally, smooth or with a few wrin kles but not annulated, terminating below tentacles on hydranth. Hydranth elongate-oval, reaching 1 mm long, 0.3 mm wide; tentacles arranged in whorls of four to six each, with those of one whorl alternating with those of adjacent whorls; oral tentacles capitate, with terminal knobs 65-85 p,m in diameter; aboral tentacles capitate, in one to four whorls, with terminal knobs of lowest whorl 55- 60 p,m in diameter; basal whorl of reduced filiform ten tacles present or absent. Hypostome conical, moderately elongate. Nematocysts— Hydroids: stenoteles (small) 11.7-12.2 pim X 6.7- 7.3 (xm; stenoteles (large) 19.9-20.8 (xm X 14.1-15.1 (xm. Gonophores presumably fixed sporosacs, arising from hydranth in axils of proximal tentacles. REMARKS This species has been referred to Coryne Gaertner, 1774, rather than to Sarsia Lesson, 1843, or Dipurena McCrady, 1859a, because gonophores in paratype material (romiz B490) appear to be fixed sporosacs arising from the axils of the tentacles. It also corresponds with the genus Coryne as defined by Petersen (1979). Coryne sargassicola, sp. nov., is regarded as conspe cific with the corynids reported earlier from Sargassum by Fraser (1912) and Morris and Mogelberg (1973) as Syn coryne mirabilis (L. Agassiz, 1849), and by Burkenroad in Parr (1939) as Syncoryne sp. Following Kramp (1928), S. mirabilis is now generally regarded as a synonym of Sarsia tubulosa (M. Sars, 1835). It is improbable that the boreal S. tubulosa ranges into the warm waters where pelagic Sargassum is normally found, and where the pres ent species occurs. Hydroids of Sarsia tubulosa and Coryne sargassicola are similar in colony form and in having smooth perisarc. However, filiform tentacles, observed on some hydranths of C. sargassicola (romiz B156, romiz B302), are lacking in S. tubulosa (e.g., see Edwards, 1978, 1983; Brinckmann- Voss, 1985). Moreover, the hydroid of S. tubulosa pro duces free medusae instead of fixed sporosacs. It is unclear whether Fraser (1912) actually observed liberation of gon ophores in the hydroid he identified as Syncoryne mira bilis, or whether he simply assumed they would be released. 65 He noted that the "medusae" of his specimens were sex ually mature before liberation. L. Agassiz (1849), Mayer (1910), and Berrill (1953) reported that free medusae were produced by hydroids of Sarsia tubulosa at certain times of year, and fixed eume- pintneriSchneider, 1898a, and of C.filiformis (Rees, 1936), but are smaller and lack annulations on the perisarc. They dusoids at other times. Edwards (1978) found no such C. sargassicola. variation in gonophore type in cultures of S. tubulosa main tained in the laboratory. Hydroids of that species cultured at various temperatures by Edwards all released their me dusae, and none of the medusae bore gonads at liberation. Likewise, Edwards demonstrated that temperature varia tion had no influence on the type of gonophore produced by either Sarsia occulta Edwards, 1978, or Sarsia lovenii (M. Sars, 1846). In the former species, gonads were par tially developed in the medusa prior to its liberation. In S. lovenii, gonophores ripened on the hydranth and were Observations on live material are needed to clarify the life cycle of this hydrozoan, and to confirm its generic not released as free medusae. Edwards concluded that L. Agassiz, Mayer, and Berrill had confused more than one valid taxon for a single species. Hydroids of Coryne sargassicola resemble those of C. also resemble the cold-water C. hincksii Bonnevie, 1898, but dimensions of the latter given by Rees (1956b) and Calder (1972) indicate that it is a more robust species than identity. ETYMOLOGY The specific name is derived from a combination of the generic name Sargassum plus the Latin suffix cola, a dweller, in reference to the algal substrate of this species. KNOWN RANGE Bermuda: on pelagic Sargassum (Burkenroad in Parr, 1939, as Syncoryne sp.). Elsewhere: western Atlantic (Fraser, 1912, as Syncoryne mirabilis). Family Cladonematidae Gegenbaur, 1857 Cladonemiden Gegenbaur, 1857:220 [emended to Cla donematidae by Poche, 1914:70]. Stauridiidae Hincks, 1868:61. Dendronemidae Haeckel, 1879:107. DIAGNOSIS Hydroid colonies with creeping stolons and short, un branched or sparingly branched hydrocauli with terminal hydranths. Perisarc of moderate thickness, terminating at hydranth base. Hydranths clavate, with an oral whorl of capitate tentacles, usually with an aboral whorl of reduced closely related, the Eleutheriidae are regarded as distinct from the Cladonematidae largely on the basis of differ ences in the medusa. These differences include the pres ence of a nematocyst ring around the umbrella margin, the lack of oral tentacles, the location of the gonads, and the reduced umbrella in the Eleutheriidae. Hydroids of this family resemble those of the Corynidae Johnston, 1836. Their medusae share with those of the Corynidae characters such as the presence of ocelli and the location of the gonad in a ring around the manubrium, but are otherwise rather highly specialized. filiform tentacles. Gonophores free medusae, arising from hydranth just distal to filiform tentacles; medusae creeping or swim ming. Manubrium with radial pouches; these pouches short and enclosed within subumbrellar cavity or moderately long and extending beyond velar opening; mouth with oral tentacles bearing nematocyst clusters. Radial canals bi furcated or simple, number variable. Marginal tentacles hollow, equal in number to radial canals, branching, bear ing organs of adhesion. Ocelli present. Gonads surround ing manubrium. Genus Cladonema Dujardin, 1843a stauridie Dujardin, 1843a: 1133. Cladonema Dujardin, 1843a: 1134. Stauridia Forbes, 1848:81 [emendation of stauridie Du jardin, 1843a]. Stauridium Krohn, 1853a: 137 [emendation of stauridie Dujardin, 1843a]. Stauridia Wright, 1858b:284 [invalid junior homonym of Stauridia Forbes, 1848 (Hydrozoa)]. REMARKS Prevot (1959), Naumov (1960), and Millard (1975) be DIAGNOSIS lieved that differences between the Cladonematidae Ge With the characters of the family. genbaur, 1857, and the Eleutheriidae Stechow, 1923a, were insufficient to warrant recognition of both families. Although the two families are acknowledged here to be 66 TYPE SPECIES Cladonema radiatum Dujardin, 1843a, by monotypy. REMARKS The genus Cladonema and its type species, C. radiatum, were described in two separate but essentially identical papers by Dujardin (1843a, 1843b). One of these papers (Dujardin, 1843a) appeared in the January-June issue of the Comptes Rendus Hebdomadaires des Seances de VAcademie des Sciences. The other (Dujardin, 1843b) was published in the December issue of the Annales des Sciences Naturelles. Accordingly, the former is taken as the first publication of the generic name Cladonema and of its type species, C. radiatum. The name "stauridie," applied by Dujardin (1843a, 1843b) to the hydroid of the medusa Cladonema radiatum, is not available [Art. 11]. The name of Dujardin's (1843a, 1843b) hydroid, stauridie, was emended to Stauridia by Forbes (1848). Wright (1858b), apparently unaware that Forbes had emended the name, also proposed the name Stauridia for this hydroid genus and included a new spe cies, S. producta, in it. However, S. producta produces medusae referable to the family Corynidae Johnston, 1836, as first shown by Hincks (1862), and is not congeneric with the hydroid described by Dujardin (1843a, 1843b). Mayer (1910) removed S. producta to a new genus, Stauridiosarsia, a name considered synonymous with Sarsia Lesson, 1843. Dendronema Haeckel, 1879, is similar to Cladonema, but was recognized as a valid genus by Bouillon (1985). Cladonema radiatum Dujardin, 1843a Fig. 50 stauridie Dujardin, 1843a: 1133. Cladonema radiatum Dujardin, 1843a: 1134 [medusa]. Coryne stauridia Gosse, 1853:260. Coryne stauridiai Gosse, 1853, pi. 16, figs. 1-5 [lapsus]. Syncoryne stauridium Krohn, 1853a:420. Cladonema radiatum gegenbauri Haeckel, 1879:109 [medusa]. Cladonema radiatum krohnii Haeckel, 1879:109 [medusa]. Fig. 50. Cladonema radiatum, part of hydrocaulus, and hy dranth with medusa bud, romiz B144. Scale equals 0.25 mm. Cladonema krohnii—Perkins, 1908:140 [medusa]. Stauridia radiatum—Mayer, 1910:100. Eleutheria radiata—Lengerich, 1922:210; fig. 1 [medusa]. Eleutheria perkinsii—Lengerich, 1922:211 [medusa]. not Cladonema radiatum var. mayeri Uchida, 1925:81; fig. 7 [= Cladonema uchidai Hirai, 1958]. Cladonema dujardini—Bedot, 1925:486 [medusa] [incor rect subsequent spelling]. Cladonema perkinsi—Bedot, 1925:486 [medusa] [incor rect subsequent spelling]. TYPE LOCALITY Mediterranean Sea. MATERIAL EXAMINED Stauridium cladonema Haeckel, 1879:109. Ferry Reach, St George's Island, opposite Whalebone Bay, on Thalassia, - 1.5 m, 26 July 1982, two hydranths, with medusa buds, romiz B144; medusa, newly liberated, ob tained in laboratory from one of the hydranths above, Cladonema sp. Fewkes, 1883:87 [medusa]. Cladonema sp. Perkins, 1902:25 [medusa]. romiz B151. Whalebone Bay, on Thalassia, -2 m, 8 September 1977, one hydranth, romiz B157. Cladonema radiatum dujardinii Haeckel, 1879:109 [medusa]. Cladonema radiatum allmani Haeckel, 1879:109 [medusa]. Cladonema radiata—Johansen and Levinsen, 1903:278 [medusa]. Cladonema perkinsii Mayer, 1904:18; pi. 4, fig. 35 [medusa]. Cladonema mayeriPerkins, 1906:118 [medusa and hydroid]. Cladonema allmani—Perkins, 1908:138 [medusa]. Cladonema dujardinii—Perkins, 1908:138 [medusa]. Cladonema gegenbauri—Perkins, 1908:140 [medusa]. DESCRIPTION Hydroid colonies with unbranched pedicels arising from a creeping hydrorhiza. Pedicels up to 4 mm long but usu ally much shorter, 75 (xm wide, supporting a terminal hydranth. Perisarc smooth, of moderate thickness, ter minating a short distance below filiform tentacles on hy dranth. Hydranths clavate, reaching 638 |xm long, 128 |xm 67 wide, with an oral whorl of four capitate tentacles and an aboral whorl of four filiform tentacles. Capitate tentacles solid, 326 \xm long, 47 |xm wide at base, tapering distally, with seven to nine endodermal cells; terminal knobs about 75 |xm wide, button-shaped. Filiform tentacles solid, 168 |xm long, 33 jxm wide, with four to six endodermal cells, alternating with capitate tentacles. Hypostome rounded, with an oral, ectodermal cavity. Medusa buds developing on short stalks just distal to filiform tentacles. Newly liberated medusa dome-shaped, 319 |xm high and 426 jxm wide in contracted, formalinpreserved specimen; mesoglea thin; manubrium short, wide, nearly spherical in longitudinal section. Radial canals nine. Tentacle bulbs nine, each with a single, reddish abaxial ocellus. Marginal tentacles nine, each with an adhesive organ and several bulbous clusters of nematocysts. REMARKS Hydroids of the various nominal species of Cladonema Dujardin, 1843a, are similar morphologically. Those of C. myersi Rees, 1949, and C. uchidai Hirai, 1958, ap parently both lack the whorl of filiform tentacles found in other species of the genus (Rees, 1949; Hirai, 1958; Nau mov, 1960), and Rees (1979) suggested that the two might be conspecific. Cladonema radiatum Dujardin, 1843a, and C. californicum Hyman, 1947, each possess both capitate and filiform tentacles and are virtually inseparable from one another in existing descriptions. However, Rees (1979), who first described the hydroid of C. californicum, did not include macrobasic mastigophores as part of the cnidome of this species. Nematocysts of this category have been reported from C. radiatum by Brinckmann and Petersen (1960), although they are not always present (Bouillon, that C. radiatum differed from D. reesi in several respects, including the shape of the knobs and number of endo dermal cells in the capitate tentacles, the presence of a slight terminal swelling on the filiform tentacles, the po sition of these tentacles on the hydranth, the shape of the stenoteles, and the presence of macrobasic mastigophores in addition to stenoteles. Calder (1970) reported that D. strangulata resembled C. radiatum in a number of these characters, such as the location of the filiform tentacles and the number of endodermal cells in the capitate ten tacles, but differed in lacking the slight terminal swelling of the filiform tentacles. Macrobasic mastigophores were also lacking in D. strangulata. Bouillon (1971) found that the three species were distinguishable by the structure of the hypostome. Ectodermal gland cells in the hypostome form a dome in D. strangulata and several related species; a button in D. reesi; and a cavity in Cladonema as well as species of Staurocladia Hartlaub, 1918, and Eleutheria de Quatrefages, 1842. Weill (1936) found medusae of Cladonema radiatum in Bermuda, and examined their nematocyst complement. Later, Weill (1937b) published observations on 106 speci mens collected near the Bermuda Biological Station. These medusae encompassed the characteristics of C. radiatum, C. perkinsii, and C. mayeri. Weill concluded that the three nominal species were no more than varieties of one rather variable species, as Mayer (1910) had suggested earlier. In Bermuda, the hydroid of Cladonema radiatum was observed only during the warmer months of the year. KNOWN RANGE Bermuda: no specific locality given (Weill, 1936); near the Bermuda Biological Station (Weill, 1937b); on Tha 1971). Bouillon observed that when macrobasic masti lassia (Calder, 1986). gophores are present in C. radiatum, they occur principally in the stolons, rarely in the body of the hydranth, and Elsewhere: western Atlantic (Mayer, 1910); eastern At lantic (Brinckmann-Voss, 1970). In his monographs on medusae, Kramp (1961, 1968) regarded Cladonema uchi dai as conspecific with C. radiatum. If this interpretation is correct, the known range of C. radiatum extends to the never on the tentacles. Hydroids of Cladonema radiatum also resemble those of Dipurena reesi Vannucci, 1956, and D. strangulata McCrady, 1859a. Brinckmann and Petersen (1960) found northwestern Pacific Ocean. Family Zancleidae Russell, 1953 DIAGNOSIS of specialized nematocyst-bearing tissue. Radial canals four. Marginal tentacles, when present, two or four, solid, each bearing abaxial cnidophores. Ocelli lacking. Gonads inter- Hydrorhiza creeping, with or without perisarc. Hydranths radial, on manubrium. Zancleidae Russell, 1953:98. with tentacles scattered. Tentacles capitate, or virtually filiform, or of both types. REMARKS Gonophores free medusae, arising from hydranths. Me dusa bell-shaped with a simple, circular mouth; exumbrella Several names were proposed for this family before Russell (1953) established the Zancleidae, namely the Orthocorynidae by A. Agassiz (1865) and the Corynipteridae and with or without oval or clavate patches or elongate tracks 68 Clavipteridae by Weill (1934). However, theselatterthree DIAGNOSIS names do not meet the criteria of availability for family- group names [Art. 1If (i)(l)], and do not take authorship Hydroid colonies stolonal, with perisarc covering hydro rhiza and hydrocaulus. Hydranthelongate. Tentaclescapi and date [Art. 10a]. If the nominal families Zancleidae tate, scattered except for those in an oral whorl. and Pteronematidae Haeckel, 1879, are eventually shown Gonophores free medusae, arising either from hydranth proximal to or among lower tentacles or, rarely, from hydrorhiza; hydranths with medusa buds occasionally re to be identical, as believed by Picard (1955, 1957, 1958) but disputed by Rees (1957), Bouillon (1974), and others, he maintained the Zancleidae and Pteronematidae as duced to blastostyles. Medusa bell-shaped, with exum brellar nematocysts; mouth simple, circular. Radial canals four. Marginal tentacles, when present, numbering two or four, with abaxial cnidophores. Ocelli lacking. Gonads separate families. Bouillon (1985) included the genus interradial. the latter name would have priority. Vervoort (1966:390) suggested that Pteronema darwinii Haeckel, 1879, might be "a mistreated and misjudged zancleid medusa," but Pteronema Haeckel, 1879, in the family Asyncorynidae Kramp, 1949, rather than in the Zancleidae. TYPE SPECIES Bouillon (1974) broadened the definition of this family to include a new genus and species of hydroid and medusa, Teissieramilleporoides, from the Seychelles. Unlike other Zanclea costata Gegenbaur, 1857, by monotypy. representatives of the Zancleidae, hydroids of T. mille poroides possessan encrustingskeletoninsteadof creeping stolons, and the polyps are polymorphic, with both gas The genus-group name Zanclea Gegenbaur, 1857, exten sively used in the literature of both hydroids and medusae for more than a century (e.g., Hincks, 1868; Mayer, 1910; trozooids and dactylozooids. The medusa generally resem bles those of other genera within the Zancleidae except in having ocelli. Bouillon included four genera in this revised Fraser, 1944; Russell, 1953; Kramp, 1961; BrinckmannVoss, 1970; Millard, 1975; Petersen, 1979; Bouillon, 1985; Calder, 1986), is a junior synonym of the virtually for family, Zanclea Gegenbaur, 1857, Pteroclava Weill, 1931, gotten name Acrochordium Meyen, 1834. In the influential works of L. Agassiz (1862), Bedot (1905), and Stechow (1923a), Acrochordium was mistakenly regarded as a jun ior synonym of Coryne Gaertner, 1774. A re-examination of Meyen's (1834) account of Acrochordium here indicates that it is congeneric with Zanclea rather than with Coryne. Mnestra parasites Krohn, 1853b, and Zanclea costata Gegenbaur, 1857, type species of Mnestra Krohn, 1853b, and Zanclea respectively, are regarded as conspecific (Rees, Rosalinda Totton, 1949, and a new genus, Teissiera. Mil lard (1975) adopted this revised definition of the family. Later, Bouillon (1978c) erected a new family, the Teissieridae, to accommodate T. milleporoides and two new species of medusae referred to Teissiera, namely T. australe and T. medusifera. The definition of the Zancleidae was thus returned by Bouillon essentially to that outlined by Russell (1953) and Kramp (1959, 1961, 1968). Rosa linda, which shares many characteristics with Teissiera (Bouillon, 1974; Petersen, 1979), cannot be retained in the Zancleidae. Instead, the new family Rosalindidae was constituted for the genus by Bouillon (1985). As currently defined, the Zancleidae once again includes the genera Zanclea and Pteroclava, and possibly also Ctenaria Hae ckel, 1879, and Oonautes Damas, 1936. Genus Zanclea Gegenbaur, 1857 Acrochordium Meyen, 1834:165. Mnestra Krohn, 1853b:281. Zanclea Gegenbaur, 1857:229. Gemmaria McCrady, 1859a: 151. Halocharis L. Agassiz, 1862:239. Gymnocoryne Hincks, 1871:75. Gemellaria Allman, 1871, pi. 7 [incorrect subsequent spelling]. Zanlcea Allman, 1872:290 [incorrect subsequent spelling]. Guentherella Weill, 1934:417. Zanklea Riedl, 1963:126 [incorrect subsequent spelling]. REMARKS 1953; Picard, 1957; Martin and Brinckmann, 1963). Thus, Zanclea is also a junior synonym of the infrequently used name Mnestra, a nominal genus with a single nominal species based on a parasitically deformed medusa (Ankel, 1952; Martin and Brinckmann, 1963). Picard (1957) re marked that he planned to submit an application to the iczn, requesting that its plenary powers be used to suppress the name Mnestra in favour of Zanclea. To my knowledge, however, the application was never published in the Bulletin of Zoological Nomenclature. In the interests of nomenclatural stability, the commis sion will be requested, in a future submission, to use its plenary powers to suppress the genus-group names Acro chordium and Mnestra for the Principle of Priority, and to place the name Zanclea on the Official List of Generic Names in Zoology. Zanclea alba (Meyen, 1834), comb. nov. Figs. 51, 52 Acrochordium album Meyen, 1834:165; pi. 28, fig. 8. Coryne sessilis Gosse, 1853:208; pi. 14, figs. 1-3. 69 Mnestra parasites Krohn, 1853b:281 [parasitized medusa]. Tubularia implexa Alder, 1856b:439. Zanclea costata Gegenbaur, 1857:229; pi. 8, fig. 4 [medusa]. Coryne pelagica Alder, 1857:103; pi. 7, figs. 1,2. Zanclea gemmosa McCrady, 1859a: 151; pi. 8, figs. 4,5 MATERIAL EXAMINED Atlantic Ocean, 2 km southeast of Castle Roads, on float ing Sargassum, 24 July 1982, one colony, up to 3 mm high, with medusa buds, romiz B145. Whalebone Bay, on floating Sargassum, 2 September 1977, one colony, up [medusa]. to 2 mm high, with medusa buds, romiz B155. Whale Coryne briareus Allman, 1859a:54. Coryne implexa—Wright, 1859:107. Coryne {margarica) implexa—Wright, 1859:108. Halocharis spiralis L. Agassiz, 1862:239; pi. 20, figs. bone Bay, on floating Sargassum, 27 February 1982, one colony, up to 2 mm high, with medusa buds, romiz B168. Natural Arches Beach, on stranded Sargassum, 8 March 1982, several colonies, up to 3 mm high, with medusa 10,10a-c. buds, romiz B170. St George's Island, beach near Fort St Catherine's, on stranded Sargassum, 15 June 1983, several medusae, liberated in laboratory from hydroid, romiz B332. Green Bay, Harrington Sound, on shell of Cerithium litteratum from Cladophora bed, -2 m, 26 September 1986, one colony, up to 2 mm high, without gonophores, romiz B359. Zanclea implexa—Allman, 1864a:357 [medusa and hydroid]. Halocharis {Corynitis) spiralis—Allman, 1864a:358. Corynitis agassizii—A. Agassiz, 1865:183 [part]. —Allman, 1872:287 [part]. —Bumpus, 1898:857. —Murbach, 1899:354; pi. 34, fig. 12. —Nutting, 1901:329; fig. 4. — Hargitt, 1901a:307. --Hargitt, 1901b:584; fig. 48. —Hargitt, 1904b:42. [medusa] [not Corynitis agassizii Mc Crady, 1859a] Gemmaria gemmosa—A. Agassiz, 1865:184; fig. 306 [medusa]. Gemmaria cladophora A. Agassiz, 1865:184; figs. 307310 [medusa]. Gymnocoryne coronata Hincks, 1871:76; pi. 5, figs. 1,1a. Gemellaria implexa—Allman, 1871, pi. 7, figs. 1-10 [in correct subsequent spelling]. Gemmaria implexa—Allman, 1872:290. Gemmaria sagittaria Haeckel, 1879:103;*pi. 7, figs. 3,4 [medusa]. Corynetis agassizii—Brooks, 1883a: 136 [incorrect sub sequent spelling] [not Corynitis agassizii McCrady, 1859a]. Zanclea inflexa—Pennington, 1885:51 [incorrect subse quent spelling]. Zanclea hargitti Hartlaub, 1907:119; fig. 109. Zanclea sagittaria—Hartlaub, 1907:124. Zanclea cladophora—Hartlaub, 1907:121; figs. 112, 113 [medusa]. Gemmaria sagittata—Hargitt, 1908:119 [incorrect sub sequent spelling]. Gemmaria costata—Fraser, 1912:346, fig. 2. Gemmaria implexa var. neapolitana Bruckner, 1914:460; figs. 7-24. Halocharis gemmosa—Stechow, 1923b:2. Gemmaria sp. Timmermann, 1932:296. DESCRIPTION Hydroid colonies with creeping hydrorhiza and upright, unbranched pedicels. Pedicels up to 1.3 mm long, about 70 |xm wide basally, expanding distally, bearing a ter minal hydranth. Perisarc of variable thickness, annulated at base of hydrocaulus, smooth distally, terminating at hydranth base. Hydranth nearly cylindrical, reaching 1.6 mm long, 150 |xm wide; hypostome dome-shaped. Tentacles as many as 40 or more, all capitate, scattered except for about 5 to 7 in an oral whorl, solid, with about six en dodermal cells each, short and relatively stout in preserved material, about 75 |xm long, 50 |xm wide at base, tapering distally, terminating in a knob of nematocysts; knobs 3550 jxm wide, nearly spherical. Nematocysts— Hydroids: stenoteles (small) 6.7-7.0 |xm x 4.9-5.3 |xm; stenoteles (large) 9.6-10.4 fxm x 8.3-8.6 |xm. Medusa buds arising in clusters on short stalks among proximal tentacles on hydranth. Well-developed medusa buds bell-shaped, with thin mesoglea; exumbrella with nematocyst patches; manubrium short, conical to tubular. Radial canals four. Marginal bulbs four; opposite two well developed and bearing tentacles with cnidophores. Nematocysts— Medusae: macrobasic euryteles 6.4-7.1 |xm x 3.64.4 (xm; stenoteles 6.8-9.2 |xm x 5.8-7.5 |xm. Guentherella implexa—Weill, 1934:417. REMARKS Zanclea sessilis—Russell and Rees, 1936:124. These Sargassum-borne specimens are almost certainly identical with Acrochordium album Meyen, 1834, a small athecate hydroid found on "Fucus natans L." off the Azores. Meyen's (1834) hydroid, a stolonal species with numerous capitate tentacles scattered over an elongate hy dranth, is not referable to Coryne Gaertner, 1774, as in dicated in earlier literature (see p. 64). Instead, it conforms to the current concept of the genus Zanclea Gegenbaur, Mnestra implexa—Picard, 1958:188. Zanklea costata—Riedl, 1963:126 [incorrect subsequent spelling]. TYPE LOCALITY Atlantic Ocean, in the vicinity of the Azores, on Sargas sum natans. 70 Fig. 51. Zanclea alba. Scale for a equals 0.5 mm; scale for b equals 0.25 mm. a, Part of colony with hydrorhiza, hydrocaulus, and hydranth, romiz B145. b. Newly liberated medusa, romiz B332. Fig. 52. Zanclea alba, nematocysts of hydranth and young medusa. Scales equal 10 |j.m. a, Macrobasic eurytele of medusa bud, romiz B332. b, Small stenotele of hydranth, romiz B170. c, Large stenotele of hydranth, romiz B170. d, Stenotele of medusa bud, romiz B332. 71 1857, and to the species commonly named Z. costata Ge genbaur, 1857. Uncertainty lingers about the scope of Z. costata (e.g., see Picard, 1957), and this name is predated by four other names as given in the synonymy list above. Accordingly, the species-group name album (as used in the binomen Acrochordium album by Meyen, 1834) is retained, notwithstanding its infrequent use in the litera ture, and the name Zanclea alba, comb, nov., is recog nized here as the valid name of the species. Russell and Rees (1936) cultured hydroids ascribed to Zanclea implexa (Alder, 1856b) in the laboratory and reared medusae, albeit with difficulty. They showed that a num ber of differences, formerly used to separate what were believed to be species and even genera, were based on characters that varied with stage of development. In the hydroid, presence or absence of visible perisarc on the hydrorhiza and hydrocaulus was found to vary with age and development of the colony. Likewise, the exact lo cation of medusa-bud formation was shown to be highly variable. In the medusa, arrangement of the nematocyst armature on the exumbrella and thickness of the mesoglea were shown to vary during development. Russell and Rees concluded that the North American Zanclea gemmosa McCrady, 1859a, should be united with the European Z. implexa (Alder, 1856b), and they provided a list of syn onyms. They provisionally retained Zanclea costata Ge genbaur, 1857, as a separate species because of the presence of four marginal tentacles in the medusa instead of two. However, Z. implexa and Z. costata were later united by Russell (1953). Coryne sessilis, a hydroid described by Gosse (1853) as having tentacles in whorls, was listed by Russell and Rees (1936) as questionably conspecific with Z. implexa. Brinckmann-Voss (1970) noted a tendency for the tentacles to occur in verticils in hydroids ascribed to Z. costata from the Mediterranean; it seems likely that Gosse exaggerated this in his illustrations and description of Z. sessilis. Picard (1957) believed that Z. costata and Z. gemmosa were distinct from Z. sessilis, but I have followed Brinckmann-Voss (1970) in regarding the three as conspecific. Zanclea sessilis was regarded as conspe cific with Coryne pusilla Gaertner, 1774, by Vervoort (1946b), but this seems quite unlikely considering the small size of Gosse's hydroids (about 1.5 mm high) and the large number of tentacles present (at least 45). Hargitt (1908) and Bedot (1925) noted that this species had been misidentified on numerous occasions as Corynitis agassizii McCrady, 1859a (= Linvillea agassizii). Bedot provided an extensive synonymy list to sort out the no menclatural confusion. Weill (1934) reported that the cnidome of a medusa identified as Gemmaria gemmosa (McCrady, 1859a) in cluded atrichs as well as macrobasic euryteles and steno teles. No atrichs were observed in newly liberated medusae of Zanclea alba from Bermuda. This species, listed under the name Zanclea costata, was reported to be common on pelagic Sargassum by Mor ris and Mogelberg (1973). Hydroids of Z. alba were com mon to abundant on pelagic Sargassum at Bermuda during this study. KNOWN RANGE Bermuda: on pelagic Sargassum (Calder, 1986). Elsewhere: apparently circumglobal in tropical and tem perate waters (Russell and Rees, 1936; Fraser, 1944; Ya mada, 1959; Kramp, 1959, 1961, 1965, 1968; BrinckmannVoss, 1970; Bouillon, 1978c). Family Milleporidae Fleming, 1828 Milleporadae Fleming, 1828:528 [emended to Millepori dae by Milne Edwards and Haime, 1849]. DIAGNOSIS Gonophores free but reduced medusae, arising from coenosarc within ampullae in coenosteum. Medusae with exumbrellar nematocyst patches; velum and tentacles lack ing. Gonads on manubrium. Hydroid colonies forming massive, calcareous exoskeletons of varied shape. Coenosteum with a complex network REMARKS of coenosarcal tubes internally, covered externally by a thin epidermal layer, with surface perforated by pores; margins of pores not protruding from surface of coenos teum; larger gastropores surrounded by smaller dactylopores, forming indistinct cyclosystems. Gastrostyles and dactylostyles lacking. Polyps polymorphic; gastrozooids relatively short and stout, with four to seven short capitate tentacles, and arising within gastropores; dactylozooids long, slender, mouthless, with scattered capitate tentacles, and arising from dactylopores. 72 Species of the family Milleporidae Fleming, 1828, have been recognized as hydrozoans since the work of L. Agas siz (1858), but their systematic position within the class Hydrozoa has been a matter of longstanding debate. Moseley (1880) referred this family, along with the Stylasteridae Gray, 1847, to the suborder Hydrocorallinae. The great differences separating milleporids and stylasterids were pointed out by S. J. Hickson (in a note in Delage and Herouard, 1901), and he placed the former in a new order, the Milleporina. Separation of these two families was upheld by Broch (1914) and Stechow (1923a), but they maintained that milleporids were capitate hydroids related to the Corynidae Johnston, 1836. According to a number of recent authors, including Bouillon (1974, 1985) and Petersen (1979), available evidence from hydranth morphology and nem atocyst complement suggests that the family has affinities with the Teissieridae Bouillon, 1978c, Zancleidae Russell, 1953, and Cladocorynidae Allman, 1872. Further details on history of the classification of the Milleporidae, together with a taxonomic assessment of the fossil species referred to the family, are given by Boschma (1951, 1956). Genus Millepora Linnaeus, 1758 Millepora Linnaeus, 1758:790. Palmipora de Blainville, 1830:356. DIAGNOSIS With the characters of the family. forms" were indeed distinct species. Boschma (1948), in an extensive and influential review of the species problem in Millepora, recognized 10 rather well defined species and discussed the characters used for specific distinction. De Weerdt (1981) noted that the species of Millepora are currently distinguished principally on the growth form of the corallum and geographic distribution, but growth form is highly influenced by environmental factors such as water movement and turbidity. The taxonomic value of characters such as texture of the corallum surface, size and shape of the gastropores and dactylopores and their distribution over various parts of the corallum, extent to which cyclosystems are isolated, presence or absence of ampullae, morphology of the soft parts, and stinging prop erties has largely been discounted. However, de Weerdt concluded that such characters, including the shape and size of ampullae, may be of more value taxonomically than supposed and merit re-examination. Millepora alcicornis Linnaeus, 1758 Figs. 53-55 TYPE SPECIES Millepora alcicornis Linnaeus, 1758:791. Millepora alcicornis Linnaeus, 1758, by subsequent des ignation by Apstein (1915). Millepora alcicornis digitata Esper, 1790:197; pi. 5, figs. REMARKS The genus Millepora Linnaeus, 1758, originally included a conglomeration of species only one of which, M. alci cornis Linnaeus, 1758, is referred to the genus as it is understood at present (Boschma, 1948). There is even some uncertainty about the identity ofM. alcicornis, which may have been a species of scleractinian and not a hydro zoan. However, Boschma (1948) indicated that there is 1,2. Millepora alcicornis corniculata Esper, 1790:197; pi. 6. Millepora alicornis—Esper, 1790:197 [incorrect subse quent spelling]. Millepora alcicornis ramosa Esper, 1790:198; pi. 7. IMillepora alcicornis Crustacea Esper, 1790:200 [not Mil lepora Crustacea Linnaeus, 1758]. Millepora alcycornis—Bosc, 1802:288 [incorrect subse quent spelling]. Palmipora alcicornis—de Blainville, 1834:391; pi. 58, support for the belief that the M. alcicornis sensu Linnaeus fig. 2. was the branched West Indian hydrozoan currently as signed that name. The colony form of this hydrozoan often Millepora moniliformis Dana, 1848:544 [not Millepora bears a resemblance to an elk's horn, as reflected in the etymology of the specific name. Moreover, the definition of M. alcicornis by Linnaeus (1767) almost certainly refers to the hydrozoan genus Milleporaas understood at present, and not to a scleractinian. Boschma (1948) recounted the differing views concern ing species limits in Millepora. These views have ranged from that of Duchassaing and Michelotti (1864), who re garded almost every different growth form as a distinct species, to that of Hickson (1898a, 1898b), who believed that the genus contained a single species, M. alcicornis. Hickson's concept of a single species displaying different growth forms or "fades" was widely, although on oc casion somewhat grudgingly, accepted through the early decades of the present century. The opinion was gradually superseded by the conclusion that some of the "growth moniliformis Rafinesque, 1820]. Millepora ramosa—Dana, 1848:544. Millepora pumila Dana, 1848:545; pi. 52, fig. 4 [not Mil lepora pumila Pallas, 1766]. Palmipora fasciculata Duchassaing, 1850:18. Palmipora parasitica Duchassaing, 1850:18. Millepora forskali Milne Edwards, 1860:228. Millepora fasciculata—Milne Edwards, 1860:228 [not Millepora fasciculata Lamarck, 1816]. Millepora gothica Duchassaing and Michelotti, 1860:84; pi. 10, figs. 9,10. Millepora schrammi Duchassaing and Michelotti, 1864:100; pi. 11, fig. 9. Millepora esperi Duchassaing and Michelotti, 1864:100. Millepora crista-galli Duchassaing and Michelotti, 1864:101; pi. 11, fig. 7 [not Milleporacristagalli Morren, 1828]. 73 Millepora delicatula Duchassaing and Michelotti, 1864:101; pi. 11, fig. 10. Millepora Candida Duchassaing and Michelotti, 1864:101. Millepora digitata—Duchassaing and Michelotti, 1864:102. Millepora carthaginiensis Duchassaing and Michelotti, 1864:102; pi. 11, fig. 6. Millepora trinitatis Duchassaing and Michelotti, 1864:102. Milleporafenestrata Duchassaing and Michelotti, 1864:103; pi. 11, fig. 1. Millepora nitida Verrill, 1868:362. Millepora alcicornis var. cellulosa Verrill, 1868:363. Millepora alcicornis var. fenestrata—Verrill, 1868:364. Montipora gothica—Dollfus, 1936:515. TYPE LOCALITY Not specified by Linnaeus (1758); subsequently given as "O. Indiae utriusque" (Linnaeus, 1767). MATERIAL EXAMINED Whalebone Bay, on ledges at entrance, -4 m, 4 March 1982, one fragmentary colony, 1.5 cm high, romiz B175. Hall's Island, Harrington Sound, on rocks, - 1 m, 13 September 1977, one fragmentary colony, 6.2 cm high, liberating medusae when alive, coll. W. Sterrer, romiz B180. Whalebone Bay, on ledges at entrance, -2 m, two fragmentary colonies, 2.0-4.5 cm high, romiz B181. Flatts Inlet, on shells and rubble, -4 m, 5 March 1982, one colony, 7.0 cm high, romiz B304. Flatts Inlet, on rubble, -3 m, 9 July 1983, one colony, 1.7 cm high, romiz B312. DESCRIPTION Colonies initially consisting of an encrusting base and fin gerlike upright projections; older colonies with extensive upright branches of varied shape. Branches irregularly re branched, generally in one plane; branchlets fused to vary ing degrees basally, with tips usually free and digitate. Coenosteum consisting of a framework of anastomosing calcareous trabeculae, interstices occupied by a network of coenosarc tubes, surface covered with an epidermal layer and perforated by pores. Gastropores 0.35 mm in diameter; dactylopores 0.25 mm in diameter; cyclosys- Fig. 53. Millepora alcicornis, dactylozooid and gastrozooid extending above coenosteum, romiz B312. Scale equals 0.25 mm. 6.6 fjim; stenoteles (medium) 15.9-17.6 |xm x 12.914.2 |xm; stenoteles (large) 21.6-24.7 jim x 15.918.7 jxm. Dactylozooids: stenoteles (small) 8.3-8.6 |xm x 5.96.5 |xm. tems distinct to indistinct, with five to nine dactylopores REMARKS surrounding each gastropore. Gastrostyles and dactylostyles absent. Gastrozooidsrelatively stout, reaching about The synonymy of this species is long and complex. The simplified list given here largely follows that of Boschma 1 mm above surface of coenosteum when extended, with (1948). an oral whorl of four to seven short, capitate tentacles. capitate tentacles grouped at distal end and scattered Millepora alcicornis Linnaeus, 1758, is the only species of its genus known to occur in Bermuda, although two other species, M. complanata Lamarck, 1816, and M. squarrosa Lamarck, 1816, have been reported elsewhere elsewhere. in the western North Atlantic. Colonies of M. alcicornis Dactylozooids long and slender when extended, reaching 1.0-1.5 mm above surface of coenosteum, with short, Nematocysts— Gastrozooids: microbasic mastigophores 30.0-31.9 fxm x 24.5-26.3 >jim; stenoteles (small) 8.3-8.9 (xm x 5.774 are highly varied in shape, but they are distinguishable from those of M. complanata and M. squarrosa in being composed of branches rather than of vertical plates 1 i *' # #..•' ^p I\^L ^^s> ^ * A •v^flC ££*S! . mt wis i»! '. lT **^ Fig. 54. Millepora alcicornis, coenosteum. Scale for a equals 5 cm; scale for b equals 100 |xm. a, Young colony encrusting bivalves and coral rubble, romiz B304. b, SEM micrograph of part of coenosteum, showing a cyclosystem, romiz B304. Fig. 55. Millepora alcicornis, nematocysts, romizB312. Scales equal 10 p.m. a, Microbasic mastigophore of gastrozooid. b, Large stenotele of gastrozooid. c, Medium stenotele of gastrozooid. d. Small stenotele of gastrozooid. e, Small stenotele of dactylozooid. 75 (Boschma, 1948). The vertical plates of M. complanata are truncated distally and united only at their bases, while those of M. squarrosa are frilled distally and often united nis, in common with other species of the genus, is ven above their bases. by Mayer (1910). Verrill (1907) reported colonies of Millepora alcicornis as large as 4-6 feet (1.2-1.8 m) across and 1-2 feet (0.30.6 m) high at Bermuda. He commented that it was the most abundant "coral" on outer reefs and on inner rocks and ledges around the islands, ranging from the shallows to a depth of 5-8 fathoms (9-15 m). When alive, the species is brownish in colour because of the presence of symbiotic zooxanthellae in the tissues. Millepora alcicor omous to humans (Verrill, 1907). A description of the medusastageof this speciesis given KNOWN RANGE Bermuda: on reefs, ledges, rocks, and other substrates around the entire Bermuda Platform (Nelson and Duncan, 1876; Moseley, 1876, 1879, 1880; Rice, 1878; Quelch, 1886; Verrill, 1900, 1902a, 1902b, 1907; Moore, 1969; Calder, 1986). Elsewhere: western Atlantic (Boschma, 1948). Family Porpitidae Goldfuss, 1818 Porpitae Goldfuss, 1818:1012 [emended to Porpitidae by Guilding 1828:403]. Velellidae Eschscholtz, 1829:165. Discalidae Haeckel, 1888a:29. Porpalidae Haeckel, 1888b:57. Porpitellidae Haeckel, 1888b:63. DIAGNOSIS Hydroids polymorphic, highly specialized, adapted for life at sea surface. Float and mantle with or without upright sail; undersurface with a large central gastrozooid, a ring of gonozooids, and a peripheral ring of dactylozooids. Gonophores free medusae. Medusae thimble-shaped with perradial exumbrellar nematocyst rows; mouth simple, cir cular. Radial canals four. Tentacle bulbs four. Ocelli lack ing. Gonad not completely encircling manubrium. REMARKS The family name Porpitidae is frequently attributed to Brandt (1835), but the name was apparently first employed by Goldfuss (1818) as the Porpitae. This name, emended to Porpitidae by Guilding (1828), thus predates the name Velellidae Eschscholtz, 1829, and must be given priority when the two nominal families are combined. I concur with Brinckmann-Voss (1970) and others that the Porpi tidae and Velellidae can be contained in a single family. Chamisso and Eysenhardt (1821) included various nom inal species of Porpita Lamarck, 1801, and Velella La marck, 1801, under the name Chondrophorae (as ' 'Medusae Chondrophorae"), but it is unclear whether they intended Chondrophorae to be a family-group name. In any case, the name Chondrophorae does not meet the criteria of availability for family-group names [Art. 1If (i)(l)], either in its original form or in any of its subsequently emended spellings. Members of this family have been variously treated as siphonophores (e.g., Eschscholtz, 1829; Huxley, 1859; L. 76 Agassiz, 1862; Haeckel, 1888a, 1888b; Bigelow, 1911; Moser, 1925), as athecate hydroids (e.g., Kolliker, 1853; Vogt, 1854; McCrady, 1859a; A. Agassiz, 1883; Ed wards, 1966b; Brinckmann-Voss, 1970), and as a separate order of Hydrozoa, the Chondrophora (e.g., Totton, 1954; Rees, 1957). The concensus among contemporary workers is that they are highly specialized athecate hydroids (see Edwards, 1966b; Brinckmann-Voss, 1970; Fields and Mackie, 1971; Bouillon, 1974, 1985; Petersen, 1979; Arai and Brinckmann-Voss, 1980; Kirkpatrick and Pugh, 1984; Calder, 1986). Opinions have differed widely concerning their system atic position within the athecate hydroids. Leloup (1929, 1954), Garstang (1946), Totton (1954), Mackie (1959, 1960), and Daniel (1976), among others, have suggested that they are most closely related to the Corymorphidae Allman, 1872, and Tubulariidae Fleming, 1828. Fields and Mackie (1971) regarded Velella as a large, floating tubulariid hydranth and placed the nominal family Velel lidae, together with the Corymorphidae, Tubulariidae, and Margelopsidae Uchida, 1927, in the superfamily Tubularioidea Fleming, 1828. Picard (1955, 1957) and Prevot (1959) included Velella and Porpita, along with the Zan cleidae Russell, 1953, in the Pteronematoidea Haeckel, 1879. Brinckmann (1964) believed that their affinities were with the tubularians and placed them in a superfamily, the Chondrophoroidea, in the order Anthomedusae. Later, she suggested that they should be placed between the Capitata and Filifera (Brinckmann-Voss, 1970). Based on the mor phology and histological structure of their polyps and me dusae, as well as their nematocyst complement, Bouillon (1974) argued that Velella and Porpita had undeniable affinities with the Zancleidae. Petersen (1979) treated them as a superfamily within the Zancleida, a suborder encom passing the Cladocorynidae Allman, 1872, Zancleidae, Teissieridae Bouillon, 1978c, and Milleporidae Fleming, 1828. Most of the available evidence points to the Porpitidae as having an affinity with taxa of the superfamily Zancleoidea Russell, 1953. If such an interpretation is correct, it follows that Porpita and Velella should be interpreted as colonies rather than as individual floating hydranths and that their zooids are true polyps, as argued by Edwards plication referred to above to use its plenary powers to suppress the seldom-used genus-group name Phyllidoce for the purposes of the Principle of Priority, and to place that name on the Official Index of Rejected and Invalid Generic Names in Zoology. The name Phyllidoce was first used by Browne (1789) for the hydroid known today as Velella velella (Linnaeus, 1758), but Browne's work has (1966b). Genus Porpita Lamarck, 1801 Porpita Lamarck, 1801:355. Polybrachionia Guilding, 1828:403. Ratis Lesson, 1830:60. Acies Lesson, 1830:61. Chrysomitra Gegenbaur, 1857:232. Disconalia Haeckel, 1888a:30. Porpitella Haeckel, 1888a:30 [invalid junior homonym of Porpitella Pomel, 1883 (Echinodermata)]. been suppressed for nomenclatural purposes by the iczn (Opinion 89). Bigelow (1911) recognized Porpema Haeckel, 1888a, as distinct from Porpita, while Totton (1954) believed that the two were congeneric. The taxa appear sufficiently dis tinct, based on present understanding of their morphology (Bigelow, 1911), to warrant recognition of both as valid genera. Bigelow (1911) regarded Porpalia Haeckel, 1888a, as a synonym of Porpema. Acting as first reviser, he chose Porpema as the valid name of the genus. Discalia Haeckel, 1888a, is probably a synonym of Porpema as well. Hae ckel 's (1888a) nominal genera Disconalia and Porpitella were regarded by Bigelow as congeneric with Porpita. DIAGNOSIS Porpitid hydroids with disc-shaped float and mantle; float flat or with central bulge; sail lacking. Dactylozooids with three vertical rows of short, capitate tentacles. Medusae with small manubrium. Juvenile specimens without marginal tentacles; adults with one to two slender, decidedly capitate tentacles; endodermal cells along radial canals bearing algal symbionts. Porpita porpita (Linnaeus, 1758) Figs. 56, 57 Medusa porpita Linnaeus, 1758:659. Holothuria denudata Forskal, 1775:103. Holothurio denudata Forskal, 1776, pi. 26, figs. L,l [lapsus]. Medusa umbella O. F. Muller, 1776b:297; pi. 9, figs. 2,3. TYPE SPECIES Holothuria nuda Gmelin, 1790:3143. Medusa porpita Linnaeus, 1758, by absolute tautonomy. Phyllidoce denudata—Modeer, 1790:201. Phyllidoce porpita—Modeer, 1790:203. Porpita indica Lamarck, 1801:355. Porpita appendiculata Bosc, 1802:155; pi. 18, figs. 5,6. REMARKS Bigelow (1911) retained the name Porpita Lamarck, 1801, for this genus instead of employing Medusa Linnaeus, 1758. Apparently, a type species has never been desig nated for Medusa, a nominal genus originally containing species of Hydrozoa, Scyphozoa, and Ctenophora, and the name has been scarcely used this century. The widely used name Porpita Lamarck, 1801, is a junior homonym of Porpita Soldani, 1789 (Protozoa), a name apparently all but abandoned in the recent zoological literature. Application will be made to the iczn to place Porpita Lamarck, 1801, on the Official List of Generic Names in Zoology, and Porpita Soldani, 1789, on the Official Index of Rejected and Invalid Generic Names in Zoology. The names Porpita Lamarck, 1801, and Velella La marck, 1801, are threatened by Phyllidoce Modeer, 1790, Porpita radiata Bory de St Vincent, 1804:99; pi. 5, figs. 2A-D. Porpita gigantea Peron and Lesueur, 1807, pi. 31, figs. 6,6a-e. Porpita forskalea Oken, 1815:111 [name published in a work rejected for nomenclatural purposes by the iczn (Opinion 417)]. Porpita nuda—Lamarck, 1816:484. Porpita glandifera Lamarck, 1816:485. Porpita granulata Cranch, 1818:418. Porpita coerulea Eschscholtz, 1825:744. Porpita globosa Eschscholtz, 1825:744. Porpita ramifera Eschscholtz, 1825:745. Porpita chrysocoma Lesson, 1826, pi. 7, figs. 1,1'. Porpita atlantica Lesson, 1826, pi. 7, fig. 2. Porpita pacifica Lesson, 1826, pi. 7, figs. 3,3'. a nominal genus originally including three nominal spe cies, P. denudata (Forskal, 1775), P. porpita (Linnaeus, Porpita moneta Risso, 1826:304. 1758), and P. velella (Linnaeus, 1758). In the interests of Medusa nuda—Bory de St Vincent, 1827:139;pi. 90, figs. nomenclatural stability, the iczn will be asked in the ap- 3-5. 77 Medusa glandifera—Bory de St Vincent, 1827:139; pi. 90, figs. 6,7. Polybrachionia linnaeana Guilding, 1828:404; pi. 10, figs. 1,2. Porpita forskahli de Haan, 1827:493. Porpita reinwardtii de Haan, 1827:493. Porpita kuhlii de Haan, 1827:494. Porpita mediterranea Eschscholtz, 1829:177. Porpita umbella Eschscholtz, 1829:179. Ratis medusae Lesson, 1830:60. Acies palpebrans Lesson, 1830:61. Porpita lutkeana Brandt, 1835:41. Porpita linnaeana—Lesson, 1843:588. Chrysomitra striata Gegenbaur, 1857:232; pi. 7, figs. 10,11 [medusa]. Wiscalia primordialis Haeckel, 1888a:30 [nomen nudum]. Disconalia pectyllis Haeckel, 1888a:30 [nomen nudum]. Disconalia gastroblasta Haeckel, 1888a:30 [nomen nudum]. Porpitella caerulea—Haeckel, 1888a:30 [incorrect sub sequent spelling]. Porpitella radiata—Haeckel, 1888a:30. IDiscalia primordialis Haeckel, 1888b:46. Disconalia pectyllis Haeckel, 1888b:48. Disconalia gastroblasta Haeckel, 1888b:48; pi. 49, figs. 7-12; pi. 50, figs. 1-10. Porpitella pectanthis Haeckel, 1888b:64; pi. 46. Porpita fungia Haeckel, 1888b:67; pi. 45. Disconalia ramifera Haeckel, 1888b:357. Porpitella coerulea—Haeckel, 1888b:358. Porpita porpita—Schneider, 1898b: 194. Phyllodice denudata—Bigelow, 1911:352 [incorrect sub sequent spelling]. TYPE LOCALITY "Habitat in India" (Linnaeus, 1758). MATERIAL EXAMINED Building's Bay, St George's Island, washed ashore, 7 Oc tober 1984, one colony, 7 mm in diameter, romiz B347. Atlantic Ocean, 10 km south of Nonsuch Island, 7 May 1984, one colony, rataria stage, 3 mm in diameter, coll. Paul Bennett, romiz B350. DESCRIPTION Hydroid pleustonic, with disc-shaped mantle and internal float; upper surface of mantle and float slightly convex, with a central pore and numerous peripheral stigmata. Mande 7 mm wide, with radiating gastrodermal canals; margin soft, flexible; central region firm, with an internal chitinous float consisting of a series of concentric air chambers; a disc-shaped reservoir of nematocytes and nematocysts lying between float and central gastrozooid. Undersurface with a single large, central gastrozooid, a medial band of gono 78 zooids, and a peripheral band of dactylozooids. Central gastrozooid short and broad with a terminal mouth; ten tacles and prominent nematocyst batteries lacking. Dac tylozooids tentaculate, with a distal whorl of four capitate tentacles;body with varying numbers of short, small, cap itate tentacles in three vertical rows; mouth lacking. Gon ozooids clavate, lacking tentacles but with prominent nematocyst batteries encircling mouth, additional nema tocyst batteries scattered over body; medusa buds given off in clusters basally. Nematocysts— Hydroids: atrichous isorhizas (on gastrozooids, dacty lozooids, gonozooids; in nematocyst reservoir) 9.3-13.2 |^m x 4.2-5.0 |xm; haplonemes (on gastrozooids, dactylo zooids) 10.8-14.1 |xm x 5.6-6.6 |xm; stenoteles (small) (on gastrozooids, dactylozooids, gonozooids; in nemato cyst reservoir) 15.6-16.7 |xm x 10.5-11.5 (xm; steno teles (large) (on gastrozooids, gonozooids; in nematocyst reservoir) 27.5-30.0 |xm x 21.8-25.3 |xm; stenoteles (bulbous) (on medusa buds) 15.2-16.2 (Jim x 12.713.9 |xm. Rataria "larva" planktonic, with swollen, subspherical central gastrozooid, a medial ring of gonozooids, a pe ripheralring of tentaculate dactylozooids, and a small disc shaped float and mantle. Medusae not seen. REMARKS Porpitid taxonomy has advanced relatively little since Bigelow's (1911) insightful study of the group. The syn onymy list here is taken in part from Bigelow's work, although Moser (1925) and Totton (1954) have been fol lowed in regarding Porpita porpita (Linnaeus, 1758) from the Indian Ocean, P. umbella (O. F. Muller, 1776b) from the Atlantic Ocean, and P. pacifica Lesson, 1826, from the Pacific Ocean as conspecific. The development of Porpita porpita is known to pass through stages somewhat resembling those of Velella ve lella (Linnaeus, 1758). Various stages in the life history of this species, including the medusa, rataria "larva," and young pleustonic hydroid, have been described by A. Agassiz (1883), Bigelow (1911), Delsman (1923), and Bouillon (1984b). The medusa, at an advanced stage of its development, has euryteles as part of its cnidome (Bouillon, 1984b, 1985). A. Agassiz (1883) commented that few specimens of Porpita porpita are stranded on beaches compared with Velella velella. Only one specimen was found on Bermuda beaches during this study, although considerable time was spent looking for the species during four field trips. Porpita porpita has been reported previously from Bermuda by Verrill (1900, as P. linnaeana), and by Totton (1936, as P. umbella). a Fig. 56. Porpita porpita. Scale for a equals 2.5 mm; scale for b equals 1 mm; scales for c and d equal 0.5 mm. a, Upper surface of float and mantle, romiz B347. b, Rataria larva, romiz B350. c, Gonozooid, with medusa buds, romiz B347. d, Dactylozooid, romiz B347. 79 KNOWN RANGE Bermuda: no specific locality given (Verrill, 1900; Calder, Vellela Bory de St Vincent, 1827:139 [incorrect subse quent spelling]. 1986); 14 km southeast of Nonsuch Island (Totton, 1936). Rataria Eschscholtz. 829:166. Elsewhere: circumglobal, tropical and temperate waters Armenistarium Costa 1841:187. (Moser, 1925; Brinckmann-Voss, 1970). Velaria Haeckel, 1 Armenista Haeckel Genus Velella Lamarck, 1801 :31. 83. DIAGNOSIS Phyllidoce Browne, 1789:387 [invalid name, published in a work suppressed under the plenary powers for nomen clatural purposes by the iczn (Opinion 89)]. Porpitid hydroids with oval float and mantle and with up right, triangular sail set diagonal to long axis of float. Dactylozooids with nematocyst patches. Medusae with short, conical manubrium. Tentacle bulbs four, one opposite pair lacking tentacles, each bulb of re Velella Lamarck, 1801:355. in J ••: / a b , /. Fig. 57. Porpita porpita, nematocysts of hydroid and medusa bud. romiz B347. Scales equal 10 u.m. a, Atrichous isorhiza of dactylozooid. b, Haploneme of gastrozooid. c, Small stenotele of dactylozooid. d, Large stenotele of gonozooid. e, Bulbous stenotele of medusa bud. 80 maining pair with one or two capitate tentacles; zooxan thellae concentrated near radial canals and ring canal. TYPE SPECIES Medusa velella Linnaeus, 1758, by absolute tautonomy. REMARKS Numerous nominal species of Velella Lamarck, 1801, have been described over the years from Atlantic, Pacific, and Indian oceans. All of these are currently believed to belong to a single species, V. velella (Linnaeus, 1758) (Schneider, 1898b; Bigelow and Sears, 1937; Totton, 1954; Daniel Velella Velella Velella Velella spirans—Eschscholtz, 1829:172. caurina Eschscholtz, 1829:173; pi. 15, fig. 2. tropica Eschscholtz, 1829:174; pi. 15, fig. 3. indica Eschscholtz, 1829:175; pi. 15, fig. 5. Velella antarctica Eschscholtz, 1829:175. Velella patellaris Brandt, 1835:38. Velella oxyothone Brandt, 1835:39. Velella oxyothone var. brachyothone Brandt, 1835:39. Velella oxyothone var. oxyothone Brandt, 1835:39. Armenistarium velella—Costa, 1841:187; pi. 13, fig. 3. Rataria cristata Haeckel, 1888a:31 [nomen nudum]. Velaria mutica—Haeckel, 1888a:31. and Daniel, 1963; Brinckmann-Voss, 1970; Daniel, 1976; Arai and Brinckmann-Voss, 1980; Kirkpatrick and Pugh, Velaria oblonga—Haeckel, 1888a:31. 1984). Rataria cristata Haeckel, 1888b:79; pi. 44. Velella patella—Haeckel, 1888b:83 [incorrect subsequent spelling]. Armenista sigmoides Haeckel, 1888b:84; pi. 43. Velella velella (Linnaeus, 1758) Figs. 58, 59 Medusa velella Linnaeus, 1758:660. Holothuria spirans Forskal, 1775:104. Holothurio spirans Forskal, 1776, pi. 26, figs. K,kl-3 [lapsus]. Phyllidoce velella—Modeer, 1790:194. Velella mutica Lamarck, 1801:355. Velella tentaculata Lamarck, 1801:355. Velella scaphidia Peron and Lesueur, 1807; pi. 30, figs. 6,6a. Medusa pocillum Montagu, 1815:201; pi. 14, fig. 4. Velella limbosa Lamarck, 1816:482. Velella pyramidalis Cranch, 1818:419. Velella sinistra Chamisso and Eysenhardt, 1821:363; pi. 32, fig. 1. Velella oblonga Chamisso and Eysenhardt, 1821:364; pi. 32, figs. 2A-C. Velella lata Chamisso and Eysenhardt, 1821:364; pi. 32, figs. 3A-B. Velella emarginata Quoy and Gaimard, 1824:586; pi. 86, fig. 9. Velella cyanea Lesson, 1826, pi. 6, figs. 3,4. Velella australis de Haan, 1827:489. Velaria indica—Haeckel, 1888a:31. Armenista Armenista Armenista Armenista Armenista mutica—Haeckel, 1888b:84. antarctica—Haeckel, 1888b:84. indica—Haeckel, 1888b:84. lata—Haeckel, 1888b:84. lobata Haeckel, 1888b:84 [nomen nudum]. Velella meridionalis Fewkes, 1889:112; pi. 1, figs. 1-3; pi. 2, fig. 3. Velella velella—Schneider, 1898b: 194. Velella spiralis—Martin, 1904:27 [incorrect subsequent spelling]. Velella subemarginata—Stephens, 1905:65 [incorrect subsequent spelling]. Phyllodice velella—Bigelow, 1911:353 [incorrect subse quent spelling]. TYPE LOCALITY 4'Habitat in Pelago. Loefling. In Mari Mediterraneo. Brander" (Linnaeus, 1758). MATERIAL EXAMINED Warwick Long Bay, washed ashore, 7 March 1982, 30 left-sailing forms, 5-22 mm long, 3-14 mm wide, and three right-sailing forms, 11-31 mm long, 9-20 mm wide, romiz B174. Velella pacifica de Haan, 1827:490. Velella radackiana de Haan, 1827:490. Velella sandwichiana de Haan, 1827:491. Vellela limbosa—Bory de St Vincent, 1827:139; pi. 90, figs. 1,2 [incorrect subsequent spelling]. Velella pocillum—Fleming, 1828:500. Rataria cordata Eschscholtz, 1829:167; pi. 16, fig. 1. Rataria pocillum—Eschscholtz, 1829:168. Rataria mitrata Eschscholtz, 1829:168; pi. 16, fig. 2. Velella aurora Eschscholtz, 1829:171. Velella septentrionalis Eschscholtz, 1829:171; pi. 15, fig. 1. DESCRIPTION Hydroids pleustonic, with flattened, oval mantle and in ternal float; upper surface of mantle and float with upright, triangular sail. Specimens occurring in two mirror-image forms, with sail lying along either of two diagonals of mantle and float. Sail with branched and anastomosing radial canals, rather rigidly erect, supported internally by slender, triangular, chitinous extension of float. Mantle coursed by radiating gastrodermal canals having short, lateral diverticulae; margin soft, flexible, bearing large numbers of algal symbionts; central region firm, with an 81 Fig. 58. Velella velella, romiz B174. Scale for a equals 3 mm; scales for b-d equal 2 mm. a, Lateral view of float, mantle, and sail, b, Gastrozooid. c, Dactylozooid. d, Gonozooid, with medusa buds. internal chitinous float consisting of a series of concentric air chambers; an elongate-oval reservoir of nematocytes and nematocysts lying between float and central gastro zooid. Undersurface with a single central gastrozooid, a medial band of gonozooids, and a peripheral band of dac tylozooids. Central gastrozooid long and tubular distally with terminal mouth, broad and oval basally, lacking both tentacles and prominent nematocyst batteries; gastrodermis with a series of longitudinal folds. Gonozooids clavate, occurring in various developmental stages, lacking ten tacles but with prominent batteries of nematocysts about mouth, additional nematocyst batteries scattered over body; blastostyles given off-proximally, each bearing clusters of 82 medusa buds. Dactylozooids tentacle-shaped, oval in crosssection, each bearing a band of nematocysts along the two narrow edges; nematocyst band typically extending up one side and across distal end of dactylozooid, and continuing down opposite side; band often becoming broken up into discontinuous patches proximally; mouth lacking. Nematocysts— Hydroids: atrichous isorhizas (on gastrozooids, dacty lozooids, gonozooids; in nematocyst reservoir) 8.5-13.6 (xm x 3.8-4.7 |xm; haplonemes (on gastrozooids) 12.714.1 |xm x 6.8-7.6 jxm; stenoteles (small) (on gastro zooids, dactylozooids, gonozooids; in nematocyst reser voir) 12.2-15.1 |xm x 9.2-11.4 p,m; stenoteles (large) Fig. 59. Velella velella, nematocysts of hydroid, romiz B174. Scales equal 10 u,m. a, Atrichous isorhiza of gastrozooid. b, Haploneme of gastrozooid. c, Small stenotele of dactylozooid. d, Large stenotele of dactylozooid. e. Bulbous stenotele of gonozooid. (on gastrozooids, dactylozooids, gonozooids; in nemato cyst reservoir) 18.8-21.2 p.m x 13.3-15.0 |xm; steno teles (bulbous) (on gastrozooids, dactylozooids, gonozooids, medusa buds; in nematocyst reservoir) 18.8-23.4 [im x 16.0-21.5 (xm. Medusae not seen. REMARKS Hydroids of Velella velella (Linnaeus, 1758) occur in two enantiomorphic forms. Edwards (1966b) noted that dif fering and sometimes conflicting terminology has been used for these two. Edwards has been followed here in designating these as left-sailing and right-sailing forms. The left-sailing form, which drifts to the left in the down wind direction, corresponds to A. Agassiz's (1883) lefthanded form, Chun's (1897a) "SW" form, and Totton's (1954) "NW" form. The right-sailing form, which drifts to the right in the downwind direction, corresponds to Chun's (1897a) "NW" form and Totton's (1954) "SW" form. Both left-sailing and right-sailing forms were ob served washing ashore at Bermuda on 7 March 1982 after several days of strong southerly winds. The development of the hydroid of Velella velella was described by Woltereck (1904, 1905) and reviewed by Garstang (1946). Woltereck found young larvae of this species in deep water off Villefranche, the youngest of 83 which possessed a rudimentary, fluid-filled float; two short, solid tentacles; and a rudimentary mouth. Garstang re garded this larva as an actinula, homologous with that of tubularians. Brinckmann-Voss (1970) questioned this, not ing that the only similarity between this larva and an ac tinula was its possession of two aboral tentacles. Growth of this larva, known as a conaria, was accompanied by the development of a nettle-ring (the incipient nematocyst reservoir known as the "centradenia") and a crimson aboral cone of endoderm. Oil droplets, serving as a temporary method of flotation, are secreted by the crimson cone, and the conaria rises to the surface. Upon reaching the surface, fluid in the rudimentary float of the conaria is expelled and replaced by air, and the crimson cone disappears. At this stage, the larva is known as a rataria. In the rataria, the float enlarges and becomes lined with chitin, the ne matocyst reservoir or centradenia becomes solid, zooids begin development, and the sail appears. Continued growth of the rataria leads to the familiar hydroid of this species. Despite the abundance of the hydroid stage and the large numbers of medusa buds produced by each colony, me dusae of this species have seldom been collected in nature. Although they have zooxanthellae and are probably epipelagic, the medusae are small and likely easily over looked. The largest specimen collected to date was only 2.8 mm high and 2.0 mm wide (Larson, 1980). Brinck mann (1964) and Brinckmann-Voss (1970) described the development of the medusa in the laboratory. Rhythmic synchronous contractions, referred to as "concerts" by Fields and Mackie (1971), occur in hy droids of Velella velella (Vogt, 1854; Chun, 1897b; Fields and Mackie, 1971). During a contraction, dactylozooids are flexed towards the central gastrozooid, gonozooids shorten, and the mantle is contracted downwards. Im mediately after a contraction, the flexed and contracted parts relax and return to their original state. Fields and Mackie (1971) noted that such contractions may occur singly or in a series, and that concert periodicity in V. velella varied from one to three minutes. The function of such behaviour remains enigmatic. Garstang (1946) sug gested that it might facilitate the spreading of mucus threads used in prey capture. Fields and Mackie did not attribute such movements either to feeding, as suggested by Gar stang, or to locomotion. Because of the presence of large numbers of zooxanthellae in the tissues, they suggested that Chun's (1897b) hypothesis that it is a respiratory movement warranted further evaluation. KNOWN RANGE Bermuda: Castle Harbour (Fewkes, 1883); 14 km south east of Nonsuch Island (Totton, 1936); no specific locality given (Calder, 1986). Elsewhere: circumglobal, tropical and temperate waters (Totton, 1954; Edwards, 1966b; Brinckmann-Voss, 1970). Acknowledgements I am indebted to Dr Wolfgang Sterrer, former director of the Bermuda Biological Station, for encouraging me to study the hydroids of Bermuda, and for facilitating my research there in many ways. Funds covering the costs of fieldwork were initially provided by Exxon Corporation, and later by the Canadian Associates of the Bermuda Bio logical Station and by the Royal Ontario Museum. I am particularly grateful to Dr Anita Brinckmann-Voss, research associate at the Royal Ontario Museum, for ad vice, encouragement, and constructive criticism of this work. Special thanks are also extended to Dr P. F. S. Cornelius, of the British Museum (Natural History), for his thorough review of the manuscript, for many kind nesses during my visit there in 1986, and for confirming the identity of the type of Sertularia pennaria Linnaeus, 1758, as a species of thecate rather than an athecate. I also acknowledge the helpful comments of an anonymous re viewer. Drs Tom Iliffe, John Markham, and Wolfgang Sterrer provided specimens and information on collecting locales. Jack Lightbourn was most generous in inviting me on offshore collecting trips aboard North Star and 84 Polaris. Harry Barnes, Warren Smith, Michael Rhodes, and Sue Jickells of the Bermuda Biological Station staff provided help in collecting specimens. Loans of study material were provided by Ardis Johnston, Agassiz Mu seum of Comparative Zoology; Peter Davis, The Hancock Museum; Peter Frank, National Museum of Natural Sci ences; and David Rackley, United States Fish and Wildlife Service. I wish to thank Barbarann Ruddell, Maureen Mahler, and Stephen Campbell for assisting with the preparation of the manuscript. Librarians at the University of Toronto, the Royal Ontario Museum, and especially the British Mu seum (Natural History) provided help in locating and ob taining obscure references. 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Atlantide Report 5:211-325. 1964 Note on the distribution of Garveiafranciscana (Torrey, 1902) and Cordylophora caspia (Pallas, 1771) in the Netherlands. Zoologische Mededelingen 39:125- 1966 Skeletal structure in the Solanderiidae and its bearing 1844a Sur les genres Eleutherie et Synhydre. Bulletins de I'Academie Royale des Sciences et Belles-Lettres de Bruxelles 11:305-314. 1844b Recherches sur l'embryogenie des tubulaires, et l'his toire naturelle des differents genres de cette familie qui habitent la Cote d'Ostende. Nouveaux Memoires de I'Academie Royale des Sciences et Belles-Lettres de Bruxelles 17(6): 1-72. 1867 146. on hydroid classification. In Rees, W. J., ed., The Cnidaria and their evolution. London, Academic Press, pp. 373-396. 1967 The Hydroida and Chondrophora of the Israel South Recherches sur la faune littorale de Belgique (po lypes). Memoires de I'Academie Royale des Sciences, des Lettres et des Beaux Arts de Belgique 36:1-207. Red Sea Expedition, 1962. Israel South Red Sea Ex pedition, 1962, Reports 25:18-54. Reporton a collectionof Hydroidafrom the Caribbean region, including an annotated checklist of Caribbean hydroids. Zoologische Verhandelingen 92:1-124. Hydroids from the Theta, Vema and Yelcho cruises 1968 VANHOFFEN, E. 1911 Die Anthomedusen und Leptomedusen der Deutschen Tiefsee-Expedition 1898-1899. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer "Valdivia" 1898-1899 19(5): 191-233. vannucci ,M. 1951 1956 Hydrozoa e Scyphozoa existentes no Institute Paulista de Oceanografia. Boletim do Institute Paulista de Oceanografia 2(l):67-98. Biological notes and description of a new species of Dipurena (Hydrozoa, Corynidae). Proceedings of the Zoological Society of London 127:479-487. 1972 of the Lamont-Doherty Geological Observatory. Zoologische Verhandelingen 120:1-247. vogt, c. 1854 M6moires de l'lnstitut National Genevois 1:1-164. WALLACE, W. S. 1909 vannucci, m. and w. j. rees 1961 A revision of the genus Bougainvillia (Anthomedu sae). Boletim do Institute Oceanografico 11(2):57100. A collection of hydroids madeatthe Tortugas, during May, June, and July, 1908. Carnegie Institution of Washington, Year Book 7:136-138. warren, E. 1906a VERRILL, A. E. 1868 Recherches sur les animaux inferieurs de la Mediter- ranee. I. Sur les siphonophores de la Mer de Nice. On Halocordyle cooperi sp. nov., a hydroid from the Natal coast. Annals of the Natal Government Museum Notice of the corals and echinoderms collected by 1:73-81. Prof. C. F. Hartt, at the Abrolhos reefs, Province of Bahia, Brazil, 1867. Transactions of the Connecticut Academy of Arts and Sciences-1:351-371. 1906b 1900 Additions to the Anthozoa and Hydrozoa of the Ber mudas. Transactions of the Connecticut Academy of 1907 On Parawrightia robusta gen. et sp. nov., a hydroid from the Natal coast; and also an account of a sup posed schizophyte occurring in the gonophores. An 1902a Comparisons of the Bermudian, West Indian, and 1908 On a collection of hydroids, mostly from the Natal 1:83-96. Arts and Sciences 10:551-572. Brazilian coral faunae. Transactions of the Connect icut Academy of Arts and Sciences 11:169-206. 1902b The Bermuda Islands: their scenery, climate, produc tions, physiography, natural history, and geology; with sketches of their early history and the changes due to 100 On Tubularia solitaria sp. nov., a hydroid from the Natal coast. Annals of the Natal Government Museum nals of the Natal Government Museum 1:187-208. coast. Annals of the Natal Government Museum 1:269355. WATSON 1978 E. New species and new records of Australian athecate hydroids. Proceedings of the Royal Society of Vic toria 90:301-314. 1985 1937 The genus Eudendrium (Hydrozoa: Hydroida) from Australia. Proceedings of the Royal Society of Vic toria 97:179-221. Okologische Untersuchungen an Hydroiden des Felslitorals von Santa Marta (Kolumbien). Helgolander Wissenschaftliche Meeresuntersuchungen 27:324-363. WEDLER, E. and R. LARSON 1986 29B(7):l-6. 1947 1981 1904 Athecate hydroids from Puerto Rico and the Virgin Islands. Studies on Neotropical Fauna and Environ ment 21:69-101. 1905 1931 gen der Deutschen Zoologischen Gesellschaft 15:106— 122. WRIGHT, T. S. Le genre Pteroclava n.gen., 1'interpretation systematique des Pteronemidae (hydraires) et la valeur taxonomique du cnidome. Comptes Rendus Hebdomadaires des Seances de I'Academie des Sciences 182:60-62. Contribution a 1'etude des cnidaires et de leurs ne- matocystes. II. Valeur taxonomique du cnidome. Travaux de la Station Zoologique de Wimereux 11:351- 1858b 701. 1859 1936 Le cnidome des cladonemides Eleutheria dichotoma et Cladonema radiatum, son cycle et son interpreta tion. Comptes Rendus Hebdomadaires des Seances de 1861 I'Academie des Sciences 203:816-818. 1863a polype {Pennaria! tiarella McGr.) a tetracnidome. Comptes Rendus Hebdomadaires des Seances de I'A 1863b cademie des Sciences 204:1749-1752. 1937b Cladonema radiatum aux lies Bermudes. Contribu tion k l'6tude des m6duses Cladonema. Bulletin Bio- logique de la France et de la Belgique 71:438-465. weis, J. s. 1968 Fauna associated with pelagic Sargassum in the Gulf Stream. American Midland Naturalist 80:554-558. 1863c WEISMANN, A. 1883 Observations on British zoophytes.—(1.) On Atrac tylis (new genus); (2.) On the fixed medusoids of Laomedea dichotoma (living specimens were exhib ited); (3.) On the reproductive organs of the medusoid of Laomedea geniculata; (4.) On the reproduction organs of Laomedea lacerata. Proceedings of the Royal Physical Society of Edinburgh 1:447-455. Observations on British zoophytes. Edinburgh New Philosophical Journal, n.s., 7:282-287. Observations on British zoophytes. Edinburgh New Philosophical Journal, n.s., 10:105-114. Observations on British Protozoa and zoophytes. An nals and Magazine of Natural History, 3rd series, 8:120-135. Existence d'un monocnidome dans le medusoide d'un 1937a Ueber die Entwicklung der Velella aus einer in der Tiefe vorkommenden Larve. Zoologische Jahrbucher, Supplement 7:347-372. Bemerkungen zur Entwicklung der Narcomedusen und der Zool. Station in Villefranche s.m.). Verhandlun- 1858a WEILL, R. Phylogenetics; the theory and practice of phylogenetic systematics. New York, John Wiley. 439 pp. Siphonophoren. (II. Planktologische Mitteilungen aus Transplantation experiments with Caribbean Mille pora species (Hydrozoa, Coelenterata), including some ecological observations on growth forms. Bijdragen tot de Dierkunde 51:1-19. 1934 Notes on hydroids. Arkiv for Zoologi 39A(5):l-23. WOLTERECK, R. WEERDT, W. H. de 1981 Boreohydra simplex n.gen., n.sp., ein Solitarpolyp von der norwegischen Kiiste. Arkiv for Zoologi WILEY, E. O. WEDLER, E. 1975 WESTBLAD, E. Die Entstehung der Sexualzellen bei den Hydromedusen. Zugleich ein Beitrag zur Kenntnis des Baues und der Lebenserscheinungen dieser Gruppe. Jena, Gustav Fischer. 295 pp. Observations on British zoophytes. Proceedings of the Royal Physical Society of Edinburgh 2:59-63. Observations on British zoophytes.—(1.) On the re production of Tunis neglecta. (2.) On the develop ment of Hippocrene {Bougainvillea) Britannica{!) from Atractylis {Eudendrium) ramosa. (3.) On the devel opment of Hydra Tuba {Strobila) from Chrysaora. Proceedings of the Royal Physical Society of Edin burgh 2:34-36. Observations on British zoophytes and Protozoa. 1. Clava nodosa. 2. Acharadria larynx. 3. Zooteirea religata. 4. Freya {Lagotia) obstetrica, Freya stylifer. 5. Chaetospira maritima. 6. Oxytricha longicaudata. Proceedings of the Royal Physical Society of Edin burgh 2:378-381. WERNER, B. 1961 Morphologie und Lebensgeschichte, sowie Temperaturabhangigkeit der Verbreitung und des jahreszeitlichen Auftretens von Bougainvillia superciliaris (L. Agassiz) (Athecatae-Anthomedusae). Helgolan der Wissenschaftliche Meeresuntersuchungen 7:206- YAMADA, M. 1959 Hydroid fauna of Japanese and its adjacent waters. Publications from the Akkeshi Marine Biological Sta 1964 Rhizogeton ezoense n.sp., a new hydroid from Hok kaido, Japan. Journal of the Faculty of Science, Hok kaido University, series 6, Zoology 25:395-397. tion 9:1-101. 237. 1984 Stamm Cnidaria, Nesseltiere. In Hartwich, G., F. Kilian, K. Odening, and B. Werner, eds., Lehrbuch der Speziellen Zoologie, Band I: Wirbellose Tiere. 2. Teil: Cnidaria, Ctenophora, Mesozoa, Plathelminthes, Nemertini, Entoprocta, Nemathelminthes, Priapulida. Stuttgart, Gustav Fischer, pp. 11-305. YAMADA, M. and K. KONNO 1973 Polyp and medusa of the hydroid Sphaerocoryne mul titentaculata (Warren) from Japan. Publications of the Seto Marine Biological Laboratory 20:103-109. 101 Index Acharadia, 52 muscus, 24 Acharadria, 52, 53 ramosa, 24 larynx, 53 BOUGAINVILLEAE, 12 Bougainvillia, 19, 21, 23-24, 27, 36 CLADOCORYNIDAE, 73, 76 Cladonema, 66-67, 68 allmani, 67 Acrochordium, 64, 69 album, 69, 70, 72 benedenii, 24 californicum, 68 dujardini, 67 dujardinii, 67 gegenbauri, 67 britannica, 24, 27 krohnii, 67 Actigia, 63, 64 Actinogonium, 63, 64 pusillum, 64 Aglaophenia, 57 Aglatophenia pinnaria, 56 /fflvWfl, 24, 27 frondosa, 28 fruticosa, 24 gtofai ( 24 longicirra, 28 mayeri, 67, 68 myersi, 68 perkinsi, 67 perkinsii, 67, 68 macloviana, 23, 24 muscus, 1, 24-28 radiatum, 66, 67-68 radiatum allmani, 67 /i/ofo, 28 radiatum dujardinii, 67 radiatum gegenbauri, 67 Acharadrium, 52 Acies, 11 pa Ipebrans, 78 Amalthaea, 48 AMALTHAEIDAE, 48 AMPHINEMIDAE, 35 Anisocalyx pinnahum, 56 Archa radio, 52 Armenista, 80 antarctica, 81 indica, 81 autumnalis, 24, 27, 28 autumnalis var. magna, 25 platygaster, 28 pyramidata, 24, 27 ramosa, 1, 24, 27 ramosa f. fruticosa, 25 ramosa var. minima, 25 radiatum krohnii, 67 radiatum var. mayeri, 67 sp., 67 uchidai, 67, 68 /efrata, 81 ramosa ramosa ramosa ramosa mutica, 81 superciliaris, 27, 28 CLADONEMATIDAE, 66 CLADONEMIDEN, 66 Ctevfl, 10, 12, 52 parasiticum, 6 Clavactinia, 32, 33 sigmoides, 81 triestina, 25 Clavatella fata, 81 Armenistarium, 80 ve/<?//tf, 81 Aselomahs, 15, 19, 23 michaeli, 19 ASYNCORYNIDAE, 69 f. musca, 25 var. wwa, 25 f. ramosa, 25 f. vanbenedenii, 25 radiata, 67 vanbenedeni, 24 ww benedeni, 24 vanbenedenii, 27 ww benedenii, 25 v. benedenii, 24 Atractilis, 23 BOUGAINVILLIIDAE, 12, 13, 14, ATRACTYLIDAE, 12, 21 Atractylis, 23, 24 BOUGAINVILLIINAE, 12, 13, 15, arenosa, 19 ramosa, 24 £flte//tf, 13, 15 mirabilis, 13 BALELLIDAE, 13, 15 Bibrachium, 63 Bicorona, 63 Bimeria, 21, 23 humilis, 21, 23 umilis, 21 vestita, 21-23 V6\s7/'ta f. rttfflfl, 21 BIMERIDAE, 12 BIMERIIDAE, 12, 21 BIMERIINAE, 12, 18, 21 Boreohydra, 48 BOREOHYDRIDAE, 48 BOREOHYDRINAE, 48 Bougainvilla, 23 Bougainvillea, 23, 24 15, 18, 19, 23 18,23 BOUGAINVILLIOIDEA, 12 Bouganvilleia, 23 Bourgainvillea, 23 britannica, 24 Bourgainvillia, 23 BRANCHIOCERIANTHIDAE, 48 BRANCHIOCERIANTHINAE, 48 Branchiocerianthus, 48 BYTHOTIARIDAE, 35 multitentaculata, 61 CLAVIDAE, 5, 12, 13, 15, 19 CLAVINAE, 5 CLAVIPTERIDAE, 69 Clavopsella, 13, 14, 15, 17, 19 annulata, 15, 16 quadranularia, 15 weismanni, 15, 16 CLAVOPSELLIDAE, 12, 13, 15 CLAVOPSELLINAE, 13 Clavopsis, 32, 33 adriatica, 33 Clavula, 7, 8 gossii, 8, 9 Cnidostoma, 28 CODONIDAE, 63 CODONIIDAE, 63 Codonium, 63 Cordylophora, 5, 6, 15, 19 Calamella, 38, 52 CALYCOPSIDAE, 36 CALYCOPSIDI, 35 CANDELABRIDAE, 48 Capsularia, 63, 64 CHONDROPHORAE, 76 CHONDROPHOROIDEA, 76 Chrysomitra, 11 striata, 78 Cionistes, 32 annulata, 15 neapolitana, 15 CORDYLOPHORINAE, 5 Corina, 63, 64 Corine, 63, 64 CORYDENDRIINAE, 1, 5 Corydendrium, 5-6, 7, 10 dendriforme, 6, 7 dendriformis, 6 flabellatum, 6, 7 103 fruticosum, 1 uchidai, 31 distichum, 44 exiguum, 44 nutricula, 8 Cytaesis, 29 parasiticum, 6-7 parasticum, 6 Cwm, 29 eximium, 44 Cytheis, 29 fruticosum, 44 glomeratum, 40 gracile, 44 gr/#/w, 46 hargettii, 46 hargitti, 46 hyalinum, 43 infundibuliforme, 40 sessile, 6, 7 splendidum, 56 Corymbogonium, 39 capillare, 41 Corymorpha, 48, 49 solitaria, 49 Dendroclava, 1 Dendronema, 67 DENDRONEMIDAE, 66 DICODONIINI, 63 Dicodonium, 63 CORYMORPHIDAE, 48-49, 76 CORYMORPHINAE, 48, 49 Dicoryne, 23 capillare, 41 Coryna, 63, 64 DICORYNIDAE, 12 laxum, 44 motzkossowskae, 38, 40 Ctfryw. 6, 39, 52, 63-64, 65, 69, 70 Dicyclocoryne, 63 parvum, 41, 43 briareus, 70 DICYCLOCORYNIDAE, 63 filiformis, 66 Dipurena, 63, 65 hincksii, 66 rees/, 68 implexa, 21, 70 margarica, 21 (margarica) implexa, 70 strangulata, 68 Discalia, 11 primordialis, 78 ramosum, 24, 27, 41, 43, 44 tenellum, 42, 43, 44 te/we, 41, 43 EUDENDROIDAE, 38 Euphysa, 48 globator, 56 multitentaculata, 61 DISCALIDAE, 76 EUPHYSIDAE, 48 pelagica, 70 pintneri, 66 pw.s///tf, 64, 72 Disconalia, 11 EUPHYSINAE, 48 sargassicola, 1,3, 64-66 sessilis, 69, 72 stauridia, 67 stauridiai, 67 vanbenedenii, 64 agassizii, 70 CORYNIDAE, 60, 63, 66, 67, 73 CORYNIPTERIDAE, 68 agassizii, 70, 72 Corynopsis, 27 gastroblasta, 78 pectyllis, 78 ramifera, 78 Ectopleura, 52-53 franciscana, 21 larynx, 55 humilis, 21 minerva, 53, 54, 55 nutans, 21 pacifica, 53-55 sp., 53, 54 robusta, 19 Edendrium, 39 Eleutheria, 68 multitentaculata, 61 perkinsii, 67 Ctenaria, 69 Cubogaster, 29 Erudendium, 39 CwwVifl Eucoryna, 55 elegans, 56 octonaria, 10 Eucoryne, 55, 56, 63, 64 bougainvillii, 24 Cybogaster, 29 gemmascens, 29 Cytaeis, 29 CYTAEIDAE, 28 CYTAEIDIDAE, 12, 28-29, 31, 32 Cytaeidium, 29 Cy/am, 28, 29, 31 imperialis, 31 japonica, 29, 31 Garveia, 21, 23 dumortierii, 53 radiata, 67 ELEUTHERIIDAE, 66 a/den, 24, 27 Fistulana, 1, 39, 63, 64 Fistularia, 63, 64 elegans, 56 Eucorynus, 56 EUDENDRIIDAE, 12, 18, 38, 39, 46 Eudendrium, 23, 27, 38-39, 42, 43, 44, 46, 64 amboinensis, 46 angustum, 40 attenuatum, 44 Gemellaria, 69 implexa, 70 Gemmaria, 69 cladophora, 70 costata, 70 gemmosa, 70, 72 implexa, 70 implexa var. neapolitana, 70 sagittaria, 70 sagittata, 70 sp., 70 Globiceps, 55, 56 globator, 56 tiarella, 55, 56 Guentherella, 69 implexa, 70 Gymnocoryne, 69 coronata, 70 Gymnogonos, 48 aylingae, 40 bermudense, 1, 39-41 HALECIIDAE, 29 Halecium, 39 niotha, 31 capillare, 41-43 capillare var. mediterranea, 41 mu/a, 31 carbarn, 19, 22, 25, 43-45, 49, nassa, 31 50 pusilla, 29 sp., 29-31 tetrastyla, 29, 31 cochleatum, AA uchidae, 29, 31 currumbense, 40 104 cunninghami, 43 Halerella, 32 HALIMEDUSIDAE, 35, 36 Halobotrys, 63 Halocharis, 69 gemmosa, 70 spiralis, 70 Halocordile, 55 disticha, 56 Halocordyle, 23, 55-56 australis, 56, 58 cooperi, 56 disticha, 56-60 Leuckartiara w'r/dtf, 74 vestita f. /?#/?#, 21 pumila, 73 Linvillea, 60, 63 agassizii, 60, 61, 72 Lizusa, 23, 24 disticha var. australis, 56 8-ciliata, 24 frag His, 56 octocilia, 24 pennaria var. australis, 56 tiarella, 56 wilsoni, 56 HALOCORDYLIDAE, 55 Halybothrys, 63 Halybotrys, 63, 64 octociliata, 24 LIZUSIDAE, 12 Lizuza, 23 Lizzia, 29 blondina, 29 LIZZIINAE, 12 Hansiella, 32 Hermia, 63, 64 Hippocrene, 23 moniliformis, 73 vestita, 21 Manicella, 21 /wsctf, 21, 22, 23 HIPPOCRENIDAE, 12 MARGELIDAE, 12 Holothuria Margelis, 23, 24 ramosa, 73 schrammi, 13 squarrosa, 74, 76 trinitatis, 74 MILLEPORADAE, 72 MILLEPORIDAE, 72-73, 76 Mnestra, 69 implexa, 70 parasites, 69, 70 Modeeria multitentacula, 8, 9 multitentaculata, 8 nutricola, 8 nutricula, 8 MOERISIIDAE, 60 MONOCAULIDAE, 48 denudata, 11 autumnalis, 24 Monocaulus, 48 nuda, 11 principis, 24 Montipora gothica, 74 Myrionema, 38, 46 amboinense, 40, 46-48 spirans, 81 Holothurio denudata, 11 spirans, 81 HYBDOCORIDAE, 52 ramosa, 24 MARGELOPSIDAE, 76 Medusa, 11 duodecilia, 27 glandifera, 78 Hybocodon, 52 /zwdtf, 77 HYBOCODONIDAE, 52 ocilia, 27 Hydractinia, 32, 33 arge', 33 pruvoti, 33 octocilia, 24, 27 pocillum, 81 porpita, 11 HYDRACTINIDAE, 31 HYDRACTINIIDAE, 12, 14, 29. 31-32, 33 HYDRACTINIINAE, 32 HYDRACTINIOIDEA, 32 Millardiana, 13 Hydractomma, 32, 33 Hydranthea, 28, 29 longitentaculata, 13-14 Millepora, 2, 73 HYDRICHTHYIDAE, 36 HYDRICHTHYINAE, 35, 36 Hydrichthys, 36 mirus, 36 Hydrissa, 32, 33 Hydrocorella, 32, 33 Hydrocoryne, 63 HYDROCORYNIDAE, 63 Hydronema, 32 sexdecilia, 27 umbella, 11 velella, 81 amboinensis, 46 griffini, 46 hargitti, 46, 48 MYRIONEMIDAE, 38 MYRIOTHELIDAE, 48 NEMOPSIDAE, 12 Nemopsis, 23 Nigritina, 29 NIOBIIDAE, 35, 36 Merona, 5 alcicornis, 1, 73-76, 84 alcicornis var. cellulosa, 74 alcicornis corniculata, 73 alcicornis Crustacea, 73 alcicornis digitata, 73 alcicornis var. fenestrata, 14 alcicornis ramosa, 73 alcycornis, 73 Oceam'tf nutricula, 8 polycirrha, 8, 9 Oonautes, 69 ORTHOCORYNIDAE, 68 Pachycordile, 14 weismanni, 15 Pachycordyle, 1, 13, 14-15, 17, 19 annulata, 16 napolitana, 14, 15-16 neapolitana, 15 HYPOLYTIDAE, 48 alicornis, 73 Candida, 74 Hypolytus, 48 Hyppocrene, 23 carthaginiensis, 74 complanata, 74, 76 PACHYCORDYLINAE, 12, 13, 15, crista-galli, 73 PACHYCORDYLINI, 1, 12, 13 Palmipora, 73 Janaria, 32, 33 JANARIIDAE, 31 Kinetocodium, 32, 33 Koellikerina, 21 fasciculata, 21 Crustacea, 73 delicatula, 74 digitata, 14 esperi, 73 fasciculata, 73 fenestrata, 74 forskali, 73 gothica, 73 weismanni, 15, 16, 19 18 alcicornis, 73 fasciculata, 73 parasitica, 73 PANDAEIDAE, 35 PANDEIDAE, 35, 36 PANDEOIDEA, 35-36 Paracytaeis, 28 105 PARAGOTOEIDAE, 48 Parawrightia, 1, 18-19 robusta, 19-20 Parigonimus, 23 Parvanemus, 14, 15 degenerates, 15 Pelagiana, 36 trichodesmiae, 35, 36-38 Pennaria, 55, 56 adamsia, 56 "americana," 56 australis, 56, 58 Podocoryna, 27, 32, 33 fucicola, 32 simplex, 31 Podocoryne, 32 fl/<ten\ 1, 24, 27 borealis, 27 sarsii, 32 tubulariae, 27 PODOCORYNIDAE, 31 Polybrachionia, 11 mitrata, 81 pocillum, 81 /tar/*, 77 medusae, 78 Rhizodendrium, 1, 5, 10 ezoense, 11, 12 nudum, 11, 12 sterreri, 1, 10-12 Rhizogeton, 10 ezoense, 10 POLYPI, 52 fusiformis, 10, 11 nematophorus, 10 australis var. cooperi, 56 Porpalia, 11 nudum, 12 caulini, 56 PORPALIDAE, 76 nudus, 10 cavolina, 56 cavolini, 56 cavolinii, 56 disticha, 55, 56, 57 disticha var. australis, 56 Porpema, 11 Porpita, 2, 76, 77 appendiculata, 11 Rhizohydra, 32 distyeha, 56 "europea," 56 gibbosa, 56 inornata, 56 pacifica, 56 parasitica, 6 pennaria, 56 rosea, 56 symetrica, 56 symmetrica, 56 tiarella, 56, 57 wilsoni, 56 PENNARIIDAE, 55 Perarella, 28, 29, 32 linnaeana, 78 atlantica, 11 RHIZORHAGIINAE, 1, 12, 18 Rhizorhagium, 18, 19, 20 napolitanum, 15 chrysocoma, 11 /ttw's, 15 coerulea, 11 robustum, 19 forskahli, 78 forskalea, 11 fungia, 78 gigantea, 11 glandifera, 11 globosa, 11 granulata, 11 indica, 11 kuhlii, 78 linnaeana, 78 roseurn, 19 RHYSIIDAE, 32 Rosalinda, 69 ROSALINDIDAE, 69 RUSSELLIIDAE, 12 Stfrs/tf, 63, 64, 65, 67 lovenii, 66 mirabilis, 64 occulta, 66 moneta, 11 tubulosa, 64, 65, 66 SARSIADAE, 63 Sarsiella, 63 nuda, 11 Sertolara, 56 lutkeana, 78 mediterranea, 78 Perigominus, 23 Perigommus, 23 Perigonella, 32 Perigonemus, 23 pacifica, 11, 78 porpita, 1, 77-80 Perigonimus, 23, 24, 36 ramifera, 11 halecina, 39 reinwardtii, 78 parasitica, 6 pennaria, 56, 57, 84 cidaritis, 21 muscoides, 24 napolitanus, 15 neapolitanus, 15 vestita, 21 vestitus, 21 Perigonismus, 23 Perigonymus, 23 radiata, 11 umbella, 78 PORPITAE, 76 Porpitella, 11 caerulea, 78 coerulea, 78 pectanthis, 78 radiata, 78 parassita, 6 pennara, 56, 57 Sertularia, 52, 56 ramosa, 24, 27 Sertulariam parasiticam, 6 Silhouetta, 13, 15, 16-17, 19 puertoricensis, 17, 18 uvacarpa, 13, 17-18 Soleniopsis, 5, 6 dendriformis, 6 Sphaerocoryne, 60-61, 63 muscus, 24 ramosus, 24 Perinema, 64 PORPITELLIDAE, 76 PORPITIDAE, 76-77 PROTIARIDAE, 35, 36 Phyllidoce, 77, 80 Pteroclava, 69 Pteronema, 69 tedtff/, 60, 61-62 multitentaculata, 60, 61, 62 darwinii, 69 PTERONEMATIDAE, 69 PTERONEMATOIDEA, 76 PTILOCODIIDAE, 32 peterseni, 61 sp., 61 denudata, 11 porpita, 11 velella, 77, 81 Phyllodice denudata, 78 producta, 67 ve/W/a, 81 Plumularia pennaria, 56 Podocorella, 32 106 SPHAEROCORYNIDAE, 60, 63 Stauridia, 64, 66, 67 Rataria, 80 cordata, 81 cristata, 81 radiatum, 67 STAURIDIIDAE, 66 Stauridiosarsia, 63, 64, 67 Stauridium. 66 TE1SS1ER1DAE, 69. 73. 76 cladonema, 67 Staurocladia, 68 Thamnostoma, 21 russelli, 21 australis, 81 cattrina, 81 cyanea, 81 emarginata, 81 Staurocoryne, 63, 64 Steenstrupia, 48 THAMNOSTOMIDAE, 12 STEENSTRUPIINI, 48 THAMNOSTOMINAE, 12, 21 /((to, 81 Stipula, 63, 64 77w«, 39 limbosa, 81 Stomotoca, 36 pterophylla, 36 sp.. 21 savignii, 39 indica, 81 meridionalis, 81 Tiara, 36 mutica, 81 STOMOTOCINI, 35 T1AR1DAE, 35, 36 Stylactaria. 1, 32-33 Timoides oblonga, 81 oxyothone, 81 o.xyothone var. brachyothone, 81 oxyothone var. oxyothone, 81 pacifica, 81 patella, 81 patellaris, 81 pocillum, 81 pyramidalis, 81 arctica, 33 <•(/•£?, 33-35 agassizii, 36 T1MOID1DAE, 35, 36 carcinicola, 33 Tregoubovia, 32 clavifori lis, 33 TRICHORH1ZIN1. 48 hooperiu 35 TR1CHYDR1DAE. 35, 36 inermis, 33 TR1CHYDR01DEA. 36 ingolfi, 33 pisicola 33 Tubiclava, 5. 10 \w/7, 33 Stylactella 28, 29, 32 abyssict la, 32 spongicola, 32 vennico'a, 32 annulata, 15 radackiana, 81 fruticosa, 16 sandwichiana, 81 TUBIDENDR1DAE, 13 Tubularia, 49, 52 scaphidia, 81 septentrionalis, 81 dumortierii, 53 sinistra, 81 implexa, 70 spiralis, 81 spiralis, 81 subemarginata, 81 STYLACTIDAE, 28, 31, 32 muscoides, 1, 39, 64 Stylactis, 28, 32. 33 pacifica, 53 arge, 33 ramea, 39 tentaculata, 81 fuciola, 32 ramosa, 24, 27, 39, 64 tropica, 81 hoopei, 33 hooperi, 33 hooperii, 33 spongicola, 50 indica, 31 inermis, 32, 33 sp.. 33. 35 solitaria, 1, 49, 50 TUBULARIADAE, 52 TUBULARIAE, 52 TUBULARIIDAE, 49, 52, 76 TUBULARIOIDEA, 76 vermicola, 28 STYLASTERIDAE, 32, 72 TURR1DAE. 5 Styllactis hooperi var. minor, 33 Syncorine, 63 Syncoryna, 6, 63, 64 parasitica, 6 neglecta, 8, 9 Turritopsis, 5, 6, 7-8, 10 pusilla, 64 Syncoryne, 63 mirabilis, 64, 65, 66 77/rrfc, 5, 8 Zanc/ea, 69, 70 a/te, 69-72 cladophora, 70 hargitti, 70 implexa, 70, 72 inflexa, 70 nutricola var. pacifica, 8 nutricula, 5, 8-10 nutricula, 8 SYNCORYNIDAE, 63 Sytlactis gemmosa, 70, 72 sagittaria, 70 sessilis, 70. 72 ZANCLEIDAE, 64, 68-69, 73, 76 ZANCLEOIDEA, 77 Zanklea, 69 costata, 70 Ve/flWo, 80 indica, 81 mutica, 81 medusifera, 69 tnilleporoides, 69 Wrightia, 19 nutricola, 8 stauridium, 67 australe, 69 limbosa, 81 costata, 69, 70, 72 parasitica, 6 sp.. 64. 65, 66 Teissiera, 69 VW/e/fi. 80 fifo/wro7, 9, 10 fascicularis, 10 polycirrha, 8 polynema, 8, 9 Turrutopsis, 1 arge, 33 wfe/to, 1. 77. 78. 81-84 VELELL1DAE, 76 oblonga, 81 Vefe//a, 2, 76, 77, 80-81 antarctica, 81 aurora, 81 Zanlcea, 69 Zyzzygus, 49 Zyzzyzus, 49 solitarius, 49 spongicolus, 50 warreni, 1, 49-51, 84 Zyzzvzuz, 49 107