1 4 MAR /983 LIBRARY Bulletin of the xcgdlti British Museum (Natural History Zoology series Vol 42 1982 British Museum (Natural History) London 1983 Dates of publication of the parts No 1 25 March 1982 No 2 29 April 1982 No 3 27 May 1982 No 4 24 June 1982 ISSN 0007-1498 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Contents Zoology Volume 42 Page No 1 The tick collection (Acarina: Ixodoidea) of the Hon. Nathaniel Charles Rothschild deposited in the Nuttall and general collections of the British Museum (Natural History) By James E. Keirans 1 No 2 Hydroids and medusae of the family Campanulariidae recorded from the eastern North Atlantic, with a world synopsis of genera By P. F. S. Cornelius 37 No 3 Miscellanea A new species of free-living nematode from the Firth of Clyde, Scotland By P. J. D. Lambshead 149 Notes on Atlantic Asteroidea 2. Luidiidae By A. M. Clark 157 New and little known species of Oncaeidae (Cyclopoida) from the Northeastern Atlantic By S. J. Malt 185 Larval and post-larval development of the Slender-legged Spider Crab, Macropodia rostrata (Linnaeus), (Oxyrhyncha: Majidae: Inachinae), reared in the laboratory By R. W. Ingle 207 No 4 Miscellanea New species of marine nematodes from Loch Ewe, Scotland By H. M. Platt and Z. N. Zhang 227 The larval development of Crangon crangon (Fabr. 1 795) (Crustacea : Decapoda) By A. R. Gurney 247 A revision of the spider genus Cocalodes with a description of a new related genus (Araneae: Salticidae) By F. R. Wanless 263 Anatomy and evolution of the jaws in the semiplotine carps with a review of the Genus Cyprinion Heckel, 1843 (Teleostei: Cyprinidae) By G. J. Howes .299 Bulletin of the British Museum (Natural History) The tick collection (Acarina: Ixodoidea) of the Hon. Nathaniel Charles Rothschild deposited in the Nuttall and general collections of the British Museum (Natural History) James E. Keirans With a Foreword by the Hon. Miriam Rothschild Zoology series Vol 42 No 1 25 March 1982 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus, not. Hist. (Zool.) © Trustees of the British Museum (Natural History), 1982 The Zoology Series is edited in the Museum's Department of Zoology Keeper of Zoology : Dr J. G. Sheals Editor of Bulletin : Dr C. R. Curds Assistant Editor : Mr C. G. Ogden ISSN 0007-1498 Zoology series Vol 42 No 1 pp 1-36 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 25 March 1982 GENERAL The tick collection ( Acarina: Ixodoidea) of thV ^ 1982 \ -4 LIBRARY "^» Hon. Nathaniel Charles Rothschild deposited in ^zy the Nuttall and general collections of the BritisBT Museum (Natural History) James E. Keirans Department of Health and Human Services, Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Rocky Mountain Laboratory, Hamilton, Montana 59840, U.S.A. With a Foreword by the Hon. Miriam Rothschild Contents Synopsis 1 Foreword 3 Introduction 4 PART I Rothschild specimens donated to Professor G. H. F. Nuttall ... 5 Species of Ixodoidea represented in the Nuttall collection, British Museum (Natural History), donated by the Hon. N. C. Rothschild .... 5 Rothschild specimens in the Nuttall collection with type status ... 6 Rothschild specimens in the Nuttall collection 6 A single tick collection given by the Hon. Walter Rothschild to G. H. F. Nuttal via Guy A. K. Marshall 17 Rothschild specimens missing from Nuttall collection 17 PART II Rothschild specimens donated to British Museum (Natural History). 20 Species of Ixodoidea represented in the collection of the British Museum (Natural History), donated by the Hon. N. C. Rothschild 20 Rothschild specimens in the British Museum (Natural History) collection with type status 21 Rothschild specimens in British Museum (Natural History) collection . . 22 Rothschild specimens missing from British Museum (Natural History) collection 32 Locality list for ticks in the Rothschild collection 32 Host list for ticks in the Rothschild collection 34 Acknowledgements 35 References 36 Synopsis The Hon. N. C. Rothschild achieved worldwide fame for his study of the Siphonaptera. During the early part of this century he also made a significant collection of ticks which he gave to Professor George H. F. Nuttall at Cambridge University. Upon Nuttall's death, the collection was given to the British Museum (Natural History) and is kept as a separate entity. Bull. Br. Mus. nat. Hist. (Zool.) 42( 1 ): 1 -36 Issued 25 March 1 982 J. E. KEIRANS Male Ixodes uriae White, 1852. (Ixodoidea: Ixodidae). A tick with a circumpolar distribution in both the northern and southern hemispheres. It is ectoparasitic on a variety of seabirds and to date, twenty different arboviruses have been recovered from this species. ROTHSCHILD TICKS IN BM(NH) 3 In addition, Rothschild on numerous occasions between 1911 and 1923 deposited ticks in the British Museum (Natural History). Herewith are recorded the 197 tick collections of the Hon. N. C. Rothschild plus one of the Hon. Walter Rothschild deposited in the Nuttall collection and the 156 collections in the general collection of the British Museum (Natural History). Foreword In 1913 N. C. Rothschild presented his collection of 'Siphonaptera and other parasitic insects' to the British Museum (Natural History). One of the conditions attached to the gift read as follows: 'The Trustees shall so soon as the said collection comes into their hands or is placed under their control and as soon as the funds placed at their disposal by Parliament will allow cause to be made and published a catalogue giving the names and full number of specimens of each species of parasitic insect contained in the said collection.' Over 60 years has now elapsed since the collection was accepted, but the catalogue has not yet been completed. Although ticks are arthropods, they are not insects, but this group of animals, along with a number of mites, was included in the collection. During my period of Trusteeship of the British Museum (Natural History) I had the opportunity of discussing this problem with Harry Hoogstraal, and he suggested that James Keirans might undertake to catalogue the Rothschild ticks. To my great delight he agreed to do this, although the task proved more time-consuming than we had at first anticipated. This was because the Rothschild Collection had not been preserved as a single unit, and the ticks, along with the Nycteribiidae, Cimicidae, Hippoboscidae and other smaller groups, had been distributed in the general collection and, in addition, had not all received Museum accession numbers. Although these circumstances added enormously to James Keirans' labours, they made the compilation of this Catalogue even more necessary and valuable. Until those who are engaged in ecological and medical research come to use museum material such as this, no one can really appreciate the immense boon of well-catalogued individual collections. Since, between 1900 and 1910, N. C. Rothschild had given Professor Nuttall the bulk of his tick material*, it was considered more appropriate to merge the two catalogues into one, and the manuscript was therefore divided into two parts: I, the specimens collected by Rothschild, now in the Nuttall collection; and II, the specimens donated by Rothschild to the British Museum (Natural History). The ticks in the general collection have been added — a course we followed when cataloguing the fleas (G. H. E. Hopkins & M. Rothschild, Vols. I-V, 1953-1 971) and the Nycteribiidae (Oscar Theodor, 1967). It will be noted that Walter Rothschild contributed one specimen to this collection. It has not usually been appreciated that N. C. Rothschild's brother also added considerably to the flea material, although he did not describe any species himself. The N. C. Rothschild donations of Nycteribiidae to the British Museum contained 56 undescribed species, and the Ixodoidea collections 10. The two brothers and their collaborators described over 5000 new species, but they probably collected twice that number of animals new to science. James Keirans has added greatly to the value of this compilation by providing information concerning the viruses and in certain cases rickettsiae of which the Rothschild ticks are known vectors. The illustration depicts a male specimen of Ixodes uriae White, one of the most spectacular species, adorned with a handsome fringe along the posterior margin, which has the dubious distinction of carrying at least 20 known viruses in five different serogroups. *After the sudden death of Professor Nuttall in December 1937 his collection was presented 19 the British Museum (Natural History) by the Molteno Institute. In accordance with the terms of the gift this collection is maintained as a discrete entity. 4 J. E. KEIRANS Altogether the meticulous care and accuracy with which James Keirans has completed this catalogue is beyond praise. Miriam Rothschild Introduction Early in the twentieth century, Nathanial Charles Rothschild acquired a large tick collection, primarily from collectors who, knowing of his great interest in fleas, would send him not only that group but also other ectoparasites including flies, bugs, and ticks. He deposited his tick collection in two institutions; the Molteno Institute for Research in Parasitology, Cambridge University, where Professor G. H. F. Nuttall made the original determinations of all tick material, and the British Museum (Natural History), where Mr A. S. Hirst made the tick determinations. As mentioned earlier, the Nuttall collection is now at the BM(NH). Because the Nuttall collection is kept as a discrete entity, separate from but housed with the BM(NH) collection, parts of Rothschild's tick collection are in each of the above collections. This catalogue is also divided into two sections: Part I, Rothschild tick donations to the Nuttall collection; Part II, Rothschild tick donations to the BM(NH). The systems for numbering tick collections are different for the two sections. In Part I, the number used is that given by Professor Nuttall to a collection. The first entry is N (Nuttall) 72 followed by N73, N74, etc., in ascending sequence, first for existing specimens, then for specimens which are now missing. Each Nuttall number is followed by a Rothschild collection number or the word 'None' if the collection received no Rothschild number. A Nuttall number refers to a handwritten entry in Professor Nuttall's tick catalogue, which is now kept in the Arachnida and Myriapoda Section of the BM(NH). The locality and host data are cited in their modern forms, but where these could not be established Nuttall's own entries are quoted. One additional point should be made relating to the Nuttall collection forming Part I. Nathaniel C. Rothschild donated all specimens recorded in Part I with the single exception of N888 which was donated by his brother, the Hon. Walter Rothschild, to the Entomo- logical Research Committee and subsequently to Professor Nuttall. Collections in Part II are entered alphabetically by genus, first Argasidae, then Ixodidae. All information is presented as in Part I with the exception that replacing a Nuttall and Rothschild number will be a British Museum (Natural History) registration number. Where no registration number was given to a collection, the word 'None' will appear. I have given each collection a set of Keirans numbers, K1-K156. These numbers, along with the Nuttall numbers from Part I, will allow the reader to use the tick species, locality, and host lists and refer back to individual collections within Parts I and II. Throughout the text, citations are given to publications which mention a particular taxon. My comments are presented within square brackets [ ]; those comments by Nuttall or some other investigator are cited within square brackets enclosed by quotation marks '[ ]'. ROTHSCHILD TICKS IN BM (NH) PARTI Rothschild specimens donated to the Nuttall collection Species of Ixodoidea represented in the Nuttall collection, British Museum (Natural History), donated by the Hon. N. C. Rothschild Nuttall numbers follow each species. Numbers in parentheses refer to missing collections. Thus, validity of original determinations is unverified. ARGASIDAE Argas boueti Robaud and Colas-Belcour 1 164 Argas persicus (Oken) 589, 590, (3536)) Argas reflexus (Fabricius) (3 1 59) Argas vespertilionis (Latreille) 591, 592, 593, 1151,1152,1153, 1163 Argas sp. 11 54, (1388) Ornithodoros capensis Neumann group 594 Ornithodoros moubata (Murray) 595 Otobius megnini (Duges) (596) IXODIDAE Amblyomma albolimbatum Neumann (78), 567, 569 Amblyomma australiense Neumann 547 Amblyomma cajennense (Fabricius) 557, 648% Amblyomma clypeolatum Neumann (75) Amblyomma compressum Macalister 564 Amblyomma cyprium Neumann 73, 565 Amblyomma decoratumC. L. Koch (93) Amblyomma dissimile C. L. Koch 568 Amblyomma gemma Donitz 558b Amblyomma geoemydae (Cantor) 545 Amblyomma hirtum Neumann (397) Amblyomma longirostre(C. L. Koch) 560 Amblyomma marmoreum C. L. Koch (77), 570 Amblyomma ovaleC. L. Koch 562, 566 Amblyomma pecarium Dunn 557 Amblyomma tholloni Neumann 559 Amblyomma triguttatum C. L. Koch 546, (551), 552,553 Amblyomma varium C. L. Koch (3351) Amblyomma sp. 74, (76), (544), 633, 1 166, 1386, 1391, 1392, 1393, 1394, 1396, 1397 Aponomma decorosum (L. Koch) 550 Aponomma gervaisi (Lucas) (92), (548) Aponomma gervaisi var. lucasi (Warburton) (90), (91) Aponomma hydrosauri (Denny) 556 Aponomma varanensis (Supino) 554 Aponomma sp. (57 1 ), 3 1 38 Boophilus decoloratus (C. L. Koch) (3157) Boophilus microplus (G. Canestrini) 588 Dermacentor albipictus (Packard) 666, 1105, 1497, 1498, 1499, 1500, 1501, 1502, 1503 Dermacentor andersoni Stiles 1387, 1390,3502 Dermacentor reticulatus (Fabricius) 3 1 60 Dermacentor rhinocerinus (Denny) 555, 563 Dermacentor sp. 3330 Haemaphysalis bispinosa Neumann 574, 582, 583,892 Haemaphysalis campanulata Warburton 579 Haemaphysalis celebensis Hoogstraal, Trapido and Kohls 72 Haemaphysalis elongata Neumann 670 Haemaphysalis erinacei Pavesi 3537, 3538, 3539 Haemaphysalis humerosa Warburton and Nuttall 669 Haemaphysalis hystricis Supino 572, 573, 575, 576 Haemaphysalis indica Warburton 575x Haemaphysalis leachi (Audouin) (539), 577, 578 Haemaphysalis punctata G. Canestrini and Fanzago 580 Haemaphysalis tiptoni Hoogstraal 670 Haemaphysalis sp. 58 1 Hyalomma aegyptium (Linnaeus) 549, 56 1 Hvalomma rufipes C. L. Koch 558a, 888 (W. Rothschild) Hyalomma syriacum C. L. Koch (3139), (3140), (3141) Hyalomma truncatum C. L. Koch 1 167 Hyalomma sp. 1398, (3 136) Ixodes australiensis Neumann 645, 646 Ixodes boliviensis Neumann 637 Ixodes brunneus C. L. Koch (396) Ixodes cookei Packard 1 504 Ixodes fecialis Warburton and Nuttall 650, 1211, 1213 Ixodesfrontalis (Panzer) 523 Ixodes hexagonus Leach (216), (217), (298), (322), (324), 597, 598, 599, 600, 601 , 602, 603, 604, 606, 607, 608, 609, 610, 611, 612, 613, (674), (746), 1067, 1159, 1160 Ixodes holocyclus Neumann 643, (644), 1212 Ixodes lividus C. L. Koch 605, 1068 J. E. KEIRANS Ixodes loricatus Neumann 638, 639, 640, 64 1 Ixodes luciae Senevet 647 Ixodes nitens Neumann 360 Ixodes putus (O. Pickard-Cambridge) (516), (5 17), (5 18) Ixodes ricinus (Linnaeus) (509), (628), (629), (630), (631), (632), (656) Ixodes rothschildi Nuttall and Warburton 634 Ixodes rubidus Neumann 826 Ixodes scapularis Say 626 Ixodes texanus Banks 1 399 Ixodes trianguliceps Birula 619, (620), 621, 622, 623,624,625,1069, 1161 Ixodes unicavatus Neumann (395), 652, 667 Ixodes uriae White 617,618 Ixodes vespertilionis C. L. Koch 649 Ixodes vestitus Neumann 642 Ixodes sp. 1066, 1 162, 1 167, 1389 Rhipicephalus evertsi Neumann (3 1 57) Rhipicephalus haemaphysaloides Supino 584, (3158) Rhipicephalus muehlensi Zumpt 585 Rhipicephalus sanguineus (Latreille) (586), (587) Rhipicephalus sp. 1 395 Rothschild specimens in the Nuttall collection with type status NUTTALL ROTHSCHILD No. 72 360 565 634 647 650 669 No. 916 None 243 159 248 203 178 Haemaphysalis celebensis Hoogstraal, Trapido, and Kohls, 1965. J. Paras it. 51: 1001, figs. 1-9. Ixodes nitens Neumann, 1904. Archs. Parasit. 8: 459. Amblyomma quasicyprium Robinson, 1926. Ticks. A Monograph of the Ixodoidea Pt. 4: 237, fig. 1 17. Ixodes percavatus rothschildi Nuttall and Warburton, 1911. Ticks. A Monograph of the Ixodoidea Pt. 2: 22 1 . Ixodes loricatus spinosus Nuttall, 1910. Parasitology, Cambridge 3: 411, fig. 5. Ixodes fecialis Warburton and Nuttall, 1909. Parasitology, Cambridge 2: 58, figs. 1-2. Haemaphysalis humerosa Warburton and Nuttall, 1909. Parasitology, Cambridge 2: 60, figs. 4-5. Rothschild specimens in the Nuttall collection N72 R916 1 9 Haemaphysalis hystricis (Haemaphysalis celebensis HOLOTYPE) ex. Sus celebensis (Sus verrucosus celebensis) Celebes. No date (Sulawesi 02WS, 121WE, Indonesia) Presented as mounted specimen No. 916 which we unmounted 4.1. 1 9 1 5, C. W. [Cecil Warburton] det. 8.1.1915. Publications: Nuttall & Warburton (1915: 425, 426); Hoogstraal el al. (19656: 1001); Hoogstraal etal. (1973: 556). N73 R915 1 d Amblyomma caelaturum [This species epithet crossed out and cyprium inserted] (Amblyomma cyprium) ex. Sus celebensis (Sus verrucosus subsp.) Celebes. No date (Sulawesi 02WS, 121°00'E, Indonesia) Presented as slide No. 9 1 5 which was unmounted, being defective 5.1.1915. 3 d1 Amblyomma cyprium mounted as slides, 2 transparent, 1 opaque — see slides No. 73 (i) (ii) (iii) (Nos. 917, 919, 920) (3 slides) (L. E. Robinson det. 26.1. 1915) redet. 26.111. 1919. Note: The 3 slide-mounted males were missing from the Nuttall collection in 1977. Publication: Robinson (1926: 236). N74 R918 1 N Amblyomma (? caelaturum Cooper & Robinson) (Amblyomma sp.). Unmounted ROTHSCHILD TICKS IN BM(NH) from slide labelled as off Sus celebensis (Sus verrucosus subsp.) Celebes. No date (Sulawesi 02WS, 121WE, Indonesia (L. E. Robinson det. 26.1. 1915). N360 None 1 9 Ixodes nitens COTYPE (Ixodes nitens SYNTYPE) ex. Mus macleari (Rattus macleari) Christmas Island, Pacific Ocean (10°39'S, 105°40'E) (ex. Shipley coll. No. 2) Publications: Neumann (1904: 460); Nuttall & Warburton(1911: 185). N523 None 1 9 Ixodes brunneus (minus hypostome, digits) (Ixodes fron tails) ex. Passer montanus Saint-Genies-de Malgoires (43°57'N, 04°13'E), Card, France 16.XI. 1908 Albert Hugues Publication: Nuttall & Warburton (1911:1 92). N545 R258 Amblyomma sp. [sex/stage not given] (1 9 Amblyomma geoemydae) (tentative deter- mination) Found on ground at Paquil, Luzon, Philippine Islands (locality not verified) 15.1.1895 A. Everett Publication: Robinson (1926: 48) recorded thfs as A. americanum. N546 R174 Amblyomma triguttatum [sex/stage not given] (4 9 Amblyomma triguttatum) ex. kangaroo Barrow Island (20°48'S, 115'23'E), N. W. Australia XI. 1900 C. J. T. Tanney. det. L. E. Robinson Publication: Robinson (1926: 57) indicated d's present but not in Nuttall collection in 1977. N547 R274 6 d, 1 9 Amblyomma australiense ex. Echidna aculeata(Tachvglossus aculeatus] Mt Anderson ( 1 7°58'S, 1 24;04'E), W. Australia [No date] C. J. T. Tanney Publications: Robinson (1926: 135, 136) illustrated d and 9 from this lot. Taylor (1946: 100) repeated Robinson's descriptions and illustrations. N549 R254 Hyalomma syriacum [sex/stage not given] (4 cf Hyalomma aegyptium) ex. tortoise No data Note: An additional 1 d, 1 9 of this collection sent to New York National Museum, New York City, 18.IV.1913. N550 R196 Aponomma decorosum [sex/stage not given] (2 d Aponomma decorosum) ex. Australian monitor Note: A. decorosum is found only in Australia and ranges from Queensland south to Victoria. N552 R179 Amblyomma triguttatum [sex/stage not given] (2 d, 3 9 Amblyomma triguttatum) ex. kangaroo Barrow Island (20°48'S, 115'23'E), N. W. Australia 20.XI.1900C.J. T. Tanney Publication: Robinson (1926: 57). N553 R211 Amblyomma triguttatum [sex/stage not given] (4 9, 13 N Amblyomma triguttatum) ex. kangaroo Barrow Island (20°48'S, 115'23'E), N. W. Australia 18.XI.1 900 C.J.T. Tanney Publication: Robinson (1926: 57). N554 R236 Aponomma exornatum [sex/stage not given] (2 d Aponomma varanensis) ex. Monitor strix [name not verified] ? Australia Note: Collection probably not from Australia. The only collection of A. varanensis from Australia is the type d of A. quadratus (an inornate form of A. varenensis). This single record may have been an accidental importation of tick and host. N555 R259 Dermacentor rhinocerotis [sex/stage not given] (7 d Dermacentor rhinocerinus) No other data Note: There is an additional d of this species mounted on a slide. N556 R287 Aponomma hydrosauri [sex/stage not given] (2 d, 1 9 Aponomma hydrosauri) ? Adelaide (34°55'S, 1 38°35'E), W. Australia N557 R198 Amblyomma cajennense [sex/stage not given] (1 d, 2 9 Amblyomma cajennense; 3 d Amblyomma pecarium) ex. peccary Tayassu sp. Santa Andrea, Tabasco 30. V.? '[No particulars]' Note: The above locality information was taken from the vial label. It should be San Andres, Estado de Tabasco ( 1 8WN, 92°40'W), Mexico. 8 J. E. KEIRANS Publication: Robinson (1926: 53) incorrectly gave the Nuttall number as 577. Amblyomma cajennense is a vector of Wad Medani virus and the rickettsia of Rocky Mountain spotted fever. N558a R263 1 d, 1 9 Hyalomma aegyptium (Hyalomma rufipes) ex. giraffe, Giraffa camelopardalis No other data Hyalomma rufipes is the vector of the viruses Dugbe, Tete, Matruh, and Crimean Congo hemorrhagic fever. N558b R263 2 d Amblyomma gemma ex. giraffe, Giraffa camelopardalis No other data. Amblyomma gemma is a vector of Nairobi sheep disease virus. N559 R285 1 9 Amblyomma tholloni No host Mt Ruwenzori (00°23'N, 29°54'E), on the Uganda-Zaire border, Africa 1906 A. F. R. Wollaston N560 R260 1 d Amblyomma longirostre ex. Coendou simonsi (Coendou bicolor simonsi) Charapaya(17°22'S, 66°45'W), Bolivia 22.VI.1901 P.O.Simons Publication: Robinson (1926: 140). N561 None Hyalomma syriacum [sex/stage not given] (4 d Hyalomma aegyptium) ex. Algerian tortoise N. C. Rothschild N562 R261 Amblyomma ovale [sex/stage not given] (2 d Amblyomma ovale) ex. Speothas venaticus Charapaya(17°22'S,66'45'W), Bolivia 27.VI.1901 P.O.Simons N563 R266 10 d '[small and large]' Dermacentor rhinocerotis (7 cf Dermacentor rhinocerinus) ex. rhinoceros No other data Note: Nuttall, in his notebook containing infor- mation on ticks sent to various individuals and institutions, indicated that this collection was from Nyasaland, British Central Africa [Malawi] in 1907. One additional d of this collection was sent to Dr J. G. Parham, Marromeu, Zembezia, Portuguese East Africa [People's Republic of Mozambique], 18.IV.1913 and one additional d was sent to New York National Museum, New York City, 18.IV.1913. N564 R218 Amblyomma badium [The species epithet crossed out and sublaeve inserted] (6 d Amblyomma compressum) ex. white-bellied pangolin, Manis tricuspis No other data L. E. Robinson det. 1926 N565 R243 1 9 Amblyomma quasicyprium Robinson, 1926 TYPE (Amblyomma cyprium) ex. Spider monkey Ateles melanochoerus [This species epithet is crossed out and melanochir inserted] (Ateles g. geoffroyi) Frontera, ( 1 8°32'N, 92°38'W), Tabasco, Mexico Notes: Both host and locality are incorrect for Amblyomma cyprium although this 9 appears to be true A. cyprium. Nuttall 575 and 3330 also bear Rothschild number R243. Publications: Robinson (1926: 237, 238) incorrectly gave the Nuttall number 564 for this collection. Taylor (1946: 116) repeated Robinson's error for the Nuttall number. N566 R270 1 9 Amblyomma ovale ex. Speothas venaticus Charapaya (17'22'S, 66'45'W), Bolivia Publication: Robinson (1926: 29) gave the date as 1900 and indicated that the collection contained only males. N567 R282 Amblyomma albolimbatum [sex/stage not given] (2 d1, 1 9 Amblyomma albolimbatum) ex. Morelia variegata (Morelia spilotes variegatd) Perth (3 1 °56'S, 1 1 5°50'E), W. Australia. Woodward L. E. Robinson det. 1926 N568 R231 Amblyomma [species epithet integrum crossed out and dissimile inserted]: [sex/stage not given] (7 d, 2 9, 7 N Amblyomma dissimile) ex. large snake Probably Frontera (18'32'N, 92°38'W), Tabasco, Mexico May 1 . 1 .V. 1 90 1 [The full date is on vial label but not in catalogue] Publication: Robinson (1926: 167) indicated the date for this collection was V.I 900 and that only one female was present. N569 R267 Amblyomma moreliae [sex/stage not given] [This species epithet crossed out and albolimbatum inserted] (1 9 Amblyomma albolimbatum) ROTHSCHILD TICKS IN BM(NH) ex. Diemenia superciliosa (Diemenia textilis) Herdman's Lake, near Perth (31'56'S, 1 15'15'E), W. Australia Det. Neumann & Warburton Publication: Robinson (1926: 226) figured this 9; on p. 227 he incorrectly gave the collection number as N 1569. N570 R182 2 cf Amblyomma marmoreum leopard tortoise [Africa] No other data L. E. Robinson det. Note from Mr. Robinson 17. XI. 1922: 'They originally showed the characteristic ornamen- tation, sufficient of which is still visible, especially if examined under alcohol, to leave no doubt. The coarse punctations are larger than those seen in specimens off Rhinoceros in East Africa, and the dark ornamentation is more salient, but I do not consider these differences to be of specific impor- tance. The other morphological characters are typical.' N572 R288 Haemaphysalis hystricis [sex/stage not given] (3 9 Haemaphysalis hystricis) ex. Canis familiaris Lhassia, Assam, India (coordinates for Assam — 26°00'N,93°00'E) 1907.R. A. Lorrain Note: Nuttall sent 1 cf, 1 9 of this collection to Or L. O. Howard of the U.S. Department of Agriculture. These specimens are now deposited in the Rocky Mountain Laboratory collection (RML 109001). Publications: Nuttall & Warburton (1915: 425); Hoogstraal et al. (19650: 476) gave combined collection data from Nuttall 572 and 573. N573 R293 Haemaphysalis hystricis [sex/stage not given] (2 9 Haemaphysalis hystricis) ex. Canis familiaris Lhassia, Assam, India (coordinates for Assam — 26WN, 93WE) 1907.R. A. Lorrain Publication: Hoogstraal et al. (19650: 476) gave combined data from Nuttall 572 and 573. N574 R294C Haemaphysalis bispinosa [sex/stage not given] (10 cf, 6 9, 1 N Haemaphysalis bispinosa) No other data Haemaphysalis bispinosa is a vector of Kyasanur forest disease virus. N575 R243 9 Haemaphysalis hystricis (2 9 Haemaphysalis hystricis)) Note: Nuttall 565 and 3330 also bear Rothschild number R243. N575x R163 Haemaphysalis leachft [sex/stage not given] (1 cf Haemaphysalis indica)' ex. Hyaena crocuta (Hyaena hyaena) Darie (locality not verified) (Exp. Erlanger Hilgert) Note: The Nuttall number for this collection is 575x in his catalogue but 575a on the vial label. Nuttall 1395 also bears Rothschild number R 163. N576 R294a Haemaphysalis hystricis [sex/stage not given] (1 9, 1 N Haemaphysalis hystricis) ex. Canis familiaris Lushai Hills (23'10'N, 92°50'E), Assam, India 1907.R. A. Lorrain Publications: Nuttall & Warburton (1915: 425) indicated that N576 was composed of 1 cf, 3 9. Hoogstraal et al. (1965a: 476) saw the 9 labeled N576ii. The nymph, in a separate vial bears the label N576i. N577 R294 Haemaphysalis leachi [sex/stage not given] (1 cf, 1 9 Haemaphysalis leachi) ex. Ictonyx capensis (Ictonyx striatus) Bulawayo (20°09'S, 28°35'E), Zimbabwe 3.IV.1907E.C.Chubb Notes: The genus Ictonyx is also known in the literature as Zorilla. I. Geoffrey, 1826. Nuttall 595 also bears Rothschild number R294. Publication: Nuttall & Warburton (1915:471). N578 R154 Haemaphysalis leachi (?) (not long enough) [sex/stage not given] (2 cf, 1 9 Haemaphysalis leachi) Deelfontein (20'17'S, 32°38'E), Zimbabwe 1. IV. 1 902 C.J.B. Grant Publication: Nuttall & Warburton (1915: 472) indicated the genus as Suricata. However, the collection locality appears to be out of the known range of the meerkat. N579 R212 Haemaphysalis campanulata [sex/stage not given] (2 cf, 1 9 Haemaphysalis campanulata) ex. Canis familiaris Yokohama (35°26'N, 139837'E), Japan 24. IV. 1902 Allan Owston Note: Nuttall 609 also bears the Rothschild number R212 but the data do not correspond to N579. Publication: Nuttall & Warburton (1915: 472) indicated the collection contained 2 cf, 2 9 from a dog. 10 J. E. KEIRANS N580 R160 1 cf 1 9 Haemaphysalis punctata [no host data] Cambridge (52° 1 3'N, 00'08'E), England A. E. Shipley (Nuttall det.) Haemaphysalis punctata is a vector of the viruses Bhanja, tickborne encephalitis, Crimean Congo hemorrhagic fever, and perhaps Tribec. N581 R183 1 N Haemaphysalis spinulosal [The species epithet crossed out and sp. inserted] (Probably H. leachi var. indica) (1 N Haemaphysalis sp.) (Bad condition, very queer. C. Warburton) ex. flying squirrel Bunguran (possibly Bungura 26°20'N, 84'05'E), India N582 R284 Haemaphysalis bispinosa [sex/stage not given] (2 9 Haemaphysalis bispinosa) ex. Equus caballus Lushai Hills (23°10'N, 92°50'E), Assam, India 26. VI. 1 907 R. A. Lorrain Publication: Nuttall & Warburton (1915:431). N583 R291 Haemaphysalis bispinosa [sex/stage not given] ( 1 cf, 1 9 Haemaphysalis bispinosa) ex. Talpa sp. Lushai Hills (23°10'N, 92°50'E), Assam, India 1907R. A. Lorrain Publication: Nuttall & Warburton ( 1 9 1 5: 43 1 ). N584 R294b Rhipicephalus haemaphysaloides [sex/stage not given] (3 cf, 1 9 Rhipicephalus haemaphysaloides) No data N585 R242 Rhipicephalus appendiculatus [sex/stage not given] (2 cf Rhipicephalus muehlensi) No data N588 R255 Boophilus sp. incert. [sex/stage not given] (10 9 Boophilus microplus) No host data Entre-Rios, Argentina. Probably — Provincia de Entra Rios (32WS, 59WW), Argentina. (There are also two small populated areas with the name Entra Rios at 23°18'S, 64° 1 1'W and 28°50'S, 64°11'W, which could be the collec- tion locality.) IV. 1 897 Dr C. Berg [C. Warburton det. 1918] Note: This collection also contained 1 cf of the human sucking louse Pediculus humanus. Boophilus microplus is a vector of the viruses Wad Medani, Seletar, and Crimean Congo hemorrhagic fever. N589 R290 Argas persicus [sex/stage not given] (9 cf, 6 9, 5 N Argas persicus)) No history N590 R292 Argas persicus [sex/stage not given] (11 cf , 4 9, 11 N Argas persicus) No history Note: An additional 1 cf , 1 9 of this collection were presented to J. de Meza of Zomba, Malawi, 19.11.1916. N591 R214 1 6L Argas vespertilionis ex. bat (also parasites off mouse) No particulars N592 R195 3L Argas vespertilionis (& Nycteribia) ex. Scot 'us wroughtoni (Scotophilus temmincki wroughtoni) Helwak(17°22'N, 73'44'E), Maharashtra, India. N593 R224 Argas vespertilionis [sex/stage not given] (3 L Argas vespertilionis) ex. Vesperugo pipistrellus (Pipistrellus pipistrellus) Puttenham (5 1 °49'N, 00°44'W), England 28.VII.1899 N594 R240 Ornithodoros talaje [sex/stage not given] (14 N Ornithodoros capensis group) ex. nestlings Arrows Sand? Puhekiox, Hawaii (locality not verified) 30.VI.1900H.W. Henshaw N597 R153 Ixodes hexagonus [sex/stage not given] (8 N 1 L, Ixodes hexagonus) ex. Erinaceus europaeus Belgium. E. A. Boulenger Publication: Nuttall & Warburton (1911:183). Ixodes hexagonus is a vector of tickborne encephalitis virus. N598 R206 Ixodes hexagonus [sex/stage not given] (1 9, 6 N Ixodes hexagonus) ex. otter (Lutra vulgaris) (Lutra lutra) No particulars Publication: Nuttall & Warburton (1911: 182) indicated that the locality was England. N599 R209 Ixodes hexagonus [sex/stage not given] (3 9, 4 N Ixodes hexagonus) ex. Erinaceus europaeus Cambridge (52° 1 3'N, 00°08'E), England Publication: Nuttall & Warburton (1911:182). ROTHSCHILD TICKS IN BM(NH) 11 N600 R221 Ixodes hexagonus [sex/stage not given] (3 9, 1 L Ixodes hexagonus) ex. polecat Mustela putorius Boxworth (52'16'N, 00°02'W), England 30.IV.1901 Publication: Nuttall & Warburton (1911:1 82). N601 R208 Ixodes hexagonus [sex/stage not given] (1 9, 41 L Ixodes hexagonus) ex. otter (Lutra vulgaris) (Lutra lutrd) No other data Note: One larva (RML 16476) of this collection was donated to the Rocky Mountain Laboratory by P. A. Buxton, London School of Hygiene and Tropical Medicine, in 1940. (See also Nuttall 1159). Publication: Nuttall & Warburton (1911: 182) indicated that the locality was England. N602 R222 1 9 Ixodes hexagonus ex. Mustela erminea Tring (5 1 '48'N, 00°40'W), England 19.V.1903 Publication: Nuttall & Warburton (1911:1 82). N603 R179a Ixodes hexagonus [sex/stage not given] (2 9, 1 N Ixodes hexagonus} ex. Mustela erminea Wigginton (5 1 °47'N, 00'38'W), England 4. V.I 903 Barrett coll. Publication: Nuttall & Warburton (1911:1 82). N604 R143 Ixodes hexagonus [sex/stage not given] (4 N Ixodes hexagonus) ex. Mustela vulgaris (Mustela nivalis) Tring (5 1 °48'N, 00°40'W), England 20.111.1902. Publication: Nuttall & Warburton (1911:1 82). N605 R164 Ixodes hexagonus [sex/stage not given] (3 9 Ixodes lividus) ex. Cotile riparia (Riparia riparid) Worsborough Bridge (53°32'N, 01'28'W), England 6.VII.1906A. Whitakercoll. Publication: Arthur (1963: 92) Ixodes lividus is a vector of tickborne encephalitis virus. N606 R225 d"s, 9's, & larvae Ixodes hexagonus (3 9, 6 N, 8 L Ixodes hexagonus) ex. polecat (Mustela putorius) Aberystwyth (52°25'N, 04°05'W), Wales 26.XI.1902. Publication: Nuttall & Warburton (1911: 183) gave the date as 1902. N607 R201 Ixodes hexagonus [sex/stage not given] (1 N, 7 L Ixodes hexagonus) ex. Mustela erminea Tring (5 1 °48'N, 00°40'W), England 3.IX.1902 Publication: Nuttall & Warburton (1911: 182) indicated that the collection was composed only ofnymphs. N608 R215 Ixodes hexagonus [sex/stage not given] (4 N Ixodes hexagonus) ex. Erinaceus europaeus Durnberg(48°13'N, 1 1'55'E), Federal Republic of Germany 3.IX.1898 Publication: Nuttall & Warburton (1911: 183) gave the spelling as Dernberg rather than the catalogue spelling of Dermberg. N609 R212 1 N Ixodes hexagonus ex. polecat (Mustela putorius) Aberystwyth (52°25'N, 04°05'W), Wales 23.XII.1902 Note: Nuttall 579 also bears the Rothschild number R2 1 2 but the data do not correspond to N609. Publication: Nuttall & Warburton (1911: 183) indicated that more than one nymph was in this collection. N610 R219 N's, larvae Ixodes hexagonus (1 N, 15 L Ixodes hexagonus) ex. Mustela erminea Tring (5 1 °48'N, 00°40'W), England 7.V.1902 Publication: Nuttall & Warburton (1911:1 82). N611 R145 1 N, 1 L Ixodes hexagonus ex. Mustela erminea Lyndhurst (50'52'N, 0 1 °34'W), England 20.III.1901G. Tate Publication: Nuttall & Warburton (1911: 182) indicated that multiple nymphs and larvae were in this collection. N612 R180 9's, N's Ixodes hexagonus (1 9, 7 N Ixodes hexagonus) ex. Mustela erminea Inismore (54° 1 8'N, 07'34'W), Ireland 23.111.1902 Publication: Nuttall & Warburton (1911:1 82). 12 J. E. KEIRANS N613 R202 9's, N's Ixodes hexagonus (5 N Ixodes hexagonus) ex. stoat (Mustela ermined) No other data N617 R177 1 9 Ceratixodes putus (Ixodes uriae) ex. wild duck No other data Publication: Nuttall & Warburton (1911:261). Ixodes uriae is a vector of the viruses tickborne encephalitis, Tuleniy, Oceanside, Zaliv Terpenya, Sakhalin, Tillamook, Avalon, Clo- Mor, Taggert, Bauline, Cape Wrath, Nugget, Tindholmur, Mykines, Great Island, Okhotskiy, Yaquina Head, Poovoot, Paramushir, and Runde. N618 R220 1 cf, 1 9, 2 N Ceratixodes putus (Ixodes uriae) British. E. G. Wheler No particulars N619 R147 1 d1, 2 9 Ixodes tenuirostris (2 rf, 2 9 Ixodes trianguliceps) ex. Hypudaeus glareolus (Clethrionomys glareolus) Campfer (46°28'N, 09°48'E), Switzerland 7.VII.1904 [full date on vial label]. K. Jordan. Publication: Nuttall & Warburton ( 1 9 1 1 : 248). Ixodes trianguliceps is a vector of tickborne encephalitis virus. N621 R204 N's Ixodes tenuirostris (1 N Ixodes trianguliceps) ex. Sorex vulgaris (Sorex araneus) Bevendean, (50°5 1 'N, 00°05'E), England VI.1899. Publication: Nuttall & Warburton (1911: 248), indicated that more than one nymph was in the collection. N622 R166 9 Ixodes tenuirostris (1 9 Ixodes trianguliceps) ex. Arvicola amphibius (Arvicola terrestris) Reservoir 6.IV.1892 Publication: Nuttall & Warburton (1911: 248) indicated the collection was from England. N623 R197 2 9 Ixodes tenuirostris (Ixodes trianguliceps) ex. Arvicola agrestis (Microtus agrestis) Tring (5 1 '48'N, 00°40'W), England Publication: Nuttall & Warburton (1911: 248). N624 R149 1 9 Ixodes tenuirostris (Ixodes trianguliceps) ex. Arvicola arvalis (Microtus arvalis) Campfer (46°28'N, 09°48'E), Switzerland 20. VII. 1904 K.Jordan Publication: Nuttall & Warburton (1911: 248). N625 R148 3 9 Ixodes tenuirostris (Ixodes trianguliceps) ex. Arvicola arvalis (Microtus arvalis) Campfer (46°28'N, 09°48'E), Switzerland 5. VII. 1904 K.Jordan Publication: Nuttall & Warburton ( 1 9 1 1 : 248). N626 R226 d"s, 9's Ixodes scapularis (4 rf, 4 9 Ixodes scapular is) ex. Cariacus virginianus (Odocoileus virginianus) Craven Co. (35'07'N, 77°03'W), North Carolina, U.S.A. 13.XI.1907 k[det. C. Warburton 5. 11.1909. Found with N666]' Publication: Nuttall & Warburton (1911: 158) gave the date of collection as XI. 1897. The date 1 3.XI. 1 897 is also on the vial label. N633 R169 N's Amblyomma sp. (8 N Amblyomma sp.) ex. Macropus sp. No other data Note: Nuttall 644 also bears the Rothschild number R169 and gives the additional data: W. Australia 28.VII. 1900. N634 R159 Ixodes percavatus (2 9, 2 N Ixodes rothschildi TYPES) ex. Puffin Fratercula arctica. Wrongly labelled Ix. putus '(Pocock)' On back of label; no parti- culars. (Sent to Neumann who confirmed the determination II. 1909.) No other data. Publications: Nuttall & Warburton (1911: 221) described the 9 and N of Ixodes percavatus rothschildi. Zumpt (1952: 17) raised this taxon to specific rank; Arthur (1953: 222-226) indepen- dently raised /. percavatus rothschildi to specific rank; Arthur ( 1 963 : 1 34) referred to 2 9, 3 N in the type lot. There was (IX. 1977) a note in the vial with the words '1 N Missing'. N637 R213 Ixodes bicornis [sex/stage not given] ( 1 1 9 Ixodes boliviensis) ex. Felis concolor '[American]' No other data Publication: Nuttall & Warburton (1911: 187) indicated only one 9 in the collection. N638 R276 1 cf, 1 9 Ixodes loricatus ex. Didelphia (Didelphis sp.) Sapucay. Foster coll. Note: Locality is probably Sapucai (25° 1 9'S, 56°55'W), Paraguay, which is in the range of /. loricatus. [See also Nuttall 1 39 1]. Publications: Nuttall & Warburton (1911: 269); Cooley& Kohls (1945: 190). ROTHSCHILD TICKS IN BNi(NH) 13 N639 R256 Ixodes loricatus [sex/stage not given] (1 d Ixodes loricatus) ex. 'Piquoti' No other data N640 R241 2 rf, 2 9, 2 N Ixodes loricatus (2 cf, 2 9 Ixodes loricatus) ex. A teles melanochir (A teles g. geoffroyi) Frontera (18'32'N, 92°38'W), Tabasco, Mexico. Note following locality '(Compared with Neumann's types at Toulouse, 1910; agrees fully but Nn's type larger).' Publications: Nuttall & Warburton (1911: 269); Cooley& Kohls (1945: 190). N641 R161 Ixodes loricatus [sex/stage not given] (2 rf, 1 L Ixodes loricatus) ex. Didelphys aurita (Didelphis marsupialis) Brazil 24.XI.1901 Publications: Nuttall & Warburton (1911: 269); Cooley& Kohls (1945: 190). N642 R165 1 9, 1 N, 6 L Ixodes vestitus (1 9, 5 N Ixodes vestitus) ex. Diemenia superciliosa (Diemenia textilis) Herdman's Lake, near Perth (31'56'S, 1 15'15'E), W. Australia B. H. Woodward Note: Nuttall 628 also bears Rothschild numfjer R165. Publications: Nuttall & Warburton (1911: 255) indicated larvae as did the catalogue entry. These were missing 7.X.1977. Taylor (1946: 59, 60) repeated Nuttall & Warburton's description and figures. Roberts ( 1 960: 45 1 , 452). N643 R265 2 N Ixodes holocyclus Perth (3 1 °56'S, 1 1 5° 1 5'E), W. Australia B. H. Woodward coll. Publication: Nuttall & Warburton (1911:238). N645 R275 3 9 Ixodes australiensis ex. tame dog (Canis familiar is) Cranbrook (34° 1 8'S, 1 1 7°32'E), West Australia 1. IV. 1 900 J.T. Young Publications: Nuttall & Warburton (1911: 252). Taylor (1946: 57, 58) repeated Nuttall & Warburton's descriptions and figures. Note: This collection was seen by R. A. Cooley in 1946 who made 3 slides of dissections (9 spiracular plate; 9 legs I & IV; 9 hypostome) all in the Nuttall collection at the British Museum (N.H.). He gave them the Rocky Mountain Laboratory (RML) number 22284. N646 R210 Ixodes australiensis [sex/stage not given] (8 9 Ixodes australiensis) ex. Bettongia lescuri (Bettongia lesueuri) Kojonup (33°50'S, 1 1 7°09'E), W. Australia 4.XI.1900 Publications: Nuttall & Warburton (1911: 252). Taylor (1946: 57, 58) repeated Nuttall & Warburton's description and figures. Roberts (1960: 440); Roberts (1970: 40) gave the correct spelling of the locality. N647 R248 3 9 Ixodes loricatus var. spinosus n. var. Nuttall, 1 9 1 0 (2 9 Ixodes luciae) ex. large opossum (Didelphis sp.) Frontera ( 1 8°32'N, 92'38'W), Tabasco, Mexico May. Found with 648. Note long spine on coxa I. Note: These are the TYPE SPECIMENS of /. loricatus spinosus. Publications: Nuttall (1910: 411), original de- scription. Nuttall & Warburton (1911: 269), repeated original description and figures. N648 R248 N's & larvae Amblyomma Icajennense (16 N Amblyomma cajennense) ex. large opossum (Didelphis sp.) Frontera ( 1 8°32'N, 92°38'W), Tabasco, Mexico May Note: Found with Nuttall 647 N649 R185 1 N Ixodes vespertilionis. ex. Vespertilio tricolor (Myotis tricolor) Kei Road, W. Kingwilliamstown, S. Australia [S. Australia crossed out and S. Africa inserted]. Kei Road, King William's Town (32'51'S, 27°22'E), South Africa. Purchased of Capt. H. Trevelyan. Publications: Nuttall & Warburton (1911: 277). Taylor (1946: 41) considered /. vespertilionis 'very doubtfully Australian' and 'original specimen is unfortunately lost'. Hoogstraal (1956: 569, 570) considered this collection to be of South African origin. Ixodes vespertilionis is a vector of the viruses Issyk Kul and tickborne encephalitis. N650 R203 1 9 Ixodes fecialis TYPE (HOLOTYPE) ex. Dasyurus geoffroyi Cranbrook (34° 1 8'S, 1 17'32'E), W. Australia 5.III.1900 Publications: Warburton & Nuttall (1909: 58) original description. Nuttall & Warburton (1911: 248) repeated the original description and figures as did Taylor (1946: 56). Roberts (1960: 445) incorrectly gave the collection date as March 1910. 14 J. E. KEIRANS N652 R168 2 9 Ixodes unicavatm(\ 9, 1 N Ixodes unicavatus) ex. shag (Phalacrocorax aristotelis) Plymouth (50'23'N, 04°10'W), England IX.1897 Publications: Nuttall & Warburton (1911: 266) indicated that the collection consisted of only one nymph. Arthur ( 1 963: 1 26) repeated this error. N666 R226 1 9 Dermacentor nigrolineatus (1 9 Dermacentor albipictus) ex. Cariacus virginianus (Odocoileus virginianus) Craven Co. (35807'N, 77°03'W), North Carolina, U.S.A. Found with N626 q.v. for details. C. Warburton det. 1917 Dermacentor albipictus is a vector of Colorado tick fever virus. N667 R250 9's, N's Ixodes unicavatus (39, 10 N, 3 L Ixodes unicavatus) ex. Phalacrocorax graculus (Phalacrocorax aristotelis) Cromarty (57°40'N, 04°02'W), Scotland. 10.111. 1898 W.Roy Publications: Nuttall & Warburton (1911: 266): Arthur (1963: 126). N669 R178 Haemaphy sails humerosa [sex/stage not given] TYPES (2 d1, 1 9, 1 N Haemaphysalis humerosa TYPES) ex. Parameles macrura (Isoodon macrourus) Barrow Island (20°48'S, 115°23'E), N. W. Australia Note: An additional 1 cf of this collection was sent to Dr L. O. Howard, U.S. Dept. of Agriculture, Washington, D.C., U.S.A. 2 1 .V. 1 909. Publications: Warburton & Nuttall (1909: 60) original description. Nuttall & Warburton (1915: 496, 497) and Taylor (1946: 82) repeated original description and figures. Roberts (1963: 39) dis- cussed the type series and the note inserted in the tube by Nuttall that one specimen was missing 20.X.1934. N670 R172 Haemaphysalis elongata [sex/stage not given] (1 d1 Haemaphysalis elongata, 4 d1, 2 9 Haema- physalis tiptoni) ex. Centetes ecaudatus (Tenerec ecaudatus) Madagascar Note: The male of H. elongata has been given a collection number by the Rocky Mountain Laboratory (RML 71004) and by Harry Hoogstraal(HH44251). Publications: Nuttall & Warburton (1915: 500). Hoogstraal (1953: 59, 64) discussed the two species represented in Nuttall & Warburton's (1915) figures and descriptions. N826 R189 2 9 Ixodes hexagonus (Ixodes rubidus) ex. Coypu, Myopotamus coypus (Myocaster coypus) S. America. No further particulars Publication: Nuttall & Warburton (1911:1 83). ' N892 R295 Haemaphysalis bispinosa [sex/stage not given] (7 d, 7 9 Haemaphysalis bispinosa) ex. Canis familiaris Luthaia Mts, Assam Note: The locality is questionable. There are no Luthaia Mts as written in the catalog or Southaia Mts as in Nuttall & Warburton (1915: 431) to be found in gazetteers of India. I believe the locality to be Lushai Hills (23°10'N, 92°50'E), Assam, India as in N576, (R294a) and N582 (R284). Publication: Nuttall & Warburton ( 1 9 1 5: 43 1 ). N1066 R245 5 9 Ixodes texanus var. (Ixodes sp. not /. texanus or /. hearlei) No other data N1067 R200 1 9 Ixodes hexagonus ex. fox ( Vulpes vulpes) Boxworth (52° 1 6'N, 00°02'W), England 22.1.1898 Publication: Nuttall & Warburton (1911:1 82). N1068 R190 6 d, 6 9 Ixodes canisuga (4 d1, 4 9 Ixodes lividus) Lyndhurst (50°52'N, 01°34'W), England 11. VII. 1900 Publications: Nuttall & Warburton (1911: 214). Arthur (1 963: 92) indicated 6 d, 6 9 present. There were 4 d, 4 9 in the collection as of IX. 1977. N1069 R146 1 d, 2 9, 3 N Ixodes tenuirostris (Ixodes trianguliceps) ex. Arvicola arvalis (Microtus arvalis) Campfer (46°28'N, 09°48'E), Switzerland 1 1. VII. 1 904 Dr K.Jordan Publication: Nuttall & Warburton (1911: 248). N1105 None 3 d1, 3 9, 6 N Dermacentor albipictus Blucher Hall (51'06'N, 120°01'W), British Columbia, Canada III. 1910. Rec'd 2.IV.1910 from Hon. N. C. Rothschild (London). N1151 R296 10 L Argas vespertilionis ex. P ipistrellus pipistrellus Oristano (39'54'N, 08°35'E), Sardinia May 1907 DrA. Krausse ROTHSCHILD TICKS IN BM(NH) 15 N1152 R175 6 LArgas vespertilionis ex. Vesperugo pipistrellus (Pipistrellus pipistrellus) Tring (5 1 °48'N, 00°40'W), England 1. IX. 1898 Nil 53 R244 2 L Argas vespertilionis No other data Ml 154 R271 2 L Argas vespertilionis (Argas (Carios) sp.) Perth (3 1 °56'S, 1 1 5°50'E), W. Australia Note: These specimens may represent a new species. It is not Argas australiensis, Argas daviesi or Argas dewae, the currently known representa- tives of the subgenus Carios in Australia. N1159 R142 N's Ixodes hexagonus (3 N Ixodes hexagonus) ex. Mustela erminea Tring (5 1 °48'N, 00°40'W), England 10.IV.1902. Note: One nymph (RML 16476) of this collection was donated to the Rocky Mountain Laboratory collection by P. A. Buxton, London School of Hygiene and Tropical Medicine, in 1940. (See also Nuttall 601). Publication: Nuttall & Warburton (1911:1 82). N1160 R194 L's Ixodes hexagonus (17 L Ixodes hexagonus) ex. fox Vulpes vulpes (cub) Boxworth (52° 16'N,00°02'W), England 30.111.1900 Note: 2 additional larvae from this collection were sent to Prof. R. Matheson, Cornell Univer- sity, Ithaca, N.Y., U.S.A. , 27. V. 1929. Publication: Nuttall & Warburton (1911:182). N1161 R278 9 Ixodes tenuirostris (Ixodes trianguliceps) Tarasp(46°38'N, 10'25'E), Switzerland summer 1901 Publication: Nuttall & Warburton (1911: 248) N1162 R207 2 L Ixodes sp. ex. Dromicia concinna (Cercartetus concinnus) Note: The nocturnal marsupial C. concinnus inhabits the southern part of western and central Australia. N1163 R247 L's Argas vespertilionis (1 3 L Argas vespertilionis) ex. Vesperugo pipistrellus (Pipistrellus pipistrellus} Yalding (5 1 ° 1 4'N, 00°26'E), England. 8. VIII. 1 897 [However, 1 997 was written] W. R. Ogilvie-Grant N1164 R250,251 L's Argas sp. (5 L Argas boueti) ex. either Phalacrocorax graculus, Cromarty, or Megaderma cor, Salamana, Havish, 4,500 ft (Number on cork defaced) ex. Megaderma cor. Salamana, Havish 1,372 m (locality not verified) Notes: Megaderma cor ranges in East Africa from Ethiopia to Tanzania. The correct Rothschild number for this collection is R25 1 . For R250 see Nuttall 667. N1166 R230 10 nymphs [genus & species not given] (Amblyommasp.) ex. Mus escularis (name not verified) St Aignan. Possibly Misima Island (10'41'S, 152°42'E), Louisade Archipelago. This island was formerly known as St Aignan. VIII-XI. 1897 (Meek) N1167 R186 N1167a 2 N Ixodes sp. [1 167], 1 d Hyalomma aegyptium [1 167a] (Hyalomma truncatum) ex. a 'Sassaby' a bastard hartebeest (Damaliscus lunatus) Republic of South Africa. Date? Hyalomma truncatum is a vector of the viruses Dugbe, Bhanja, Jos, and Crimean Congo hemorrhagic fever. N1211 R237 2 N Ixodes sp. (2 N, 1 L Ixodes fecialis) ex. Tarsipes rostratus or Smiothopsis murinus [vial label reads S. murinus] (Sminthopsis murina) Albany (35'02'S, 1 1 7°53'E), W. Australia 2 1. XII. 1 900 B. H. Woodward N1212 R272 5 N Ixodes sp. (Ixodes holocyclus) Perth (3 1 °56'S, 1 1 5°50'E), W. Australia B. H. Woodward No further details N1213 R216 1 9 Ixodes fecialis var. aegrifossus (Ixodes fecialis) ex. Perameles obesula (Isoodon obesulus) Bannister (32°40'S, 1 16'33'E), W. Australia 21.VIII.1900 Publication: Nuttall & Warburton (191 1: 250). N1386 R249 N's, L's Amblvomma sp. (14 N, 2 L Amblyomma sp.) ex. Cariacus virginianus mexicanus (Odocoileus virginianus) San Rafael (20'12'N, 96°5 1 'W), Veracruz, Mexico 27. III. 1903 16 J. E. KEIRANS N1387 R264 1 N ? genus (Dermacentor andersoni) ex. Lagomys princeps (Ochotona princeps) Canadian National Park, ? Ottawa, Canada 5. VIII. 1 899 J. F. Dippie Note: Ottawa, Ontario is far to the east of the ranges of D. andersoni and the pika. The Canadian National Park and the date 1899 suggests a relationship to N1390 (R280), Banff National Park, which is within the range of both O. princeps and D. andersoni. Dermacentor andersoni is a vector of the viruses Powassan and Colorado tick fever, the bacterium of tularemia, and the rickettsia of Rocky Mountain spotted fever. N1389 R155 larvae Ixodes sp. (3 N Ixodes sp.) ex. Crysomys longicaudatus (Oryzomys longicaudatus) Valparaiso (33°02'S, 7 1 °38'W), Chile Date?J. A.Wolffshon N1390 R280 2 N ? genus '(has eyes!)' (2 N Dermacentor andersoni) Banff (5 1 °38'N, 1 1 6°22'W), Alberta, Canada 19.VII.1899 N1391 R269 4 N ? genus (3 N Amblyomma sp.) ex. Didelphys sp. (Didelphis sp.) Sapucai (25'19'S, 56°55'W), Paraguay ?Date N1392 R187 N's Amblyomma sp. (10 N Amblyomma sp.) ex. Hare wallaby (Lagorchestes conspicillatus) Barrow Island (30'48'S, 115'23'E), N. W. Australia N1393 R176 larvae Amblyomma sp. (64 L Amblyomma sp.) ex. Bettongia penicillata Bokerup, W. Australia. Possibly Bokarup Swamp (34'24'S, 116°50'E). 20. IV. 1900 N1394 R227 2 N Amblyomma sp. ex. Ontalis vetula macalii (Neotoma alleni vetula) Frontera (18'32'N, 92'38'W), Tabasco, Switzerland [sic.] Mexico 18.V.- N1395 R163 N's Rhipicephalus sp. (2 N Rhipicephalus sp.) ex. Hyaena crocuta (Hyaena hyaena) Darie (locality not verified) ? Date. Von Erlanger N1396 R246 1 N Amblyomma sp. ex. Black squirrel (Sciurus sp.) Frontera ( 1 8°32'N, 92°38'W), Tabasco, Mexico 19.V.- N1397 R171 larvae Amblyomma sp. (26 L Amblyomma sp.) ex. Macropus eugenii (Macropusfuligenosus) Cranbrook (34° 1 8'S, 1 1 7°32'E), W. Australia 6.III.1910 N1398 R150 1 larvae? genus (1 LHyalommasp.) ex. Gerbillus gerbillus Zaghig* 1 .III. 1903 Hon. N. C. Rothschild ""Locality not verified. However, N. C. Rothschild was in the Nile delta early in 1903. Quite possibly the locality is Zagazig (30°35'N, 31*3 l'E), Egypt. N1399 R162b N's Ixodes sp. (4 N, 4 L Ixodes texanus) ex. Putorius energermanis (Mustela vison) Sumas, B.C., Canada* III. 1903 *Either Sumas Lake (49'07'N, 122°02'W), British Columbia, Canada or Sumas (49°00'N, 122813'W), Washington, U.S.A. N1497 None 1 N Dermacentor sp. (Dermacentor albipictus) ex. Moose (Alces alces) New Brunswick, (47WN, 66WW), Canada 30.XI.1911 J. Birrellcoll. N1498 None 1 cf Dermacentor albipictus ex. man (Homo sapiens) (Dr Walker) New Brunswick (47WN, 66WW), Canada 29.XI.1911 J. Birrellcoll. N1499 None 2 c? Dermacentor albipictus ex. Moose (Alces alces) New Brunswick (47WN, 66WW), Canada 30.XI.1911 J. Birrellcoll. N1500 None 2 cf, 1 9, 1 N Dermacentor albipictus ex. carabou [sic.] (Rangifer tarandus) New Brunswick (47WN, 66WW), Canada 25.XI.1911 J. Birrellcoll. N1501 None 1 N Dermacentor sp. (Dermacentor albipictus) ex. carabou [sic.] (Rangifer tarandus) New Brunswick (47WN, 66'00'W), Canada 25. XI. 1911 J. Birrell coll. N1502 None 1 N Dermacentor sp. (Dermacentor albipictus) ex. carabou [sic.] (Rangifer tarandus) New Brunswick (47'00'N, 66WW), Canada 25. XI. 19 11 J. Birrellcoll. ROTHSCHILD TICKS IN BM(NH) 17 N1503 None 1 d Dermacentor sp. (5 N Dermacentor albipictus) ex. carabou [sic.] (Rangifer tarandus) New Brunswick (47'00'N, 66WW), Canada 25.XI.1911 J. Birrellcoll. N1504 None 1 9 Ixodes hexagonus (Ixodes cookei) '[Not typical — bad condition; possibly /. rubidus]' ex. black cat New Brunswick (47WN, 66WW), Canada 19.XII.1911 J. Birrellcoll. Ixodes cookei is a vector of Powassan virus. N3138 R926 1 d Aponomma laeve var. capensis (Aponomma sp.) ex. Echidna sp. No further data Gift of Hon. N. C. Rothschild "(unmounted by me from a slide R926)". L. E. Robinson del. Note: This is not Aponomma concolor or A. oudemansi. Coxal spurring is similar to A. pattoni but the specimen lacks cervical pits. It is a species near A. pattoni. N3160 R238 2 d1, 1 9, 1 N Dermacentor reticulatus ( 1 d1, 1 9, 1 N Dermacentor reticulatus) Revelstoke, and Revelstoke Point, now known as Stoke Point (50° 1 8'N, 04°0 1 'W), England III.1902. Dermacentor reticulatus is a vector of the viruses tickborne encephalitis, Russian spring summer encephalitis, and Omsk hemorrhagic fever. N3330 R243 1 9 Dermacentor auratus (Dermacentor sp.) No other data Note: Nuttall 565 and 575 also bear Rothschild number R243. N3502 None d's, 9's Dermacentor andersoni (7 d, 9 9 Dermacentor andersoni) Okanagan Falls (49'21'N, 119'31'W), British Columbia, Canada V. 19 13 Rothschild coll. Gift of Seymour Hadwen, 1922 N3537 None N's Haemaphysalis leachi (ION Haemaphysalis erinacei) ex. Dipodillus campestris (Gerbillus campestris) Timgad (35°30'N, 06°35'E), Algeria 1 5. IV. 1920 N. C. Rothschild & K. Jordan, coll. Publication: Hoogstraal (1955:222, 223) (see note under 3539). N3538 None N's Haemaphysalis leachi (UN Haemaphysalis erinacei) ex. Dipodillus campestris (Gerbillus campestris) Timgad (35'30'N, 06835'E), Algeria 15.IV. 1920 N. C. Rothschild & K. Jordan coll. N3539 None N's Haemaphysalis leachi (4 N Haemaphysalis erinacei) ex. Dipodillus campestris (Gerbillus campestris) Timgad (35°30'N, 06°35'E), Algeria 1 7.IV. 1 920 N. C. Rothschild & K. Jordan coll. Publication: Hoogstraal (1955: 223) gave the Nuttall number 3739 for this collection. He indicated the total number of nymphs as 1 7 in combined collections 3537 and 3539. In 1977 the total number of nymphs in these two collections was 14. A single tick collection given by the Hon. Walter Rothschild to G. H. F. Nuttall via Guy A. K. Marshall N888 None 2 d, 1 9 Hyalomma aegyptium (Hyalomma rufipes) ex. giraffe (Giraffa camelopardalis subsp.) Ogo, Senegal ex. Riggenbach coll. given by Hon. Walter Rothschild to African Entomological Committee (rec'd 19.X.1909 from Guy A. K. Marshall). Notes: There are two populated places and one campsite with the name Ogo in Senegal: PPL. 14°35'N, 15°50'W; PPL. 15°33'N, 13°17'W; CMP. 14°45'N, 15°03'W. 18 specimens of this collection were received 11. X.I 909 by the Entomological Research Committee (Anon. 1910). Rothschild specimens missing from Nuttall collection N75 R1015, 1017, 1019 3 d Amblvomma clvpeolatum ( = atrogenatum N. &W.) ex. tortoise Zoological Society's Gardens, London, 11. X. 1895 Presented by N. C. Rothschild Mounted (i) opaque & (ii) (iii) transparent on 3 slides nos. 1015,1017,1019 L. E.Robinson det. 26.111. 19 19 18 J. E. KEIRANS N76 R964 1 $ Amblyomma sp. ex Monitor Kalao Island. No further data Mounted opaque on slide No. 964 from N. C. Rothschild N77 R1033, 1034, 1035 3 cf Amblyomma marmoreum ex. Testudo pardalis (leopard tortoise) No further data. Mounted as transparent specimens on slides Nos. 1033, 1034, 1035 Presented by N. C. Rothschild L. E. Robinson det. 26. III. 1919 N78 R996,997,999 1 cf, 2 9 Amblyomma albolimbatum ex. black & white snake 30.1.1896. No further data. Mounted as trans- parent specimens Nos. 997, 996, 999 Presented by N. C. Rothschild L. E. Robinson det. 26.111.1919. N90 R938-941 4 d1 Aponomma gervaisi var. lucasi ex. Ophiophagus bungarus (Hamadryad) Presented by N. C. Rothschild No particulars. Mounted on slides 938-941; 2 in glycerine, 2 in balsam C. Warburton det. 1.1915 N91 R900 1 cf Aponomma gervaisi var. lucasi ex. Varanus civitatus (Monitor) Presented by N. C. Rothschild No particulars. Mounted as slide 900 in dil. glycerin. C. Warburton det. 1. 1 5 N92 R962,965 2 c? Aponomma gervaisi ex. Varanus civitatus (Monitor) Presented by N. C. Rothschild No particulars. Mounted as slides 962, 965 in glycerin. C. Warburton det. 1.1915 N93 R959,963,964,966 4 d1, 1 9 Amblyomma decoratum ex. Monitor Kalas Island Presented by N. C. Rothschild No particulars. 5 slides, 3 in glycerin 2 cf in balsam: R959, 961, 963, 964, 966. C. Warburton det. 1.1915 N216 R96 1-973 9's, N's, L's Ixodes hexagonus ex. Lulra vulgaris No doubt British N. C. Rothschild 7 slides N217 R930,931 2 9 Ixodes hexagonus ex. fox (Canis vulpes) No doubt British Gift of N. C. Rothschild (2 slides R930-93 1 ) N298 R1000-1011 6 9, 5 N, 1 L Ixodes hexagonus ex. stoat (Putorius ermined) No doubt British Gift of N. C. Rothschild 12 slides N322 R929 1 9 Ixodes hexagonus ex. Mustela putorius (ferret) Probably British Gift of N. C. Rothschild. R929 mounted in glycerin in glass cell. G. H. F. Nuttall det. N324 R944-946 3 N Ixodes hexagonus ex. Erinaceus europaeus Probably British Gift of N. C. Rothschild (3 slides R944-946). G. H. F. Nuttall det. N395 R988-990 3 L Ixodes unicavatus slightly fed ex. Phalacrocorax graculus (shag) No locality etc. Probably British Gift of N. C. Rothschild. Mounted on 3 slides R988-90. G. H. F. Nuttall det. 13.11.1915 N396 R1032 1 N Ixodes brunneusl ex. Erithacus rubecula (robin) No data. Probably British Gift of N. C. Rothschild Mounted on a slide R 1032 N397 R911 1 cf Amblyomma hirtum ex. bird (in spirit) Galapagos Islds, Pacific Ocean (W. of Ecuador) Gift of N.C.Rothschild Mounted on a slide R9 1 1 6.III.1915 L. E. Robinson det. N509 R982, 984-987 4 9, 2 N Ixodes ricinus ex. Cervus capreolus (roe deer) ? British Gift of N.C.Rothschild Mounted on 5 slides R982, 984-987 N516 R901-3,905 5 N Ixodes putus ex. Diomedea salvinii (albatross) No data Gift of N. C. Rothschild 5 slides R901-3,905 N517 R934 1 9 Ixodes putus ex. Uria lachrymans (ringed guillemot) ROTHSCHILD TICKS IN BM(NH) 19 No data Gift of N. C. Rothschild 1 slide N518 R975-979 5L Ixodes putus ex. Puffin No data. Probably British Gift of N. C. Rothschild 5 slides G. H. F. Nuttall del. 12.11.1915 N539 R1036 d Haemaphysalis leachi 1 slide ex. Erinaceus albiventris Gebel Auli, White Nile, Egypt 9.V.1900S. &F. Witherby Received from N. C. Rothschild 1908 Det.G.H.F. Nuttall 27.1. 1909 N544 R253 Amblyomma sp. not varium; bad condition ex. 3-toed sloth (Bradypus tridactylus) Brazil N548 R257 Aponomma gervaisi ex. Boa Feb. 1887 Given to D. S. MacLagan 14/10/38 N551 R181 Amblyomma triguttatum ex. Kangeroo(sic.) Barrow Island, N. W. Australia 20.XI.1900C.J. T. Tanney N571 R279 Aponomma sp. ex. Diemenia superciliosa Herdman's Lake, Perth, West Australia N586 R193 d Rhipicephalus sanguineus ex. Camel Locality unknown N587 R151 2 d Rhipicephalus sanguineus Zaghig, 25.11. 1908 N.C.Rothschild coll. N595 R294 Ornithodoros moubata No history Note: Nuttall 577 also bears Rothschild number R294 N596 R157 N's Ornithodoros megnini Okanagan, British Columbia 1 2. XII. 1902 Allan Brooks Note: Otobius is the correct genus for this tick N620 R942 1 9 Ixodes tenuirostris ex. Mus minutus (Harvest mouse) No further particulars '[Mounted on a slide]' N628 R165 9 & N Ixodes ricinus No particulars Note: Nuttall 642 also bears Rothschild number R165 N629 R233 4 d Ixodes ricinus ex. stag No particulars N630 R217 3 N Ixodes ricinus ex. Sciurus vulgaris 17. IV. 1901 Timmer, Ireland N631 R223 4 9 Ixodes ricinus ex. Erinaceus europaeus Belgium E. A. Boulengercoll. N632 R173 2 9 Ixodes ricinus ex. fox No particulars N644 R169 2 9 Ixodes holocyclus ex. Macropus sp. W.Australia 28. VII. 1900 Note: Nuttall 633 also bears Rothschild number R169 N656 R205 2 L Ixodes ricinus ex. Mustela erminea Lyndhurst, Hants., England 25. VII. 1 900 G. Tatecoll. N674 R922-924 3 9 Ixodes hexagonus ex. Myopotamus coypu ? locality, host a rodent from S. America Presented by N. C. Rothschild (R922-924) '[see record of 2 other 9'$ in Ticks Part II, p. 183]' G.H.F. Nuttall.det.il. 19 15 N746 R232 larvae Ixodes hexagonus ex. polecat Aberystwyth, Wales VIII. 1900 N1388 R162 1 LArgassp. ex. Putorius energermanis Sumas, B.C., Canada III.1903 N3136 R958 1 d Hyalomma sp. 20 -J- E. KEIRANS ex. Capra falconer i (the Markhoor) No further data Gift of N. C. Rothschild (slide R958) Note: The host occurs in Kashmir, Afghanistan, Gilgit, etc. N3139 R1012-1020 Hyalomma syriacum ex. tortoise No further data Gift of N. C. Rothschild (5 slides i-v R1012-1020) N3140 R1022, 1023 2 9 Hyalomma syriacum ex. Testudo pardalis No further data Gift from N. C. Rothschild R1022— 1023 (2 slides, N3141 R906, 1031 Hyalomma syriacum ex. Araconda [sic] No further data (evidently bought slides from M. Pillischer, optician, 88 New Bond St., London W.) Gift of N. C. Rothschild (2 slides R906, 1031) N3157a R1026-1029 N(3 1 57) 3 d Rhipicephalus evertsi (a) 1 cf Boophilus decoloratus ex. Zebra No locality Gift of N.C.Rothschild 1908 Mounted on slides in balsam (R 1026-1 029) R. evertsi det. C. Warburton 18.111.1915 B. decoloratus det. C. Warburton & G. Nuttall 1915 H. F. N3158 R907 1 9 Rhipichephalus haemaphysaloides India Mounted on slide in glycerin Gift of N.C.Rothschild 1908 (Prep, bought from Fredk. Enock*) *Spelled Enock in this collection, but see N3 1 59 N3159 R908-910 3 N Argas reflexus India 3 slides, glycerine in cells G. H. F. Nuttall det. 111.1915 Gift of N.C.Rothschild 1908 (3 slides Nos. 908-9 10) Fredk. Enoch prep. Cost 2 shillings each! N3351 None 9 Amblyomma varium ex. Bradypus tridactylus (sloth) Gift of N. C. Rothschild years ago. Scutum, legs, spiracle on 3 slides L. E. Robinson det. 26. III. 1919 N3536 None 1 9 Argas persicus Found on wall Djama, Algeria 25.11.1920 N. C. Rothschild & K. Jordan coll. PART II Rothschild specimens donated to British Museum (Natural History) Species of Ixodoidea represented in the collection of the British Museum (Natural History), donated by the Hon. N. C. Rothschild K numbers follow each species. Numbers in parentheses refer to missing collections. Thus, validity of original determinations is unverified. ARGASIDAE Argas persicus (Oten)Kl, K2,(K148) Argas reflexus (Fabricius) K3 Argas sanchez i Duges K.4 Ornilhodoros talaje Guerin-Meneville K5 IXODIDAE Amblyomma albolimbatum Neumann K6 Amblyomma americanum (Linnaeus) K7 Amblyomma brasiliense Aragao K.8 Amblyomma cajennense (Fabricius) K9 Amblyomma calcaratum Neumann K10 Amblyomma compressum Macalister Kl 1 Amblyomma gemma Donitz K12, K13,(K149) Amblyomma humeraleC. L. Koch K14, K15 Amblyomma incisum Neumann K16 Amblyomma longirostre(C. L. Koch) K17, K18 Amblvomma maculatum C. L. Koch K19, K20, K2'l,K22 Amblyomma naponense (Packard) K23 Amblyomma nodosum Neumann K8, K24, K25 ROTHSCHILD TICKS IN BM(NH) 21 Amblyomma ovale C. L. Koch K8, K23, K26, K27, K28, K29, K30, K3 1 , K32 Amblyomma postoculatum Neumann K33 Amblyomma pseudoconcolor Aragao K34 Amblyomma testudinarium C. L. Koch K35 Amblyomma variegatum (Fabricius) K36, K37 Amblyomma sp. K38, K39 Aponomma concolor Neumann K40, K41, K42, K43 Aponomma decorosum (L. Koch) K44, K45, K46, K47 Aponomma exornatum (C. L. Koch) K48 Aponomma fimbriatum (C. L. Koch) K49 Aponomma gervaisi var. trimaculatus (Lucas) (K150) Aponomma latum (C. L. Koch) K50 Aponomma trimaculatum (Lucas) K5 1 Boophilus decoloratus (C. L. Koch) K52 Boophilus microplus (G. Canestrini) K53 Dermacentor atrosignatus Neumann group K54 Dermacentor reticulatus (Fabricius) K55, K56 Haemaphysalis bancrofti Nuttall and Warburton K57, K58, K59, K60 Haemaphysalis calcarata Neumann K6 1 Haemaphysalis juxtakochi Cooley K53 Haemaphysalis lagostrophi Roberts K33 Haemaphysalis leachi (Audouin) K62, K63, K64, K66, K67, K68, K69, K70, K71, K72, K73, K74, K75,(K151) Haemaphysalis leachi indica Warburton (Kl 52} Haemaphysalis leachi (Audouin) group K76 Haemaphysalis novaeguineae Hirst K77, K78, K79 Haemaphysalis parmata Neumann K80 Haemaphysalis spinigera Neumann K8 1 Haemaphysalis spinulosa Neumann K82, K83 Haemaphysalis sulcata G. Canestrini, and Fanzago K84, K85 Haemaphysalis tibetensis Hoogstraal K86 Hyalomma aegyptium (Linnaeus) K87 Hyalomma anatolicum excavatum C. L. Koch K88 Hyalomma marginatum C. L. Koch K89 Hyalomma truncatum C. L. Koch K90, (Kl 53) Hyalomma sp. K9 1 , K92, K93 Ixodes alluaudi Neumann K94, K95, K96, K97, K98 Ixodes cordifer Neumann K99, K100, K101, K102 Ixodes fecialis Warburton and Nuttall K103 Ixodes hexagonus Leach K104, K105, K106, K107, K108,K109,K110 Ixodes loricatus Neumann Kl 1 1 , Kl 12 Ixodes luciae Senevet Kl 1 3 Ixodes ornithorhynchi Lucas K 1 1 4 Ixodes cumulatimpunctatus Schulze Kl 15 Ixodes ricinus (Linnaeus) Kl 16, (Kl 54) Ixodes tasmani Neumann Kl 1 7 Ixodes thomasae Arthur and Burrow K69 Ixodes trianguliceps Birula Kl 18 Ixodes vespertilionis C. L. Koch K119, K120, K121, K122, K123, K124,(K155) Rhipicephalus haemaphysaloides Supino K125 Rhipicephalus jeanneli Neumann K126 Rhipicephalus kochi Donitz K127 Rhipicephalus kochi Donitz group K128 Rhipicephalus longicoxatus Neumann K127 Rhipicephalus pravus Donitz K127, K129, K130, K131 Rhipicephalus pulchellus (Gerstacker) K93, K132 Rhipicephalus sanguineus (Latreille) K133, K134, K135, K136, K137, K138, K139, K140, K141,(K156) Rhipicephalus simusC. L. Koch K142, K143 Rhipicephalus sp. K144, K145, K146, K147 Rothschild specimens in British Museum (Natural History) collection with type status B.M.(N.H.) Accession No. None 19 14. 1 1.1 7. 1-46. None Dermacentor reticulatus aulicus Hirst, 1916. Ann. Mag. nat. Hist. (8), 17:308. Haemaphvsalis spinigera novaeguineae Hirst, 1914. Trans, zool. Soc. Lorn/ 20: 325, fig. 16. Haemaphvsalis tibetensis Hoogstraal, 1965. J. Parasit. 51: 452, figs. 1-26." 22 J. E. KEIRANS Rothschild specimens in British Museum (Natural History) collection ARGASIDAE Kl 1912.6.21.21 1 cf Argas persicus In railway carriage El Kantara (35° 13'N,05'43'E), Algeria 17.111.1911 Hon. Walter Rothschild & Ernst Hartert Argas persicus is a vector of Crimean Congo hemorrhagic fever virus. K2 1919.8. 14.1-70 (pt) 1 $ Argas persicus No host Ouargla house (locality not verified) E. Hartert and C. Hilgert Note: The locality is probably a house in Ouargla (3 r57'N,05°20'E), Algeria. K3 1912.6.21,1-2 1 9, 1 N Argas reflexus ex. Columba livia Leipzig (51°18'N, 12°20'E), German Democratic Republic V.I 9 10 O. Fritsche Argas reflexus is a vector of the viruses Ponteves and Grand Arbaud. K4 1912.12.4.1-8 Argas persicus (1 cf , 5 9, 3 N Argas sanchezi) ex. chicken Price, Pinal Co. (33°05'N, 1 1 1'30'W), Arizona, U.S.A. 27.VII.1911 Dr R. E. Kunzr K5 None 1 cf , 3 9, 3 N Ornithodoros talaje ex. native house Colombia, S. America IXODIDAE K6 1912.6.21.304-318 Amblyomma albolimbatum (19 cf, 6 9 Amblyomma albolimbatum; 1 9 Haema- physalis sp.; 1 9 Haemaphysalis sp.) ex Trachysaurus rugosus Bernier Is. (24°52'S, 1 13°08'E), W. Australia 1910 B. H. Woodward Note: 1 cf, 1 9 included in the above total were seen by R. A. Cooley in 1946 who made six slides of dissections (RML 22306). The Haemaphysalis specimens represent either two species or a single exceptionally variable species (Hoogstraal, personal communication). K7 None Amblyomma sp. (1 N Amblyomma americanum) ex Scuirus niger Mount Pleasant (32°47'N, 79°52'W), South Carolina, U.S.A. 1909 A. T. Wayne Amblyomma americanum is a vector of Lone Star virus. K8 None Amblyomma nodosum (6 cf, 1 9 Amblyomma brasiliense; 25 cf, 5 9 Amblyomma nodosum; 1 9 Amblyomma ovale) Host unknown Joinvile (26°18'S, 48'50'W), Humboldt, Estadode Santa Catharina, Brazil 25.VI.1912 W. Ehrhardt K9 1912.6.21.326-337 6 cf, 1 1 9, 3 N Amblyomma cajennense ex. cattle, Canis familiaris & Homo sapiens St. Ann's ( 1 8°2 1 'N, 77° 1 6'W), Jamaica 22-29. III.1911 F. E. Sherlock KlONone Amblyomma concolor! '9 etc.' (79, 1 N, 4 L Amblyomma calcaratum) ex. Tapir americanus ( Tapirus terrestris) Joinvile (26° 1 8'S, 48°50'W), Humboldt, Estado de Santa Catarina, Brazil W. Ehrhardt 15. IX. 19 12 Kll 1912.6.21.299-302 Amblyomma cuneatum (Species epithet crossed out and compressum inserted) (1 cf, 4 9 Amblyomma compressum) ex. Pangolin (or Scaly Ant-Eater) Manis sp. Bibianha, Gold Coast. Bebianiha (05°43'N, 00°27'W) or other populated places of this name nearby. Ghana 9. XII. 1911 H. G. F. Spurrell Note: Vial label gives accession number as 1912.6.21.299-303 K12 1911. 12.9.1-320 pt 2 9 Amblyomma gemma ex. Oxen Voi (03°23'S, 28°34'E), Kenya 25. IV. 1910 Robin Kemp K13 191 1.12.9. 1-320 pt 5cf , 4 9 Amblyomma gemma ex. Oxen Voi (03°23'S, 38°34'E), Kenya 30.IV. 1910 Robin Kemp ROTHSCHILD TICKS IN BM(NH) 23 K14 1912.10.31.18-20 1 cf, 3 9 Amblyomma humerale ex. donkey (Equus asinus) Locality unknown K15None 4 cf Amblyomma humerale ex. turtle Guyana K16None 1 cf Amblyomma incisum ex. tapir ( Tapirus sp.) Guyana Note: There is 1 9 in this collection which is tentatively being called A. incisum but which has a 3/3 hypostome. K17 1912.6.21.338 1 cf Amblyomma longirostre ex. Galera barbara (Eira barbara) Mar de Espanha (21'52'S, 43'00'W), Minas Gerais, Brazil Z. F. Zikan K18 1912.6.21.339-343 2 cf, 3 9 Amblyomma longirostre ex. Synetheres prehensilis (Coendou prehensilis) Mar de Espanha (21'52'S, 43WW), Minas Gerais, Brazil 6.III.1911 Z. F. Zikan K19 1912.10.31.13 1 cf Amblyomma maculatum ex. Canis sp. Buenos Aires (34°36'S, 58°27'W), Argentina 30.VI.1912 Miss Idina G. Runnael K20 1912. 12.4.12-14 1 cf, 1 $ Amblyomma maculatum ex. Small Chubut Fox BahiaBlanca(38°43'S, 62° 17'W), Argentina 15.11.1911 E. Weiske K21 1919.8. 14.1-70 pt 4 cf, 5 9 Amblyomma maculatum ex. Blastocerus dichotomus (B. bezoarcticus) Parana Delta (vicinity of 33°43'S, 59'15'W), Argentina 25.11.1917 Robin Kemp K22 None 3 cf Amblyomma maculatum ex. Pseudalopex gymnocerus (Dusicyon gymnocerus) Laguna Alsina (36°49'S, 62'13'W), to Buenos Aires (34°36'S, 58°27'W), Argentina V.1916 Robin Kemp K23 1919.8. 14. 1-70 pt Amblyomma mantiquirense (1 cf Amblyomma naponense) ex. Hydrochaerus hydrochaeris Eastern Ecuador This collection also contains 1 9 of Amblyomma ovale or a closely related species. K24 None 1 cf, 1 $ Amblyomma nodosum ex. Anteater Colombia, S. America 1912 K25 1912.6.21.325 1 cf, 3 N, 3 L Amblyomma nodosum ex. Myrmecophaga tetradactla (Myrmecophaga tridactyld) Misantla (19°56'N, 96°50'W), Mexico 1.1911 W. Engelmann Note: Immature specimens tentatively this sp. K26 1912.6.21.344-352 8 cf, 4 9, 3 N Amblyomma ovale ex. Nasua socialis (Nasua nasud) Mar de Espanha (21°52'S, 43'00'W), Estado de Minas Gerais, Brazil 15.X.1910 Z. F. Zikan Note: Also present were 1 cf, 2 9 of the mallophagan Neotrichodectes pallidus K27 1912.6.21.353-358 7 cf , 3 9 Amblyomma ovale ex. Galera barbara (Eira barbara) Mar de Espanha (20'52'W, 43WW), Minas Gerais, Brazil Z. F. Zikan K28 1921.12.9.10-11 1 cf, 1 9 Amblyomma ovale ex. Galictes vittata (Grison vittatd) 9.VIII.1916 W. Ehrhardt K29 1921.12.9.12-20 8 cf , 2 9 Amblyomma ovale ex. Nasua familiar is (Nasua nasua) Joinvile(26°18'S,48°50'W), Humboldt, Estado de Santa Catharina, Brazil 5.VIII.1918 W. Ehrhardt K30 None 3 cf , 2 9 Amblyomma ovale ex. Procyon cancrivorus Joinvile (26'18'S, 48'50'W), Humboldt, Estado de Santa Catharina, Brazil 24.VII.1912 W. Ehrhardt 24 J. E. KEIRANS K3 1 None 1 rf A mblyomma ovale ex. Nasua socialis (Nasua nasua) Mar de Espanha (21'52'S, 43WW), Estado de Minas Gerais, Brazil 18.VIII.1911 Z. F. Zikan K32 None 1 2 rf, 5 9 Amblyomma ovale ex. Nasua socialis (Nasua nasua) Mar de Espanha (21'52'S, 43WW), Estado de Minas Gerais, Brazil 18.VIII.1911 Z. F. Zikan K33 1912.6.21.319-324 Amblyomma albolimbatum (2 9 Amblyomma postoculatum; 5 9 Haemaphysalis lagostrophi) ex. Lagostrophus fasciatus Bernier Island (24°52'S, 1 1 3°08'E), W. Australia 1910 B. H. Woodward K34 None Amblyomma concolor (2 rf, 7 N, 3 L Amblyomma pseudoconcolor) ex. Armadillo La Rioja (29°26'S, 66° 1 5'W), Argentina E. Giacomelli K35 1912.12.4.15-16 2 d Amblyomma testudinarium ex. Sheep Sri Lanka K36 None 1 N Amblyomma variegatum ex. Raven Harar (09° 1 9'N, 42°09'E), Ethiopia 17.11.1912 G. Kristensen Amblyomma variegatum is a vector of the viruses Dugbe, Thogoto, Jos, Crimean Congo hemorrhagic fever, Nairobi sheep disease, Bhanja and Somone. K37 None 1 d Amblyomma variegatum Host unknown Harar (09° 1 9'N, 42°09'E), Ethiopia 9.II.1912 G. Kristensen K38 None 1 N Amblyomma sp. ex. Canis familiar is Harar (09° 1 9'N, 42°09'E), Ethiopia 4.XII.1911 G. Kristensen K39 None 2 N Amblyomma sp. ex. Homo sapiens Gola National Forest (06°56'N, 10°45'W), Liberia 26.V.1910 R. H. Bunting K40 1912.6.21.370-380 Amblyomma concolor (3 9, 23 N Aponomma concolor) ex. Echidna aculeata (Tachyglossus aculeatus) Gippsland District (38°00'S, 147WE), Australia 12.IX.1910 A. Coles K41 None Amblyomma concolor (11 N Aponomma concolor) ex. Echidna sp. (Tachyglossus aculeatus) Australia A. Coles K42 1912.6.21.381-383 Amblyomma concolor (3 9, 3 N Aponomma concolor) ex. Echidna aculeata (Tachyglossus aculeatus) Traralgon (38°12'S, 146°32'E), Australia 29.X.1910 A. Coles K43 1912.6.21.384-386 Amblyomma concolor (3 9 Aponomma concolor) ex. Echidna sp. (Tachyglossus aculeatus) Australia 27.11.1911 A. Coles coll. K44 None Aponomma decorosum (Species epithet crossed out and undatum inserted) (22 cf, 3 9, 3 N Aponomma decorosum) Ex. Varanus sp. Kelsey Creek (20°26'S, 148°27'E), Queensland, Australia 1911 A. Fontaine K45 1912.6.21.391^01 Aponomma decorosum (Species epithet crossed out and undatum inserted) (22 cf , 6 9, 18 N, 2 L Aponomma decorosum) ex. Brown Iguana ( Varanus sp.) Kelsey Creek (20°26'S, 148°27'E), Queensland, Australia A. Fontaine Publication: Kaufman (1972: 369) K46 19 12.6.2 1.402^10 Aponomma decorosum (Species epithet crossed out and undatum inserted) ( 1 5 cT, 1 3 9, 2 1 N, 2 L Aponomma decorosum) ex. Black Iguana ( Varanus sp.) Kelsey Creek (20°26'S, 148°27'E), Queensland, Australia ROTHSCHILD TICKS IN BM(NH) 25 A. Fontaine Publication: Kaufman (1972: 369) K47 None Aponomma decorosum (Species epithet crossed out and undatum inserted) (1 d1, 1 9, 4 N Aponomma decorosum) ex. Iguana ( Varanus sp.) Australia 11.1.1911 Publication: Kaufman (1972: 370) K48 1912.10.31.21-25 2 cf Aponomma exornatum ex. Egyptian monitor (Varanus sp.) Bibianha [Bebaianiha (05'43'N, 00°27'W) or other populated place of this name nearby], Ghanaa 17.XII.1911 H. G. F. Spurrell K49 1912.6.21.387-390 Aponomma ecinctum (2 d1, 2 9 Aponomma fimbriatum) ex. Tiger snake (Notechis scutatus) Melbourne (37°49'S, 144°58'E), Australia 12.VII.1911 A. Coles Note: Very poor specimens; hypostomes lacking in all specimens. Capitulum present in only one 9 but from porose areas and scutal configuration of 9 and scutum and tarsi of cf these appear to be good A. fimbriatum. K50 1912.6.21.416 1 d Aponomma latum ex. Hystrix sp. Mduna River, Hlabisa District, (28°10'S, 32°10'E), Zululand Region, Republic of South Africa 9.III.1910 F. Toppin Received from Natal Museum, Pietermaritzberg H. Hoogstraal det. 1952 'unusual host' K51 1912.6.21.359-369 Aponomma gervaisi var. trimaculatus (11 d, 2 9, 1 N Aponomma trimaculatum) ex. Python sp. Satte'lberg, Huon Gulf (07°10'S, 147'25'E), Papua New Guinea 1911 Prof. F. Forster K52 None 2 9 Boophilus decoloratus Host unknown Tanzania Dr E. J. Baxter Boophilus decoloratus is a vector of the viruses Dugbe, Thogoto, Bhanja, Jos, Somone and Crimean Congo hemorrhagic fever. K53 1912.10.31.26-29 Haemaphysalis kochi(\ 9 Boophilus microplus; 2 d, 1 9, 1 N Haemaphysalis juxtakochi) ex. Deer Caparo (10'27'N, 61°20'W), Trinidad Island, Trinidad and Tobago 22.V.1912 S. M. Klages K54 1912.12.4.17 1 cf Dermacentor atrosignatus group ex. Sheep Sri Lanka K55 None Dermacentor reticulatus var. aulicus SYNTYPE ( 1 9 Dermacentor reticulatus) Host unknown Valescure, nr. St. Raphael (43°26'N, 06°46'E), France 2.IV.1910 Publication: Hirst (1916: 308) original description. K56 1912.6.21.77-97 2 d, 8 9 Dermacentor reticulatus ex. Canis familiaris Kuybyshev (55°40'N, 78°12'E), Tomsk, U.S.S.R. 27.VIII.1911 W. M. Thomas K57 1912.6.21.69-71 1 9 Haemaphysalis bancrofti ex. Horse Equus caballus Kelsey Creek (20°26'S, 148°27'E), Queensland, Australia 1911 A. Fontaine K58 1912.6.21.76 1 d Haemaphysalis bancrofti ex. Black Wallaby [Macropus dorsalis, the black- striped wallaby or Wallabia bicolor, the black, or black-tailed wallaby, both of which occur in Queensland]. Kelsey Creek (20°26'S, 148°27'E), Queensland, Australia 1911 A. Fontaine K59 None 2 d Haemaphysalis bancrofti ex. Wallaby ( Wallabia sp. or Macropus sp.) Kelsey Creek (20°26'S, 148°27'E), Queensland, Australia 1911 A. Fontaine K60 None 2 d Haemaphysalis bancrofti ex. Canis familiaris 26 J. E. KEIRANS Kelsey Creek (20'26'S, 148°27'E), Queensland, Australia 1911 A. Fontaine K61 None 2 rf Haemaphysalis calcarata Host unknown Voi (03°23'S, 38°34'E), Kenya 14.IV.1910 Robin Kemp K62 None Haemaphysalis leachi leachi (2 d1, 2 9 Haema- physalis leachi) Host unknown Tanzania Dr E. J. Baxter K63 1912.10.31.30-35 Haemaphysalis leachi leachi (9 d1, 2 9 Haema- physalis leachi) ex. Felis sp. Lake Suai, Ethiopia (locality not verified) [Hirst original det. Haemaphysalis sp.] K64 1912.6.21.33-35 2 rf, 2 9 Haemaphysalis leachi ex. Hyaena (Hyaena hyaena) Harar (09° 1 9'N, 42°09'E), Ethiopia 31.XII.1911 G. Kristensen K65 1912.6.21.36^4 1 0 d1, 8 9 Haemaphysalis leachi ex. Felis catus Harar (09° 1 9'N, 42°09'E), Ethiopia 22.1.1912 G. Kristensen K66 19 12.6.2 1.4 5-49 5 cf Haemaphysalis leachi ex. Marten Harar (09° 1 9'N, 42'09'E), Ethiopia 18.1.1912 G. Kristensen K67 1912.6.21.50-55 5 cf, 3 9 Haemaphysalis leachi ex. 'Badlington' dog Gola National Forest (06'56'N, 10°45'W), Liberia 17.VI.1910 R. H. Bunting K68 1912.6.21.56-58 1 d1, 2 9 Haemaphysalis leachi ex. Felis serval Mpwapwa (06°2 1 'S, 36°29'E), Tanzania 15.X. 1911 Marschner. Rec'd. from W. Fasius K69 None Haemaphysalis leachi (4 d Haemaphysalis leachi; 2 9 Ixodes thomasae) Host unknown Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 7.VI.1910 Robin Kemp K70 None 7 rf, 109 Haemaphysalis leachi Host unknown Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 6.VI.1910 Robin Kemp K71 None 4 rf, 4 9 Haemaphysalis leachi Host unknown Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 8.VI.1910 Robin Kemp K72 None 2 cf Haemaphysalis leachi ex. Domestic Rabbit (Oryctolagus sp. probably.) The genus Oryctolagus does not exist south of the Sahara unless bred in captivity. Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 20. VI. 1910 Robin Kemp K73 None 1 1 d1, 8 9, 1 N Haemaphysalis leachi Host unknown Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 11.VI.1910 Robin Kemp K74 None 25 d1, 1 1 9 Haemaphysalis leachi Host unknown Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 9.VI.1910 Robin Kemp K75 None 1 cf Haemaphysalis leachi ex. Genetta stuhlmanni Fennek, Mutoragwa Aberdare Mts, Kenya (locality not verified*) 23. HI. 19 10 Robin Kemp *There is a Mutarakwa Hill (00°47'S, 35°14'E) in the Aberdares that is a possible locality for this collection. ROTHSCHILD TICKS IN BM(NH) 27 K76 1912.6.21.59-65 Haemaphysalis leachi! (7 cf, 4 9 Haemaphysalis leachi group) ex. Paradoxurus niger (Paradoxurus hermaphroditus) Hoshangabad District (22°35'N, 77°40'E), India 1912 C. A. Crump Received from Bombay Nat. Hist. Soc. K77 1914.11.17.16-35 Haemaphysalis spinigera var. novae-guineae TYPE (19 cf, 4 9, 1 N Haemaphysalis novae- guineae paralectotypes) ex. Kangaroo Sattelberg, Huon Gulf (07°10'S, 147'25'E), Papua New Guinea 1911 Prof. F. Forster Publications: Hirst (1914: 328), original descrip- tion; Nuttall & Warburton (1915: 452); Roberts (1963:57) K78 1914.11.17.1-15 Haemaphysalis spinigera var. novae-guineae TYPE (1 cf Haemaphysalis novaeguineae lecto- type; 9 cf, 1 1 9 Haemaphysalis novaeguineae paralectotypes) ex. Perameles sp. ? Sattelberg, Huon Gulf (07'10'S, 147°25'E), Papua New Guinea Prof. F. Forster Publications: Hirst (1914: 328) original descrip- tion; Nuttall & Warburton (1915: 452); Roberts (1963: 57) selected lectotype and paralectotypes. K79 1914.11.17.36-46 Haemaphysalis spinigera var. novae-guineae TYPE (6 cf, 10 9, 1 N Haemaphysalis novae- guineae paralectotypes) ex. Hydromys sp. Sattelberg, Huon Gulf (07°10'S, 147°25'E), Papua New Guinea 1911 Prof. F. Forster Publications: Hirst (1914: 328), original descrip- tion; Nuttall & Warburton (1915: 452); Roberts (1963: 57) designated paralectotypes. K80 1912.6.21.66 1 cf Haemaphysalis parmata ex. Neotragus pygmaeus Bebaianiha (05°43'N, 00°27'W) or other populated places of this name nearby, Ghana 10.XI.1911 H. G. F. Spurrell K81 None 13 N Haemaphysalis spinigera ex. Lepussp. Hoshangabad District (22°35'S, 77°40'E), India 1912 C. A. Crump Received from Bombay Nat. Hist. Soc. Haemaphysalis spinigera is a vector of the viruses Kaisodi and Kyasanur forest disease. K82 1912.10.31.36-46 Haemaphysalis sp. (28 cf, 7 9 Haemaphysalis spinulosa) ex. Wild cat Harar (09° 1 9'N, 42°09'E), Ethiopia 6. 11.1912 G. Kristensen K83 None Haemaphysalis leachi] (1 9 Haemaphysalis spinulosa) ex. Canis familiar is Himo River (03°33'S, 37°30'E), Tanzania 30.V.1910 Robin Kemp K84 19 12.6.21. 22-32 pt Haemaphysalis cinnabarina var. punctata (2 cf, 3 9 Haemaphysalis sulcata) ex. Sheep Larnaca (34°55'N, 33°38'E), Cyprus XI.1911 C. B. Horsbrugh K85 19 12.6.21. 22-32 pt 2 cf , 3 9 Haemaphysalis sulcata ex. Sheep Larnaca (34°55'N, 33°38'E), Cyprus XI.1911 C. B. Horsbrugh Note: A portion of this collection was originally entered under Haemaphysalis cinnabarina var. punctata. K86 None Haemaphysalis warburtonil (1 cf, 2 9, 2 N Haemaphysalis tibetensis TYPES 1 cf (holotype) 2 9, 2 N (paratypes) ex. Canis familiar is Yatung (27°25'N, 89 WE) (ca. 3896 m) mouth of Chumbi Valley, Tibet D. Macdonald Publication: Hoogstraal (1965: 452), original description. K87 1912.6.21.247-250 Hyalomma syriacum (3 cf, 1 9 Hyalomma aegyptium) ex. Testudo graeca ibera Hammam Rirha (36°26'N, 02°28'E), Algeria 24. V. 1911 Hon. Walter Rothschild & Ernst Hartert 28 J. E. KEIRANS K88 1912.6.21.251-253 Hyalomma syriacum (Species epithet crossed out and aegyptium inserted) (4 9 Hyalomma anatolicum excavatum) Host unknown North West Morocco Received from Hon. W. Rothschild K89 1912.6.21.240-6 Hyalomma aegyptium (2 cf, 1 9 Hyalomma marginatum) Host unknown Hammam Rirha (36°26'N, 02°28'E), Algeria 1911 C. Hilgert Hyalomma marginatum is a vector of the viruses Dhori, West Nile, tickborne encephalitis, Bhanja, Crimean Congo hemorrhagic fever, Tamdy and Batken. K90 19 11. 12.9. 1-320 (pt) Hyalomma aegyptium (2 9 Hyalomma truncatum) ex. Aged donkey (Equus asinus) Voi (03°23'S, 38'34'E), Kenya 29.IV.1910 Robin Kemp K91 None 8 N Hyalomma sp. Host unknown Biskra (34°50'N, 05°40'E), Algeria 25.111.1909 Hon. L. W. Rothschild K92 19 19.8. 14. 1-70 (part) 3 N Hyalomma sp. ex.? Dipodillus (gerbil) (Gerbillus (Dipodillus) sp.) ('host similar to 108') Guelt-es-Stel (35°09'N, 03°02'E), Algeria IV.1912 Hon. Walter Rothschild and Karl Jordan K93 1912.6.21.202 Rhipicephalus sp. (4 N Hyalomma sp.; 1 9 Rhipicephalus pulchellus; 10 N Rhipicephalus sp. ex. Lepus sp. Harar (09° 1 9'N, 42°09'E), Ethiopia 9.1.1912 G. Kristensen Rhipicephalus pulchellus is a vector of the viruses Dugbe, Crimean Congo hemorrhagic fever, and Nairobi sheep disease. K94 None Ixodes var. offecialis (2 9 Ixodes alluaudi) ex. Lophuromysjlavopunctatus zena Aberdare Mts (00°25'S, 36°38'E), Kenya 14.11.1910 Robin Kemp K95 None Ixodes var. offecialis ( 1 9 Ixodes alluaudi) ex. Lophuromys fJavopunctatus zena Aberdare Mts (00°25'S, 36°38'E), Kenya 1 1 ,000 ft (3,353 m) 21.11.1910 Robin Kemp K96 None Ixodes tenuirostris (2 9 Ixodes alluaudi) ex. Otomys tropicalis elgonis Mt Ilkinangop (00°38'S, 36°42'E), Aberdare Mts, Kenya 24.11.1910 Robin Kemp K97 None Ixodes tenuirostris (1 9 Ixodes alluaudi) Host unknown Rombo, Kilimanjaro (03°05'S, 37°20'E), Tanzania 15.IV.1910 Robin Kemp Note: This 9 has been previously pinned. K98 None [undetermined genus & species] (1 N Ixodes alluaudi) ex. Mus jacksoni (Possibly Anomalurus derb ianus Jackson i) Mutaragwa, Aberdare Mts (locality not verified*), Kenya 13.111.1910 R. Kemp *There is a Mutarakua Hill (00°47'S, 35°14'E) in the Aberdares that is a possible locality for this collection. K99 1912.6.21.280-2 4 9 Ixodes cordifer ex. Pseudoircornia (probably Pseudocheirus sp.) Sattelberg, Huon Gulf (07°10'S, 147°25'E), Papua New Guinea 1911 Prof. F. Forster K100 1912.6.21.283 1 9 Ixodes cordifer ex. Dactylopsida trivirgata Sattelberg, Huon Gulf (07°10'S, 147°25'E), Papua New Guinea Prof. F. Forster Note: label in vial — seen by Prof. R. A. Cooley 1946. 3 slides of dissections (RML 22285). K101 1912.6.21.284-8 4 9 Ixodes cordifer ex. Phalanger maculatus Sattelberg, Huon Gulf (07°10'S, 147'25'E), Papua New Guinea 1911 Prof. F. Forster ROTHSCHILD TICKS IN BM(NH) 29 K102 1912.6.21.279 1 d1 Ixodes cordifer ex. Perameles sp. Sattelberg, Huon Gulf (07°10'S, 147°25'E), Papua New Guinea 1911 Prof. F. Forster K103 1912.12.4.11 1 9, 4 N Ixodes fecialis cx Fclis so Studley Park, Melbourne (37°49'S, 144°58'E), Australia 19.V.1911 A. Coles Note: Reverse of the vial label bears the inscription — Native cat. K104 1912.6.21.254— 6 5 9, 2 N Ixodes hexagonus ex. Mustela vulgaris (Mustela nivalis) Tring (5 1 °47'N, 00°39'W), England 12.111.1912 J. F. Cox K105 1912.6.21.259-61 1 9, 2 N, 7 L Ixodes hexagonus (1 9, 2 N Ixodes hexagonus) ex. Mustela erminea Tring (5 1 °47'N, 00°39'W), England III.1912 J. F. Cox K106 1912.6.21.262-3 4 N Ixodes hexagonus (4 N, 8 L Ixodes hexagonus) ex. Meles taxus (Meles meles) Senj (45WN, 14°54'E), Yugoslavia 5.XII.1911 F. Dobiasch K107 None 22 N, 6 L Ixodes hexagonus ex. Mustela erminea Loch Tay (56°28'N, 04° 1 8'W), Lawes, Scotland 2.III.1911 L. G. Esson KlOSNone 1 9 Ixodes hexagonus Host unknown Faugeres*, S. France 8.X.1913 W. Davy N. C. Rothschild *Three possible localities for Faugeres, France: 43°34'N, 03'11'E; 44°28'N, 03°58'E; 44°58'N, 04°35'E. K 109 None 2 N Ixodes hexagonus ex. Mustela putorius Carmarthen (5 1°52'N, 04° 1 8'W), Wales 1914 F. W. Frohawk Kl 10 1919.8. 14.1-70 (part) 1 9 Ixodes hexagonus ex. dog Canis familiaris Bridgnorth (52°32'N, 02°25'W), England 28.X.1916 Miss Frances Pitt Kill 1912.12.4.9-10 1 rf, 1 9 Ixodes loricatus ex. Didelphis marsupialis aurita Mar de Espanha (20°52'S, 43'00'W), Estado de Minas Gerais, Brazil 6.X.1910 Z. F. Zikan K112 1919.8. 14.1-70 pt 2 0-5 mm long, with lines running downwards from rim (sometimes absent in one species) 22 Hydrotheca < 0-5 mm long, without lines 23 22 Lines meeting tips of cusps, which are roundly pointed, not notched; hydrotheca 2+ times long as broad [probably not south of Newfoundland and Spitzbergen] Campanularia crenata (p. 52; Fig. 2) Lines meeting bottoms of embayments; cusps flat-topped with notch; hydrotheca up to 1*5 x long as broad [widespread] Campanularia hincksii (p. 53; Fig. 3) 23 . Hydrotheca usually much thickened [probably Mediterranean southwards] Orthopyxis crenata (p. 58; Fig. 5) - Hydrotheca unthickened [Mediterranean northwards] . Campanularia volubilis (p. 55; Fig. 4) 24 Hydrothecal cusps bimucronate [S England southwards] . Clytia paulensis (?. 88; Fig. 14) - Hydrothecal cusps simple [widespread] Clytia hemisphaerica (p. 73; Fig. 9; also C. mccradyi, ?S France only; p. 87; Fig. 13) N.E. ATLANTIC CAMPANULARIID HYDROZOANS 47 Taxonomic section Family CAMPANULARIIDAE Johnston, 1836 DIAGNOSIS. Colonial Thecata (sens. Millard, 1975); hydroid stage stoloniferous or erect, stolon when present may be branched or unbranched; hydrotheca bell-shaped, radially symmetrical, pedicellate, with diaphragm and an associated annular thickening, or with annulus alone and without diaphragm; no operculum; hydranths radially symmetrical, usually with prominent hypostome; no caecum; one ring of tentacles; nematophores absent; d" & 9 gonothecae usually externally identical. Medusa generation when present variable; reduced in Obelia and (?) facultatively retained in Orthopyxis; typical leptomedusan in Clytia and Gastroblasta; always retained as gonomedusa in Gonothyraea; identifiable in more reduced form in at least some other genera and species. REMARKS. The family was first proposed by Johnston (1836, 1847, but not 1838). Originally Lafoea dumosa (Fleming, 1 820) was included in its scope but was removed to the Lafoeidae by Hincks (1868). The limits of the family have remained unchanged ever since and the redefinition by Millard (1975) seems sound. Ralph (1957) also provided a detailed appraisal. Much has been written about generic limits within the family but it is convenient to go no further back in the literature than the works of Broch (1905, 1910) and Goette (1907). Broch recognized two broad genera. These he called Campanularia Lamarck, 1816, which had no hydrothecal diaphragm, and Laomedea Lamouroux, 1812, which had one. He divided his concept of Campanularia into the subgenera Eucampanularia Broch, 1910, having sessile gonophores, and Clytia Lamouroux, 1812, with free medusae. Broch split his other broad genus concept, Laomedea, into the three subgenera Eulaomedea Broch, 1910, with sessile gonophores and no medusoid structures; Gonothyraea Allman, 1864a, with retained 'eumedusoids' (now called gonomedusoids, p. 93); and Obelia Peron & Lesueur, 1810a, with free medusae. Thus Broch's basic division within the family was on a hydrothecal character; while within each of the two main divisions his classification was on the state of reduction of the medusa generation (following and elaborating on the interpretation of these structures by Goette, 1 907). Splettstosser (1924 : 424^425) followed Broch's system but further split Broch's subgenus Eulaomedea into one group with intracapsular gonophores ('Laomedea gelatinosa\ L. flexuosa and L. calceolifera] and a second, in which the mature gonophores were extra- capsular (L. neglecta). Splettstosser acknowledged that the classification might be criticized since just a small number of species was considered. But Broch (1928) gave it support when he later introduced the subgeneric name Paralaomedea for the 'L. neglecta group', comprising that species alone, in the combination 'Laomedea (Paralaomedea) brochi Splettstosser (= Laomedea neglecta Alder)'. [Splettstosser's restriction of the subgenus concept was thus cited as indication by Broch; but the authority for the associated subgenus name Paralaomedea was Broch (1928).] Finally Hummelinck (1936) redefined the subgenus rather tightly, again to include only L. neglecta. As Splettstosser had commented, very few species were considered in his classification and it is questionable whether so many infra-generic divisions were justified. (Some additional sub-divisions of Eulaomedea' proposed by Splettstosser were not given names, and are not mentioned here.) His own work on L. neglecta and other species, and that for example of Goette (1907) and Miller (1973), emphasized that the gonophore 'types' identified by Splettstosser form part of a series in which the medusa is progressively reduced. The gonophore of L. neglecta seems simply to fit into this series. Further, Broch's (1910) primary division of the family into two was on the basis of a single hydrothecal character, and this division too might be challenged. The extent to which the medusa-medusoid-gonomedusoid-gonophore series should be classified into genera will perhaps be debated for as long as the series is regarded valid. But today as in Broch's time, most is known about the life-cycles of the western European 48 P. F. S. CORNELIUS species. Until more information is available on species from other parts of the World it seems unwise to split Laomedea into subgenera. Aside from the taxonomic debate, there are some nomenclatural problems which need solution. These I have considered in a submission to the International Commission on Zoological Nomenclature (Cornelius, 1981). The aims of the proposals are provisionally included in the present paper. The subgenera proposed by Broch, Splettstosser and others need not be recognized. But the evolutionary fate of the medusa generation is still reflected in the classification adopted here. In Orthopyxis the medusa is reduced and lacks several normal adult characters, functioning simply as an ephemeral gamete carrier. It is thought to be facultatively released in some or all of the Orthopyxis species (see p. 63); while in the closely related Orthonia it is still further reduced, to a retained acrocyst. In Campanularia, Rhizocaulus and Silicularia the gonosome has become intracapsular, with no obvious indications of a medusoid ancestry. A parallel series showing progressive retention of the medusa can be demonstrated within the genera Obelia, Gonothyraea, Laomedea and Hartlaubella. In Obelia the medusa is released. The extra-capsular gonomedusoids of Gonothyraea are easily identified as retained and vestigial medusae; and in Laomedea it has been shown that the gonosomes of several species represent reduced medusae. Indeed, so reduced has the medusa generation of Laomedea become that until the work of Miller (1973; see also Goette, 1907) the medusoid nature of the gonophore was not appreciated. The same confusion prevailed also in interpretations of the reproductive structures in Orthopyxis Integra, in which the medusa is sometimes retained. It was thought until quite recently that the retained examples had 'sporosacs' in place of medusae, and that they might therefore be a different species (O. caliculata', p. 65-66)! The genera Clytia, Gastroblasta and Tulpa, which have a true hydrothecal diaphragm and sub-hydrothecal spherules, apparently form another group but their relation to the rest of the family is not clear. The three series recognized are shown in Figure 1 . The groupings seem natural and are here given subfamily status: Campanulariinae (p. 50), Clytiinae nom. nov. (p. 69) and Obeliinae Haeckel, 1 879 (p. 9 1 ). See also page 49. The generic limits suggested by Millard (1975) are slightly modified, as is her nomenclature. Orthopyxis is here separated from Campanularia sens. Millard. Eulaomedea sens. Millard is here called Laomedea. Sertularia gelatinosa Pallas, 1766, not in Millard's faunal area, is referred to the monotypic genus Hartlaubella; and another species not in her list, S. verticillata Linnaeus, 1758, is here referred to the nearly monotypic Rhizocaulus. To promote stability of nomenclature I have attempted to include all extra-limital generic synonyms. The valid genera found outside the NE Atlantic are treated briefly. They are Eucalix, Orthonia and Silicularia from the Campanulariinae (p. 50); and Gastroblasta and Tulpa from the Clytiinae (p. 70). The problem genus Hypanthea is discussed along with Silicularia (p. 50). From the medusa stage, Kramp (1961) recognized only five genera World-wide: Agastra, Eucopella, Gastroblasta, Obelia and Phialidium\ but of these only Gastroblasta and Obelia can now be recognized. Reference to discussions of these genera can be made using the index.. . I have previously commented (Cornelius, 1975a) on the genera Medusa Linnaeus, 1758 (part); Schizocladium Allman, 1871; Obelaria, Obeletta & Obelissa, all Haeckel, 1879 (Obelaria Hartlaub, 1897, is a junior homonym and is discussed here under Hautlaubelld)', and Monosklera von Lendenfeld, 1885; all except the first of which fall into the synonymy of Obelia Peron & Lesueur, 1810#. I then mentioned also Thaumantias Eschscholtz, 1829, a junior subjective synonym of Clytia Lamouroux, 1812 (see below, p. 71). I overlooked (p. 254) that Mayer (1910: 262) had designated Sertularia volubilis sens. Ellis & Solander, 1786 (non Linnaeus, 1758) type species of Clytia (see p. 70 below). Lastly, I have reversed my opinion on the use of the genus name Laomedea. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 49 Medusa entirely suppressed, intracapsular Medusa reduced to external acrocyst Medusa vestigial and retained Medusa released Hartlaubella, Laomedea (part) Laomedea negleata Gonothyraea Siliaularia Campanularia, Tulpa Rhizocaulus Orthopyxie (part) Orthopyxie (part) Clytia, Gastroblasta OBELIINAE CAMPANULARI INAE CLYTI INAE Fig. 1 Affinities within the Campanulariidae. The genus Eucalix is not included since its method of reproduction is unknown, but vegetative characters suggest it is close to Orthopyxis (discussion on pp. 50-51). The diagram shows present-day similarities, not phylogenies, but the general direction of evolutionary advance is up the pagejand towards the right. Knowledge of the group is incomplete and the chart should be regarded as provisional. Some of the species described herein are known only from either hydroid or medusa stages, and others were formerly so. Most species in which the two stages have been reconciled are now known by appropriate combinations, based on application of the International Code of Zoological Nomenclature; but those with incompletely known life-cycles cannot yet have their names confidently derived. Current knowledge in this family seems adequate for the Code to be applied to the two stages simultaneously, but this is arguably not so in all hydro- medusan families and in some there may still be a case for retaining the dual system. The subfamily divisions and their nomenclature Although the limits of the family Campanulariidae have been agreed for nearly a century and a half (p. 47) only three authors (Haeckel, 1879; Mayer, 1910; Russell, 1953) have sought to group the genera into formal subfamilies. Indeed, until some quite recent studies of the range of reproductive structures found within the family had appeared (Miller, 1973; but also Splettstosser, 1 924) interpretation and grouping had been difficult. Miller's important work showed that the structures which had once been called fixed gonophores in for example Laomedea spp. were vestigial, retained medusae; and that the curious externally-held 'meconidium' ofGonothyraea loveni is similarly to be regarded as a retained medusa. Happily, this new interpretation of the dispersive generations (planulae and medusae) corroborates the broad divisions of the family based long ago solely on the vegetative characters of the hydroid stage (Broch, 1905, 1910; Goette, 1907; see p. 47, above). Hence the subfamily divisions adopted here, which draw on both groups of characters, might seem soundly based. But some problems remain and further refinement will no doubt be achieved when more is known of the non-European members of the family. 50 P. F. S. CORNELIUS Subfamily CAMPANULARIINAE Campanulariadae Johnston, 1836 : 107 (part). Obelidae Haeckel, 1879 : 163 (part). Obelinae: Mayer, 1910 : 231 (part). Orthopyxinae Russell, 1953 : 303. non Campanularinae: Russell, 1953 : 284 ( = Clytiinae nom. nov.; see p. 69). NOMENCLATURE. The spelling Campanulariinae takes as its root the genus name Campanularia, and Campanularinae is wrong. DIAGNOSIS. Campanulariidae with colony usually replant, secondarily erect and poly- siphonic in Rhizocaulus; no true hydrothecal diaphragm; medusa absent except in Orthopyxis, where reduced. TYPE GENUS. Campanularia Lamarck, 1816, the nominate genus. SCOPE. The genera Campanularia Lamarck, 1816; Silicularia Meyen, 1834; Orthopyxis Agassiz, 1862; Rhizocaulus Stechow, 19196; Orthonia Stechow, 19230; and probably Eucalix Stechow, 192 la. REMARKS. Of the included genera only Campanularia, Orthopyxis and Rhizocaulus are fully treated in this paper. The others have not been recorded from the eastern North Atlantic and are discussed only in this section. Millard (1975:201) united Campanularia and Orthopyxis because she had seen Orthopyxis colonies having some unthickened hydrothecae; but I feel the remaining characters justify a separation. The genus Silicularia Meyen, 1834, was proposed to include two species, S. rosea and S. gracilis, both being described as new. The early date of Silicularia and inadequacies in the descriptions of the two species make detailed comments necessary. The type species of Silicularia is S. rosea, designated by Stepanyants (1979 : 33). The species was based on Ethiopian and South African material. It was redescribed by Nutting (19 1 5) and Stepanyants (1979), and Blanco (19670) provided useful notes. The second species, S. gracilis, was based on infertile hydroid material from the Sargasso Sea and the Azores. It was probably a Clytia species. However, the figures and description do not include details of the hydrothecal rim or reproductive structures and I agree with Bedot (1905 : 171) that the species cannot be confidently assigned (see also p. 118). Nutting (1915 : 66) referred 'S. gracilis' to the rather dubious species Orthopyxis clytioides (Lamouroux, in Freycinet, 1824, as Tubularia). He wrongly quoted Meyen as using the combination Silicularia clytioides. Meyen actually used S. gracilis. Rees & White (1966) made the same error when citing Meyen's Azores record. There seem no other reports of 'S. gracilis' from the eastern North Atlantic. I provisionally refer T. clytioides Lamouroux to Obelia dichotoma, under which it is discussed further (p. 1 1 8). S. gracilis was mentioned recently by Stepanyants ( 1 979), as Campanularia. Nutting (1915), Bedot (1925), Broch (1929) and Stepanyants (1979) all regarded as congeneric with Silicularia the later genus Hypanthea Allman, 18760 (type species H. repens Allman, 18760, by monotypy; type locality of the type species, Kerguelen I.), and I agree. Hypanthia Nutting, 1915 : 22, was a lapsus. Both Allman's (18760, 1888) concept of Hypanthea and Nutting's and Stepanyants' of Silicularia included thick, asymmetrical hydrothecae, pedicels and stolons reminiscent of Orthopyxis Agassiz, 1862, to which the original concepts of the two genera come close. I have not located the type material of H. repens, but later material referred to Hypanthea species by Allman (1888) had an anastomosing stolon and other orthopyxine features. So far as I can determine no Silicularia species has been recorded from the eastern North Atlantic. S. atlantica (Marktanner-Turneretscher, 1890, as Hypanthea), was based on material said to have come from 6° S, 38° W, but this position is on the mainland of South America! The genus Eucalix Stechow, 19210 : 254, was proposed to accommodate the sole species N.E. ATLANTIC CAMPANULARIID HYDROZOANS 51 Campanularia retroflexa Allman, 1888, type locality Honolulu. Stechow maintained that the unusual hydrotheca of E. retroflexus justified generic separation. Both the type series (BMNH reg. no. 1888.11.13.14) and the original illustration (Allman, 1888 : pi. 11, figs 1, la) show the hydrothecal characters regarded important by Stechow. These characters, together with the anastomosing stolon of the type material, suggest that Stechow was justified in proposing the new genus. Millard (1957 : 196; 1975 : 212, as Campanularia morgansi) listed relevant literature. (I am grateful to Professor W. Vervoort for discussing the characters of this species; and to Dr D. M. Devaney for identifying the substrate of the type material. The substrate is a coralline alga, Halimeda sp., not a millepore as Allman stated. Dr Devaney informs me millepores have not been recorded from Hawaii.) The genus Orthonia Stechow, 19230 : 94, 107, was proposed to accommodate a single orthopyxine species, Campanularia everta Clarke, 1876:253-254, pi. 39, fig. 4, type locality San Diego. Nutting had subsequently assigned to this species material having acrocysts and it was this character on which Stechow distinguished the genus from Orthopyxis. An element of subjectivity was thereby introduced since Stechow assumed that Nutting had material of the same species as had Clarke; and Stechow had no proof. Further revision of the orthopyxine species seems necessary before Orthonia is evaluated further. Genus CAMPANULARIA Lamarck, 1816 Campanularia Lamarck, 1816:112 (part); Hincks, 1868 : 160 (part); Nutting, 1915 : 27 (part). Campanula Westendorp, 1843 : 23 (lapsus pro Campanularia}. Campanulata Agassiz, 1862 : 354 (lapsus pro Campanularia). Campanularia (Eucampanularid) Broch, 1910 : 184 (part). Paracalix Stechow, 1923c : 3. TYPE SPECIES. Provisionally to be taken as Sertularia volubilis Linnaeus, 1758 : 811 (non Ellis & Solander, 1786, see p. 70) as designated * by Naumov (1960: 249). Nutting (1 9 1 5 : 28) earlier designated S. verticillata Linnaeus, 1758 : 8 1 1 , as type species but applica- tion has been made to the International Commission on Zoological Nomenclature for this designation to be set aside (Cornelius, 1981; see Remarks). Broch (1905 : 10) proposed that 'Campanularia calyculata Hincks, 1853' should be type species, but 'calyculata' was not among the species originally included in the genus and so is not eligible. The correct spelling is of course caliculata (p. 65). DIAGNOSIS. Stoloniferous and colonial Campanulariidae, stolon not anastomosing; hydro- thecae borne on pedicels inserted on the stolon at irregular intervals; true diaphragm absent; sub-hydrothecal spherule present; no medusa stage. REMARKS. The species Sertularia verticillata Linnaeus, 1758, was designated type species of Campanularia by Nutting (1915). But some authors, with whom I agree, have sought to remove verticillata to a distinct genus (Stechow, 19 \9b, c; Naumov, 1960, 1969). This would leave the name Paracalix Stechow, 1923c, available for the present genus; so that Paracalix would become applied for example to the common hydroids widely known as Campanu- laria hincksii (p. 53) and C. volubilis (auct.; p. 55). The genus Paracalix Stechow, 1923c, was proposed to accommodate only Campanularia pulcratheca Mulder & Trebilcock, 1914: 11, pi. 2, figs 1-2, a species based on sterile material from Torquay, Victoria, Australia. (The generic name was actually misprinted Cmpanularia in Mulder & Trebilcock's heading.) The hydrotheca was sigmoid in lateral view and Stechow was impressed by the resulting bilateral symmetry. This was the main character on which the species, and subsequently Stechow's proposed genus, were based; but the specimen seems simply to have been a deformed specimen of C. volubilis or a closely related species. On this interpretation the species pulcratheca and the genus Paracalix are referred to Campanularia. I have applied to the International Commission on Zoological Nomenclature for Nutting's designation of S. verticillata as type species of Campanularia to be set aside (Cornelius, 1981). If approved, this will validate Naumov's (1960) designation of S. volubilis Linnaeus, 52 P. F. S. CORNELIUS 1758, as type species of Campanularia. As a consequence the genus name Rhizocaulus Stechow, 19196, can then be applied to the species verticillata (in the combination R. verticillatus, p. 67). See also the notes under Rhizocaulus (p. 67). The subgenus Eucampanularia Broch, 1910, was introduced to embrace the five species Sertularia volubilis Linnaeus, 1758, Campanularia Integra Macgillivray, 1842, C. groen- landica Levinsen, 1893, C. speciosa Clarke, 1877 and S. verticillata Linnaeus, 1758. The subgenus name has hardly been used in the literature. I designate S. volubilis Linnaeus, 1758, as its type species; so that Eucampanularia can be regarded a junior objective synonym of Campanularia (subject to my proposals to the ICZN being accepted; see also Cornelius, 1981). Campanularia crenata Allman, 1 8766 (Fig. 2) Campanularia crenata Allman, 1 8766 : 258-259, pi. 11, figs 1-2. Campanularia speciosus Clarke, 1877:210 (lapsus pro speciosa). Campanularia speciosa (Clarke, 1877:214-215, pi. 9, fig. 11; Linko, 1911 : 185-187, fig. 34 (syn. C. crenata Allman); Broch, 1912a : 17-18, fig. 3; Nutting, 1915 : 48, pi. 8, fig. 5 (syn. C. crenata Allman); Broch, 1918 : 158-159 (syn. C. magnifica Eraser); Calder, 1970 : 1519, pi. 4, fig. 3. Campanularia magnifica Eraser, 1913 : 164, pi. 1 1, figs 1-3. NOMENCLATURE. The widely used trivial name speciosa was introduced in a paper published on 2 January, 1877, and not in 1876 as usually assumed. Hence crenata, genuinely introduced in 1 876, has priority (see note on page 1 29 under Clarke, 1877). The combination Campanularia crenata has been applied also to the species here called Orthopyxis crenata; but in that species the original binominal was Eucopella crenata, and primary homonymy has not occurred. There is secondary homonymy, however, and this is discussed under O. crenata (p. 60). TYPE LOCALITY AND MATERIAL. Infertile colony on 'Thuiaria crassicaulis* (Sertulariidae), ?Tsuger Straits, Japan, 183m (lOOfms); 1877.4.12.8 (previously unpublished data with specimen). OTHER MATERIAL EXAMINED. All BMNH material is listed. 45 m, Store Hellefiskebanke, Greenland, fertile colony on Sertularia mirabilis (Verrill, 1873), coll. G. M. R. Levinsen, exch. Copenhagen Mus.; 1896.8.15.2 (Fig. 2; ?mentioned, Broch, 1918). 'Greenland', infertile colony on Sertularia mirabilis, ex D'Arcy Thompson colln, pres. Univ. Dundee; 1957.1.1.12. Infertile colony on Sertularella sp., Norman St, Labrador, Canada, ex D'Arcy Thompson colln, pres. Univ. Dundee; 1956.10.23.69. Infertile colony on Symplectoscyphus sp., Bell I, Newfoundland, Canada, 17 Apr 1892, ex D'Arcy Thompson colln, pres. Univ. Dundee; 1957.1.3.24. Infertile fragment, Bel Sund, Spitzbergen, 14 Jul 1898, 20m, coll. Spetsberg Expedn, exch. Stockholm Mus.; 1960.8.29.33. No locality, fertile colony on sertulariid hydroid, exch. Copenhagen Mus.; 1 9 1 2. 1 2.2 1 .44. DESCRIPTION. Colony reptant. Stolon tortuous, branched, rugose. Hydrothecae on usually long pedicels, at irregular intervals; large, narrowest c. \ from rim, bulging out basally; rim much flared, with c. 10 rounded cusps, usually with striations running proximally from apex of each cusp; sub-hydrothecal spherule present; pedicel usually longer than hydrotheca, up to c. 3x length, spirally grooved throughout. Gonotheca ?rf = 9, elongate-ovoid, with or without long neck, borne on stolon. Variation. The short necked gonotheca illustrated was apparently mature, indicating that the long necks usually regarded distinctive are not invariably present. DISPERSIVE STAGE. Planulae, which develop within the gonotheca. Present material (1896. 8. 15. 2) has just one in each gonotheca but there may have been more in life. REPRODUCTIVE SEASON. No information. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 53 DISTRIBUTION. A circumpolar Arctic Ocean and cold water species recorded in the Atlantic as far south as Newfoundland (present material), W Greenland and Spitzbergen (Broch, 19120;Calder, 1970). HABITAT. Usually reported epizoic on other hydroids. Naumov (1960, 1969) found an extreme depth range of 3 m to 600 m in Russian seas, most of his records being between 20 m and 200 m. REMARKS. This species has been widely known by the combination Campanularia speciosa. The long gonothecal neck often thought characteristic of the species is not present in all the material listed here, and is evidently variable in length. Broch's (1918) attack on the validity of Campanularia magnifica Eraser, 1913, left no doubt that it is conspecific. Campanularia hincksii Alder, 1856a (Fig. 3) Campanularia hincksii Alder, 1856a:360, pi. 13, fig. 9; Hincks, 1868: 162-163, pi. 24, fig. 3; Goette, 1907 : 189-193, pi. 15, figs 307-312; Broch, 1933 : 87-93 (syn. C. aha Stechow); Vervoort, \946a : 276-277, fig. 122 (syn. C. aha Stechow); Patriti, 1970 : 33-34, fig. 41 (syn. C. brachycaulis Stechow, 1919a, here referred to Clytia hemisphaerica, see p. 82; C. macrotheca Leloup); Millard, 1975 : 208, fig. 67b-e. Campanularia aha Stechow, 1919a : 54-57, fig. P. Campanularia rara Stechow, 1 9 1 9a : 60-6 1 , fig. R. Campanularia macrotheca Leloup, 1930a : 101-102, figs 1-3. TYPE LOCALITY AND MATERIAL. Coast of Northumberland, England (Alder, 1856a; Millard, 1975). The syntype series is preserved jointly in the Hancock Museum, Newcastle upon Tyne, Northumberland (several colonies in spirit, epizoic on sertulariid hydroids) and the BMNH [small dry colony, 1857.8.3.58, epizoic on Lafoea dumosa (Fleming, 1820)]. It has been catalogued by Cornelius & Garfath (1980). TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia aha Stechow, 19190, infertile fragment of syntype on microslide, Naples; Munich Zoological Museum. C. rara Stechow, \9\9a, infertile fragment on microslide, Marseille; MZM. OTHER MATERIAL EXAMINED. BMNH collection, c. 50 specimens. The following, collected by W. J. Rees, had fertile d" gonothecae: Hjeltefjord, nr Bergen, Norway, 40-90 m, 9 Apr 1962, 1962. 10.7.20; I of Cumbrae,W Scotland, 90m, 11 Jul 1966; 1967.12.1. 10-12. DESCRIPTION. Colony a tortuous stolon bearing unbranched hydrothecal pedicels at irregular intervals. Hydrotheca large, campanulate, truncate basally; length : breadth ratio variable (1-3-2-25:1, Millard, 1975); rim castellate, 8-15 blunt cusps each usually notched, occasionally deeply; main embayments deep, curved, often conspicuous, with characteristic folds trailing down from centres. Hydrothecal pedicel long, with spherule distally; shaft smooth to sinuous, usually with several annuli basally, sometimes also 1 - several annuli along length (Vervoort, 1946a). Hydranth ?undescribed, 18-24 tentacles visible in contracted BMNH material. Gonothecae cf = 9, borne on stolon; sub-cylindrical, sometimes asymmetrical; broadest near base, truncate below, tapering gradually above; sides smooth to irregularly sinuous in a loose succession of rings; truncated and sometimes slightly flared distally; aperture wide, terminal; planula development probably internal; gonothecal pedicel short, ringed; colonies dioecious. Variation. The BMNH series shows variation in the following features: size and length : breadth ratio of hydrotheca, height and number of cusps, depth of notch in cusps, presence or absence of longitudinal folds in hydrothecal wall; sinuosity of perisarc of hydrothecal pedicel, length of pedicel, number of basal annulations (may be absent), shape of proximal cavity in hydrotheca; sinuosity of gonothecal wall, amount of flaring below gonothecal aperture. Billard (1934) reported that the notch in the tips of the hydrothecal 54 P. F. S. CORNELIUS cusps may be absent, when the hydrothecae sometimes resemble those of Campanularia volubilis(p. 55). DISPERSIVE STAGE. Planulae, which probably develop within the female gonotheca. Develop- ment of the male gonomedusoid was described by Goette (1907). REPRODUCTIVE SEASON. Jun-Oct in NW France (Teissier, 1965). BMNH fertile material has collection dates within these limits except a male specimen from near Bergen, dated 9 Apr 1962. DISTRIBUTION. Nearly cosmopolitan in shallow waters. Although not the most abundant hydroid C. hincksii can be expected almost throughout the eastern North Atlantic, local conditions permitting. Notable records include: N & S Iceland, Lofoten Is & Norway (Kramp, 1938); Mediterranean (Picard, 19586); Italy (Rossi, 1971); Portugal (Da Cunha, 1950); Cap Spartel, Tangier & Cap Blanc, Morocco (Billard, 1907); Azores (Rees & White, 1966); Mauritania (Billard, 193 la); South Africa ('rare', Millard, 1975). The species is widespread in parts of temperate western Europe, including the British Isles (Hincks, 1868), but is scarce in Dutch and Belgian waters (Vervoort, 19460; Leloup, 1952). There are several records from the Skagerrak and Kattegat (Kramp, 1935) and W Sweden (Jagerskiold, 1971), but no records from the Baltic Sea (Stechow, 1927; Broch, 1928; Naumov, 1960, 1969) or Black Sea (Naumov). HABITAT. Usually recorded between 20 m and 200 m but occasionally deeper: 'a few metres down to 800m' (Kramp, 1938); 20-100 m, SW England (Marine Biological Association, 1957); c. 20 m, SW Wales (Crothers, 1966); 25-50 m, Scilly Is (Robins, 1969); below 20 m, NW France (Teissier, 1965); 1 12-120 m, Strait of Gibraltar & Morocco (Billard, 1907); 27-98 m, Azores (Rees & White, 1966); 86-210 m, southern Africa (Millard, 1975). Shallow records include: 15 m, NW Wales (Knight-Jones & Jones, 1956); 10-1 12 m, Faeroes (Kramp, 1929); 9-5-80 m, W Sweden (Jagerskiold, 1971). Apparently no intertidal records. The species seems unrecorded from brackish waters and may be stenohaline. REMARKS. Millard (1975) summarized the doubts concerning the shape of the d gonotheca, which it seems has not been reported before now. The BMNH series includes several colonies in which the gonothecal contents are preserved. The d1 and 9 gonothecae are identical, and are borne on separate colonies. The contents of the cf were described by Goette (1907) and are clearly gonomedusoid in Miller's (1973) terminology. The name applied to the distinct but closely related nominal species Campanularia laevis Hartlaub (1905:565-567, pi. 1, based on Chile material) is a junior homonym of Campanularia laevis Couch, 1844 (see p. 65). I propose the name Campanularia agas nom. nov. for the Hartlaub species. C. agas was recently redescribed by Vervoort (1972 : 85-87, as Campanularia laevis). Both Hartlaub and Vervoort discussed similarities between C. agas (= C. laevis Hartlaub) and C. hincksii. Hickson & Gravely (1907) referred additional material to 'C. laevis Hartlaub', but Totton (1930) considered their material distinct. He referred it to a third nominal species, Campanularia hicksoni Totton, 1930. This was a species proposed to accommodate the material described by Hickson & Gravely, and also some collected by the Terra Nova'. It was discussed briefly by Rees & Thursfield (1965:90, as Campanularia laevis sensu Hickson & Gravely) and in detail by Stepanyants (1979 : 29). Campanularia aha Stechow, 19190, was based partly on new material from Villefranche and partly on some accounts of earlier authors. Stechow's material had young male gonothecae characteristic of C. hincksii, but he illustrated a hydrotheca more typical of Clytia hemisphaerica. However, the earlier descriptions included (i.e. those of Billard, 1907; Goette, 1907; Broch, 19126) seem undisputedly of C. hincksii. Hence I concur with Broch (1933) and Vervoort (19460) in regarding C. aha conspecific, and not with Picard (19510, 1955) who maintained it distinct. Campanularia macrotheca Leloup, 19300, based on material from Monaco, was justifiably referred to the present species by Patriti (1970). N.E. ATLANTIC CAMPANULARIID HYDROZOANS 55 Figs 2-4 Fig. 2 Campanularia crenata. (a) hydrotheca and (b) 9 gonotheca, Greenland, 1896.8.15.2. Scale 500 //m. Fig. 3 Campanularia hincksii. (a) hydrothecal pedicel and part of stolon. The pedicel is unusually short for the species. Note the irregular thickening. W Norway, 30-40 m. (b) 9 and (c) d gonothecae, sexes identified from contents. W Scotland, 90 m, July 1 966; 1967.12.1.16 and 10 respectively. The 9 gonotheca is unusually long. Scale (a-c) 500 //m. Fig. 4 Campanularia volubilis. (a) hydrotheca and pedicel, 30^0 m, nr Bergen, 1 5 August 1962; 1962. 1 1 .7.6. Scale 500 um. (b) vertical optical section through (a), showing flexible region. Scale 50 urn. (c-d) two gonothecae, one with ova, from a single colony, Shetland; 1912.12.21.55. Scale as (a). Campanularia volubilis (Linnaeus, 1758) (Fig. 4) Corallina minima scandens, vesiculas campaniformes in summo caule lineari contorto gerens. Ellis, 1755: 24-25, pi. 14, figs A, a. Sertularia volubilis Linnaeus, 1758:811; Linnaeus, 1767:1311; (non Pallas, 1766 : 122, junior homonym, = Calycella syringa (Linnaeus, 1767), see also Cornelius, 1978; non Ellis & Solander, 1 786 : 5 1 , pi. 4, figs E, e, F, f, = Clytia hemisphaerica, see p. 70). Sertularia uniflora Pallas, 1766 : 121-122 (nom. nov. pro S. volubilis Linnaeus, 1758; see pp. 77-78); (non Ellis, 1768 : 434, pi. 19, fig. 9, = Clytia hemisphaerica, see p. 78). Campanularia volubilis: Alder, 1857: 125-126, pi. 4, fig. 7; Hincks, 1868: 160-162, pi. 24, fig. 2 (non Hincks, 1 852, nee Du Plessis, 1871, = Clytia hemisphaerica, see p. 70). Campanularia groenlandica Levinsen, 1893: 168, pi. 5, figs 10-12; Naumov, 1960:252-253, fig. 139; Naumov, 1969 : 273-274, fig. 139; see Remarks, non Clytia volubilis: Hargitt, 1909 : 373-374 ( = C. hemisphaerica, see p. 78). Clytia mollis Stechow, 1919a : 44^45, fig. L (?syn. Clytia iaevis Weismann, 1883). Campanularia brachycaulis Stechow, 1919a : 62-63, fig. T. NOMENCLATURE. Further synonymies were given by Bedot (1901-1925), Vervoort (\946a) and Naumov ( 1 960, 1 969) among others. TYPE MATERIAL AND LOCALITY. Linnaeus (1758) gave only Ellis' (1755) illustration as indication. As with some other hydroids (Cornelius, 1979:309, notes 11-14) Linnaeus apparently based the designation on Ellis' plate and not on specimens. Almost certainly the material now in the Linnaeus collection in the Linnean Society of London (Savage, 1945 : 206) reached Linnaeus after the original description was published and cannot be 56 P. F. S. CORNELIUS regarded as type (Cornelius, 19750 : 273, footnote). The material collected and described by Ellis [infertile colony on Hydrallmaniafalcata (Linnaeus, 1758); Brighton, Sussex, England, June, 1754; illustrated, Ellis, 1755 : pi. 14, figs A, a] can thus be considered type. Although some hydroid material of John Ellis survived until recently it seems that only a single speci- men (of Nemertesia sp.) escaped destruction during World War II (Cornelius, \915a : 267, footnote) and the specimen illustrated by Ellis can be assumed lost. The type locality is Brighton. TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia groenlandica Levinsen, 1893, infertile syntype material on two pieces of Lafoea dumosa (Fleming, 1 820), in spirit, exch. Copenhagen Mus., Davis Strait, '80 frns'; 1 896.8. 1 5.1. Campanularia brachycaulis Stechow, 1919a, infertile fragments on 2 microslides, Villefranche; Munich Zoological Mus. Clytia mollis Stechow, 19 19a, small fertile colony on weed, Sete, S France; MZM. OTHER MATERIAL EXAMINED. BMNH collection, c. 60 specimens. DESCRIPTION. Colony comprising creeping stolon bearing irregularly spaced, erect, straight pedicels each supporting a hydrotheca. Stolon smooth to irregularly spirally grooved. Pedicels apparently always unbranched, smooth to spirally grooved throughout; sub- hydrothecal spherule present. Hydrotheca tubular, tapering abruptly basally; rim with 10-12 shallow blunt cusps; sometimes with fine longitudinal striae associated with the cusps. Gonothecae scarce, 20 in O. Integra: Ralph, 1957; but see p. 40). Other reported differences seem invalid (but see Dispersive stage). Fig. 5 Orthopyxis crenata. Hydrotheca, Port Phillip, Australia, intertidal; 1959. 10. 1.1. Scale lO^m. Variation. Ralph (1957) and Millard (1975) indicated that the variation in O. crenata parallels that in O. Integra (p. 63). DISPERSIVE STAGE. A medusa. By homology with O. Integra it might be expected that the medusa is short lived and does not feed. Hirohito (1969) described newly released medusae. The umbrella was sub-spherical (0'5 mm high, 0'6 mm wide). There was a distinct velum, 4 broad radial canals and 8 statocysts; but no tentacles or stomach. Published descriptions suggest that the medusa of O. Integra differs in being proportionately taller. REPRODUCTIVE SEASON. Fertile material recorded early March near Marseille (Stechow, \9\9a). DISTRIBUTION. From N coast of Spain (Garcia Corrales et al., 1978, as Campanularia everta) and Mediterranean Sea southwards (S France, Stechow, 1919a as C. intermedia; Picard, 195 la, 19586; Millard, 1975; Algeria, Picard, 1955; S Spain, Garcia Corrales et al.}. Widely 60 P. F. S. CORNELIUS distributed in warmer parts of all oceans (Millard). Cape Verde Islands (Rees & Thursfield, 1965). HABITAT. On Posidonia (eel grass) and Bryozoa (Millard & Bouillon, 1973, Seychelles); intertidal to about 3 m (Millard, 1975, southern Africa); 1-20 m, Spain (Garcia Corrales et al., 1978). Campanularia africana sens. Buchanan (1957), possibly conspecific, came from 14 m ofTGhana (see Remarks). REMARKS. Authors who have placed this species in the genus Campanularia have apparently overlooked the senior homonym Campanularia crenata Allman, 18766 (see p. 52). If the present species is again referred to Campanularia another specific name would be required, and one of the names discussed by Hartlaub ( 1 90 1 ) might be available. Hartlaub thought O. crenata (Hartlaub) close to Eucopella Campanularia von Lendenfeld, 1883a, 6, and 'identical with' Campanularia bilabiata Coughtrey, 1875. Ralph (1957), however, treated 'Orthopyxis crenata' and 'Silicularia bilabiata' under different genera. In this Ralph was unwise since crenata Hartlaub was, in part, a nom. nov. for bilabiata. The name bilabiata might prove to be available for the present species but to avoid further confusion crenata is retained pending a review of the whole genus. See also the notes on Eucopella (p. 58). Picard (19586) recorded the nominal species Orthopyxis everta (Clarke, 1876, as Campanularia, based on Californian material) from 'the Mediterranean'. The original description resembles the present species, and the two might prove conspecific; but I have seen type material of neither. Ralph (1957) separated them on the structure of the gonotheca. If a synonymy were propsed everta might take priority for the present species but C. bilabiata Coughtrey is still older. Vervoort (1972 : 87) redescribed 'O. everta' recently and gave further synonymy. The nominal species Campanularia ?intermedia Stechow, 1919a, was based on material from Marseille. I have not located type material but the vegetative characters given in the description seem identical with those of the present species as currently understood. As suggested by Stechow, and also by Garcia Corrales et al. (1978), C. lennoxensis Jaderholm, 19046, is probably conspecific. Campanularia africana Stechow (1923d: 104, nom. nov. pro C. tincta sensu Warren, 1908, from Natal; non C. tincta Hincks, 186 la, from 'Australia', see p. 58) was recorded from Takoradi, Ghana at 14m depth by Buchanan (1957). O. africana has been distinguished from O. crenata by Millard (1975), who redescribed both, mainly on gonothecal characters; and from the several reportedly endemic South African species she recognized on variations in these characters alone. However, the relatively poor original descriptions of most of the nominal species and the general taxonomic confusion in the genus make it unwise to accept Buchanan's record without further evidence. It is the only record of 0. africana from north of the equator. Discussion of the non- Atlantic nominal species of Orthopyxis having crenate hydrothecal margins was provided by Ralph (1957). Millard (1975) provisionally referred Orthopyxis delicata Trebilcock, 1928, to the present species; and it seems likely that O. delicata sensu Garcia Corrales et al. (1978; N & S Spain) is similar. Orthopyxis Integra (Macgillivray, 1842) (Fig. 6) IClytia undulata Lamouroux, in Freycinet, 1 824 : 6 1 7-6 1 8, pi. 94, figs 4-5. Campanularia Integra Macgillivray, 1842 : 465; Johnston, 1847 : 109, pi. 28, fig. 2 (syn. C. laevis: Saunders, in Johnston, 1847); Hincks, 1868 : 163-164, pi. 31, fig. 1; Levinsen, 1893 : 168-169, pi. 5, figs 14-18 (syn. C. caliculata Hincks; C. gracilis Allman, 18766); Broch, 1918 : 159-162 (syn. C. compressa Clarke; C. ritteri Nutting, 1901a); Vervoort, 1946^:274-276, figs 120-121 (syn. C. laevis Couch; C. caliculata Hincks; C. breviscyphia Sars; Clytia (Orthopyxis) poterium Agassiz; Laomedea repens Allman); Millard, 1975:208-211, fig. 69 (syn. C. caliculata Hincks; C. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 61 compressa Clarke; Agastra mira Hartlaub; Agastra rubra Behner; ?Campanularia gracilis: Stechow, 1925). Clytia ryckholtii Westendorp, 1 843 : 23-24, pi. 1 , figs e, f. Campanularia laevis Couch, 1844:42; Gosse, 1855:25; (non C. laevis Hartlaub, 1905=junior homonym). Capsularia Integra: Gray, 1848 : 86 (?syn. Campanularia laevis Couch). Capsularia laevis: Gray, 1848 : 87. Campanularia caliculata Hincks, 1853 : 178-179, pi. 5, fig. B; Hincks, 1868 : 164-167, pi. 31, fig. 2 (syn. C. breviscyphia Sars; Clytia (Orthopyxis)poterium Agassiz. Campanularia breviscyphia Sars, 1857 : 158-159, pi. 1, figs 12-13. Clytia (Orthopyxis) poterium Agassiz, 1862 : 297-304. Clythia poterium Agassiz, 1862 : pi. 28, figs 1-20, pi. 29, figs 1-5. Orthopyxis poterium Agassiz, 1862 : 355. Clytia posterior Wright, 1862 : 308 (lapsus pro poterium Agassiz). Laomedea repens Allman, 1871: 49, fig. 20. ?Eucopella Campanularia von Lendenfeld, 1883a : 186-189. Campanularia compressa Clarke, 1877:214, pi. 8, figs 5-6; Patriti, 1970:34-35, fig. 43 (syn. C. platycarpa Bale). Campanularia borealis Marktanner-Turneretscher, 1890 : 206. Campanularia integriformis Marktanner-Turneretscher, 1890 : 207, pi. 3, fig. 2. 'A leptomedusan' Browne, 1897 : 832, pi. 49, figs 3, 3a. Agastra mira Hartlaub, 1897:452, 504-506, pi. 22, figs 5, 8-10; Mayer, 1910:234 (syn. Campanularia caliculata Hincks); Russell, 1953:303-306, pi. 19, fig. 1, text-figs 186-188 (syn. ' Leptomedusa sp.' Browne; Campanularia caliculata Hincks). Agastra caliculata: Browne, 1900 : 714-715 (syn. A. mira Hartlaub; Leptomedusa Browne). Leptomedusa gen.? sp.? Browne, 1900 : 714. Campanularia calyculata: Goette, 1907:193-204, pi. 15, figs 313-325 (syn. Clytia poterium Agassiz). , ?Agastra rubra Behner, 1914 : 393-398, pi. 7, fig. 6, text-figs 8-10. Orthopyxis compressa: Stechow, 1919a : 69, fig. Wa-b; Picard, 19516 : 1 10; Picard, 1958a : 2. Orthopyxis asymmetrica Stechow, 1 9 1 9a : 71-72, fig. Xa-e. Clytia rijckholtii Leloup, 1947 : 22 (unjust, emend, pro C. ryckholtii Westendorp). Orthopyxis caliculata: Ralph, 1957:838, text-figs 6a-f (syn. O. macrogona von Lendenfeld); Picard, 1958& : 191 (syn. Campanularia integriformis auct.; see Remarks). Orthopyxis integral Rees & Thursfield, 1965 : 103-104. Eucopella caliculata: Hirohito, 1969 : 6-7, fig. 6. NOMENCLATURE. An unjustified emendation of the nominal species name caliculata, to calyculata, was followed by several authors (listed in Bedot, 1918, 1925). Browne (1 897) described the medusa of the present species but did not identify it, calling it simply 'A Leptomedusa gen.? sp.?'. Evidently Browne did not then regard Leptomedusa a generic name; but he later (Browne, 1900) used it thus: 'Leptomedusa gen.? sp.?', in a formal synonymy under 'Agastra caliculata (Hincks, 1853)'. Nevertheless it seems in keeping with Browne's intentions not to regard Leptomedusa as part of nomenclature. The widely used species name Integra may prove to be threatened by an older but obscure name, undulata. TYPE LOCALITY AND MATERIAL. Mouth of River Don, Aberdeen, Scotland; on Tubularia indivisa Linnaeus, 1758; material not located. TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia laevis Couch, r844, neotype, proposed herein. Campanularia caliculata Hincks, 1853, colony on Laminaria sp. and the red alga Phycodrys rubens (L.) Batt. (det. J. H. Price), in spirit, nr Old Head of Kinsale, Co Cork, Eire, coll. R. Allman, syntype; 1853.4.7.16. Remainder of type series, from Pegwell Bay, E. Kent, England, coll. R. S. Boswell, not located. The type locality of C. caliculata was restricted to Pegwell Bay by Ralph (1957) but the Co Cork material remains part of the syntype series. Orthopyxis asymmetrica Stechow, 1919a, infertile material on 2 microslides, Marseille; Munich Zoological Museum. 62 P. F. S. CORNELIUS OTHER MATERIAL EXAMINED. BMNH collection, c. 50 specimens, mostly from the British Isles. DESCRIPTION OF HYDROID STAGE. Colony a creeping hydrorhiza with single, irregularly spaced hydranths and hydrothecae on long pedicels and, separately, subsessile gonophores and gonothecae. Hydrorhiza smooth but sinuous, walls almost unthickened but often (Ralph, 1957; Millard, 1975) with a flat lateral flange of perisarc; branched frequently and (e.g. BMNH 1922.3.6.170, 1962.11.7.9) occasionally anastomosing. Hydrothecal pedicels usually narrower than hydrorhiza; walls usually much thickened; typically grooved with a smooth spiral; often 2-3 widely spaced shallow annuli near top; sub-hydrothecal spherule present; hydrotheca cup-shaped, length : breadth ratio variable; base wide to narrow, walls almost straight and diverging in narrower-based specimens; often flared near rim; walls thin to very thick, but rim region almost always unthickened; amount of thickening varying between adjacent hydrothecae and within a single hydrotheca (Fig. 6); rim even; small Fig. 6 Orthopyxis Integra, (a-e) 15-25 m, Espegrend, W Norway, 13 April 1962; 1962.10.7.1 1. (a-b) adjacent hydrothecae with differing pedicel lengths, (c) gonotheca, sex unknown, (d-e) sub-hydrothecal spherule, (f-h) Knysna, Cape Province, Republic of South Africa, 1922.3.6.170. Scales: (a-c, 0 500 //m; (d-e) 10 //m; (g) 10 //m; (h) 10 //m. spherical chamber formed basally within hydrotheca by internal ring of perisarc. Hydranth with 20-30 tentacles (histological details in Agassiz, 1862; Stefani, 1956, 1959; Kawaguti, 1966; anatomical details of a possibly conspecific form in von Lendenfeld, 18836). Gonotheca ^ = 9; broad, roughly parallel sided; truncated and slightly narrowing above, tapering more or less abruptly below; length usually l^-2x breadth, occasionally 5-6x (e.g. Allman, 1871 : fig. 20, as Laomedea repens; Vervoort, 1946a : fig. 120); usually laterally flattened but sometimes circular in transverse section; walls of gonotheca often thickened, sometimes much so; smooth through sinuous to deeply grooved spirally; aperture distal, N.E. ATLANTIC CAMPANULARIID HYDROZOANS 63 nearly as wide as maximum diameter of gonotheca. Pedicel short to absent, usually unringed and grading into base of gonotheca (but see Fig. 6). Blastostyle with one well developed medusa, whether retained or released, and a second basal bud the fate of which seems unrecorded (see Dispersive stage). Present evidence suggets that medusae of either sex are sometimes retained. Nematocysts described by Ostman (1979). Variation in hydroid stage. See also the comments of Ralph (1957 : 838) and Millard (1975:209). Even among the Campanulariidae O. Integra is unusually variable in morphology, and is unusual also in its habit of sometimes releasing and at other times retaining the medusa. Some of the variation may be genotypic, but the controlling factors are hardly known. Naumov (1969) referred colonies with thick walled hydrothecae to a variety, caliculata Hincks, 1853, which he considered grew only in strong currents; but although this relation seems logical he offered no proof. In his introductory sections (p. 123) he reported that hydrothecae of this species grow larger in cool water than in warm. DISPERSIVE STAGE. Basically a short lived medusa. But this is often retained (as a gonomedusa), when the planula is the only motile stage. The free medusa was perhaps first described by Hartlaub (1897) from Helgoland, and shortly after by Browne (1897) working independently in SW Ireland. But von Lendenfeld (1883a, b) had earlier described a closely similar nominal species which may prove identical, from Australia ('Eucopella campanu- laria"; see Remarks); and Agassiz (1862, as Clytia poterium) had still earlier described planula release from retained medusae. The medusa, when released, is degenerate and ephemeral. It lacks organs of feeding and survives only a few days. Umbrella height c. 1 mm, width c. 0*65 mm; jelly thick, velum broad; stomach, manubrium and mouth absent; four narrow radial canals each with lobed gonad midway along; no tentacles or marginal cirri (after Russell, 1953). Apparently only one medusa at a time is produced from each blastostyle. The medusae swim actively (Hartlaub, 1897) but are probably short lived since they are presumed not to feed. They are sexually mature on release and do not develop further (Russell, 1953). Giard (1898) has often been thought the first to have linked the medusa to its hydroid but von Lendenfeld's (18836) earlier work might have been on this species (see Remarks). Giard was certainly the first to record that the medusa is not always released. He has been misquoted but his paper was quite explicit. Giard thought that time of year influenced medusa release, and so did Behner (1914) who worked on the probably conspecific Mediterranean medusa Agastra rubra Behner, 1914. But Stefani (1959) recorded liberation in turbulent water and retention under calmer conditions. Millard (1975; pers. comm.), however, stated that medusa release had not yet been recorded in southern African popu- lations (see also Remarks). The factors influencing release are still unclear. The female gonophore was recorded by several of the earlier workers but the male was not described until the work of Stefani (1956) and Hamond (1963), again excepting the much earlier and largely overlooked work of von Lendenfeld (18836) on the possibly identical Australian populations. Some authors (Russell, 1953, quoted in Rees & Thursfield, 1965; Hamond, 1963) have interpreted the retained medusae as sporosacs, but current knowledge of the life cycle confirms that they are medusoid. Following Miller (1973) they can be called gonomedusae. The often reported 'second medusa bud' near the base of the blastostyle [e.g. von Lendenfeld, 18836; Giard, 1898; Hamond, 1963 (c?); Hirohito, 1969 (9); Millard, 1975 (9); BMNH 1915.3.6.12 (d1); also in congeneric species, Ralph, 1957] corroborates Miller's theory of descent from a gonophore producing medusae. Evidently the ancestral form produced several medusae on each blastostyle but today only one is produced at a time. REPRODUCTIVE SEASON. Free medusae recorded May-November in British waters (Russell, 1953); June-September in NW France (Teissier, 1965); December-February & July at Naples (Lo Bianco, 1909). Some authors, from Giard (1898) onwards, have considered that medusa release occurs only towards the end of the reproductive season and that gamete release from sessile medusae occurs earlier in the year; but precise dates are unrecorded. 64 P. F.S.CORNELIUS Teissier (1965) found reproductive structures on the hydroid stage from May to October in NW France. DISTRIBUTION. Nearly cosmopolitan, occurring in all oceans from the intertidal to a little below Continental Shelf depths (at least in cold seas); and from the tropics to latitudes as high as 76° 40' N (Greenland). The species is one of the most widely distributed of all hydroids. Noteworthy records from the eastern North Atlantic include: E & W Greenland (Kramp, 1929, 1943), N Norway (Mathiesen, 1928), Greece (Yamada, 1965), Black Sea (Manea, 1972; possibly also Naumov, 1960, 1969, as Campanularia integriformis, see Remarks), Morocco (Patriti, 1970, as Campanularia compressa), Ghana (Buchanan, 1957), Senegal (Leloup, 1939), Cape Verde Is (Ritchie, 1907); and the range of the species extends at least to the southernmost tip of Africa (Millard, 1975). However, there are gaps in this wide distribution. Broch (1928) thought the species to be absent from the Kattegat, Skagerrak and Baltic, and Stechow (1927) from the Baltic alone; but Kramp (1935) and Jagerskiold (1971) recorded it from W Sweden. Apparently there are still no records from the Baltic Sea. The species evidently did not occur in the Zuider Zee (Hummelinck, 1936) and has probably never been reliably recorded from Dutch waters (Vervoort, 19460). However, it is sometimes washed ashore on the Belgian coast (Leloup, 1952). Similarly, Irish Sea and W Scottish records are few: Bardsey I, Wales (Knight-Jones & Jones, 1956), Isle of Man (Bruce, Colman & Jones, 1963), Isle of Cumbrae, Clyde Sea (Chopin, 1894; Rankin, 1901), 10m depth in Cregan Narrows, Loch Creran, Argyll (C. Edwards colln, pers. comm.). Chumley (1918) recorded no Clyde Sea material; and Stephens (1905) gave only a few Irish localities: Belfast, Dublin and Co Cork, the last including some of the syntypes of Campanularia caliculata. Possibly the only record from the west coast of Ireland is from Valencia I, Co Kerry (Browne, 1900), incidentally one of the earliest descriptions of the medusa. Hincks (1868) and Russell (1953) similarly listed no records from the western coasts of Eire and Scotland but the species is small and may have been over- looked. Broch (1918) included the NW Irish and W Scottish coasts in the North Atlantic distribution but did not cite material and may have been guessing. Lastly, Arai & Brinckmann-Voss (1980: 103) thought the species might not occur in British Columbia and Puget Sound. HABITAT. World depth data range from intertidal (e.g. Hincks, 1853, British Isles) through 300 m (Broch, 1918, Davis Strait) exceptionally to 470 m (Kramp, 1929, W Greenland). The deepest records are from cold waters. The species has been recorded on a wide variety of algae, hydroids, other animals and inorganic substrates, and there is no regular association. Mathiesen (1928) recorded O. integra on Laminaria sp. to depths of c. 100 m off Norway. Broch (1918) regarded the species as stenohaline, a view supported by the lack of records from the Baltic Sea and Zuider Zee. REMARKS. Several species closely related to O. integra were recognized by Ralph (1957), Millard (1975) and Gow & Millard (1975), and a world revision of the genus would be timely. Much discussion was provided by Arai & Brinckmann-Voss (1980). O. integra itself is both variable and nearly cosmopolitan, and has consequently been described under many species names (cf. Clytia hemisphaerica, p. 73). The above synonymy includes only North Atlantic synonyms and main ones from other areas when they enter the discussion. The following notes on them are arranged in date order where possible. Baster (1762 : pi. 2, fig, 7A, a) published some unidentified illustrations which Maitland (1876) referred to the present species. Vervoort ( 1 946a : 276) regarded them as indeterminate, however, and I concur. They are discussed further under Clytia hemisphaerica (p. 78). Although Pallas (1766) arguably applied an earlier species name partly to Baster's illustra- tions, the later name integra is not threatened (but see the paragraph after next). Clytia urnigera Lamouroux (1816: 203, pi. 5, fig. 6), based on 'Australasian' material, was N.E. ATLANTIC CAMPANULARIID HYDROZOANS 65 discussed by Couch (1844) when proposing Campanularia laevis Couch, a nominal species considered below. Couch noted a resemblance between the hydrothecae of urnigera and those of laevis but rightly pointed out that the narrow gonothecal aperture of urnigera contrasted with the wide aperture in laevis, and held them distinct. C. urnigera is here referred to Clytia hemisphaerica (p. 78); and C. laevis to O. Integra (see below). Clytia undulata Lamouroux, in Freycinet, 1824, was based on a fertile colony growing on 'marine plants' at Port Jackson, New South Wales. The species was regarded by Lamouroux as close to 'Clytia urnigera Lamouroux', here referred to C. hemisphaerica. It was mentioned again only twice in the literature according to Bedot (1905), in 1824 and 1836, but no further taxonomic features were mentioned. C. undulata seems to be closer to the present species than to 'C. urnigera' and C. hemisphaerica. It is mentioned here as it predates other Orthopyxis species and might prove conspecific with O. Integra; but before it can be fully assessed more information is needed about the Australian populations of Orthopyxis. Clytia ryckholtii Westendorp, 1843, was based on material from Ostend, Belgium. It was referred to O. Integra by Billard (1914); and also by Leloup (1947) as 'C. Rijckholtii Slab.\ Both Leloup's spelling and his reference to Slabber are wrong: Slabber (1769-1778) did not treat O. Integra or anything similar. I have not located the type material; but Westendorp's illustrations show a reptant colony with long hydrothecal pedicels spirally grooved top and bottom each with an intervening smooth portion, and an even rimmed hydrotheca. They are the earliest illustrations of O. integra yet identified — but those of Clytia undulata Lamouroux, in Freycinet, 1824, which might prove conspecific, are earlier. The description of 'ryckholtii' states the pedicels to be 3 mm long and mentions an even rimmed hydrotheca, confirming the identification. A rare and later work by Westendorp (1853) on Belgian zoophytes was illustrated by actual specimens. Had C. ryckholtii been included the speci- mens might have been types; but it was not (see note 1, p. 123), and I agree with Billard (1914) that the type material is probably lost. Campanularia intertexta Couch, 1844, was based on a mixed type series comprising Lafoea dumosa (Fleming, 1 820) and an unidentified campanulariid which might have been O. integra. C. intertexta is now referred to L. dumosa (see p. 122) but the original illustration, which included the campanulariid, was one of the earliest of O. integra. Campanularia laevis Couch, 1844, type locality Polperro, Cornwall, was implicitly referred to the present species by Johnston (1847). The type material may be have been preserved for a long time in the Royal Institution of Cornwall, Truro, but if present would have been destroyed by a flood in 1953 (Curator, Roy. Inst. Cornwall, pers. comm.). Johnston (1847) mentioned material sent to him by W. W. Saunders (BMNH reg. no. 1847.9.24.65, on a herbarium sheet). The material is labelled in Johnston's hand 'Campanularia laevis! W. Wilson Saunders, Hastings, 1840' and a label has been glued on later, also in Johnston's hand, on which is written 'Campanularia integra'. I concur with Johnston's later identification as O. integra. In the absence of the original type material I designate the specimen 1847.9.24.65 neotype of C. laevis Couch, 1844; and extend the type locality to comprise coastal waters of the south of England. I agree also with Johnston's suggestion that the original description of C. laevis Couch should be referred to O. integra; and with the tentative but similar opinion of Bedot (1905 : 1 57) that the two are conspecific. Gray (1848 : 86) too referred both the species C. laevis and the material just mentioned to O. integra; but, inconsistently, on the next page gave C. laevis Couch full specific rank. He did this under the genus name Capsularia Cuvier, 1797, now regarded a junior objective synonym of Coryne Gaertner, in Pallas, 1774 (see Cornelius, \915b : 378). Turning to a later work, it seems that Hincks' (1868 : 164) Hastings record of Saunders' material refers to the same specimen. Since Johnston (1847) did not publish the locality it seem likely Hincks deduced if from Gray (1848), who did. The junior homonym Campanularia laevis, Hartlaub, 1905, is discussed under Campanularia hincksii, which that nominal species most resembles (p. 54). Campanularia caliculata Hincks, 1853, is the main European synonym to have been applied to the hydroid stage. When proposing the nominal taxon Hincks in fact provided the 66 P. F. S. CORNELIUS first good description of the present species. Bedot (1918) recorded that Levinsen (1893) was the first to refer caliculata to Integra. Several senior authors have accepted this synonymy (e.g. Broch, 1918; Kramp, 1935; Vervoort, 1946a; Millard, 1975) and examination of type and non-type BMNH material supports their view. Hincks (1853) stated clearly the differences from Integra as he saw them. They were simply: a 'double' hydrotheca and pedicel (inaccurate observation of thick walled material) and a more gradual tapering of the hydrotheca in caliculata than in Integra (a variable feature). Neither character is reliable (see the above description and the discussion by Millard, 1975 : 209-210). Russell (1953) drew attention to the gonothecal contents later illustrated by Hincks (1868 : pi. 31, figs, la, b) as Integra. They seemed to be sporosacs, and Russell was not entirely ready to accept the synonymy. It then seemed plausible that there were two species, one with sporosacs and the other with facultatively released medusae. The question was resolved when Millard (1975) illustrated structures similar to those shown by Hincks and described release of gametes from them. Millard had not recorded medusa release in her area (southern Africa). Still, she commented 'In partly spent gonophores the medusoid structure can sometimes be seen and is best observed by dissecting the gonophore out of the gonotheca' (op. cit., p. 209). Evidently the sessile eumedusoids (called gonomedusae by Miller, 1973) resemble sporosacs closely and their medusoid nature is not easy to see. Hence Russell's point is answered, and synonymy seems in order. Campanularia breviscyphia Sars, 1857, was referred to C. caliculata Hincks, 1853, by Hincks (1868), and to the present species by Vervoort ( 1 946a). Clytia (Orthopyxis) poterium Agassiz, 1862, was a nominal species apparently based on North American material. Agassiz' description was exceptionally detailed and beautifully illustrated, but he failed to consider the several related species already described from European waters (integra, ryckholtii, laevis, caliculata, breviscyphia). His account suggests strongly that his material was merely O. integra. Hincks (1868) and Bale (1914) summarily dismissed poterium as conspecific; but Agassiz' account remained the most detailed of the species for many years. It was the earliest of the medusa, albeit of retained specimens. The caption to the plates included the spelling ^Clythia\ an unjustified emendation of Clytia. Laomedea repens Allman, 1871, was referred to the present species by Bedot (1910) and Vervoort (1946a), and I concur. The originally illustrated material was said to have come from Scotland (Allman, op. cit., p. 48) and comprised only a female gonophore of O. integra. The nominal species was apparently never described again. Allman later applied the same specific name to another taxon, Hypanthea repens Allman, 1876a, type locality Kerguelen Island (see also p. 50). Campanularia gracilis Allman (18766 : 260, pi. 12, figs 5-6), from Japan, was referred to the present species by Levinsen (1893) but this was quite unjustified. The type material (BMHN reg. no. 1877.4.12.5) is clearly distinct and not closely related to O. integra. Stechow (1925 : 423, fig. 6) described new material of C. gracilis Allman but Millard (1975 : 208) provisionally referred it to O. integra. However, Stechow's illustration closely resembles the type material and Millard, like Levinsen, was apparently mistaken in uniting the two taxa. Yamada (1959 : 35) evidently regarded C. gracilis as valid but recorded no material apart from the type. Campanularia compressa Clarke, 1877, based on the hydroid stage, was referred to the present species by Broch (1910, 1918) and Millard (1975), although Arai & Brinckmann- Voss (1980) regarded it distinct. Other material of O. integra was recorded under the species name compressa by Stechow (1919a, Villefranche), Picard (19516, Senegal), Picard (1958#, Israel) and Patriti (1970, Morocco). See also the comments on Agastra rubra Behner, 1914, the corresponding nominal species based on the medusa, below. Eucopella Campanularia von Lendenfeld, 1 883#, is discussed above (p. 60). Campanularia borealis Marktanner-Turneretscher, 1890, was based on hydroid material from Spitzbergen. No illustration was provided. Linko (191 1 : 164) referred the species to O. integra and I concur. Campanularia integriformis Marktanner-Turneretscher, 1890, based on hydroid material N.E. ATLANTIC CAMPANULARIID HYDROZOANS 67 from Trieste, seems the same as O. Integra. Naumov (1960, 1969) briefly described speci- mens from the Black and Adriatic Seas as C. integriformis but this too was probably O. Integra. If so, Naumov's report of 0. Integra in the Black Sea predates that of Manea (1972), who had claimed the first record. Picard (1958&) referred integriformis to caliculata Hincks, 1853, without comment but caliculata, too, is now referred to Integra. Agastra mira Hartlaub, 1897, based on Helgoland material, has been widely regarded as the first description of the free medusa of O. Integra. Browne (1897) described the medusa from SW Eire in the same year, as an unidentified leptomedusan; but a footnote in Browne's paper comments on Hartlaub's account, which had presumably already appeared. Both accounts of the medusa might be predated, however, if the closely similar Australian nominal species Eucopella campanularia von Lendenfeld, 1883#, b, proves conspecific. Several authors listed by Bedot (1918, 1925) applied the combination Agastra caliculata to the medusa stage. Agastra rubra Behner, 1914, a medusa from the Mediterranean, was referred to O. integra by Stefani (1959) and Millard (1975) among others. Its nominal hydroid, Campanularia compressa Clarke, 1877 (see above), was referred to O. integra by Broch (1910, 1918); and both stages were so treated by Millard (1975). Kramp (1961), however, considering the medusa stage alone, held A. rubra distinct on the shape of the gonads. Further work seems necessary to resolve these different views, but the majority opinion is that A. rubra is invalid. Finally, the type material of Orthopyxis asymmetrica Stechow, \9\9a, from Marseille, was examimed and found merely to be O. integra. Philbert (1935a) described growth forms under this name. Genus RHIZOCAULUS Stechow, \9\9b Sertularia Linnaeus, 1758 : 81 1 (part). * Campanularia Lamarck, 1816 : 112 (part); Hincks, 1868 : 160 (part). Rhizocaulus Stechow, 19 196 : 852; Stechow, 1919c: 16. Verticillina Naumov, 1960 : 9, 1 15, 122, 269; Naumov, 1969 : 6, 1 15, 123, 291. TYPE SPECIES. Sertularia verticillata Linnaeus, 1 758; by original designation. DIAGNOSIS. Campanulariidae forming erect, polysiphonic colonies; hydrothecae in whorls; no true hydrothecal diaphragm; sub-hydrothecal spherule present; no medusa stage. REMARKS. Nutting (1915 : 28) designated Sertularia verticillata Linnaeus, 1758, type species of the genus Campanularia Lamarck, 1816, but for the reasons given under that genus (p. 5 1) I have applied to the International Commission on Zoological Nomenclature for that designation to be set aside (Cornelius, 1981). Rees & Thursfield (1965) opposed the erection of a distinct genus to accommodate verticillata on the grounds that in Lafoea Lamouroux, 1821, family Lafoeidae, there are both stolonal and erect colonies; but colony habit does not have the same taxonomic value in all families, and the separation is upheld here. Stechow (1919/7) listed several nominal species in the genus but they may prove conspecific with R. verticillatus. Rhizocaulus verticillatus (Linnaeus, 1758) (Fig. 7) Corallina ramosa, ramis singulis equisitiformibus in summis capillamentis contortis et verticillatim dispositis . . . Ellis, 1 755 : 23-24, pi. 1 3, figs A, a. Sertularia verticillata Linnaeus, 1758 : 81 1. Campanularia verticillata: Lamarck, 1816: 113; Hincks, 1868: 167-168, pi. 32, fig. 1, la; Goette, 1907: 179-189, pi. 14, figs 294-304, pi. 15, figs 305-306; Linko, 1911 : 188-200, fig. 35 (syn. Clytia olivacea Lamouroux, 1 82 1 ). Clytia olivacea Lamouroux, 1821 : 13, pi. 67, figs 1-2. Campanularia verticellata Couch, 1842 : 49 (lapsus). 68 P. F. S.CORNELIUS Campanulata verticillata: Agassiz, 1862 : 354, footnote (unjustified emendation of Campanularia). Rhizocaulm verticillatus: Stechow, 19196 : 852: Stechow, 1919c : 16; Stechow, 1923a : 105-106. Verticillina verticillata: Naumov, 1960 : 269-270, fig. 159; Naumov, 1969 : fig. 159. TYPE MATERIAL AND LOCALITY. Linnaeus (1758) provided only a diagnosis of this species, implying that he did not see material (cf. Cornelius, 1979 : 309). Indeed, there is none in the Linnaeus collection held by the Linnean Society of London (Savage, 1945). Linnaeus (1758) cited only the illustration of Ellis (1755 : pi. 13, fig. A, but not fig. a), and the colony Ellis illustrated can be regarded as holotype. It was collected from the coast of Cumberland, NW England, to which the type locality can be restricted. The specimen is almost certainly now lost (Cornelius, 1975a : 267, footnote). MATERIAL EXAMINED. BMNH collections, about 40 specimens. Some northerly material deserves mention: Barents Sea, 74° 8' 50" N, 30° 31' 28" E, 375m, 1882, ex D'Arcy Thompson collection; 1956.1 0.23 . 1 80. DESCRIPTION. Colony large, erect, occasionally and irregularly branched; stems polysiphonic with pedicels roughly in whorls, recalling the terrestrial horse-tail plants (Equisetum L.). Component perisarc tubes straight, parallel, each bearing straight, smooth or spirally grooved hydrothecal pedicels (sometimes annulated, e.g. Fig. 7) at approximately regular intervals. Hydrotheca bell shaped, sub- hydrothecal spherule present; rim with c. 12 blunt cusps. Gonotheca ?rf = 9, flask shaped, with neck of varied length; on short pedicel. Fig. 7 Rhizocaulus verticillatus. (a) part of colony, Isle of Man, 25 March 1894; 1948.10.1.21. (b) gonotheca with long neck, 30-40 m depth, nr Bergen, 15 August 1962; 1962.11.7.8. (c) gonotheca with short neck, 45 m depth, W Scotland; 1 888.4.2.39. Scale (a-c) 500 //m. DISPERSIVE STAGE. Planulae, probably developing within the gonotheca. Early gonophore development was described by Goette (1907). No medusa. REPRODUCTIVE SEASON. Fertile material recorded May, 1934, off NE England (H. O. Bull, in Evans, 1978); July, NW France (Teissier, 1965); 15 August, 1962, nr Bergen, Norway (BMNHreg.no. 1962.11.7.8). N.E. ATLANTIC CAMPANULARIID HYDROZOANS 69 DISTRIBUTION. Widespread in sublittoral and coastal waters from Tromso, Norway (Mathiesen, 1928) and Danmarks Havn, Greenland (Kramp, 1943) at least to Roscoff, NW France (Teissier, 1965). The species is well known from offshore areas of Britain, the Netherlands, Belgium, Denmark and Sweden. See also Material examined. The status of JR. verticillatus along the W coast of France and the Bay of Biscay is unclear. Vervoort (\946a) and Naumov (1969) recorded it but the species was not listed in several local faunal surveys (Billard, 1927; Nobre, 1931; Da Cunha, 1944, 1950; Fey, 1969) and there is no BMNH material from south of the English Channel. Picard (19586) did not record the species from the Mediterranean Sea; but Manea (1972) provided an acceptable record from the Black Sea, possibly the farthest south the species has been found on European coasts. HABITAT. Naumov (1969) gave a usual depth range of 50-200 m, with extremes of 15 m and 680 m. Mathiesen (1928) gave a range of 20 m to 600 m in S Norway. Hamond (1957) associated the species with 'sandy grounds' off the Norfolk coast, but Teissier (1965) recorded it on algae in NW France. Hincks (1868) and Leloup (1952) found the species on pebbles and shells. R. verticillatus seems to occur typically on occasional solid substrates in otherwise sandy areas. REMARKS. There seems no need for revision of this distinctive species. Rees & Thursfield (1965) drew attention to the superficial similarities between it and Campanularia volubilis (but see p. 57). The development and structure of the compound stem of R. verticillatus was studied by Schach(1935). The combination Campanularia verticillata var. grandis Hickson & Gravely, 1907, related to an antarctic species now known as Stegella grandis (Hickson and Gravely) and assigned to the Campanulinidae (e.g. Totton, 1930; Stepanyants, 1979). The similarity in colony form is striking but S. grandis lacks a sub-hydrothecal spherule and the hydrothecal rim is quite different. Subfamily CLYTIINAE nom. nov. Phialidae Haeckel, 1879 : 163 (part). Phialinae: Mayer, 1910 : 232 (part). Campanularinae: Russell, 1953 : 284. DIAGNOSIS. Campanulariidae with a usually reptant, branched but not anastomosing stolon; with true hydrothecal diaphragm; usually without subhydrothecal spherule (but present in Clytia hummelincki); medusa present in some genera, a typical leptomedusa with prominent velum. SCOPE. The genera Clytia Lamouroux, 1812; Gastroblasta Keller, 1883; and Tulpa Stechow, 192 la. Gastroblasta and Tulpa have not been recorded in the eastern North Atlantic and are discussed only under Remarks, but Clytia is treated below. TYPE GENUS. Clytia Lamouroux, 1812. REMARKS. Previous subfamily names applied in part to this group (Phialidae Haeckel, 1879; Phialinae Mayer, 1910) have been derived from Phialium Haeckel, 1897, the name of a genus now referred to the Lovenellidae (Mayer, 1910; Kramp, 1961). Phialium can be taken as type genus of the nominal subfamily Phialiinae; and there seems no available name for the present group. There has no doubt been confusion between Phialium Haeckel and Phialidium Leuckart, 1856. Phialidium is today referred to Clytia Lamouroux, 1812, and basing a new subfamily name on Phialidium might be confusing. Haeckel (1879 : 186) indicated that he understood the relation between Phialidium and Clytia by including Clytia johnstoni sensu Bohm in the 70 P. F. S. CORNELIUS synonymy of P. variable. This relationship had only just been understood (p. 76) and Haeckel might have been reticent to consider all the nomenclatural implications. The subfamily name Clytiinae is to be considered a nom. nov. applied to the present sub- family as restricted by Russell (1953) under the preoccupied name Campanulariinae. It should not be confused with family group names derived from Clytus, a coleopteran genus name. These are spelt Clytinae, Clytini and so on (C. R. Smith, pers. comm.), being derived from the root Clyt. The root from which Clytiinae is derived is Clyti. The genus Gastroblasta Keller, 1883 : 622, is now restricted to a single, distinctive medusa species based on Red Sea material (Kramp, 1961). The name was once applied to species now referred to one or other of the Clytia species treated below (discussion on p. 72). The genus Tulpa Stechow, 1 92 la : 254, was proposed to embrace three species none of which has been recorded from the eastern North Atlantic. The type species is Campanularia tulipifera Allman, 1888, by original designation (= C. tulpifera lapsus auct.). The genus was accepted by Totton (1930) who described another species. Tulpa was reviewed by Ralph (1957) and redefined by Stepanyants (1979); and discussed also by Vervoort (1972) and Gravier-Bonnet (1979) who gave systematic notes. Genus CL YTIA Lamouroux, 1812 Medusa: Linnaeus, 1767 : 1096 (part). Sertularia Ellis & Solander, 1786:51 (part). Oceania Peron & Lesueur, 18100 : 343 (part). Clytia Lamouroux, 1812 : 184. Campanularia auct., part (see Remarks under Clytia hemisphaerica, p. 77). Thaumantias Eschscholtz, 1829 : 102; Forbes, 1848 : 52 (part). Silicularia Meyen, 1 834 : 206 (part; see p. 50). Eucope Gegenbaur, 1856 : 241 (part). Phialidium Leuckart, 1856 : 18; Kramp, 1961 : 164. Epenthesis McCrady, 1857 : 191; Haeckel, 1879 : 182. Clytia (Trochopyxis) Agassiz, 1862 : 304. Clytia (Platypyxis) Agassiz, 1862 : 306. Clytea Wright, 1862 : 308 (lapsus pro Clytia). Clythia van Beneden, 1 866 : 166 (lapsus pro Clytia). Gastroblasta auct., part (non Keller, 1 883, s. str.). Pseudoclytia Mayer, 1900 : 53. Multioralis Mayer, 1900 : 54. non Clythia Agassiz, 1862 : pi. 28 (lapsus pro Clytia, but referred to Orthopyxis, p. 57). TYPE SPECIES. There were three species originally included in the genus, listed thus: 'Sertularia volubilis Ellis, S. syringa Ellis, S. verticillata Ellis'. The descriptions cited are presumably those of Ellis & Solander (1 786), in which book binominals were employed, and not those of Ellis (1755) which lacked them. The point is important since 'S. volubilis sens. Lamouroux, 1812' was subsequently designated type species (see next paragraph). Confusion was caused when Lamouroux himself (in Lamouroux, Bory de Saint-Vincent & Deslongchamps, 1 824 : 202) later applied the name S. volubilis to another species, citing therein the Ellis, 1 755, illustration which in fact shows the species here called Campanularia volubilis (p. 55). Lamouroux' (1812) earlier citation, then, refers to Sertularia volubilis Ellis & Solander, 1786 [ = the nominal hydroid species Clytia johnstoni (Alder, 1856#), usually regarded the same as the medusa Clytia hemisphaerica Linnaeus, 1767; see p. 77]; and the later citation (Lamouroux et al., 1824), employing the same combination, refers to Campanularia volubilis (Linnaeus, 1 758) (see p. 77 for further discussion). The type species of Clytia might appear to be Sertularia volubilis Ellis & Solander, 1 786 (non Linnaeus, 1758, see above), as designated by Mayer, 1910 : 262; but as just explained the combination is a junior homonym. This would be unimportant to current nomenclature if the hydroid Sertularia volubilis Ellis & Solander, 1786, were definitely known to be the same species as Medusa hemisphaerica Linnaeus, 1767. A relation is usually assumed (see N.E. ATLANTIC CAMPANULARIID HYDROZOANS 71 notes under Clytia hemisphaerica, p. 79), and Linnaeus indicated Gronovius' (1760 : pi. 4, fig. 7) illustrated specimen, collected from Belgian waters (the type locality of hemi- sphaerica}. The illustration shows a strongly convex exumbrella surface, suggesting C. hemisphaerica sens. auct. (e.g. p. 73); but the lingering doubts over the identity of the hydroid C. gracilis (Sars, 1850; see p. 78) make the relation of the two nominal species uncertain. Therefore, it is useful to establish a soundly based name for the type species of the genus Clytia. The earliest available name which can be unequivocally linked with S. volubilis sens. Ellis & Solander, 1786, is Campanularia johnstoni Alder, 18560. Hence the correct name for the type species of Clytia is C. johnstoni. The often quoted synonymy with C. hemisphaerica, repeated below, is subjective. See also Cornelius (1981), and page 78 concerning the very similar 'Laomedea gracilis' Sars, 1850. DIAGNOSIS. Colonial Campanulariidae with free medusa stage; hydroid generation forming umbranched stoloniferous or branched upright colonies; hydrotheca with true diaphragm, rim sinuous to deeply indented with round to sharp clefts and cusps; diaphragm transverse; no sub-hydrothecal spherule; medusa hemisphaerical to flat, with hollow marginal tentacles, velum well developed; manubrium short; 4-8 marginal tentacles on release (in species so far studied), many tentacles in adult. REMARKS. A summary of the species of Clytia in which the life cycles have been worked out was given by West & Renshaw (1970); and Roosen-Runge (1970) and Arai & Brinckmann-Voss (1980) gave detailed appraisals of several North American species. Early generic names applied to the type species were broad in concept (Sertularia volubilis = Medusa hemisphaerica = Campanularia johnstoni) and the diagnosis of them by successive workers often overlapped. Hence it is not unexpected that species now included in Clytia s. str. should at one time or another have been included in one or more other genera. There is little value in discussing past uses of generic names such as Medusa, Sertularia, Oceania and Campanularia for species now referred to Clytia. The name Thaumantias Eschscholtz, 1829, was also once widely used. It is a junior subjective synonym of Clytia, since Forbes (1848 : 41) nominated Medusa hemisphaerica Linnaeus, 1767, its type species. This medusa is, of course, widely held to come from the hydroid which is type species of Clytia; but the link is neverthless subjective and is likely to remain so as the original description of the medusa was brief. The genus Silicularia Meyen, 1834, is discussed on page 50. Availability of the generic name Eucope Gegenbaur, 1856, was discussed by Haeckel (1879), Mayer (1910), Rees (1939) and Russell (1953) among others. The four originally included species were referred to Obelia and Clytia (or Phialidium) by Mayer and subsequent authors; but so far as I can determine no type species has been designated. Russell (1953) referred three of the originally included species (E. thaumantoides, E. campanulata & E. affinis) to Clytia hemisphaerica (as Phialidium); and the remaining one (E. polystyla) is Obelia (e.g. following Cornelius, 19750). I designate E. affinis Gegenbaur, 1856, type species of Eucope, which falls in the subjective synonymy of Clytia (see also Bedot, 1910 : 414 and Hincks, 1868: 143). The genus Phialidium Leuckart, 1856, was based on a single species, P. viridicans Leuckart (1856 : 18-19, pi. II, figs 12, 14) which is therefore type by monotypy. Mayer (1910) confirmed its type status, and Kramp's (1961) designation of 'P. hemisphaericum L.' is incorrect. Russell (1953) drew attention to the close similarity between P. viridicans and Clytia hemisphaerica, including them in the same species synonymy. Thus the case for regarding Clytia and Phialidium congeneric is strong and Phialidium, being the later name, need no longer be used. The genus Epenthesis McCrady, 1857, was referred to Clytia by Mayer (1910 : 261) and Nutting (19 15 : 1) among others. The name is clearly a junior synonym of Clytia. The subgenus Platypyxis Agassiz, 1962, was referred to Clytia by Bedot (1910), and I concur (see Remarks under C. hemisphaerica). 72 P. F. S. CORNELIUS The genus Gastroblasta Keller, 1883, now includes just one species, from the Red Sea. All other uses of the generic name have related to abnormal medusae of Clytia species (Kramp, 1 961; also Mayer, 1910, Kramp, 1957,1965). The genera Multioralis Mayer, 1900, and Pseudoclytia Mayer, 1900, were referred by Kramp ( 1 957, 1 96 1 ) to Phialidium Leuckart, 1856, and hence fall into Clytia. Luminescence has long been known in the genus (e.g. Darwin, 1860 : ch. 2, hydroid stage; other references in Forbes, 1848, medusa stage). Light emission was probably first recorded in Clytia by Macartney (1810). In October, 1804, he demonstrated to an invited audience at Herne Bay, Kent, that flashing in the medusa of C. hemisphaerica (which he called Medusa lucidd) was induced by raised temperature, electric shock and alcohol. His published illustration of the medusa is among the earliest of the genus. Clytia discoida (Mayer, 1900) (Fig. 8) Oceania discoida Mayer, 1900 : 5 1 , pi. 20, figs 53-55. Phialidium discoidum Mayer, 1910:272, pi. 33, figs 9-11; Kramp, 1959: 148, fig. 187; Kramp, 1961 : 1 65-1 66; Schmidt & Benovic, 1977:637. TYPE LOCALITY. Tortugas, Florida; in plankton. MATERIAL EXAMINED. None. DESCRIPTION (after Mayer, 1910; Kramp, 1959; Schmidt & Benovic, 1977). Adult medusa 'quite flat' (Mayer), about 4 mm diameter; 1 6 short marginal tentacles with large basal bulbs; usually 3 statocysts between tentacle bases; velum well developed; 4 straight radial canals; gonads along almost whole length, eggs in 9 unusually large and prominent; manubrium 'urn-shaped' (Mayer) with 4 recurved lips. Hydroid stage not recorded. Fig. 8 Clytia discoida. Adult medusa. Diameter c. 4 mm. Redrawn after Mayer (1910 : pi. 33, fig. 10). DISPERSIVE STAGE. The species is known from the medusa alone. See also notes under Dispersive stage of C. linearis. REPRODUCTIVE SEASON. Summer; June-August (Mayer, 1900; Schmidt & Benovic, 1977). DISTRIBUTION. Recorded from: Florida, Mexico, Brazil and W. Indies (references in Kramp, 1961); southern Adriatic Trough (once only, Schmidt & Benovic, 1977). Several records from the Pacific Ocean were discredited by Kramp ( 1 96 1 ). HABITAT. Coastal plankton. REMARKS. The species is distinguished by its small size at maturity and by the extension of the gonads along almost the entire length of the radial canals. Kramp (1959, 1961) seems mistaken in stating the umbrella to be 'almost hemispherical'. Others have reported it nearly flat in the adult (Mayer, 1900, 1910; Schmidt & Benovic, 1977). However, younger stages are reportedly less flat (Mayer), but Schmidt & Benovic reported a young specimen only N.E. ATLANTIC CAMPANULARIID HYDROZOANS 73 1*5 mm in diameter which was already 'more flat than hemispherical'. The adult diameter is 4 mm. The species is known in the NE Atlantic from a single Adriatic specimen (Schmidt & Benovic, 1977). Kramp (1961) provided a literature synopsis. Clytia hemisphaerica (Linnaeus, 1 767) (Fig. 9) Medusa hemisphaerica Linnaeus, 1767 : 1098; Miiller, 1776 : 233. Sertularia uniflora:E\\\&, 1768 : 434, pi. 19, fig. 9. Sertularia volubilis: Ellis & Solander, 1 786 : 5 1 , pi. 4, figs E, e, F, f (non S. volubilis Linnaeus, 1 758; see p. 76 and Remarks). Oceania flavidula Peron & Lesueur, 1810a : 345; Peron & Lesueur, 718106 : 33. Oceania hemisphaerica: Peron & Lesueur, 18100 : 347; Peron & Lesueur, 18106 : 35. Clytia urnigera Lamouroux, 1816 : 203, pi. 5, fig. 6 (see Remarks under Orthopyxis Integra). Thaumantias inconspicua Forbes, 1848 : 52, pi. 8, fig. 3 a-d; Ritchie, 1911:31. ILaomedea gracilis Sars, 1850 : 138; Sars, 1857 : 160, pi. 2, figs 1-3, 5 (but not fig. 4, = Gonothyraea loveni; see Stechow, 19230 : 111); (non Dana, 1846 = Obelia dichotomd). Campanularia volubilis: (sens. Ellis & Solander) Hincks, 1852 : 84-85, pi. 3, fig. 5 (see Dispersive stage and Remarks). Campanularia sp. Gegenbaur, 1854 : 154, 189, pi. 1, figs 8, 8a, 9. Campanularia johnstoni Alder, 18560 : 359-360, pi. 8, fig. 8 (nom. nov. pro Sertularia volubilis Ellis & Solander, 1786). IPhialidium viridicans Leuckart, 1856 : 18-19, pi. 1, figs 12, 14 (see pp. 71, 86). IClytia noliformis McCrady, 1857 : 194-195, pi. 11, fig 4; Fraser, 1944 : 144-145, pi. 26, fig 1 17 (syn. C. simplex Congdon). Campanularia gegenbauriSars, 1857 : 48-49. Campanularia volubiliformis Sars, 1857 : 156 (nom. nov. pro Campanularia sp. Gegenbaur, 1854). Campanularia raridentata Alder, in Hincks, 18616 : 292; Hincks, 1868 : 176-177, fig. 2. Clytia (Trochopyxis) bicophora Agassiz, 1862 : 304-305, pi. 29, figs 6-9. ?Clytia (Platypyxis) cylindrica Agassiz, 1 862 : 306-307, 354, text-figs 4 1-44, pi. 27, figs 8-9. Clytea vicophora Wright, 1862 : 308 (unjustified emendation of Clytia bicophora). Campanularia ?gigantea Hincks, 1866:297; Hincks, 1868:174-175, pi. 35, fig. 1; Jaderholm, 1909: 19, 33, 69, pi. 7, figs 1-3. Clytia johnstoni: Hincks, 1868 : 143-146, pi. 24, figs 1, la (syn. Sertularia volubilis Ellis & Solander; Eucope campanulata Gegenbaur; E. thaumantoides Gegenbaur; E. affinis Gegenbaur; Campanularia gegenbauri Sars; Clytia bicophora Agassiz; see Remarks); Bohm, 1878 : 167-171, pi. 2, figs 1-9 (syn. Platypyxis cylindrica Agassiz; Eucope picta Keferstein & Ehlers, 1861; E. exigua Keferstein & Ehlers, 1861; E. variabilis Claus; E. gemmifera Keferstein; Thaumantias dubia Kolliker; T. thompsoni Forbes; T. convexa Forbes); Billard, 1928: 456-457 (syn. C. raridentata Hincks); Russell, 1953: 293, fig. 179; Ralph, 1957: 823-824, figs Ih-u, 3a-f(syn. C. compressa Totton, 1930). Clytia volubilis: DM Plessis, 1871 : 167-170, pi. 2; Hargitt, 1909 : 373-374. Clytia laevis Weismann, 1 883 : 1 58-1 59 (see Remarks under Campanularia volubilis, p. 57). Clytia flavidula: Metschnikoff, 18860 : 241-243, 257, 260, pi. 22, figs 9-10, 15. Campanularia? serrulata Bale, 1888 : 757, pi. 12, fig. 4. Campanularia raridentata var. Marktanner-Turneretscher, 1890 : 205, pi. 3, figs 3a-b. Thaumantias hemisphaerica: Browne, 1896 : 480-482 (syn. Medusa cymbaloidea Slabber, 1775; M. campanella Shaw & Nodder, 1795; M. lucida Macartney, 1810; Thaumantias lucida: Lesson, 1843; Epenthesis cymbaloidea Haeckel, 1879). Campanularia inconspicua: Calkins, 1899 : 349. ^Campanularia attenuata Calkins, 1899 : 350, pi. 2, figs 9, 9a-c, pi. 6, fig. 9d. ^Campanularia edwardsi Nutting, 19016 : 346, fig. 28. ^.Campanularia pelagica Van Breemen, 1905 : 205-209, fig. 18. Clytia simplex Congdon, 1907 : 471-472, figs 14-15. Clytia obeliformis Stechow, 1914 : 128-129, fig. 6. IClytiapelagica: Billard, 1917 : 539-542, fig. 1. ?Clytia serrulata: Stechow, 19190 : 46-47, fig. M. 74 P. F. S. CORNELIUS Campanularia acuta Stechow, 1919a:54 (nom. nov. pro C. raridentata var. Marktanner- Turneretscher). Campanularia ?attenuata Stechow, 1919a : 61-62, fig. S (non Calkins, 1899). Campanularia brachycaulis Stechow, 1919a : 62-63, fig. T. Orthopyxis volubiliformis: Stechow, 1 9 1 9a : 70. Campanularia villafrancensis Stechow, 1919a : 157 (nom. nov. pro C. attenuata Stechow, 1919a (sic) : 6 1 , preoccupied). Clytia uniflora: Stechow, 1923a : 1 1 1 (non Sertularia uniflora Pallas). Thaumantias raridentata: Stechow, 1923a: 107-1 08, fig. 17. Clytia villafrancensis: Stechow, 1923a : 109-1 10. Clytia compressa Totton, 1930 : 146-148, text-fig. 6. tPhialidiumbicophorum.-T'hiel 1935 : 172; Kramp, 1959 : 149; Kramp, 1961 : 164-165. ILaomedea pelagica: Vervoort, 19460 : 285-288; Vervoort, 1959:313-315, fig. 55b-c; Vervoort, 1968 : 15-17, fig. 5; Vervoort, 1972 : 91-92, fig. 26c. Laomedea gigantea: Leloup, 1952 : 161, fig. 93. Phialidium hemisphaericum: Billard, 1928 : 457 (syn. Thaumantias inconspicua Forbes); Kramp, 1919 : figs 16-17 (syn. P. temporarium Browne, 1896; Thaumantias buskiana Gosse, 1853); Russell, 1953 : 285-294, pi. 16, fig. 1, pi. 17, fig. 6, text-figs 172-179 (syn. Thaumantias pileata Forbes, 1841; T. sarnica Forbes, 1841; T. thompsoni Forbes, 1841; T. punctata Forbes, 1841; T. lineata Forbes, 1848; T. hemisphaerica: Forbes, 1848; T. inconspicua Forbes, 1848, Hincks, 1868; 7". buskiana Gosse, 1853; Eucope affinis Gegenbaur, 1856; E. campanulata Gegenbaur, 1856; E. thaumantias Gegenbaur, 1856; Phialidium viridicans Leuckart, 1856; ?T. typica Green, 1857; Clytia johnstoni: Hincks, 1868; T. leucostyla Spagnolini, 1876; Campanulina acuminata Bohm, 1878; Epenthesis cymbaloidea Haeckel, 1879; P. variabile Claus, 1881; Clytia flavidula Metschnikoff, 1886; C. viridicans Metschnikoff, 1886; P. buskianum Browne, 1896; P. temporarium Browne, 1896; T. forbesi Johansen & Levinsen, 1903; Clytia volubilis: Sverdrup, 1921; for these references see Russell, 1953); Kramp, 1955 : 256-257 (syn. P. variabile Haeckel, 1879; ^Oceania languida Agassiz, 1862). Clytia hemisphaerica: Rees & Thursfield, 1965 : 95-96; Millard, 1966 : 478-480, fig. 14a-f; Vervoort, 1968 : 16-17;Calder, 1975 : 300-302, fig. 4a-b; Millard, 1975 : 217-218, fig. 72a-d. Clytia gigantea: Rees & Thursfield, 1965 : 96-97. Thaumantias raridentata: Teissier, 1965 : 17. ILaomedea (Clytia) pelagica Garcia Corrales et al., 1978 : 28-29, fig. 1 1 . ^Campanularia gracilis: (sensu Sars, 1850) Stepanyants, 1979 : 32, pi. 5, fig. 3 (syn. Clytia serrulata: Vanhoffen, 1910). IClytia sarsi Present paper, p. 78 (nom. nov. pro Laomedea gracilis Sars, 1 850, preocc; see p. 78). For further synonmy see Bedot (1901-1925), Russell (1953) and Kramp (1961). Kramp cited just the following combinations from the World literature not included in Russell's British list: Gastroblasta raffaelei, Clythia johnstoni, ^Phialidium languidum (provisionally retained distinct by Kramp), Pseudoclytia pentata f. hexaradiata. See also Clytia incertae sedis (p. 9 1 ). NOMENCLATURE. Millard (1966) has shown that the apparent use of the species name hemisphaerica by Gronovius (1760) was not strictly binominal. Neither also was that by Houttuyn (1770 : 423), cited by Bedot (1901 : 486) as Medusa hemisphaerica, despite the implication of Bedot's entry. Bedot dated Houttuyn's work as 1761-1773; but the volume in which the hemisphaerica reference appeared was published in 1770, three years after Linnaeus' (1767) introduction of the genuine binominal Medusa hemisphaerica. Linnaeus cited Gronovius' illustration as indication. The combination Clytia hemisphaerica results from the subjective linking of hydroid and medusa, and was probably first used by Rees & Thursfield (1965). The next year, Millard (1966) too discussed the combination. Many authors have placed hemisphaerica in the 'medusa' genus Phialidium Leuckart, 1856; but Clytia Lamouroux, 1812, is older and now the life cycle is known should be used instead. The reason why Phialidium need not be used for any species is given above (P. 71). Browne (1896) incorrectly ascribed the combination Medusa hemisphaerica to Miiller, whose work Browne dated as 1766. The correct date of Miiller's work was 1776, and Linnaeus ( 1 767) has clear priority. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 75 TYPE LOCALITY AND MATERIAL. The type material of C. hemisphaerica Linnaeus, 1767, was the medusa described and illustrated by Gronovius (1 760 : 38, pi. 4, fig. 7). I have not tried to find it. Linnaeus gave the type locality as 'Belgian seas', from whence Gronovius' material came. TYPE MATERIAL OF OTHER SPECIES EXAMINED. I have examined the type series of the hydroids Campanularia johnstoni Alder, 18560, and C. raridentata Alder, in Hincks 1861&, and consider them to be C. hemisphaerica (see Cornelius & Garfath, 1980; see also Remarks). The holotype colonies of the hydroids C. ?gigantea Hincks, 1866 (BMNH reg. no. 1899.5.1.106, Lamlash, Arran, W Scotland) and Clytia obeliformis Stechow, 1914 (Munich Zoological Museum, fertile colony on microslide, Bergen) are also Clytia hemisphaerica. OTHER MATERIAL EXAMINED. This species is well represented in the BMNH collections. The following material, from other Museums, deserves mention: 53° 01' N, 4° 22' E, numerous colonies in spirit, coll. Lightship 'Texel', 1961, cf. 'Laomedea pelagica' (sensu Van Breemen), det. W. Vervoort; Rijksmuseum van Natuurlijke Historic, Leiden. 10°40'N, Fig. 9 Clytia hemisphaerica. (a-b) 'pelagic' form, from microslide preparation in Institut Royal des Sciences Naturelles de Belgique, det. 'Laomedea gracilis" by A. Billard (mentioned, Billard, 1917; 'probably ofTOstend'). (a) 'primary attachment disc' with four hydrothecal pedicels. Note absence of stolon, (b) hydrotheca. (c) 'pelagic form', gonotheca, 53° 01' N, 4° 22' E, det. 'Clytia pelagica" by W. Vervoort; colln Rijksmuseum van Natuurlijke Historic, Leiden, (d-e) 'benthic form'; (d) hydrotheca and (e) gonotheca, S England; 1934.8.17.19. (f) 'benthic form', base of pedicel branch from extremely sheltered locality, Caol Scotnish, Loch Sween, Argyll, W Scotland, 1 m, 30 May 1962; 1962.6.19.21. (g) 'benthic form1, base of pedicel branch, nr Bergen, 40-90 m, 9 April 1962; 1962.10.7.19. (h-j) three hydrothecal rims from same colony, Scotland; 1964.8.7.72. (k) hydrotheca, W Sweden, 20-30 m; 1962.11.8.10. (1) primary hydrotheca and attachment disc, reared from medusa ("Phialidium hemisphaericum* sens. Russell, 1953) by W. J. Rees, 2-17 March 1936; 1969.12.2.2. (m) medusa, Naples, redrawn after Mayer (1910: text-fig. 1 44). Scales: (a-c, d-1) 500 //m; (m) unknown. See also Fig. 1 4(a). 76 P. F. S. CORNELIUS 6° 44' W, 65 m, many colonies in spirit, coll. 'Atlantide' expedition, sta. 151, 16 Apr 1946, cf. 'L. pelagica\ det. W. Vervoort; RMNH, Leiden. Sete, S France, infertile material on three microslides, det. Clytia serrulata by Stechow, 1919a : 46; Munich Zoological Museum. DESCRIPTION OF HYDROID STAGE. Among the most variable of all Atlantic hydroids. Detached colonies can continue growing in the plankton and form one extreme of variation, while benthic colonies are more typical of the species and form the other. 1. Benthic colonies. Colony comprising a creeping stolon from which short to long hydrothecal pedicels arise at irregular intervals. Hydrotheca thin walled, campanulate, length : breadth ratio 1-3, rim with 7-16 pointed to rounded or (?rarely) flat topped cusps, embayments typically pointed but sometimes rounded; hydrotheca usually round in cross- section but sometimes (Vervoort, 1968) sinuous near the top. Hydranth with the wide range of 20-30 tentacles (Hincks, 1868; cf. Obelia geniculata, p. 120). Pedicel straight, erect; annulated, usually with smooth central region and 2-10 annuli each end, rarely annulated throughout; sometimes branched, each branch having characteristic upward-curved basal region (similar to that in C. paulensis), branches often parallel with main pedicel; branch annulated basally and distally, central portion smooth. Gonotheca 3 = 9, broad, tubular, sometimes asymmetric, walls smooth to deeply concertinered, often with slight sub-terminal constriction; tapered below; aperture large, terminal; gonothecal pedicel short, on hydro- thecal pedicel or on stolon. Medusa released at four-tentacle stage. Nematocysts described by Ostman(1979). 2. 'Planktonic' colonies. Floating colonies have been described by several authors (e.g. Van Breemen, 1905; Billard, 1907; Vervoort, \946a, 1959, 1968, 1972). Available evidence suggests that they are benthic colonies which have grown on sand and become detached (see Remarks). Colonies comprise one to several pedicels arising from a basal disc which often (Leloup, 1933) envelops a sand grain, stolons apparently being absent. Pedicels exceptionally long, branching repeatedly, branches often aligned parallel with primary pedicel. Hydrotheca long, narrow, thin walled, terminal cusps as in benthic form. Gonotheca long, smooth walled; rugose and concertinered gonothecae apparently not recorded from free floating material. Variation. Ralph (1957) found that cooler water increased the number of hydrothecal cusps. Although the species is highly variable there have been few studies of the factors involved. DISPERSIVE STAGE. A medusa, released at the four-tentacle stage. Umbrella hemispherical or slightly flatter; up to 25 mm in diameter; jelly fairly thin. Velum narrow; stomach short, four-sided; manubrium about 1/6 bell diameter in length, four-lipped. Radial canals 4; gonads elongate to oval, on distal 1/4-3/4 of radial canal but stopping just short of ring canal. Marginal tentacles 16-32 [58 recorded, Kramp, 1919 (=64)], short, hollow, smooth, with spherical bases. 1-3 closed marginal vesicles between each marginal tentacle and next. Tentacle bases and stomach yellow, yellowish brown, greenish or purple; gonads yellowish. (After Kramp, 1919, 1961; Russell, 1953.) The relation between hydroid and medusa has not always been understood, and there are still some unsolved problems relating to the nominal species described from the hydroid stage as 'Laomedea gracilis Sars'. Bohm (1878), working at Helgoland, worked out the life cycle of C. hemisphaerica and his account was quoted by Haeckel (1879 : 187) in his World synopsis; but many years later Mayer (1910 : 267) wrote 'the mature hydroid is not known with certainty, but is probably a Campanulina (see Hincks, 1868 : 179)'. Mayer should have quoted Hincks as indicating "Campanularia raridentata\ now regarded as conspecific with hemisphaerica. Hincks' citation in fact referred back to Wright's (1862) description of the hydroid stage under the name Thaumantias inconspicua; but Hincks (1852) himself had still earlier seen medusae released (identifying his material as "Campanularia volubilis Ellis & Solander'). Hincks thus seems to have been the first to record medusa release in the present species. Du Plessis (1871) also saw medusa release at an early date, but used the combination Clytia volubilis. (Notes on N.E. ATLANTIC CAMPANULARIID HYDROZOANS 77 the widespread confusion between the species names volubilis and hemisphaerica are given below.) Mayer's oversight is the more surprising since he quoted MetschnikofTs (1886&) description of a polyp reared from the Mediterranean medusa Clytia flavidula (Peron & Lesueur, 1810a), which Mayer thought conspecific. The identity of C. flavidula is discussed under Remarks. The later but independent suggestions of Browne (1896 : 488; 1900 : 725-726; Browne & Vallentin, 1904:125, 127) that the hydroid of 'Phialidium hemisphaericum' was 'Campanularia Johnston? were supported by Kramp in 1914 (Kramp, 1919 : 93, footnote), some forty years after Bohm and Haeckel wrote and even longer after the suggestions of Du Plessis, Hincks and Wright. The delay parallels that which occurred in the working out of the Obelia life cycle (Cornelius, \911a, b). REPRODUCTIVE SEASON. Medusae occur all through the year in British waters but are most numerous from spring to autumn [Hincks, 1852, as C. volubilis (in February); Russell, 1953; J. H. Robson, in Evans, 1978, as C. gracilis)]. Lo Bianco (1909, as Clytia flavidula and C. johnstoni) recorded medusae off Naples from August to October and from January to March. DISTRIBUTION. Nearly cosmopolitan in coastal waters (Ralph, 1957; Kramp, 1961), occurring in most of the present area. In European waters the medusa is one of the commonest (Russell, 1953). However, the species was said to be scarce in N Norway by Mathiesen (1928) who cited only two certain records, from Bergen and Hammerfest. It has been reported absent from Greenland (Kramp, 1943; Calder, 1970) although present in Iceland, Spitzbergen, the Faeroes and the Bering Sea (Calder, 1970). 'Planktonic' hydroid colonies have been reported as common in the southern North Sea and off much of the coast of W Africa (Vervoot, 19460, 1959). HABITAT. Usually benthic; intertidal to 150+ m (Crawshay, 1912; Marine Biological Association, 1957; Kramp, 1959; Millard, 1975; BMNH collections). Naumov (1969) reported a lower limit of 300 m but did not cite material. Rees & White (1966) listed a record from 1250 m off the Azores as C. gigantea, a species here regarded conspecific; but I have not checked this report. The species has been found on a wide variety of invertebrate and algal substrates, and there is no regular association. Among the unusual recorded substrates are sand grains (see Remarks) and pelagic cirripedes. There are several records on parasitic copepods themselves on fish (on Lernaeocera on Gadus by Leloup, 1930&; on Peniculus on Mullus, on Lernaeenicus on Clupea, and on Dinematura on Cetorhinus (the Basking Shark), all by Debouteville & Nunes, 1951, 1952). As in Obelia, long distance transport would seem to result (pp. 45, 120). REMARKS. Two factors have contributed to the profusion of redescriptions of this species and to the consequent number of synonyms: it is nearly cosmopolitan, and it is highly variable. The combination Clytia hemisphaerica was introduced only some fifteen years ago and there is still much use in a detailed analysis of the taxonomic history of the species. There has been confusion with other species and with other genera, and some of the problems have yet to be solved. The nominal species described from the NE Atlantic are considered below, so far as possible in chronological order. The notes by Stechow (192 la, \923a) and Rees & Thursfield (1965) on the nominal species Sertularia uniflora Pallas, 1766, were partly misleading. Pallas included in his synonymy Ellis' (1755) plate 14, figure A, Linnaeus' (1758) citing of that plate under the name Sertularia volubilis, and Baster's (1762) plate 2, figures 2a, b, d, 3, 4c, e, 7a-c. Ellis' illustration shows a campanulariid hydroid growing on Hydrallmania falcata (Linnaeus, 1758). The pedicels were spirally grooved throughout, not annulated top and bottom as in the present species, and had a spherule at the upper end. These two characters indicate that Ellis' material was the species here called Campanularia volubilis. Hargitt (1909), Stechow and Rees & Thursfield wrongly supposed Ellis' material to be C. hemisphaerica. The name Sertularia uniflora Pallas, 1766, was in fact a nom. nov. for Sertularia volubilis Linnaeus, 78 P. F. S. CORNELIUS 1758, and since both were based on Ellis' illustration uniflora is a junior objective synonym ofvohibilis. The other illustrations cited by Pallas under S. uniflora, those of Baster, show in one place (fig. 2A, b) a sharply cusped hydrothecal rim, but nowhere show details of the pedicel. Hence it cannot be determined whether Baster's material was volubilis or hemisphaerica. Gonothecae were not shown. Maitland (1876, quoted in Vervoort, 19460 : 276) referred part of Baster's illustrations (pi. 2, fig. 7A, a) to Orthopyxis Integra (Macgillivray, 1842), an opinion disagreed with by both Vervoort (19460) and myself as the hydrotheca of Integra does not have a cusped margin. Nevertheless, I here designate the material shown by Ellis (1755 : pi. 14, fig. A) lectotype of 5". uniflora Pallas, 1766, to provide formal protection for the name Integra from possible threat from uniflora should Maitland's opinion ever be upheld. Rees & Thursfield (1965) were further confusing in stating that 'earlier references by Ellis (17550, b) [sic] do not distinguish' between hemisphaerica and volubilis. They omitted the Ellis references from their paper: indeed, I cannot trace relevant papers by Ellis in that year. His only 1755 publication mentioning campanulariids was his book. In this Ellis (1755 : 25) stated clearly that there were two species and illustrated both on his plate 14. Linnaeus and Pallas (op. cit.) were each careful to cite only one of the species when referring Ellis' illustration respectively to their synonymies of volubilis and uniflora. The other illustrated species (Ellis, 1755 : pi. 14, B) was named Sertularia syringa by Linnaeus (1767 : 1311) and is now known as Calycella syringa, family Calycellidae (e.g. Cornelius, 1978). Thus there is little confusion in the early literature; but Rees & Thursfield (1965) were probably correct in stating that the 'hydroid [stage of C. hemisphaerica] is recognisable for the first time under the name Sertularia uniflora: Ellis, 1 768 : pi. 19, fig. 9'. And in this paper Ellis clearly did confuse the two species he had previously illustrated separately. His use of the name uniflora in that paper can be regarded as misidentification, not as homonymy. Ellis' (1768) illustrations show clearly the annulated, not spiralled, pedicels of hemi- sphaerica— incidentally slightly unusual in lacking a smooth central portion — and a characteristic 'concertinered' gonotheca. Further confusion might have resulted from Ellis & Solander's (1786) application of the name Sertularia volubilis to illustrations clearly of the present species, but the slightly earlier literature just discussed prevents it. However, Hargitt (1909) was not alone among later authors in applying the combination Clytia volubilis to the present species. C. hemisphaerica was placed in the genus Oceania Peron & Lesueur, 18100 (as O. flavidula), by those authors. Clytia urnigera Lamouroux, 1816, is referred to C. hemisphaerica but is discussed here under Orthopyxis Integra (pp. 64-65). The combination Laomedea gracilis Sars, 1850, has plagued the literature (see also Remarks under Gonothyraea loveni); but as noted by Vervoort (19460 : 285) the species name should not be used as it is preoccupied by "Lomedea gracilis'' C. Pickering, in Dana, 1 846 : 689 (lapsus pro Laomedea gracilis), which I refer to Obelia dichotoma (p. 117). I propose the name Clytia sarsi nom. nov. in place of L. gracilis Sars, 1850 (preocc.). Lectotype material of L. gracilis Sars is designated here on page 94. Nevertheless, the name gracilis has been widely applied to colonies having long, narrow hydrothecae and smooth gonothecae. But following Ralph's (1957) account of variation in New Zealand populations most authors have regarded '50r$/-type' colonies as an extreme variation of hemisphaerica. Further evidence was provided by W. J. Rees (in Rees & Thursfield, 1965) who reared a medusa from a sarsi ('gracilis'') colony and found it the same as the medusa hemisphaerica s. str. Kiihl (1967) regarded 'sarsi-type' colonies conspecific with those of hemisphaerica on more subjective grounds. But Ostman (1979) separated the species on the fine structure of the nematocysts (see also p. 42) and the problem is not yet resolved. There has been some further debate over the degree of affinity between C. sarsi (=L. gracilis Sars, 1850), and Campanularia pelagica Van Breemen, 1905. Some authors have regarded them identical (e.g. Leloup, 1933, 1952; Vervoort, 19460; Naumov, 1960, 1969; N.E. ATLANTIC CAMPANULARIID HYDROZOANS 79 Garcia Corrales et al., 1978); but the validity of C. pelagica s. str. has been difficult to assess. (Campanularia attenuata Calkins, 1899, from Puget Sound, seems similar.) Among recent authors Vervoort's (1946a, 1959, 1968, 1972) descriptions of the pelagica 'form' have been important but the possibility has remained open that pelagica was simply based on sarsi or hemisphaerica material which had detached and grown in the plankton. Billard (1917) also had failed to resolve the question but E. T. Browne, A. Kemna and E. Leloup (in Leloup, 1933) thought sarsi (gracilis) just to be detached, floating colonies of hemisphaerica. Vervoort (1959, as Laomedea pelagica) reported that 'floating' colonies occurred in the North Sea and along much of the west coast of Africa, but Millard (1966) referred Vervoort's African coast material to C. hemisphaerica without comment. Subsequently Vervoort (1968) reaffirmed his view that pelagica was distinct. He had not seen 'intermediate material'; but presumably there is no intermediate environment. Vervoort separated pelagica on its slender, gradually narrowing hydrothecae, an undulating cross-section just below the hydrothecal rim and a smooth walled gonotheca. He cautiously noted that two species of Clytia medusae had been recorded from the North Sea by Russell (1953) and that these might relate to the two hydroids, pelagica and hemisphaerica. He subsequently (Vervoort, 1972) retained specific status for pelagica but Millard (1975) again referred Vervoort's African coast material to hemisphaerica. So the experts were divided. Naumov (1960, 1969) meanwhile had considered pelagica distinct but extended its limits slightly further than Vervoort to include sarsi-lype material with smooth walled gonotheca. Garcia Corrales et al. (1978) similarly regarded sarsi + pelagica distinct from hemisphaerica, based on sharper and longer hydrothecal cusps and longer hydrotheca in the former; and like Vervoort (1968) had seen no intermediate material. Other features on which pelagica has been distinguished are the frequency of branching and the tendency of the branches to run parallel with the primary stem, noted by Vervoort (1959) but not apparent in all planktonic material; and a smooth, asymmetric gonotheca (e.g. Billard, 1917). Vervoort (1972 : 91, footnote) drew attention to the report by Sars (1850, 1857; ?repeated in Leloup, 1952) that 'pelagica' had sessile gonophores. Vervoort cited observations that 'pelagica-type' material released its medusae and (pers. comm.) now considers that the contrary observations of Sars might have been made on Gonothyraea loveni material. Old and new evidence for uniting pelagica with hemisphaerica runs as follows. Ralph (1957) showed that smooth gonothecae are not unusual in hemisphaerica, and that the length of the hydrotheca and shape of the cusps on the rim vary widely, to an extent which encompasses the three nominal taxa. The basal discs (=Haftplatte of Kuhn, 1913, and disque de fixation of Houvenaghel-Crevecoeur, 1973; there seems no English equivalent already) of attached C. hemisphaerica examined by me seem identical with the 'partie basale' or "pied . . . globuleuse' described in planktonic colonies by Billard (1917) and Leloup (1933) under the name pelagica (Fig. 9). Also, Leloup showed that the basal discs of these free-floating colonies often contain a sand grain, indicating a benthic origin. Next, the distinctive upward-swept pedicel bases in hemisphaerica s. str. seem identical with those described as pelagica. Finally, it might have been expected that a hydroid which was habitually planktonic would have some obvious modification to that end; but there seems none. The relation between the three nominal forms hemisphaerica, sarsi and pelagica is certainly close, and the relation tojohnstoni is still uncertain (see below). Whether the differ- ences are phenotypic or genotypic cannot yet be decided; but on skeletal and medusa characters it seems best now to interpret the variation as representing a single 'morphological species'. More detailed studies, such as that of Ostman (1979) on the nematocysts, might shed further light. Campanularia volubilis (Ellis & Solander, 1786) sens. Hincks (1852) was the present species. Hincks' account was probably the earliest record of medusa release in C. hemisphaerica. Campanularia johnstoni Alder, 1856a, is apparently the earliest available name for the hydroid stage most usually (but subjectively) connected with the medusa on which the 80 P. F. S. CORNELIUS present species is based (see Nomenclature, above). It is also the correct species name for the type species of the genus Clytia (p. 71). The type series of Campanularia johnstoni is mentioned under Material, above, and by Cornelius & Garfath (1980). See also Cornelius (1981). Phialidium viridicans Leuckart, 1 856, is discussed above (p. 7 1 ) and on page 86. Clytia noliformis (McCrady, 1857, as Campanularia) was founded on a hydroid which released a Clytia medusa lacking gonads when liberated. Kramp (1959, 1961) regarded the lack of gonads on release an important character; and the species has been widely recognized from both hydroid and medusa stages in warm waters throughout the World (references in Fraser, 1944; Kramp, 1961; comment in Rees & Thursfield, 1965). Mammen (1965) gave this name to a medusa he reared but it closely resembled Russell's (1953 : pi. 17, fig. 6) illustration of C. hemisphaerica. Mammen's medusa differed only in not showing gonads till 48 hrs old. C. noliformis has not otherwise been redescribed since Kramp (1961) wrote and may prove conspecific. Picard (1949) referred medusa and hydroid material from Ville- franche to noliformis but his specimens, like Mammen's, seemed to differ from hemi- sphaerica only in the short delay in acquiring gonads. Later he included the species in a Mediterranean faunal list, regarding C. jlavidula Metschnikoff, 1886a, and C. mollis Stechow, 1919a, as conspecific (Picard, 19586). These two species have both been referred to hemisphaerica by at least some senior authors; and are discussed in their chronological place below. Rees & Thursfield (1965) referred some Cape Verde material from James Ritchie's collection to C. noliformis. They regarded Campanularia ptychocyathus Allman, 1888, from the Azores, as conspecific; but Stechow (1925 : 521) treated ptychocyathus as dubious. Campanularia gegenbauri Sars, 1857, was based on a fertile hydroid colony illustrated by Gegenbaur (1854: pi. 1, figs 1, la, as Campanularia n. sp.). I concur with Hincks (1868 : 145), Bohm (1878 : 168) and Bedot (1910 : 254) that the species should be referred to Clytia hemisphaerica. Campanularia volubiliformis Sars, 1857, was a name applied to material described as Campanularia sp. by Gegenbaur (1854). The shape of the hydrotheca, pedicel and gonotheca illustrated indicate the present species and not Campanularia hincksii as suggested by Carus (1844) and Broch (19126). Heller (1868) and Picard (195 la) reported further material without description; while Stechow 1919#:70) referred the species to Orthopyxis\ Campanularia raridentata Alder, in Hincks, 18616, was referred to Clytia hemisphaerica by Billard (1928) and Rees & Thursfield (1965), and provisionally by Vervoort (1968). The synonymy was agreed by Cornelius & Garfath (1980), who alone saw the holotype. The specimen was simply a young colony of C. hemisphaerica. Rees & Thursfield wrongly ascribed the original description to 'Alder, 1857'. Clytia bicophora Agassiz, 1862, originally based on hydroid material from New England, was recorded as a medusa in the Gulf of Trieste by Thiel (1935 : 172, as Phialidium bicophorum). But Agassiz' detailed description and clear illustrations appear to represent C. hemisphaerica, and bicophora can be regarded conspecific. Kramp (1959 : 149) listed other records of bicophora as 'uncertain', and indeed Bohm (1878) referred bicophora to C. johnstoni, which is also regarded conspecific. Kramp (1959, 1961) nevertheless described a medusa under the name bicophora and listed references to that species; but the supposed differences from hemisphaerica seem unimportant. Clytia (Platypyxis) cylindrica Agassiz, 1862, was based on both hydroid and medusa material from Massachusetts Bay and Buzzards Bay, North America. The species was referred to C. hemisphaerica by Bohm (1 878), but to C. noliformis (McCrady, 1 857) by Bedot (1910:348) and Kramp (1961 : 170). (Bedot clearly regarded Platypyxis a synonym of Clytia, and I concur; see p. 71.) Agassiz' description of cylindrica includes no important differences from hemisphaerica and like Bohm I regard them conspecific. Vervoort (1968) described new material, and commented on the similarity of C. elsaeoswaldae Stechow, 19 14. 1 agree with Vervoort in regarding the Stechow species conspecific. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 81 The combination Phialidium languidum (Agassiz, 1862) has been applied to medusae caught off Senegal, but Kramp (1955) referred both the material involved and the nominal species itself to C. hemisphaerica (see Kramp, 1933, 1955, 1 96 1 , for discussion). Campanularia ?gigantea Hincks, 1866, was probably based on large hemisphaerica hydroid material. The eleven hydrothecae of the holotype well enough preserved for study measured, in mm from diaphragm to tips of hydrothecal cusps, M8, 1'26, l-29 (3), 1'53, 1-62 (2), 1*78, 1'88, 1-91. The wide range is striking, as is the sheer size of the largest; but Hincks was rightly cautious to introduce the name as ?gigantea. Several authors have referred large material to gigantea solely on the basis of size. The pedicels too are long, but the upward curve of the pedicel bases recalls hemisphaerica s. str. and continued separation seems uncalled for. Gonothecae have never been reported in this large material. Hincks' (1868) later illustration of the holotype is accurate; and that of Jaderholm (1909), of non-type material, is similar. The material fron N Scotland listed by Rees & Thursfield (1965; BMNH 1964.8.7.75) is C. hemisphaerica and is smaller than the holotype of ?gigantea. The material from Belgium cited by Leloup (1952) seems to have been the same, as does the North Sea material listed by Billard (1928, as Clytia johnstoni). Gastroblasta raffaelei Lang, 1886, was based on a medusa from Naples. The species was provisionally referred to C. hemisphaerica by Kramp (1959 : 148). Clytia flavidula: Metschnikoff, 1886<2, was a comb. nov. for Oceania flavidula Peron & Lesueur, 1810a, originally described from medusae collected from Nice. Mayer (1910) referred the species to C. hemisphaerica and Russell (1953) concurred. Lo Bianco (1909 : 540) also used the combination Clytia flavidula. Kramp (1961 : 65) listed another use of the binominal O. flavidula, for O. armata, a clavid medusa. The Mediterranean material referred by Stechow (191 9a) to Campanularia serrulata Bale, 1888 (first described from Sydney), was examined here and found to be simply Clytia hemisphaerica; as indeed seems Bale's species, which was described from infertile material. In general agreement, Stepanyants (1979) referred C. serrulata to 'Campanularia gracilis' Sars, 1850, itself regarded conspecific (see above). Picard (1958/7) recorded the species as Mediterranean without comment. Campanularia attenuata Calkins, 1899, originally described from Puget Sound, resembles 'Laomedea pelagica Van Breemen, 1905', and like it should probably be regarded conspecific with Clytia hemisphaerica. Material was recorded from Ghana by Buchanan (1957) and it is well known that 'pelagica-type' colonies occur along the African coast (see above). Although West & Renshaw (1970) recognized the species on the basis of its branched colonies as grown in vitro at certain temperatures (see p. 40) further proof of validity seems necessary. C. attenuata Stechow, \9\9a, from Villefranche, is a junior homonym and later in the same paper Stechow (1919a:157) introduced the nom. nov. Campanularia villafrancensis instead. However, villafrancensis too seems conspecific with hemisphaerica. Campanularia edwardsi Nutting, 190 \b, based on material from Massachusetts, was redescribed at length by Kubota (1978, as Clytia). He provisionally referred C. edwardsi to the present species, and I concur. Clytia simplex Congdon, 1907:471-472, figs 14-15, based on material taken off Sargassum weed near Bermuda, was reported south of the Azores by Vanhoffen (1910). Fraser (1944), however, referred the species to C. noliformis McCrady, 1857, here regarded conspecific with C. hemisphaerica; and it seems probable that C. simplex too is conspecific. Clytia obeliformis Stechow, 1914, was based on material from Bergen, Norway. The original illustration shows both hydrotheca and gonotheca typical of C. hemisphaerica and the type material, examined here, comprises simply a rather elongate colony of the present species. There is no reason to maintain a separation. Campanularia acuta Stechow, 1919a, was a nom. nov. applied to C. raridentata: var. Marktanner-Turneretscher, 1890, based on material from NW France. The variety was originally held to differ from C. raridentata s. str. in lacking annulations around the base of the gonotheca; and like that species seems to have been based on Clytia hemisphaerica material. 82 P. F. S. CORNELIUS Campanularia brachycaulis Stechow, 1919a, was based on a small, infertile colony from Villefranche. The illustrated hydrotheca does not seem to differ from that of Clytia hemisphaerica; and I must disagree with Patriti (1970) who referred the species to Campanularia hincksii. Clytia compressa Totton, 1930, was proposed on the basis of the shape of the hydrothecal cusps and of the gonotheca. Ralph (1957) and Vervoort (1968) regarded it conspecific, and I concur. Stolon growth in the hydroid stage was described by Hale (1964, \913a, b). Clytia hummelincki (Leloup, 1935) (Fig. 10) Laomeda hummelincki Leloup, 1935 : 19, fig. 7; Buchanan, 1957 : 360, fig. 1 1 a-b. Campanularia hummelincki: Fraser, 1944 : 122, pi. 21, fig. 93; Vervoort, 1966 : 96. Clytia hummelincki: Deevey, 1954 : 270; Millard, 1966 : 480-481, fig. 14g-l; Millard, 1975 : 218-220, fig. 72 f-h. TYPE LOCALITY AND MATERIAL. Isle de Bonaire, West Indies, 0'7 m depth, on dead coral, infertile material on microslide (Fig. 10); Mus. Roy. Hist. Nat. Belg. collection. OTHER MATERIAL EXAMINED. Fragments of fertile colonies on two microslides, Aghulas Bank, off South Africa (34° 43' S, 25° 40' E); South African Museum reg. no. H 2967 (mentioned, Millard, 1966, 1975). DESCRIPTION OF HYDROID STAGE (partly after Leloup, 1935; Buchanan, 1957; Millard, 1975). Colony a tortuous stolon bearing long hydrothecal pedicels at irregular intervals; Fig. 10 Clytia hummelincki. (a-b) hydrotheca from syntype series, two magnifications, (c) 9 gonotheca (after Millard, 1975 : fig. 72H). Scales: (a) 50 /zm; (b-c) 500 //m. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 83 gonothecae subsessile on stolon. Hydrothecae short, wide, unthickened, length = maximum breadth, tapering sharply towards base which is usually slightly constricted; rim often sloping, even to slightly and irregularly sinuous; diaphragm delicate, oblique in lateral view, either parallel with hydrothecal aperture or sloping in opposite direction; basal chamber small. Hydrothecal pedicel long, smooth or with several basal annulations and up to c. five groups of 3-5 annuli along length; sub-hydrothecal spherule present, below which often a slight constriction (see Fig. 10). Hydranth with spherical hypostome and c. 20 tentacles. Gonotheca (?rf = 9) sessile to shortly stalked, truncate; widest above, tapering below; sometimes asymmetric; aperture probably broad as end; with 1-2 medusa buds, develop- ment successive; buds with 4 radial canals and 4 tentacle rudiments (South African material); mature gonotheca and medusa undescribed. MEASUREMENTS. See Table 2. Table 2 Clytia hummelincki Measurements in /zm. Caribbean Ghana South Africa (Leloup, 1935) (Buchanan, 1957) (Millard, 1966, 1975) Hydrotheca Length 200-240 250 250-^00 Breadth (max) 250 240-250 200-420 Length/breadth 0-8-0-96 1-00-1-04 0-7-1-36 Pedicel Length 1500-2000 2000 1920^730 Breadth (max) 40-60 90-150 730-1260 280-450 DISPERSIVE STAGE. Probably a medusa. Millard (1966, 1975) noted medusa buds with 4 marginal bulbs and so referred the species to Clytia. See also Description, Remarks under C. hemisphaerica and Dispersive stage under C. gravieri. REPRODUCTIVE SEASON. The only recorded fertile material was collected from Aghulas Bank, South Africa, on 10 February, 1962 (Millard, 1966). The gonothecae contained well developed medusae. DISTRIBUTION. A little known species, recorded in the NE Atlantic only from Ghana (Buchanan, 1957). Other Atlantic records indicate a wide distribution: West Indies (Leloup, 1935; Vervoort, 1966), Florida Keys and Woods Hole (Deevey, 1954, possibly northernmost record of species) and South Africa (Millard, 1966, 1975). HABITAT. The few collected colonies have been on Lepas (Cirripedia) attached to a buoy, on sublittoral coral debris and on intertidal Sargassum weed (Leloup, 1935; Buchanan, 1957; Millard, 1966), indicating a wide substrate range. Recorded depths have so far ranged only from intertidal (Buchanan) to less than 1 m (Millard). REMARKS. This rarely reported species is clearly widespread in the warmer parts of the Atlantic Ocean. It is retained in Clytia following Millard's notes on the developing medusa. Although the species has a sub-hydrothecal spherule it is not transferred to Campanularia since that character might be due to convergence (see p. 41). But the affinities of the species remain uncertain and the identity of the medusa might provide better indication. 84 P. F. S. CORNELIUS Clytia islandica (Kramp, 1919) (Fig. 11) Phialidium islandicum Kramp, 1919 : 95, pi. 4, figs 1 1-13, pi. 5, figs 1-2; Russell, 1953 : 294-296, text-figs 180-181; Kramp, 1959: 149, 215, 218, 221, 222, fig. 190; Kramp, 1961 : 169 (?syn. Staurostoma laciniatum var. hybridum Le Danois). TYPE LOCALITY. Coastal waters of Iceland. DESCRIPTION OF MEDUSA STAGE (Hydroid unknown). Diameter of adult 35-40 mm, umbrellar saucer-shaped, jelly thin; stomach small, cruciform; manubrium reduced, mouth square surrounded by crenulated lips; gonads narrow, along almost whole length of radial canals; tentacles c. 200, alternating with statocysts. Variation. See Russell ( 1 953) and Kramp ( 1 959). Fig. 1 1 Clytia islandica. Redrawn after Kramp ( 1 959 : fig. 1 90). Diameter 35-40 mm. DISTRIBUTION. Reported at least as far south as SW Ireland but records generally more northerly (Russell, 1953). Kramp ( 1 96 1 ) cited unconfirmed records from the Bay of Biscay. REMARKS. Distinguished from the medusa stage of Clytia hemisphaerica by: larger diameter (up to 40 mm, not up to 20 mm), flat (not hemispherical) shape, more numerous tentacles [up to c. 200, not up to (rarely) 58], usually rather larger gonads and one (not 1-3, usually 2) statocysts between tentacles. The hydroid is not known. Disuse of the genus name Phialidium is discussed above (p. 74). Kramp (1961) referred the genus Staurostoma Haeckel, 1879:130, to Staurophora Brandt, in the family Laodiceidae. Clytia linearis (Thornely, 1 899) (Fig. 12) Obelia linearis Thornely, 1899 : 453, pi. 44, fig. 6. Campanularia gravieri Billard, 1904a : 482, fig. 1; Billard, 1907 : 171-172. ?Clytia geniculata Thornely, 1904 : 1 12-1 13, pi. 3, figs 4, 4a. Campanularia ?obHqua Clarke, 1907 : 9, pi. 5, figs 1—4. Clytia linearis: Stechow, 1913 : 66-69, figs 23-25; Hirohito, 1977 : 14-20, fig. 4a-j (syn. Campanularia gravieri Billard; Clytia hendersonae Torrey, 1904; C. alternata Hargitt; Laomedea bistriata Leloup). Clytia alternata Hargitt, 1924 : 483, pi. 2, fig. 7. tClytia (?)foxi Billard, 1926 : 93-94, fig. 9A-B. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 85 Laomedea (Obelia) bistriata Leloup, 193 la : 4, figs 8-1 1 . Clytia gravieri: Billard, 1938 : 429-432, figs 1-3, ?fig. 4 (syn. C. alternata Hargitt, Laomedea bistriata Leloup); Picard, 1955 : 185-186; Millard & Bouillon, 1973 : 51-54, fig. 7e-g (syn. Obelia striata Clarke, 1907; C. serrata Millard, 1958); Millard, 1975 : 215-217, fig. 71 F-H (syn. Obelia striata Clarke, 1907); Garcia Corrales, Inchaurbe & Mora, 1978 : 29-30, fig. 12. Clytia obliqua: Picard, 1950 : 51-52. Campanularia (Clytia) gravieri: Vervoort, 1967:50-52, fig. 16 (syn. Clytia alternata Hargitt; Laomedea bistriata Leloup). TYPE LOCALITY. Blanche Bay, New Britain, Bismarck Archipelago. Material not located. TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia obliqua Clarke, 1907, infertile colonies on sertularian hydroid, in spirit, 'Perico Island', Gulf of Panamar, coll. r.v. 'Albatross'; Smithsonian Institution cat. no. 29616. Clytia alternata Hargitt, 1924, fertile colonies preserved on microslide, Port Galero, Mindoro, Philippines; Smithsonian Institution cat. no. 42644 (Fig. 12). OTHER MATERIAL EXAMINED. None. DESCRIPTION OF HYDROID STAGE (partly after Thornely, 1899; Billard, 1904> _£- 6 c ta o 2 c/) o •a S3 3 "O ^ to £* "O O o-S 8 8 S w t S 6 i- c _^<2T3 0 0 1 I II C "^ M-^ O O 2i c 0 0 ^ tt2 «- c JS •- included. M O 'S 1 45 b 8 "1 1 c «« o5>,o^uiS £ C ~ * "5 ^ o ."g 1 ll!| CO J? u 03 to a> o o . i |B LL O *-• * f» r>.a V CO (75 O U UJ CX i/j «/•> UJ o -J tr c/) E 1 nllS ~ CO U • «• CO ^^ - 18 W < CO "2 o o SO o ro UN Os 53 13 •j2 10 0 § -a -a •o-o •^otrno^-o rsT^ C^l , O (N <^ O — OS c fli ^^ ftj M rf^ ,.f^ f^ «^ C Os <""] Os Os — w C •o" "" 5 S •- •- t/3 t/n V3 *-T - ~ *-T t/5 — — 1 0 2 Sill 1 £ IS '-J a ex ex •§•§ | o §" 0-3 a ex c ^ o ^ ex "2 §• ^3 o "H "H-- c •<-> CO O &o "^ C C C C 3 cD D D D X > J > D 03 -J 03 03 H < • TT (N ^, CO SO so 2 "" — — • «>• o- T3 C CO r^4 O "o c^ C c . 3 •**, PJ **-> 1 IS rr, o CO o ,0 13 {2 >, 3 X) C 6 >, U w- O C .t?C/3 3 C ^ a | ^ 01 •a -a .Ji — o aj O ^ »^ Q M Jj 3 CO CO (/5 • 22 > T3 2 C/5 "™ ^ _O — -^ 'o "ob U "o U = 'sj|i 2 (-.<-• e- W CU U 'C CO 03 03 H < S^E so" O >> C ^ r* r^ ^^3?.^^Z|| (U (/5 c ^ -c 9 I c/5 uoi H u z 00 u» N.E. ATLANTIC CAMPANULARIID HYDROZOANS 1 1 7 HABITAT. Substrates not recorded in my \915a paper include an isopod, Anilocra physodes (Linnaeus), from Naples (Stechow, 19216, 1925, as Obelia bifurca) and a commercial shrimp, Pennaeus aztecus Ives, from Florida (Kruse, 1959; via A. A. Fincham, pers. comm.); ships hulls, hermit crabs gastropod shells and algae (Millard, 1975). O. bidentata is highly tolerant of brackish water. It has been recorded at salinities of 18'6%o (Netherlands; Vervoort, 19460), 10-20%o (Italy; Morri, 19796) and even so low as l%o (S Carolina, U.S.A.; Calder, 1976). The species has often been recorded in shallow water and Millard (1978) listed it as intertidal. REMARKS. A detailed, world-wide treatment has appeared recently (Cornelius, \915a) and the notes given here are mainly intended to supplement that account. O. bidentata is a nearly cosmopolitan species, and a possible explanation is given above (pp. 44-45). Differences from the superficially similar Hartlaubella gelatinosa are given under that species, and from the hydroid stages of other Obelia species by Cornelius (\915a : table 1). Mayer (1910), Stechow (1925) and recently Stepanyants (1979) regarded Obelia austro- georgiae Jaderholm, 1904#, as conspecific. I do not disagree (cf. Cornelius, 1975a) but have no fresh evidence. However, I cannot yet agree with Stepanyants that Clytia paulensis Vanhoffen, 1910, is identical with O. bidentata. The young medusae of both are sufficiently well described to indicate that two species are involved (see notes on Dispersive stage under the two species, pp. 89, 113). It could be argued that since Vanhoffen's material was infertile it was merely young O. bidentata: but this was not Stepanyants' argument. I believe neither Stepanyants nor I have seen the type material of C. paulensis. If it did prove to be young O. bidentata then a new name would have to be found for C. paulensis auct. (see synonymy in this paper). In that event it would be useful to apeal to the International Commission on Zoological Nomenclature to validate the widely used name paulensis sens, auct. Clytia arborescens Pictet, 1 893 : 34-35, pi. 2, figs 30-3 1 , originally based on material from Indonesia, was reported south of Madeira at 100 m by Billard (1907); but Stechow (1925) included no further records in his synoptic list of warm water W Atlantic hydroids. Billard's material lacked gonothecae but, like the type material, had a compound stem and long hydrotheca (560-700 /zm) with sharply pointed cusps. It is likely that his material was O. bidentata; and it is relevant that he had wrongly identified O. bidentata from the English Channel (see Distribution, above). O. bidentata was not well known at the time he published the arborescens record. The type material of arborescens was probably not O. bidentata, however, since Pictet reported hemispherical medusa buds in the gonotheca, indicating Clytia sp. Obelia dichotoma (Linnaeus, 1758) (For illustrations see Cornelius, 1975a) Sertularia dichotoma Linnaeus, 1758 : 812. ICymodocea simplex Lamouroux, 1816 : 2 1 6, pi. 7, fig. 2 (see p. 121). ITubularia clytioides Lamouroux, in Freycinet, 1824 : 620, pi. 95, figs 6-8. Lomedea gracilis C. Pickering, in Dana, 1846 : 689, -pi. 61, figs 7, 7a-b (lapsus pro Laomedea; see p. 78). Obelia dichotoma: Hincks, 1868: 156-157, pi. 28; Cornelius, 1975a : 265-272, figs 3-4. Obelia plicata Hincks, 1868 : 159, pi. 30, figs 1, la. Campanularia cheloniae Allman, 1 888 : 22, pi. 11, figs 2, 2a. Obelia dischotoma Billard, 1927 : 332 (lapsus pro dichotoma). lObelia commensuralis: Gudger, 1937: 1-6 (?lapsus pro 0. commissuralisMcCrady, 1857). Obelia hyaliana Vannucci, 1955 : 56. Laomedea plicata: Rasmussen, 1973 : 30, fig. 9. Eulaomedajlexuosa: Rees & Thursfield, 1965 : 102-103 (lapsus pro Eulaomedea\ redet.; see p. 105). For further synonymy see Remarks, Table 4 and Cornelius ( 1 975a). 118 P. F. S. CORNELIUS DISPERSIVE STAGE. Medusae of O. dichotoma and O. geniculata reared from the hydroids in the laboratory still cannot be distinguished and some taxonomic confusion continues (Russell, 1953; Cornelius, 1975a; Arai & Brinckmann-Voss, 1980; see also Remarks). Russell's book includes the best account of the biology of Obelia spp. medusae. The alternation of generations characteristic of medusoid coelenterates was actually first appreciated in scyphozoans, in 1829. It was reported in Obelia as early as 1836, first among the hydrozoans; but several still earlier workers had come close to its discovery in this genus. This and other historical aspects have been reviewed recently (Winsor, 1976; Cornelius, \911a, b). A paper by Desor (1849) was missed by these reviewers. Desor provided an early confirmation of medusa release in Obelia, probably O. dichotoma. However, like several contemporary workers (see p. 97) he assigned his material to Hartlaubella gelatinosa which has no medusa. He probably identified his material using the well illustrated work of van Beneden (1844), who made the same error. Several other workers have assigned an Obelia-\ike medusa to H. gelatinosa and, despite the early clarification by Couch (1844), there was confusion for many years between O. dichotoma, O. geniculata, H. gelatinosa and Gonothyraea loveni (see notes under H. gelatinosa (p. 97), and Cornelius, 1977a). Couch (1844) was in addition the first to note medusa release in Obelia geniculata (p. 120). Other early records in that species were contributed by F. W. L. Thomas (in Johnston, 1847: 467), Hincks (1852 : 85)andGosse(1853 : 84-90, pi. 4). Illustrations of the medusae of 0. dichotoma were first provided by Baster (1762, pi. 5, fig. 7), but the earliest of 0. geniculata appeared nearly a century later, in Gosse's (1853) book. Naumov (1969, as O. longissima, here referred to O. dichotoma) reported that 'the medusae were apparently not liberated from' Black Sea populations. But the species of Obelia are exceptionally well documented and since there is no other report of medusa retention in the genus a repeat of the observations would be desirable. REPRODUCTIVE SEASON. Medusae of Obelia spp. have been reported around the British Isles in all months but are commonest from 'spring to late autumn' (Russell, 1953). Fertile colonies were recorded on the coast of Egypt between 6 Sep and 4 Oct, 1933 (Billard, 1936). DISTRIBUTION. Found throughout European and African coastal waters with the exception of the Black Sea, from which there are no confirmed records (but see last paragraph of Dispersive stage). The species is remarkable in being nearly cosmopolitan in coastal waters and is one of the most widely distributed of hydroids. Aspects of this are discussed on page 45; and a detailed summary of locality records has appeared elsewhere (Cornelius, 1975a). HABITAT. Tolerance of 12%o salinity recorded by Calder (1976) in S Carolina. Unusual substrates noted in the literature included a sponge and the fins of a shark (Couch, 1 844 : XV, as 'the sea thread of Ellis'), a turtle (Allman, 1 888, as Campanularia cheloniae), a crab (Rasmussen, 1973: the colony was thought to be feeding on debris in the exhalent current); on the parasitic copepod Lernaeocera on the gills of a cod (Gadus); and on the back and claws of an anomuran 'crab', Lithodes maia (in Evans, 1978). Probably the record by Gudger (1937), of 'Obelia commensuralis' on the blueback herring (Alosa aestivalis), referred to this species; but his account suggested that the hydrorhizae entered the muscles of the fish and the identification is not certain. REMARKS. A revision of this species has appeared recently (Cornelius, 1975a) and only additional notes are given here. Identification of the hydroid stage is discussed above (p. 1 13). Nematocysts were described by Ostman (1979) and by others whom she cited. Tubularia clytioides Lamouroux, in Freycinet, 1824, was based on live material collected from algae in the Azores. Probably it was Obelia dichotoma. Rees & White (1966) evidently thought Silicularia gracilis Meyen, 1834, also described from Azores material, to be conspecific but while the clytioides type specimen was an erect colony that of gracilis was reptant. Rees & White presented a muddled synonymy, referring to the 'species' (clytioides + gracilis) as Orthopyxis clytioides (Lamouroux); and including in its synonymy N.E. ATLANTIC CAMPANULARIID HYDROZOANS 1 19 the entry "Silicularia clytioides Meyen, 1834', evidently a lapsus for "S. gracilis Meyen'. Nutting (1915) had made the same error fifty years earlier (p. 50). The once widely recognized nominal species O. commissuralis McCrady, 1857, based on the hydroid stage, was referred to O. dichotoma first by Bohm (1878), and again by me (19750). Bohm's paper has been overlooked; and I know of no other works giving this synonymy. Obelia plicata Hincks, 1868, described from infertile hydroid material, type locality 'Shetland', was recently referred to the present species without comment (Cornelius, 1975a). I have not been able to locate type material. The species deserves close evaluation as it has been recognized by several authors (Marktanner-Turneretscher, 1890; Nutting, 1915, as ?O. plicata; Billard, 1927; Kramp, 1935, as Laomedea plicata; Fraser, 1944; Teissier, 1965; Rasmussen, 1973) although not by Broch (19 126). O. plicata was said by Hincks to resemble H. gelatinosa in having compound stems but the hydrothecal rims were even, not castellated. The hydrotheca was said by Hincks 'very much [to] resemble in form that of O. dichotoma". Gonothecae were first assigned to O. plicata by Marktanner-Turneretscher (1890) who said they were the same as those of O. dichotoma. Indeed, the two species seem conspecific. Some older colonies of O. dichotoma (e.g. BMNH 1948.9.8.14, Plymouth, 21 Apr 1899, coll. E. T. Browne) exhibit compound stems resulting from overgrowth of up to 10 young, pale stems along the older, thicker and darker ones. The specimen mentioned resembles closely the colony illustrated by Hincks as O. plicta, and synonymy seems justified. Godeaux (1941, as H. gelatinosa) described similar O. dichotoma material, from the North Sea; while Billard (1927) maintained O. plicata distinct on the basis of the original characters. Broch (19126) alone raised doubts, but was non-committal. The factors causing overgrowth in occasional O. dichotoma specimens are not known, most colonies being solitary. However, such overgrowth has been described in other hydroids and called 'auto-epizoism' (Millard, 1973). If an overgrown specimen results from second and subsequent planulae settling on the original colony then the whole should be regarded as an aggregation of colonies, and not as a single colony. The phenomenon is discussed above (p. 40). Campanularia cheloniae Allman, 1888, was based on infertile material collected during the 'Challenged expedition from the back of a turtle; but the locality of the turtle was not recorded. The original description and what remains of the type series (BMNH 1 888. 1 1.13.15, small colony in spirit + microslide preparation) are probably O. dichotoma. Laomedea sargassi Broch, 1913, was recorded from Ghana by Buchanan (1957 : 360). The name sargassi was a nom. nov. for O. hyalina Clarke, 1879, introduced just to prevent confusion with Gonothyraea hyalina auct., and is therefore inadmissible under the current conventions of nomenclature. Congdon (1907 : 468) described material from Bermuda as O. hyalina, but Hargitt (1909) gave that material specific status, as Laomedea congdoni. I have previously (Cornelius, 1975a) referred O. hyalina Clarke and O. congdoni (Hargitt) to O. dichotoma, but overlooked the nom. nov. sargassi. Recent examination of the type material of O. congdoni (U.S. nat. mus. cat. no. 42647, microslide preparation of infertile fragments) confirms that it too is simply O. dichotoma. Obelia hyaliana Vannucci, ? in ms, was a name given to some "O. hyalina' material which Vannucci (1955) later considered conspecific. Obelia geniculata (Linnaeus, 1758) (For illustrations see Cornelius, 19750) Sertularia geniculata Linnaeus, 1758 : 812. Obelia geniculata: Hincks, 1868: 149-151, pi. 25, fig. 1, la; Cornelius, 1 9750:272-278, figs 1,5. For further synonymy see Cornelius ( 1 91 5a), Remarks and Table 4. DISPERSIVE STAGE. A medusa. See Remarks and notes under Obelia dichotoma. REPRODUCTIVE SEASON. Russell (1953) recorded the following in British waters: Obelia sp. medusae found nearly throughout year, especially 'spring to late autumn' (Russell); medusae 120 P. F. S. CORNELIUS released at Plymouth, March-November (Orton, 1920); some evidence of lunar periodicity in release times (Elmhirst, 1925). In the Mediterranean, Lo Bianco (1909) recorded medusa release at Naples March-June and October-January. DISTRIBUTION. An almost cosmopolitan shallow water species found nearly throughout the eastern North Altantic. The species was not recorded from the Black Sea by Naumov (1969). He listed the species as absent from the Baltic as well but Stechow (1927) reported it from Stockholm. It was recorded from Iceland and the Faeroes but not from Spitzbergen by Kramp (1929), although already known from Jan Meyen Island (Broch, 1918). Calder (1970) among others recorded the species from W Greenland; but I (Cornelius, 19750) misquoted him as including E Greenland in its distribution. I know of no records from E Greenland. HABITAT [see also Geographical distribution (p. 45), the habitat notes under O. dichotoma and Cornelius, 19750]. The species was collected from the fins of a spur-dog (Squalus acanthias) by Couch (1844), who deduced that planula settlement must be rapid. Other actively swimming animals on which O. geniculata has been recorded include a sea-horse (Hippocampus guttulatus, by Zirpolo, 1939, 1940) and parasitic copepods themselves on fish (Lernaeocera spp. on Gadus (cod), by Leloup, 193 1/?; on Lernaeocem on Clupea (herring), by Debouteville & Nunes, 1951, 1952; on Dinemoura on Cetorhinus (basking shark) by Debouteville & Nunes). Long distance travel on floating algae has also been recorded (Arnaud, Arnaud, Intes & Le Loeuff, 1976). Recorded at 100 m depth by Naumov (1960). However, some of these records on fish might have been of 0. dichotoma. REMARKS. A revision of this species has appeared elsewhere (Cornelius, 19750) and the present account simply adds to that. Couch (1844) gave some interesting details. He stated the number of tentacles of the hydranth to vary widely (1 1-29 on p. 2, 14-28 on p. 38). He described medusa bud develop- ment but not medusa release, which was not known till 1847 in this species (Cornelius, 19750, 1977a, b; see also p. 1 18). He recorded a range of tissue colour in the hydroid from 'white to deep red'. The nematocysts were described by Ostman (1979) and by others whom she cited. Campanularia vermicularis Van Beneden, 1866, was described without illustrations and identification has been thought impossible (Vervoort, 19460; Leloup, 1947). The species was based on Belgian material. Van Beneden kept live colonies for three weeks. He described the species as being more robust than O. geniculata, and as having larger hydrothecae and more numerous annulations. It was collected off the brown alga, Fucus vesiculosus L. The hydranth was said to have as many as 40^4 tentacles, a character which might one day help to identify his material (cf. Couch's observations, immediately above). Meanwhile I follow Vervoort in treating the species here but not including it in the synonymy. Obelia spp. indet. 1. Hydroids. Obelia longicyatha Allman (1877 : 10, pi. 7, figs 4-5) was recorded from 35° 42' N, 8° 40' W (Cap Spartel, Tangier) and the Gulf of Cadiz by Billard (1907 : 168, as Clytia longicyatha), but the identity of the species remains unclear. I have discussed it elsewhere (Cornelius, 19750:264). See also the note on Campanularia vermicularis Van Beneden, 1866 (immediately above). 2. Medusae. Some species from the eastern North Atlantic cannot be assigned to a hydroid. Obelia adriatica Neppi (1912 : 726-727, pi. 3, fig. 8, 8a) was described from Adriatic material; and Obeliopsis fabredomergui Le Danois (1913:110) was based on material from the Little Minch, NW Scotland. It seems most unlikely that either is valid. O. plana Sars (1835 : 28, pi. 5, fig. 13, as Thaumantias) was referred to O. jlabellata hydroid by Bedot (1910 : 484) and some later authors, and hence would fall in the synonymy of O. dichotoma. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 121 But there seems slim reason for this identification (cf. Cornelius, 19750:271, footnote). Other pre-1910 medusa names in this genus were listed by Bedot (1901-1925) and Mayer (1910), and are not repeated here. As explained above (p. 1 18) most cannot be confidently assigned to a hydroid stage and a new list would serve little purpose. Taxa now referred to other families A few genera and species recorded from the eastern North Atlantic are no longer referred to the Campanulariidae. As several have been included in this family in some standard works notes on them are given here to summarize the current opinions. The taxa are treated in date order. 1. The genus Capsularia Cuvier, 1797, was used by Gray (1848:85-87) to embrace several British species of Campanulariidae but should be regarded a junior synonym of Coryne Gaertner, in Pallas, 1774 (discussion in Cornelius, 1 9756 : 378). 2. Sert ularia fruticosa Esper, 71810 [pi. 34, figs 1-2 (see note 2, p. 124); also Hammer, in Esper, 1829:162-163 (syn. Sertularia laxa Lamarck, 1816:116; Laomedea sauvagii Lamouroux, 1816: 206; both nom. nov. pro S. fruticosa)] was recorded from the Adriatic Sea by Marktanner-Turneretscher (1890 : 205) as Campanularia fruticosa. The species is now referred to the genus Thyroscyphus in the family Sertulariidae (e.g. Vervoort, 1967; Millard, 1975). It is otherwise unrecorded from the eastern North Atlantic and Marktanner- Turneretscher's record may well be wrong. However, T.fruticosus occurs in the Red Sea as far north as the Gulf of Suez (Vervoort, 1 967). The exact date of publication of the species name is unusually difficult to determine and the necessary bibliographic work on Esper's important multi-part book has apparently not been done. Precise dating would be useful as the plates were issued several at a time around 1810, and they carry binominals. They thus predate the posthumous text, edited by Hammer (Esper, 1 829), from which this part of the work is often thought to date. The plates predate also several important European works on zoophytes [e.g. those by Lamarck (1816) and Lamouroux (1812, 1816, 1821; Lamouroux et al., 1824)]. Esper's work is rare, and few of the original wrappers survive; but the information on an incomplete set of wrappers in the BMNH library gives hope that dating would be simple if a complete copy were located. See also section 7, below. 3. The genus Cymodocea Lamouroux, 1816:214, originally included two species, C. simplex and C. ramosa, both described as new. C. simplex was based on material from Great Yarmouth and 'Ireland', C. ramosa on fragments from the Antilles. Johnston (1838) commented on the genus and on the identities of the two species; and Gray (1848) dismissed them as 'doubtful species' at the end of the Campanulariidae. Lamouroux (1821) and Lamouroux et al. (1824) included additional species in the genus but these do not enter the present discussion. Billard (1909), who saw some of Lamouroux' original material, referred the two species to Nemertesia antennina (Linnaeus, 1758), family Plumulariidae. I designate C. ramosa type species of the genus Cymodocea, the name of which hence becomes a junior subjective synonym of Nemertesia Lamouroux, 1812. Billard was confi- dent that the C. ramosa material he saw was type, but had doubts about that of C. simplex. His doubts are significant since in J. Fleming's opinion, quoted by Johnston (1838), the original illustrations of C. simplex might be identified as an Obelia dichotoma colony lacking hydrothecae. I concur with Fleming's opinion; and disagree with Johnston who thought simplex was a plumularid. If the illustration is actually of O. dichotoma then Billard's caution was justified and the specimen he saw was not type. Almost all the Lamouroux collections were destroyed during the liberation of Caen on 7 July, 1944, so the C. simplex material is no longer available (Redier, 1967). However, C. simplex is here provisionally referred to O. dichotoma (p. 1 1 7). 122 P. F. S. CORNELIUS The species Cymodocea comata Lamouroux, 1821, was recorded from the coast of Devon by W. E. Leach (in Johnston, 1838); but the species was referred to Nemertesia ramosa (Lamarck, 1 8 1 6) by Billard ( 1 909), who saw type material. 4. Campanularia intertexta Couch, 1844 : 41^2, pi. 11, fig. 3, was based on material then in the Museum of the Royal Institution of Cornwall, Truro. The material almost certainly no longer exists (Curator, Roy. Inst. Cornwall, pers. comm.). Johnston (1847:109-110) repeated Couch's description verbatim but regarded the species as unsoundly based. Gray (1848 : 88) listed it without comment, incidentally assigning to it Shetland material of Lafoea dumosa (Fleming, 1820), BMNH reg. no. 1847.9.24.66. At the same time Gray proposed the new genus Conchella (?lapsus pro Couchella), of which Campanularia intertexta is type species by monotypy. Hincks (1868:220) thought the species might be L. dumosa, stating that Couch's description was inadequate for identification. Later Bedot (1905 : 157) referred intertexta to 'Coppinia arcta\ an invalid species accepted by several nineteenth century authors but merely based on the coppinia or reproductive branch of L. dumosa. Couch's illustration shows an unidentifiable campanu- larid hydroid, possibly Orthopyxis Integra, growing on a coppinia of L. dumosa. Thus the type 'series' was mixed. The name intertexta is here restricted solely to the illustrated coppinia, which becomes lectotype. Hence Campanularia intertexta is to be regarded a junior synoym of L. dumosa; and the genus name Conchella a junior synonym of Lafoea Lamouroux, 1821. See also page 65. 5. Campanularia lacerata Johnston, 1847:111, pi. 28, fig. 3, is now known as Opercularella lacerata and referred to the Campanulinidae (e.g. Millard, 1975). The species has been referred to a variety of genera in the literature, including Capsularia, Laomedea, Wrightia and Calycella (references in Hincks, 1868). 6. Laomedea obliqua Johnston, 1847 : 106-107, pi. 28, fig. 1, based on British material, is today known as Monotheca obliqua and Plumularia obliqua by different authors and referred to the Plumulariidae (e.g. Millard, 1975). 7. Campanularia fruticosa Sars, 1850 : 138-139, is today referred to Lafoea dumosa (Fleming, 1820), in the Lafoeidae (e.g. Cornelius, 19756). It has often been given full specific status, as L. fruticosa. See also section 2, above. 8. Campanularia abietina Sars, 1850 : 139, based on Norwegian material, has long been known as Grammaria abietina and assigned to the family Lafoeidae (e.g. Cornelius, 19756). 9. Campanularia parvula Hincks, 1853 : 178, pi. 5a, was probably based on material of Calycella syringa (Linnaeus, 1767) lacking operculae. Several authors have referred the species to Lafoea Lamouroux, 1821, but this seems wrong (references and discussion in Cornelius, 19756). 10. Campanularia gracillima Alder, 1856a:361, pi. 14, figs 5-6, based on hydroid material from NE England, has frequently been referred to Lafoea, family Lafoeidae; but lately to the species Lafoea dumosa (Fleming, 1820) (e.g. Cornelius, 19756; Cornelius & Garfath, 1980). 11. Laomedea acuminata Alder, 18566 : 441, pi. 16, figs 5-8, based on hydroid material from NE England, is the hydroid of an Aequorea sp. medusa, family Aequoreidae. The two species recognized from British waters from the medusa stage were described earlier so the name acuminata should be regarded provisional. Although the medusa generation of the two can be separated the respective hydroids are morphologically identical, so far as is known. Hence at present it cannot be decided to which of the 'medusa species' acuminata should rightly be referred (Russell, 1953). 12. The nominal species Laomedea tenuis Allman, 1859 : 367-368, was once known as Leptoscyphus tenuis (e.g. Hincks, 1868). It is currently regarded as a campanulinid, having N.E. ATLANTIC CAMPANULARIID HYDROZOANS 123 been provisionally referred to Phialella quadrata (Forbes, 1848) by Stechow (1923a : 129) and Rees (1939:441). Browne (1896:479) summarized the mistaken observations by Allman who assigned the medusa of one species to the hydroid of another. Campanulina tennis Van Beneden, 1886 : 174, 176, pi. 13, is a different nominal species, provisionally referred to Aequorea vitrina Gosse, 1853, by Russell (1953). Bedot (1910) listed an earlier homonym of C. tenuis. 13. Campanularia fastigiata Alder, 1860 : 73-74, pi. 5, fig. 1, based on Scottish material, was known for some decades as Stegopoma fastigiatum. Edwards (1973) showed that fastigiatum was the hydroid stage of the medusa Modeeria rotunda (Quoy & Gaimard, 1827), the binominal of which takes priority. The species is assigned to the family Laodiceidae (e.g. Rees & Rowe, 1969; Edwards, 1973; see also Cornelius & Garfath, 1980). 14. Campanularia humilis Hincks, 1866 : 298, is now known as Cuspidella humilis. Its affinities are unclear but it was removed from the Campanulariidae long ago (discussion in Cornelius & Garfath, 1 980). 15. Clytia eucophora Haeckel, 1879 : 168, was a combination applied to the hydroid stage of Eucopium primordiale Haeckel, 1879 : 168 (sic), from Corsica. As Mayer (1910 : 236) pointed out, the two names are objective synonyms with the same date of publication. Mayer, as first reviser, suppressed eucophora in favour of primodiale\ but at the same time referred Haeckel's species to Eucope picta Keferstein & Ehlers, 1861. Although Kramp (1961) omitted to treat E. picta there is no doubt that it and the Haeckel species should be referred to the Phialellidae. Weismann (1883 : 158) introduced the combination Clytia eucopophora, a lapsus of Haeckel's spelling. 16. Campanularia mutabilis Ritchie, 1907 : 504, pi. 23, figs 3-5, based on Azores material, is now known as Scandia mutabilis and referred to the Lafoeidae (e.g. Millard, 1975). 17. Campanularia divisa was attributed by Bassindale (1941 : 148) to Todd (1906 : 137) whom he supposed to have reported material from Ilfracombe. Bassindale misread Todd's list, which included Campanularia species and Tubularia indivisa. The word 'indivisa' was split between the two lines (in/divisa) and evidently read wrongly by Bassindale. 18. The genus Hincksella Billard, 1918 : 22, was considered by Totton (1930) and Ralph (1957) to be closely related to the Campanulariidae, but is now referred to the Syntheciidae (e.g. by Millard, 1975). 19. The genus Billardia Totton, 1930 : 150 (type species B. novae zealandiae, by original designation) was based on southern hemisphere material. Totton assigned Billardia to the Campanulariidae but the genus is similar to Hincksella and like it comes within the scope of the Syntheciidae sensu Millard (1975), in my opinion. Blanco (\961b) and Stepanyants (1979), however, retained Billardia in the Campanulariidae. Possibly a greater under- standing of the reproductive structures will help. Totton suspected that the blastostyles of Billardia were produced in place of hydranths, within the hydrothecae, but said his material was inadequately preserved for him to be certain. If his suspicion were confirmed the Syntheciidae would be the correct family for Billardia. Notes 1 (See p. 65). G. D. Westendorp (1813-1868) apparently produced only two works involving coelenterates (Westendorp, 1843, 1853). Both were on the zoophytes of the Belgian coast. The first was a straightforward taxonomic account including some new genera and species, among them Clytia ryckholtii (here referred to Orthopyxis integrd). Probably none is valid. His second work, published in 1853, was remarkable in being illustrated by dried specimens mounted on sheets. It is rare but still important since one of 124 P. F.S.CORNELIUS the included species was described as new in the 1843 work, and the specimens used in illustration might be considered types. The 1853 work comprised a printed title page, a printed page giving a list of species and localities, and 32 hebarium sheets each bearing one species. There were 16 hydroids, 14 bryozoans and 2 sponges. Examples of the work were seen by Neviani (1903), Bedot (1910 : 200-201) and Billard (1914; also seen by Leloup, 1947 : 5); and another, imperfect copy has recently been found in the British Museum (Natural History) (Cornelius, unpublished ms in BMNH). I have not seen an intact copy. Neviani listed the 32 species but C. ryckholtii was not among them. The copy described by Billard (1914), in Brussels (Mus. roy. Hist, nat.; inv. no. 3440), similarly does not have C. ryckholtii. The BMNH example also does not include that species. The copy evidently came to the (then) British Museum library about 1867. Certainly a collection of specimens corresponding with Neviani's list was purchased from Westendorp at that time, and was given the 32 zoological accession numbers 1867.5.4.22-24 and 1867.5.7.1-29. The specimens were curated each under its own species, and the printed species list was cut up to provide labels which were in most cases glued to the herbarium sheets. Much, perhaps all, of this material survives and is still curated under the various species; but I have not found the title page of this copy. Only one type specimen of any group is included in this collection, that of the hydroid, Plumularia macleodi Westendorp, 1843 (BMNH reg. no. 1867.5.7.7). It can be considered a syntype of P. macleodi; and is in fact a specimen of Kirchenpaueria pinnata (Linnaeus, 1758). P. macleodi is thus a junior synonym of K. pinnata. Billard (1914), working on the corresponding Brussels syntype specimen, reached the same conclusion. 2 (See p. 121). Some evidence was presented by Sherborn (1922 : 1, entry under Esper; Sherborn, 1926 : 2528, entries under fruticosa, Laomedea and fruticosa, Sertularia) that at least some of the plates of Esper's (1829) 'Die Pflanzen-Thiere' appeared before 1816, possibly around 1810. Since the plates bear binominals, and might predate the synoptic works of Lamouroux (1812 onwards) and Lamarck (1816 onwards), correct dating is important. However, it seems that bibliographers have not yet dated the plates and Sherborn's (1926) provisional date of 1810 for the plate of Sertularia fruticosa Esper is the best compromise. Certainly the plate of fruticosa predated the works of Lamarck ( 1 8 1 6) and Lamouroux (1816), which referred to Esper's plate as being already published. Indeed, they proposed new names in place of fruticosa. The three volume Esper work was issued in parts, some after Esper's death in 1810; and the few copies I have seen have been collated with the undoubtedly earlier Sertularia plates intermingled with the text. But the partial set of wrappers with the BMNH copy gives much detailed information, and a full set might enable all the plates of this scarce work to be dated accurately. 3 (See p. 1 12). Dating of the two works here listed as Peron & Lesueur, 1810a, 718106, has caused confusion. A clarification is desirable since in them many genus names of medusae were introduced which are still used. The works comprised the earliest serious systematic treatment of medusae, and these authors finally disbanded the genus Medusa Linnaeus, 1758. The first of the two works was one of a series of journal papers written by the two authors, and comprised their taxonomic treatment of medusae. Several other papers in the series also dealt with marine animals but are not important here. They have been listed by Goy (1980). The second work (Peron & Lesueur, 718106), a book entitled Histoire generale des meduses, brought the series of papers together each forming a chapter of the book. The book version was repaginated from 1 on, and was presumably published after the series of papers (although it might conceivably have been issued in parts as the papers appeared). The publication date of the taxonomic paper (Peron & Lesueur, 1810#) has been taken as 1809 by most authors and this date might be inferred from the title page of the volume of the journal in which it appeared. But there is good evidence that it did not appear until January, 1810 (Sherborn, 1914; 1929 : 4455, entry under Obelia\ Cornelius, 1975a; Goy, 1980). The book, the Histoire generale des meduses, has been almost entirely overlooked. It was dated N.E. ATLANTIC CAMPANULARIID HYDROZOANS 125 1809 in the British museum catalogue of printed books. But J. Goy (pers. comm.) and I concur that the book version almost certainly would have followed the series of papers, and so would have appeared at least as late as 1810 (unless the book version were issued in parts). Hence the many names introduced by Peron & Lesueur almost certainly date from the 1810^ paper, published January, 1810. Apart from minor heading changes the book version (71810&) was apparently printed from the same 'blocks' as the series of papers. For taxonomic purposes the two versions differ only in date. The numerous plates that were to have accompanied the account of the medusae (Peron & Lesueur, 1810#) were prepared but not published immediately (cf. Cornelius, \915a : 253, footnote; 1977a:49, footnote). But most of the illustrations showing Peron & Lesueur's 'new' species were brought together in an extremely rare work usually attributed to Lesueur alone (Peron & Lesueur, 71815). This work was cited by Haeckel (1879, in several of his synonymies, as the Planches inedites) and Totton (1965 : p. 45, pi. 8) among others but almost all twentieth century medusa workers have overlooked it. Fewer than half a dozen copies are known (Totton, 1965; Goy, 1980). Goy (1980) cited evidence from the P.-v. Seanc. Acad. Sci. Paris, 1795-1831 (issue covering 14 August 1815) 5 : 532, that 1815 is the correct date; and not 71811 as given in the British museum catalogue of printed books (187 : 1 1 1) under Peron alone. She concluded that the bulk of the illustrations, showing most of the nominal species newly described by Peron & Lesueur (1810(2), were copied and published by a variety of contemporary compilers in their own works and so made public. They included such famous names as de Blainville, Cuvier, Milne Edwards, Lamarck and Lesson (references in Goy, 1980). The original Lesueur drawings are preserved in the Museum at Le Havre. Although the bulk of the rare work (Peron & Lesueur, 71815) comprised illustrations of medusae exquisitely drawn by Lesueur, the title page leaves no doubt that the authorship should be ascribed jointly to Peron & Lesueur (Mrs A. Datta, pers. comm.). Peron had died in 1810 and it can be inferred that Lesueur wished him still to be senior author, as he had been of the lengthy text (Peron & Lesueur, 1 8 1 Oa, 7 1 8 1 06) of the work. Acknowledgements I am indebted to Professor W. Vervoort, Rijksmuseum van Natuurlijke Historic, Leiden, for generous and detailed discussion on the Campanulariidae; and through him to the Stichting Jan Joost ter Pelkwijk Fonds for financial assistance while on a visit both to the Leiden Museum and the Institut voor Taxonomische Zoologie, Zoologisch Museum, Universiteit van Amsterdam, where Dr R. W. M. van Soest was my kind host. Dr J. M. Paul, Zoology Department, University of Oxford, helped trace literature about Laomedea neglecta. I am grateful also to Dr R. G. Hughes, Westfield College, University of London, for new records of Clytia paulensis and to Dr R. B. Williams of Tring for helpful discussions. Dr A. G. Long, Hancock Museum, Newcastle upon Tyne, kindly provided facilities while I worked on the Joshua Alder collection. Lastly I am grateful to Miss L. J. Wadmore for preparing the illustrations of medusae and to R. H. Harris for histological assitance. Other acknowledge- ments are made in the text. Addendum On 15 August 1981 I found Laomedea angulata (p. 98) common on the eel grass bed at Studland, Dorset. The previous record from the British mainland was dated 1906; and from Dorset was 1890, also at Studland. Whether or not the species had disappeared from that locality in the meantime might be hard to tell. On 15 September 1981 I found the same species abundant on eel grass near Misery Point, R Yealm, near Plymouth. At this locality too the status of the species during the last several decades was unclear. 126 P. F. S.CORNELIUS References Agassiz, L. 1862. Hydroidae. Contributions to the natural history of the United States of America. Second Monograph, 4 : 1 8 1-380. Boston. Alcock, A. 1892. A case of commensalism between a gymnoblastic anthomedusoid (Stylactis minoi) and a scorpaenoid fish (Minous inermis). Ann. Mag. nat. Hist. (6) 10 : 207-2 14. Alder, J. 1856a. A notice of some new genera and species of British hydroid zoophytes. Ann. Mag. nat. Hist. (2) 18 : 353-362. 1 8566. Descriptions of three new British zoophytes. Ann. Mag. nat. Hist. (2)18: 439-44 1 . 1857. A catalogue of the zoophytes of Northumberland and Durham. Trans. Tyneside Nat. Fid Cl. 3 : 93-162. (Although dated 1856 on the wrapper, a footnote on page 87 of a previous paper is dated 29 January, 1857.) 1860. Descriptions of a zoophyte and two species of Echinodermata new to Britain. Ann. Mag. nat. Hist. (3)5: 73-75. Alexander, W. B. 1932. The natural history of the Firth of Tay. Trans. Proc. Perthsh. Soc. nat. Sci. 9 : 35-42. Allman, G. J. 1859a. Notes on the hydroid zoophytes. Ann. Mag. nat. Hist. (3)4 : 137-144. 18596. Notes on the hydroid zoophytes. Ann. Mag. nat. Hist. (3)4 : 367-370. 1864a. On the construction and limitation of genera among the Hydroida. Ann. Mag. nat. Hist. (3) 13 : 345-380. 18646. Report on the present state of our knowledge of the reproductive system in the Hydroida. Rep. Br. Ass. Advmt Sci. (1863)(1) : 351-426. 1871. A monograph of the gymnoblastic or tubularian hydroids. I. The Hydroida in general. London. 1876a. Descriptions of some new species of Hydroida from Kerguelen's Island. Ann. Mag. nat. Hist. (4) 17: 113-115. 1 8766. Diagnoses of new genera and species of Hydroida. J. Linn. Soc. (Zool.) 12:25 1-284. 1887. Report on the Hydroida collected during the exploration of the Gulf Stream by L. F. de Pourtales, Assistant, United States Coast Survey. Mem. Mus. comp. Zool. Harv. 5 (2) : 1-66. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years 1873-1876. Part II. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae, and Thalamophora. Rept. sclent. Results Voy. Challenger '23 : l-90 + i-lxix. Annandale, N. 1915. Fauna of the Chilka Lake. The coelenterates of the lake, with an account of the Actiniaria of brackish water in the Gangetic delta. Mem. Indian Mus. 5 : 65-1 14. Aria, M. N. & Brinckmann-Voss, A. 1980. Hydromedusae of British Columbia and Puget Sound. Can. Bull. Fish, aquat. Sci. 204 : 1-192, i-viii. Arnaud, F., Arnaud, P. M., Intes, A. & Le Loeuff, P. 1976. Transport d'invertebres benthiques entre PAfrique du Sud et Sainte Helene par les laminaires (Phaeophyceae). Bull. Mus. Hist. nat. Paris (3) 384 (Ecol. gen. 30) : 49-55. Babic, K. 1912. Dimorphismus der Gonangien bei Laomedea angulata Hincks. Zool. Anz. 39 : 457-460. Bale, W. M. 1888. On some new and rare Hydroida in the Australian Museum collection. Proc. Linn. Soc. N.S.W. (2) 3: 745-749. 1914. Further notes on Australian hydroids. III. Proc. R. Soc. Viet. (N.S.) 27 : 72-93. Bassindale, R. 1941. Studies on the biology of the Bristol Channel. IV. The invertebrate fauna of the southern shores of the Bristol Channel and Severn Estuary. Proc. Bristol Nat. Soc. (4) 9 : 143-201 . Baster, J. 1762. Opuscula subseciva, observationes miscellaneas de animalculis et plantis. Volume I. Haarlem, Holland. Bedot, M. 1901. Materiaux pour servir a 1'histoire des hydroi'des. Ire periode. Revue suisse Zool. 9:379-515. 1905. Materiaux pour servir a Phistoire des hydroi'des. 2me periode (1821 a 1850). Revue suisse Zool. 13: 1-183. 1910. Materiaux pour servir a Phistoire des hydroi'des. 3me periode (1851 a 1871). Revue suisse Zool. 18: 189-490. 1912. Materiaux pour servir a Phistoire des hydroi'des. 4me periode (1872 a 1880). Revue suisse Zool. 20:213-469. 1916. Materiaux pour servir a Phistoire des hydroi'des. 5me periode (1881 a 1890). Revue suisse Zool. 24: 1-349. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 127 — 1918. Materiaux pour servir a 1'histoire des hydroi'des. 6me periode (1819 a 1900). Revue suisse Zool. 26 (suppl.) : 1-376. 1925. Materiaux pour servir a 1'histoire des hydroi'des. 7me periode (1901 a 1910). Revue suisse Zool. 32 (suppl.) : 1-657. Behner, A. 1914. Beitragzur Kenntnisder Hydromedusen. Z. wiss. Zool. Ill : 381-427. Beloussov, L. V. 1961. Vital observations on cell displacements in the hydroid polyp Obelia flexuosa. Dokl. Akad. Nauk. SSSR. 136 : 1490-1 493. 1973. Growth and morphogenesis of some marine Hydrozoa according to histological data and time-lapse studies. Publs Seto. mar. biol. Lab. 20 : 3 1 5-366. Beneden, P.-J. van 1843. Memoire sur les campanulaires de la cote d'Ostende, considered sous le rapport physiologique, embryogenique et zoologique. (Extrait). Annls Sci. nat. (Zool.) (2) 20 : 350-369. 1844. Memoire sur les campanulaires de la cote d'Ostende, consideres sous le rapport physiologique, embryogenique et zoologique. Nouv. Mem. Acad. R. Sci. Bruxelles, 17 (5) : 1-42. 1866. Recherches sur la faune littorale de Belgique (Polypes). Mem. Acad. r. Sci. Lett. Belg. 36(2): 1-207. Bergh, R. S. 1 879. Studien iiber die erste Entwicklung des Eies von Gonothvraea loveni (Allm.). Morph. Jb. 5: 22-61. Berrill, N. J. 1949. The polymorphic transformations of Obelia. Q. Jl microsc. Sci. 90 : 235-264. 1950. Growth and form in calyptoblastic hydroids. II. Polymorphism within the Campanu- lariidae. /. Morph. 87 : 1-26. Betencourt, A. 1888. Les hydraires du Pas-de-Calais. Bull, scient. Fr. Belg. 19 : 201-2 14. Billard, A. 1902. Les hydroi'des de la Bai de la Hougue. Bull. Mus. Hist. nat. Paris 8 : 53 1-536. 1904a. Hydroi'des recoltes par M. Ch. Gravier dans le Golfe de Tadjourah. Bull. Mus. Hist. nat. Paris 10: 480-485. 1904/7. Contribution a 1'etude des hydroi'des. (Multiplication, regeneration, greffes, variations). Annls Sci. nat. (Zool.) (8) 20 : 1-25 1 . 1906. Mission des pecheries de la cote occidentale d'Afrique. III. Hydroi'des. Act. Soc. linn. Bordeaux, 56 : 69-76. 1907. Hydroi'des. Exped. scient. Travailleur- Talisman 8 : 153-243. 1909. Revision des especes types d'hydro'i'des di la collection Lamouroux conservee a 1'Institut Botanique de Caen. Annls Sci. nat. (Zool.) (9) 9 : 307-336. 1912. Hydroi'des de Roscoff. Archs Zool. exp. gen. 51 : 459-478. 1914. Note sur les hydroi'des de la collection Westendorp du Musee Royal d'histoire naturelle de Belgique. Ann. Soc. r. zool. malac. Belg. 48 : 27-30. 1917. Note sur quelques especes d'hydro'i'des libres. Bull. Mus. Hist. nat. Paris, 23 : 539-546. 1918. Notes sur quelques especes d'hydro'i'des de 1'expedition du 'Siboga'. Archs zool. exp. gen. 51 (notes et revue 2) : 21-27. 1922. Note sur deux especes d'hydro'i'des du littoral d'Ostende. Ann. Soc. r. zool. malac. Belg. 52: 135-139. 1926. Rapport sur les Hydroi'des [of the Cambridge Expedition to the Suez Canal]. Trans, zool. Soc.Lond. 22:85-104. 1927. Les hydroi'des de la cote Atlantique de France. C. r. Congr. Socs sav. Paris. Sect. Sci. (1926): 326-346. 1928. Clytia johnstoni Alder, Campanularia raridentata Alder et Thaumantias inconspicua Forbes. Bull. Mus. nat. Hist. nat. Paris 34 : 456^157. 193 la. Hydroi'des de Mauritanie. Bull. Mus. nat. Hist. nat. Paris (2) 3 : 673-678. 193 \b. Hydroi'des des cotes de Syrie (Missions Gruvel). Bibltque Faune. Colon.fr. 3 : 389-395. 1934. Note sur quelques hydroi'des du Maroc. Bull. Soc. zool. France 59 : 227-231. 1936. Les fonds de peche pres d'Alexandrie. Hydroidea. Notes Mem. Dir. Rech. Pech., Le Caire 13: 1-11. 1938. Note sure une espece de campanularides (Clytia gravieri Billard). Bull. Mus. Hist. nat. Paris (2) 10: 429-432. Blanco, O. M. 1976a. Estudio critico sobre las especies del genro "Silicularia" Meyen. 1834. Revta Mus. La Plata, N.S. (Zool.) 9:21 7-241 . 1967^. Contribucion al conocimiento de los hydrozoarios Argentines. Revta Mus. La Plata, N.S. (Zool.) 9: 243-297. Bohm, R. 1878. Helgolander Leptomedusen. Jena Z. Naturw. 12 : 68-203. Botten-Hansen, P. & Petersen, S. 1870. Norsk Bog-Fortegnelse. 1848-1865. Oslo. 128 P. F. S. CORNELIUS Bougis, P. 1963. Cours d'oceanographie biologique. Fascicule 1. Hydromeduses. Caracteres generaux. classification. Villefranche. Brattegard, T. 1966. The natural history of the Hardangerfjord. 7. Horizontal distribution of the fauna of rocky shores. Sarsia, 22 : 1-54. Breemen, P. J. Van 1905. Plankton van Noord- en Zuiderzee. Tijdschr. ned. dierk. Vereen. (2) 9 : 145-324. Brinckmann-Voss, A. 1970. Anthomedusae/Athecatae (Hydrozoa, Cnidaria) of the Mediterranean. Part I. Capitata. Fauna Flora Golfo Napoli, 39 : 1-96. Broch, H. 1905. Nordsee-Hydroiden von dem norwegischen Fischereidampfer 'Michael Sars' in den Jahren 1903-1904 gesammelt, nebst Bemerkungen iiber die Systematik der Tecaphoren Hydroiden. Bergens Mm. Arb. ( 1 905) (6) : 1-26. 1910. Die Hydroiden der arktischen Meere. Fauna arct. 5 : 127-248. 1912a. Coelenteres du fond. In D uc d 'Orleans Campagne Arctique de 1907. Brussels, pp. 1-29. \9\2b. Hydroidenuntersuchungen III. Vergleichende Studien an adriatischen Hydroiden. K. norske Vidensk. Selsk. Skr. (1911)(l):l-65. 1913. Hydroida from the "Michael Sars" North Atlantic deep-sea expedition 1910. Rep. scient. Results Michael Sars N. Atlant. deep sea Exped. 3(1): 1-18. 1918. Hydroida (Part II). Dan. Ingolf Exped. 5 (7) : 1-206. 1928. Hydrozoa I. Tierwelt N.- u. Ostsee, 3 (b) . 1-100. 1929. Hydroidenuntersuchungen VI. Studien an Silicularia Meyen. Meddr zool. Mus., Oslo (1929)20: 1-30. 1933. Zur Kenntnis der adriatischen Hydroidenfauna von Split. Skr. norske VidenskAkad. mat.-nat.Kl.(\933)4: 1-115. Brooks, W. K. 1888«. On a new method of multiplication in hydroids. Johns Hopk. Univ. Circ. 7 : 29-30. 18886. The life history of Epenthesis mccradvi (n.sp.). Stud. biol. Lab. Johns Hopkins Univ. 4: 147-162. Browne, E. T. 1896. On British hydroids and medusae. Proc. zool. Soc. Lond. (1896) : 459-500. 1897. On British medusae. Proc. zool. Soc. Lond. (1897) : 816-835. 1900. The fauna and flora of Valencia harbour on the west coast of Ireland. Part I. The pelgaic fauna. II. Report on the medusae (1 895-98). Proc. R. Ir. Acad. (3) 5 : 694-736. 1904. Hydromedusae with a revision of the Williadae and Petasidae. Fauna Geogr. Maldive Laccadive Archipel. 2 : 722-749. Browne, E. T. & Vallentin, R. 1904. On the marine fauna of the Isles of Scilly J. R. Instn Cornwall, 16: 120-132. Bruce, J. R. 1948. Additions to faunal records, 1941^6. Rep. mar. Biol. Stn Port Erin (1945^7). 58-60 : 39-58. Bruce, J. R., Colman, J. S. & Jones, N. S. 1963. Marine fauna of the Isle of Man. L.M.B.C. Mem. typ. Br. mar. PL Anim. 36 : i-ix + lr-307. Buchanan, J. B. 1957. The hydroid fauna of the Gold Coast. Revue Zool. Bot. afr. 56 : 349-372. Calder, D. R. 1970. Thecate hydroids from the shelf waters of northern Canada. J. Fish. Res. Bd, Canada 27 : 1501-1547. 1975. Biotic census of Cape Cod Bay : hydroids. Biol. Bull. mar. biol. Lab., Woods Hole, 149:287-315. 1976. The zonation of hydroids along salinity gradients in South Carolina estuaries. In Mackie, G. O. (editor), Coelenterate ecology and behavior (Third international symposium on coelenterate biology), New York. Pp. 165-174. Calkins, G. N. 1 899. Some hydroids from Puget Sound. Proc. Boston Soc. nat. Hist. 28 : 333-367. Carus, J. V. 1884-1893. Prodromus faunae mediterraneae. Stuttgart. 2 volumes. (Vol. 1, pars I, Colenterata, Echinodermata, Vermes, 1884; pp. 1-282.) Chas Brinquez, J. C. & Rodriguez Babio, C. 1977. Fauna marina de Galicia. Contribucion al conocimiento de los hidropolipos del literal gallego. Monogr. Univ. Santiago de Compostela, 39: 1^3. Chopin, A. 1894. A visit toCumbrae: The veteran naturalist: The Millport Biological station. Trans, a. Rep. Manchr micr. Soc. (1 894) : 43-54. Christiansen, B. O. 1972. The hydroid fauna of the Oslo Fiord in Norway. Norw. J. Zool. 20 : 279-3 10. Chumley, J. 1918. The fauna of the Clyde Sea area, being an attempt to record the zoological results obtained b y the late Sir John Murra v and his assistants on board the S. Y. ' 'Medusa ' ' during the rears 1884 to 1892. Glasgow. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 129 Clarke, S. F. 1875. Descriptions of new and rare species of hydroids from the New England Coast. Trans. Conn. Acad. Arts Sci. 3 : 58-66. 1876. The hydroids of the Pacific coast of the United States, south of Vancouver Island. With a report upon those in the Museum of Yale College. Trans. Conn. Acad. Arts Sci. 3 : 249-264. 1877. Report on the hydroids collected on the coast of Alaska and the Aleutian Islands, by W. H. Dall, U.S. Coast Survey, and party, from 1871 to 1874 inclusive. Proc. Acad. nat. Sci. Philad. (1876) 28 : 209-238. (A note on p. ii of the volume states unequivocally that the publication date of this paper was 2 January, 1877.) 1879. Report on the Hydroida collected during the exploration of the Gulf Stream and Gulf of Mexico by Alexander Agassiz, 1877-78. Bull. Mus. comp. Zool. Harv. 5 : 239-252. 1907. Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer 'Albatross', from October, 1904, to March, 1905, Lieut. -Commander L. M. Garrett, U.S.N., commanding. VIII. The hydroids. Mem. Mus. comp. Zool. Harv. 35 (1) : 1-18. Congdon, E. D. 1907. The hydroids of Bermuda. Proc. Am. Acad. Arts Sci. 42 : 461^85. Cornelius, P. F. S. 1975a. The hydroid species ofObelia (Coelenterata, Hydrozoa: Campanulariidae), with notes on the medusa stage. Bull. Br. Mus. nat. Hist. (Zool.) 28 : 249-293. 19756. A revision of the species of Lafoeidae and Haleciidae (Coelenterata: Hydroida) recorded from Britain and nearby seas. Bull. Br. Mus. nat. hist. (Zool.) 28 : 373^26. 1977 a. The linking of polyp and medusa stages in Obelia and other coelenterates. Biol. J. Linn. Soc. 9 : 45-57. 1977 b. The gradual discovery of the relation between medusa and polyp in Obelia and other coelenterates. Microscopy 53 '. 172-174. — 1978. The genus names Calicella Hincks and Calvcella Hincks (Coelenterata: Hydrozoa). Bull. Br. Mus. nat. Hist. (Zool.) 33 : 233-234. — 1979. A revision of the species of Sertulariidae (Coelenterata: Hydroida) recorded from Britain and nearby seas. Bull. Br. Mus. nat. Hist. (Zool.) 34 : 243-32 1 . 1981. Clytia Lamouroux, 1812, Laomedea Lamouroux, 1812, and Campanularia Lamarck, 1816 (Coelenterata, Hydroida): proposed designations of type species by use of the plenary powers, and comments on related genera. Z.N.(S.) 2326. Bull. zool. Norn. 38 : 208-220. — (unpublished). Notes on the BMNH copy of a rare work by Westendorp, 1853. (MS in BMNH library.) & Garfath, J. B. 1980. The coelenterate taxa of Joshua Alder. Bull. Br. Mus. nat. Hist. (Zool.) 39:273-291. Couch, R. Q. 1842. An essay on the zoophytes of Cornwall. Rep. R. Cornwall polvtech. Soc. (1840:27-91. 1844. A Cornish fauna; being a compendium of the natural history of the county. Part 3, containing the zoophytes and calcareous corallines. Truro. Coughtrey, M. 1875. Notes on the New Zealand Hydroideae. Trans. Proc. N.Z. Inst. (1874) 7:281-293. Crawford, J. H. 1895. The hydroids of St. Andrews Bay. Ann. Mag. nat. Hist. (6) 16 : 256-262. Crawshay, L. R. 1912. On the fauna of the outer western area of the English Channel. J. mar. biol. Ass. U.K. 9 : 292-393. Crothers, J. H. (Ed.). 1966. Dale Fort marine fauna. Second edition. Fid Std. 2 (suppl.) : i-xviv + 1-169. Crowell, S. 1957. Differential responses of growth zones to nutritive level, age, and temperature in the colonial hydroid Campanularia. J. exp. Biol. 134 : 63-90. 1961. Developmental problems in Campanularia. In Lenhoff, H. M. & Loomis, W. F. (editors) The biology of Hydra and of some other coelenterates: 1961 Miami, pp. 297-3 16. & Wyttenbach, C. R. 1957. Factors affecting terminal growth in the hydroid Campanularia. Biol. Bull. mar. biol. Lab., Woods Hole 113 : 233-244. Cunha, A. X. da 1944. Hidropolipos das costas de Portugal. Mems Estud. Mus. zool. Univ. Coimbra 161 : 1-101. 1950. Nova contribuicjio para o estudo dos hidropolipos das costas de Portugal. Archos Mus. Bocage,2\ : 121-144. Cuvier, G. L. C. F. D. 1797. Tableau elementaire de I'histoire naturelle des animaux. Paris. (Dating follows J. Typogr. Biblphique(\191) 1 : 81-82.) Dana, J. D. 1846, 1849. United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 130 P. F.S.CORNELIUS 1842. Under the command of Charles Wilkes, U.S.N. Zoophytes. Philadelphia. Vol. 1, text (1846); vol. 2, atlas ( 1 849). (Dating after Haskell, 1 942.) Darwin, C. R. 1 860. Journal of researches into the natural history and geology of the countries visited during the voyage of H. M.S. Beagle round the world, under the command ofCapt. Fitz Roy, R.N. 2nd edition. London. Debouteville, C. D. & Nunes, L. P. 1951. Hydraires epizoi'ques sur les copepodes parasites. Vie Milieu 2:42\-432. 1952. Hydraires epizoi'ques sur les copepodes parasites. Archos Mus. Bocage 23: 1-11. Deevey, E. S. 1954. Hydroids of the Gulf of Mexico. In Galtsoff, P. S., Gulf of Mexico. Its origin, waters, and marine life. Fishery Bull. Fish. Wildl. Serv. U.S. 55 : 267-272. den Hartog, C. 1970. The sea-grasses of the world. Verh. K. ned. Akad. Wet. 59 : 1-275. Desor, M. E. 1849. Lettre sur la generation medusipare des polypes hydraires. Annls Sci. nat. (3) 12:204-217. Du Plessis, G. 1871. Evolution medusipare de Clvtia (Campanularia) volubilis. Bull. Soc. vaud. Sci. nat.ll : 167-170. Edwards, C. 1973. The medusa Modeeria rotunda and its hydroid Stegopoma fastigiatum, with a review of Stegopoma and Stegolaria. J. mar. biol. Ass. U.K. 53 : 573-600. Ehrenberg, C. G. 1834#. Beitra'ge zur physiologischen Kenntniss der Corallenthiere im allgemeinen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben. Phys. Math. Abh. K. Akad. Wiss. Berlin (1832) 1 : 225-380. (Reprinted as Ehrenberg, 18346.) 18346. Die Corallenthiere des rothen Meeres, physiologisch untersucht und systematisch verzeichnet. Berlin. (Reprint of Ehrenberg, 1 843a, in book form, paginated 1-1 56.) Ellis, J. 1 755. An essay towards a natural history of the corallines, and other marine productions of the like kind, commonly found on the coasts of Great Britain and Ireland. To which is added the description of a large marine polype taken near the North Pole by the whale-fishers, in the summer 1753. London. 1756a. 'Account of the vesicles on corallines.' [Running title at top of page.] Gentleman's Mag. 26 : 288-290. 17566. Essai sur I'histoire naturelle des corallines, et d'autres productions marines du meme genre, qu'on trouve communement sur les cotes de la Grande- Bretagne et d'Irlande; auquel on a joint une description d'un grand polype de mer, pris aupres du Pole Arctique, par des pecheurs de baleine, pendant I'ete de 1 753. The Hague. Translated by P. de Hondt. 1756c. Natuurlyke historic van de koraal-gewassen, en andere dergelyke zee-lighamen, die men gemeenelyk vind op de kusten van Groot-Brittanien en lerland: benevens eene beschryving van een grooten zee-polyp, in den zomer van 't jaar 1753 by den Noord-Pool door de walvis-vangers gevonden. The Hague. Translated by J. Tak. — 1767. Versuch einer Natur-Geschichte der Corall-Arten und anderer dergleichen Mer-Korper, welche gemeiniglich an den Kusten von Gro/3-Britanien und Inland gefunden werden; nebst der Beschreibung eines grossen Buschel-Polypen, welcher in dem Eis-Mere gefangen worden. Aus dem Englischen und Franzosischen ubersezt, und mil Anmerkungen, auch einem Anhange fiinf hieher gehoriger Abhandlungen der Herren Schlosser, Baster und Ellis begleitet von D. J. G. Kriiniz. Niirnberg. Translated by D. J. G. Kriiniz. — 1 768. An account of the Actinia sociata, or clustered animal-flower, lately found on the sea-coasts of new-ceded Islands. Phil. Trans. R. Soc. (1 767) 57 : 428^37. & Solander, D. C. 1 786. The natural history of many curious and uncommon zoophytes, collected from various parts of the globe. London. Edited by M. Watt. Elmhirst, R. 1925. Lunar periodicity in Obelia. Nature, Lond. 116 : 358-359. Eschscholtz, F. 1829. System der Acalephen. Berlin. Esper, E. J. C. 71810. (There is strong evidence that some of the plates of Esper (1829) were published in 1810, but I have not traced a perfect copy. See note 2 (p. 1 24) and next reference.) 1829. Die Pflanzen-Thiere. Volume 3, pp. 145-283. Edited by F. L. Hammer. Niirnberg. (See also previous reference.) Evans, F. O. 1978. The marine fauna of the Cullercoats district. 6. Coelenterata and Ctenophora. Rep. Dove mar. Lab. (3) 19 : 1-165. Faulkner, G. H. 1929. The early prophases of the first oocyte division as seen in life, in Obelia geniculata. Q. Jl microsc. Sci. 73 : 225-242. Faure, C. 1 965. Sur la distinction des deux especes de Laomedea. Cah. Biol. mar. 6:41 9-426. Fey, A. 1969. Peuplements sessiles de 1'archipel de Glenan. I. Inventaire : hydraires. Vie Milieu (B) 20:387-413. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 131 Fleming, J. 1 820. Observations on the natural history of the Sertularia gelatinosa of Pallas. Edinb. phil. J. 2 : 82-89. 1828. A history of British animals. Edinburgh. Forbes, E. 1848. A monograph of the British naked-eyed medusae: with figures of all the species. London. Fraipont, J. 1880. Recherches sur 1'organisation histologique et le developpement de la Campanularia angulata. Contribution a 1'histoire de 1'origine du testicule et de 1'ovaire. Archs Zoo/, exp. gen. 8 : 433-466. Fraser, C. M. 1913. Hydroids from Nova Scotia. Bull. Victoria meml Mus. 1 : 157-186. 1918. Hydroids of eastern Canada. Rep. Dept. Naval Serv. Ottawal (suppl.) : 329-367. 1 944. Hydroids of the Atlantic coast of North America. Toronto. 1946. Distribution and relationship in American hydroids. Toronto. Freycinet, L. de (editor) 1 824. Voyage autour du monde entrepris par ordre du Roi. Zoologie [by Quoy, J. R. C. & Gaimard, J. P.]. Paris. (The hydroid descriptions were written by J. V. F. Lamouroux.) Garcia Corralles, P., Inchaurbe, A. A. & Mora, D. G. 1978. Contribucion al conocimiento de los hidrozoos de las costas espanolas. Parte I : Halecidos, campanularidos y plumularidos. Boln Inst. esp. Oceanogr. 4 (253) : 3-73. Gegenbaur, C. 1854. Zur Lehre vom Generationswechsel und der Fortpflanzung bei Medusen und Polvpen. Wiirzburg. (Paginated 1-68; reprint of article with same title in Verh. phvs.-med. Ges. Wurzb.4: 154-221.) 1856. Versuch eines Systemes der Medusen, mil Beschreibung neuer oder wenig gekannter Formen; zugleich ein Beitrag zur Kenntniss der Fauna des Mittelmeeres. Z. wiss. Zoo/. (1857) 8 : 202-273. Giard, A. 1898. Sur 1'ethologie du Campanularia caliculata Hincks. (Stolonisation et allogonie). C. r. Seanc. Soc. Biol. ( 10) 5 : 1 7-20. Godeaux, J. 1941. Sur la croissance d'un hydrozoaire. C. r. Ass. franc.. Avanc. Sci. 63(1939) : 707-710. Goette, A. 1907. Vergleichende Entwicklungsgeschichte der Geschlechtsindividuen der Hydropolypen. Z. wiss. Zoo/. 87 : 1-335. Gosse, P. H. 1853. A naturalist's rambles on the Devonshire coast. London. 1855-1856. A manual of marine zoology for the British Isles. London. 2 volumes. (Volume 1, 'Zoophytes', 1855, is cited here.) Gow, C. & Millard, N. A. H. 1975. Two new species of campanularian hydroids from South Africa. Ann. S. Afr. Mus. 67 : 1-6. Goy, J. 1980. Lesmedusesde Francois Peron et Charles-Alexandre Lesueur( 1775-1 810 et 1778-1846) revelees par les velins de Lesueur. Bull, trimest. Soc. geol. Normandie Amis Mus. du Havre 67 (2): 65-78, pis 1-27. Gravier-Bonnet, N. 1979. Hydraires semi-profonds de Madagascar, (Coelenterata Hydrozoa), etude systematique et ecologique. Zoo/. Vehr. Leiden 169 : 1-76. Gray, J. E. 1848. List of the specimens of British animals in the collection of the British Museum. Part 1. Cenlroniae or radiated animals. London. Gronovius, L.-T. 1760. Observationes de animalculus aliquot marinae aquae innatantibus atque in littoribus Belgicis obviis. Acta helv. 4 : 35-40. [The author's name does not appear in the paper, which was ascribed to L.-T. Gronovius by Bedot ( 1 90 1 : 396).] Gudger, E. W. 1928. Association between sessile colonial hydroids and fish. Ann. Mag. nat. Hist. (10) 1 : 17-48. 1937. A glut herring, Pomolobus aestivalis, with an attached colonial hydroid, Obelia commensuralis. Am. Mus. Novit. 945 : 1-6. Haeckel, E. 1879. Das System der Medusen. Erster Theil einer Monographie der Medusen. Denkschr. med.-naturw. Ges. Jena, 1 : 1-360. Hale, L. J. 1964. Cell movements, cell division and growth in the hydroid Clvtia johnstoni. J. Embryol. exp.Morph. 12:517-538. 1973a. The pattern of growth of Clytia johnstoni. J. Embryol. exp. Morph. 29 : 283-309. 19736. Morphogenetic properties of the parts of the colony of Clvtia johnstoni. J. Embryol. exp. Morph. 30 : 773-789. Hammett, F. S. 1943. The role of the amino acid and nucleic acid components in developmental growth. Part one. The growth of an Obelia hydranth. Chapter one. Description of Obelia and its growth. Growth 7 : 33 1-339. Hamond, R. 1957. Notes on the Hydrozoa of the Norfolk coast. J. Linn. Soc. (Zool.) 43 : 294-324. 1 963. Further notes on the Hydrozoa of the Norfolk coast. Ann. Mag. nat. Hist. (13)6 : 659-670. 132 P. F. S. CORNELIUS & Williams, R. B. 1977. The Ctenophora, Scyphozoa and Anthozoa of Norfolk, with additional notes on the Hydrozoa. Trans. Norfolk Norwich Nat. Soc. 24 : 58-74. Hargitt, C. W. 1909. New and little known hydroids of Woods Hole. Biol. Bull. mar. biol. Lab., Woods Hole, 17 : 369-385. — 1924. Hydroids of the Philippine Islands. Philipp. J. Sci. 24 : 467-507. Hartlaub, C. 1897. Die Hydromedusen helgolands. Zweiter Bericht. Wiss. Meeresunters. Helgol. (N.F.) 2 (Heft 1): 449-536. 1901. Hydroiden aus dem Stillen Ocean. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896-97). Zoo/. Jb. (Syst.) 14 : 349-379. 1905. Die Hydroiden der magelhaensischen Region und chilenischen Kiiste. Zoo/. Jb. (suppl. 6): 497-7 14. Haskell, D. C. 1942. The United States Exploring Expedition, 1838-1842, and its publications 1844-1874. New York. Heath, H. 1910. The association of a fish with a hydroid. Biol. Bull. mar. biol. Lab., Woods Hole, 19 : 73-78. Heller, C. 1868. Die Zoophvten und Echinodermen des adriatischen Meeres. Vienna. Hickson, S. J. & Gravely, F. H. 1907. Hydroid zoophytes. Nat. Antarct. Exped. 1901-1904 Nat. Hist. 3 (Colenterata II) : 1-34. Hincks, T. 1852. Notes on the reproduction of the Campanulariadae; with a description of a new species ofLaomedea. Ann. Mag. nat. Hist. (2) 10 : 8 1-87. 1853. Further notes on British zoophytes, with descriptions of new species. Ann. Mag. nat. Hist. (2)11: 178-185. 1861fl. On new Australian Hydrozoa. Ann. Mag. nat. Hist. (3)7 : 279-281. 1861&-1862. A catalogue of the zoophytes of south Devon and south Cornwall. Ann. Mag. nat. Hist. (3) 8 (1861): 152-161,251-262, 290-297, 360-366; 9 (1862): 22-30. 1863. On some new British hydroids. Ann. Mag. nat. Hist. (3) 11 : 45-47 [plus pi. 9 from volume 10,1862]. 1866. On new British Hydroida. Ann. Mag. nat. Hist. (3) 18 : 269-299. 1868. A history of the British hydroid zoophytes. 2 volumes. London. 1871. Supplement to a "Catalogue of the zoophytes of south Devon and South Cornwall," with descriptions of new species. Ann. Mag. nat Hist. (4)8 : 73-83. 1872. Contributions to the history of the Hydroida. Ann. Mag. nat. Hist. (4) 10 : 385-395. 1889. On the Polyzoa and Hydroida of the Mergui Archipelago collected for the Trustees of the Indian Museum, Calcutta, by Dr J. Anderson, F.R.S., Superintendent of the Museum. J. Linn. Soc. 21 : 121-135. Hirohito. 1969. Some hydroids of the Amakusa Islands. Publ. Biol. Lab. Imp. Household, Tokyo (1969) (9): 1-32. 1977. Five hydroid species from the Gulf of Aqaba, Red Sea. Publ. Biol. Lab. Imp. Household, Tokyo (1977) (11): 1-26. Hoeven, J. van der, 1862. Eenige aanteekeningen over Martinus Slabber's Natuurkundige Verlustigingen; benevens opgave der systematische namen van de daarin afgebeelde diersoorten. Versl. Meded. K. Akad. wet. Amst. (Afd. Nat.) 14 : 270-285. Houttuyn, M. 1770. Natuurlyke Historic ofuitvoerige beschryving der Dieren, Planten en Mineraalen, volgens het Samenstel van den Heer Linnaeus. Eerste Deels, veertiende Stuk. Amsterdam. Houvenaghel-Crevecoeur, N. 1973. Sur le mode de fixation de la planule de H ydrall mania falcata L. (hydroi'de thecate, Sertulariidae). C. r. hebd. Seanc. Acad. Sci., Paris (D) 276 : 28 1 3-28 1 5. Hughes, R. G. 1975. The distribution of epizoites on the hydroid Nemertesia antennina (L.). J. mar. biol. Ass. U.K. 55 : 275-294. 1980. Current induced variations in the growth and morphology of hydroids. In Tardent, P. & Tardent, R. (eds) Developmental and cellular biology ofcoelenterates. Amsterdam; pp. 179-184. Hummelinck, P. W. 1930. Beitragezur Kenntnis hollandischer Hydroiden. I. Bemerkungen iibereinige Campanuliniden und Campanulariiden vom Vangdam und Nieuwediep. Tijdschr. ned. dierk. Vereen. (3)2(1) : 28^12. 1936. Hydropoliepen. Flora Fauna Zuiderzee (Suppl.) : 41-64. Jaderholm, E. 1904a. Mitteilungen ueber einige von der Schwedischen Antartic-Expedition 1901-1903 eingesammelte Hydroiden. ArchsZool. exp. gen. (4) 3 (Notes et revue) : I-XIV. 1904/?. Aussereuropaische Hydroiden im schwedischen Reichsmuseum. Ark. Zoo/. (1903) 1 : 259-3 12. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 133 1909. Northern and arctic invertebrates in the collection of the Swedish State Museum. IV. Hydroiden. K. svenska VetenskAkad. Handl. 45 (1) : 1-124. Jagerskiold, L. A. 1971. A survey of the marine bethonic macro-fauna along the Swedish west coast 1921-1938. Ada R. Soc. sclent, litt. gothoburg. (Zool.)6 : 1-146. Johnston, G. 1836. A catalogue of the zoophytes of Berwickshire. Hist. Berwicksh. Nat. Club, 1 : 107-108. (Dating follows a manuscript note by C. D. Sherborn in the BMNH copy, drawing attention to the inclusion of the annual Presidential Addresses in the journal. That in the part including pp. 107-108 was delivered in September, 1836; but there is no further proof that the part was issued before 1 837.) 1 838. A history of the British zoophytes. Dublin, Edinburgh & London. 1 847. A history of the British zoophytes. 2nd edition. 2 volumes. London. Kawaguti, S. 1966. Electron microscopy on the Hydrozoa, Orthopyx'^ caliculata. Biol. J. Okayama Univ. 12:93-103. Keferstein, W. & Ehlers, E. 1861. Zoologische Beitrdge gesammelt im Winter 1859/60 in Neapel und Messina. Leipzig. Keller, C. 1883. Untersuchungen iiber neue Medusen aus dem rothen Meere. Z. wiss. Zool. 38:621-670. Knight, D. P. 1965. Behavioural aspects of emergence in the hydroid of Campanularia jlexuosa (Hincks). Nature, Lond. 206 : 1 170-1 171. 1970. Sclerotization of the perisarc of the calyptoblastic hydroid, Laomedea jlexuosa. \. The identification and localization of dopamine in the hydroid. Tissue Cell 2 : 467-477. 1971. Sclerotization of the perisarc of the calyptoblastic hydroid, Laomedea Jlexuosa. 2. Histochemical demonstration of phenol oxidase and attempted demonstration of peroxidase. Tissue Cell 3: 57-64. Knight-Jones, E. W. & Jones, W. C. 1956. The fauna of rocks at various depths off Bardsey. Rep. Bardsey Birds Fid Obs. (1955)3: 23-30. Kramp, P. L. 1919. Medusae. Part I. Leptomedusae. Dan. IngolfExped. 5 (8) : 1-1 12. 1929. Marine Hydrozoa. Zoology Faroes, 1(1 : 5) : 1-59. 1933. Occasional notes on Coelenterata. II. Vidensk. Meddrdansk. naturh. Foren. 94 : 237-247. 1935. Polypdyr (Coelenterata). I. Ferskvandspolypper og Goplepolypper. Danm. Fauna, 41 : 1-208. 1938. Marine Hydrozoa. a. Hydroida. Zoology Iceland, 2 (5a) : 1-82. 1943. The zoology of east Greenland. Hydroida. Meddr. Gronland, 121 (11): 1-52. 1955. The medusae of the tropical west coast of Africa. Atlantide Rep. 5 : 239-324. 1957. Some Mediterranean hydromedusae collected by A. K. Totton in 1954 and 1956. Vidensk. Meddrdansk. naturh. Foren. 119 : 1 15-128. — 1959. The hydromedusae of the Atlantic Ocean and adjacent waters. Dana Rep. 46 : 1-283. — 196 1 . Synopsis of the medusae of the world. J. mar. biol. Ass. U.K. 40 : 1-469. 1965. The hydromedusae of the Pacific and Indian Oceans. [Section 1.] Dana Rep. 63 : 1-162. Kruse, D. N. 1959. Parasites of the commercial shrimps, Pennaeus aztecus Ives, P. duorarum Burkenroad and P. setiferus (Linnaeus). Tulane Stud. Zool. 7 : 123-144. Kubota, S. 1978. The life-history of Clytia edwardsi (Hydrozoa; Campanulariidae) in Hokkaido, Japan. J. Fac. Sci. Hokkaido Univ. (6) 21:31 7-354. Ktthl, H. 1967. Die Hydromedusen der Emsmiindung. Veroff. Inst. Meeresforsch. Bremerh. 10 : 239-246. Kuhn, A. 1913. Entwicklungsgeschichte und Verwandtschaftsbeziehungen der Hydrozen. I Teil : Die Hydroiden. Ergebn. Fortschr. Zool. 4(1-2) : 1-284. Lamarck, J. B. P. A. de 1 8 1 6. Histoire naturelle des animaux sans vertebres. Volume 2. Paris. 1836. Histoire naturelle des animaux sans vertebres. 2nd edition, edited by Deshayes, G. P. & Edwards, H. M. Volume 2, Histoire des polypes. Lamouroux, J. V. F. 1812. Extrait d'un memoire sur la classification des polypiers coralligenes non entierement pierreux. Nouv. Bull. Sci. Soc. philom. Paris, 3 : 181-188. 1816. Histoire des polypiers coralligenes flexibles, vulgairement nommes zoophytes. Caen. 1 82 1 . Exposition methodique des genres de I'ordre des polypiers. Paris. , Bory de Saint-Vincent, J. B. G. M. & Deslongchamps, E. 1824, Histoire naturelle des zoophytes, ou animaux rayonnes, faisant suite a I'histoire naturelle des vers, de Bruguiere. In: Encyclopedic methodique (suppl.). Paris. Lang, A. 1886. Gastroblasta rajfaelei. Eine durch eine Art unvollstandiger Theilung entstehende Medusen-Kolonie. Jena Z Naturw. 19 (N.P. 12) : 735-763. 134 P. F. S. CORNELIUS I .a rw ood. G. P. & Rosen, B. R. (editors) 1979. Biology and systematics of colonial organisms. Syst. Ass. spec. vol. 1 1 . Laverack, M. S. & Blackler, M. 1974. Fauna and flora of St. Andrews Bay. Edinburgh and London. Le Danois, E. 1913. Note sur trois nouvelles meduses et liste des coelenteres du plankton recueillis a bord du Pourquoi-Pas? dans sa croisiere dans les mers du Nord. Bull. Mus. Hist. Nat. Paris 19(1913): 110-113. Leloup, E. 1930a. Campanularia macrotheca nov. sp. Annls Soc. r. zool. Belg. (1929)60 : 101-102. 19306. Deux cas d'epibiose de 1'hydropolype Campanularia johnstoni Alder. Bull. Mus. roy. Hist. nat. Belg. 6 (\9): 1-5. 193 la. Trois nouvelles especes d'hydropolypes. Bull. Mus. r. Hist. nat. Belg. (2) 2 : 1-73. 19316. Un cas d'epibiose de 1'hydropolype, Laomedea geniculata (Linne). Bull. Mus. r. Hist. nat. Belg. 7(24): 1-3. 1933. Contribution a la connaissance des hydropolypes de la cote des Pays-Bas. Bull. Mus. r. Hist. nat. Belg. 9 (45): 1-30. 1935. Hydraires calyptoblastiques des Indes Occidentales. Mem. Mus. r. Hist. nat. Belg. (2) 2: 1-73. 1939. Hydropolypes marins et dulcicoles du Congo Beige. Revue Zool. Bot. afr. 32 : 418-423. 1940. Hydropolypes provenant des croisieres du Prince Albert ler de Monaco. Result. Camp. sclent. Prince Albert 1 104 : 1-38. 1947. Les coelenteres de la faune Beige. Leur bibliographic et leur distribution. Mem. Mus. r. Hist. nat. Belg. 107 : 1-73. 1952. Coelenteres. Faune Belg., pp. 1-283. Lendenfeld, R. von 1883a. Eine ephemere Eucopide. Zool. Anz. 6 : 186-189. 18836. Uber Coelenteraten der Siidsee. IV. Mittheilung. Eucopella Campanularia nov. gen. Z. wiss. Zool. 38 : 497-583. 1 885. Addenda to the Australian Hydromedusae. Proc. Linn. Soc. N.S. W. ( 1 884) (1)9: 908-924. Leslie, G. & Herdman, W. A. 1881. The invertebrate fauna of the Firth of Forth. Edinburgh. Lesson, R.-P. 1 843. Histoire naturelles des zoophytes. Paris. Leuckart, R. 1856. Beitrage zur Kenntniss der Medusenfauna von Nizza. Arch. Naturgesch. 22(1): 1-40. Levinsen, G. M. R. 1893. Meduser, Ctenophorer og Hydroider fra Grgmlands Vestkyst, tilligemed Bemaerkninger om Hydroidernes Systematik. Vidensk. Meddr dansk naturh. Foren. (1892) (5) 4: 143-212,215-220. Linko, A. K. 1911. Hydraires (Hydroidea). Volume I. Haleciidae, Lafoeidae, Bonneviellidae et Campanulariidae. Fauna Rossii, pp. 1-250. Linnaeus, C. 17 '58. Sy 'sterna naturae. 1 Oth edition. Holmiae. 1767. Systema naturae. 12th edition. Tom I, Pars II. Holmiae. Lloyd, R. 1907. Nudiclava monocanthi, the type of a new genus of hydroids parasitic on fish. Rec. Indian Mus. I : 28 1-294. Lo Bianco, S. 1909. Notizie biologiche riguardanti specialmente il periodo di maturita sessuale degli animali del golfo di Napoli. Mitt. zool. Stat. Neapel 19:51 3-763. Macartney, J. 1810. Observations upon luminous animals. Phil. Trans. R. Soc. 100 : 258-293. Macgillivray, J. 1842. Catalogue of the marine zoophytes of the neighbourhood of Aberdeen. Ann. Mag. nat. Hist. (1)9 : 462^69. Maitland, R. T. 1876. Determinatie der dieren, beschreven en afgebeeld in de Werken van Job Baster en Martinus Slabber. Tijdschr. ned. dierk. Vereen. 2 : 7-15. 1 897. Prodrome de la faune des Pays-Bas et de la Belgique fla mande. Leiden. Mammen, T. A. 1965. On a collection of hydroids from south India. II. Suborder Thecata (excluding family Plumulariidae). J. mar. biol. Ass. India 1 : 1-57. Manea, V. 1972. Contribujii la studiul hidroidelor (Coelenterata) din Marea Neagra. Studii Cere. Biol. (Zool.) 24: 409-4 17. Marine Biological Association, 1957. Plymouth marine fauna. 3rd edition. Plymouth, England. Marktanner-Turneretscher, G. 1890. Die Hydroiden des k. k. naturhistorischen Hofmuseums. Annln naturh. Mus. Wien, 5 : 195-286. Mathiesen, O. 1928. Hydroids from northern Norway, with a list of unreported specimens in the Tromsjar Museum. TromspMus. Aarsh. (1926)49 (4) : 1-40. Mayer, A. G. 1900. Some medusae from the Tortugas, Florida. Bull. Mus. comp. Zool. Harv. 37: 13-82. 1910. Medusae of the world. Volume II. The hydromedusae. Washington. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 1 35 McCrady, J. 1857. Gymnopthalmata of Charleston Harbor. Proc. Elliott Soc. nat. Hist. 1 : 103-221. Metschnikoff, E. 1886a. Medusologische Mittheilungen. Arbor, zool. Inst. Univ. Wiend : 237-266. 1 886/j. Embryologische Studien an Medusen. 2 vols, text & atlas. Vienna. Meyen, F. J. F. 1834. Uber das Leuchten des Meeres und Beschreibung einiger Polypen und anderer nieder Thiere. Nova Acta Acad. Caesar. Leop. Carol. 16 (suppl. 1) : 125-2 16. Millard, N..A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43 : 173-243. 1958. Hydrozoa from the coasts of Natal and Portugese East Africa. Part I. Calyptoblastea. Ann. S. Afr. Mus. 44: 165-226. 1966. The Hydrozoa of the south and west coasts of South Africa. Part III. The Gymnoblastea and small families of Calyptoblastea. Ann. S. Afr. Mus. 48 : 427-487. 1973. Auto-epizoism in South African hydroids. Publs Seto mar. biol. Lab. 20 : 23-34. 1975. Monograph of the Hydroida of southern Africa. Ann. S. Afr. Mus. 68 : 1-513. 1977. The South African Museum's Meiring Naude cruises. Part 3. Hydroida. Ann. S. Afr. Mus. 73: 105-131. 1978. The geographical distribution of southern african hydroids. Ann S. Afr. Mus. 74 : 1 59-200. & Bouillon, J. 1973. Hydroides from the Seychelles. Annls Mus. r. Afr. cent. (8°, Zool.) 206: 1-106. Miller, M. C. 1961. Distribution and food of the nudibranchiate mollusca of the south of the Isle of Man. J. anim. Ecol. 30 : 95-1 16. Miller, R. L. 1973. The role of the gonomedusa and gonangium in the sexual reproduction (fertilization) of the Hydrozoa. Publs Seto mar. biol. Lab. 20 : 367-400. Morri, C. \919a. Quelques observations sur les hydroides de la Lagune d'Orbetello. Rapp. P.-v. Reun. Commn int. Explor. scient. Mer Mediterr. 25-26 : 1 19-120. 1979/7. Contribute alia conoscenza degli idrozoi lagunari Italiani: idropolipi di alcuni laghi costieri Mediotirrenici. Annali Mus. civ. Stor. nat. Geneva (1978) 82 : 163-171. Mulder, J. F. & Trebilcock, R. E. 1914. Victorian Hydroida. With description of new species. Geelong Nat. 6: 6-15. Miiller, O. F. 1 776. Zoologiae Danicae prodromus, seu Animalium Daniae et Norvegiae indigenarum characteres, nomina et synonyma imprimis popularium. Pp. xxxii + 282. Havniae. Murdock, G. R. 1976. Hydroid skeletons and fluid flow. In Mackie, G. O. (editor), Coelenterate ecology and behaviour (Third international symposium on coelenterate biology), New York. Pp. 33-40. Naumov, D. V. 1960. Gidroidy i gidromeduzy morskikh, solonovatovodnykh i presnovodnykh basseinov S.S.S.R. Fauna S.S.S.R. 70 : 1-626. 1969. Hydroids and hydromedusae of the U.S.S.R. Fauna S.S.S.R. 70 : 1-660. Israel Program for Scientific Translations, cat. no. 5 108. & Stepanyants, S. D. 1972. Marine invertebrates from Adelie Land, collected by the Xllth and XVth French Antarctic Expeditions. 3.-Hydroida. Tethys(Supp\.)4 : 25-60. Neppi, V. 1912. Adriatische Hydromedusen. Sber. Akad. Wiss. Wien 121 (1) : 709-734. Neviani, A. 1903. Intorno ad una rara pubblicazione di G. D. Westendorp. Boll. Soc. zool. ital. (2) 4 : 96-97. Nishihira, M. 1968. Distribution pattern of Hydrozoa on the broad-leaved eelgrass and narrow-leaved eelgrass. Bull. mar. biol. Stn Asamushi 13 : 125-138. Nobre, A. 1931. Contribul'coes para o estudo dos coelenterados de Portugal. Porto. Norman, A. M. 1869. Shetland final dredging report. Part II. On the Crustacea, Tunicata, Polyzoa, Echinodermata, Actinozoa, Hydrozoa, and Porifera. Rep. Brit. Ass. Advmt Sci. (1868) (1) : 247-336. Nutting, C. C. 1899. Hydroida from Alaska and Puget Sound. Proc. U.S. natn. Mus. 21 : 741-753. 190 la. Papers from the Harriman Alaska Expedition. XXI. The hydroids. Proc. Wash. Acad. Sci. 3: 157-216. 1901/7. The hydroids of the Woods Hole region. Bull. U.S. Fish Commn (1899) : 325-386. 1915. American hydroids. Part III. The Campanularidae and the Bonneviellidae. Spec. Bull. U.S. natn. Mus., pp. 1-126. Oken, L. 1815. Okens Lehrbuch der Naturgeschichte. Dritte Theil. Zoologie. Jena. Orton, J. H. 1920. Sea-temperature, breeding and distribution in marine animals. J. mar. biol. Ass. • U.K. 12 : 339-366. Ostman, C. 1979. Two types of nematocysts in Campanulariidae (Cnidaria, Hydrozoa) studied by light and scanning electron microscopy. Zoologica Scr. 8 : 5-12. Pallas, P. S. 1 766. Elenchus zoophytorum. The Hague. 1774. Spicilegia zoologica. Tom. I, fasc. 10 : 1-42. Berlin. 136 P. F. S. CORNELIUS Parker, H. W. 1939. Turtles stranded on the British coast. Proc. Linn. Soc. Lond. 151 : 127-129. Patriti, G. 1970. Catalogue des cnidaires et ctenaires des cotes atlantiques marocaines. Rabat. Pennington, A. S. 1885. British zoophytes: an introduction to the Hydroida, Aclinozoa, and Polyzoa found in Great Britain, Ireland, and the Channel Islands. London. Peron, F. & Lesueur, C.-A. 1810a. Tableau des caracteres generiques et specifiques de toutes les especesde meduses connuesjusqu'ace jour. Annls Mus. Hist. nat. Paris (1809) 14 : 325-366. (Dating of this publication follows Sherborn, 1914.) 718106. Histoire generate des meduses. Paris. (Dating of this work is discussed in note 3, p 124). 71815. Voyage de decouvertes aux terres australes. Histoire naturelle. Histoire generale et particuliere de tous les animaux qui composent la famille des meduses, et de quelques autres radiaires mollasses. Paris. (Dating of this work is discussed in note 3, p. 124). Philbert, M. 1935a. Le phenomene de stolonisation chez trois especes d'hydraires fixees sur des posidoniesen Mediterranee. Bull. Inst. oceanogr. Monaco 663 : 1-8. 19356. Les hydraires de la region malouine. Bull. Inst. oceanogr. Monaco 673 : 1-36. Picard, J. 1949. Sur la presence en Mediterranee de Clytia noliformis (McCrady). Bull. Mus. Hist. nat. Marseille 9: 184-190. 1950. Observations sur les hydraires recoltes aux Martigues dans le canal de Carente. Vie Milieu 1:51-52. 1 95 1 a. Note sur les hydraires littoraux de Banyuls-sur-Mer. Vie Milieu 2 : 338-349. 195 16. Hydraires littoraux du Senegal recoltes par H. Sourie aux environs de Dakar. Bull. Inst.fr. Afr. noire 13: 109-115. 1955. Hydraires des environs de Castiglione (Algerie). Bull. Stat. Aquic. Peche Castiglione (NS) 7: 177-199. 1958a. Notes sur une collection d'hydroides provenant des cotes Mediterraneennes d'Israel. Bull. Sea Fish. Res. Stn Israel 15 : 1-3. 19586. Origines et affmites de la faune d'hydropolypes (gymnoblastes et calyptoblastes) et d'hydromeduses (anthomeduses et leptomeduses) de la Mediterranee. Rapp. P.-v. Reun. Commn int. Explor. scient. Mer Mediterr. (NS) 14 : 187-199. Pictet, C. 1 893. Etude sur les hydraires de la Baie d'Amboine. Revue suisse Zool. 1 : 1-64. Pieper, F. W. 1844. Erganzungen zu "Heller's Zoophyten etc. des adriatischen Meeres". Zool. Anz. 7: 148-152,164-169,185-188,216-221. Poche, F. 19 14. Das System der Coelenterata. Arch. Naturgesch. 80 (Abt. A, Heft 5) : 47-128. Prenant, M. & Bobin, G. 1956. Bryozoaires. Premiere partie. Entoproctes, phylactolemes, ctenostomes. Faune Fr. 60 : 1-398. Quoy, J. R. C. & Gaimard, J. P. 1827. Observations zoologiques faites a bord de V Astrolabe, en mai 1 826, dans le detroit de Gibraltar. Annls Sci. nat (Zool.) (1 ) 10:5-21,1 72-193, 225-239. Ralph, P. M. 1956. Variation in Obelia geniculata (Linnaeus, 1758) and Silicularia bilabiata (Coughtrey, 1875) (Hydroida, F. Campanulariidae). Trans. R. Soc. N.Z. 84 : 279-296. 1957. New Zealand thecate hydroids. Part I. Campanulariidae and Campanulinidae. Trans. R. Soc. N.Z. 84: 81 1-854. & Thomson, H. G. 1968. Seasonal changes in growth in the erect stem of Obelia geniculata in Wellington Harbour, New Zealand. Zool. Publs Victoria Univ. Wellington 44 : 1-21. Rankin, J. 1901. Hydroida. In Elliot, G. F. S., Laurie, M. & Murdoch, J. B. (editors). Fauna, flora & geology of the Clyde area; Glasgow [British Association for the Advancement of Science]; pp. 369-371. Rasmussen, E. 1973. Systematics and ecology in the Isefjord marine fauna. Ophelia, 11 : 1-495. Ray, J. 1 724. Synopsis methodica stirpium Britannicarum. Edn. 3. London. Redier, L. 1967. Revision de la collection du Museum des hydraires de Lamouroux. Bull. Mus. natn Hist. nat. Paris (2) 39 : 381^10. Rees, W. J. 1939. A revision of the genus Campanulina van Beneden, 1847. Ann. Mag. nat. Hist. (11) 3 : 433^47. & Rowe, M. 1969. Hydroids of the Swedish west coast. Ada. R. Soc. Scient. litt. gothoburg. (Zool.) 3 : 1-24. & Thursfield, S. 1965. The hydroid collections of James Ritchie. Proc. R. Soc. Edinb. (B) 69 : 34-220. — & White, E. 1966. New records and fauna list of hydroids from the Azores. Ann. Mag. nat. Hist. (13)9:271-284. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 137 Riedl, R. 1959. Die Hydroiden des Golfes von Neapel und ihr Anteil an der Fauna unterseeischer Hohlen. Publ. Staz. zool. NapoliSO (suppl.) : 591-755. Rioja y Martin, J. 1906. Datos para el conocimiento de la fauna marina de Espana. Boln R. Soc. esp. Hist. not. 6: 275-281. Ritchie, J. 1907. On collections of the Cape Verde Islands marine fauna, made by Cyril Crossland, M.A. (Cantab.), B.Sc. (Lond.), F.Z.S., of St. Andrews University, July to September, 1904. The hydroids. Proc. zool. Soc. Lond. (1907) : 488-514. 1910-1911. Contribution to our knowledge of the hydroid fauna of the west of Scotland. Ann. Scot. nat. Hist. 76 (1910) : 220-225; 77 (191 1) : 29-34; 79 (191 1) : 158-164; 80 (191 1) : 217-225. Robins, M. W. 1969. The marine flora and fauna of the Isles of Scilly. Cnidiaria and Ctenophora. /. nat. Hist. 3: 329-343. Robson, J. H. 1914. Catalogue of the Hydrozoa of the north-east coast (Northumberland and Durham). Rep. Dove mar. Lab. (NS)3 : 87-103. Roosen-Runge, E. C. 1970. Life cycle of the hydromedusa Phialidium gregarium (A. Agassiz, 1862) in the laboratory. Biol. Bull. mar. biol. Lab., Woods Hole, 139 : 203-22 1 . Rossi, L. 1950. Celenterati del Golfo di Rapallo. Boll. Musei Zool. Anat. comp. R. Univ. Torino 2: 193-235. 197 1 . Guid a cnidari e ctenofori della fauna Italiana. Quad. civ. Sta. idrobiol. Milano 2 : 5-101 . Russell, F. S. 1953. The medusae of the British Isles. Anthomedusae, Leptomedusae, Limnomedusae, Tr achy medusae and Narcomedusae. Cambridge. Sars, M. 1835. Beskrivelser og Jagttagelser over nogle maerkelige eller nye i havet ved den Bergenske Kyst levende Dyr af Polypernes, Acalephernes, Radiaternes, Annelidernes og Molluskernes Classer, med en kort Oversigt over de hidtil af Forfatteren sammesteds fundne Arter og deres Forekommen. Bergen. 1846. Fauna littoralis Norvegiae. Erstes Heft. I. Ueber die Fortpflanzungsweise einiger Polypen (syncoryna — Podocoryna — Perigonimus — Cytaeis). Christiania. 1850. Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Lofoten og Finmarken. Nyt. Mag. Naturvid. 6 : 121-211. [The date of this paper has been given as 1851 (e.g. by Bedot, 1910; Vervoort, 1972), and that date appears on the title page of the volume. However, Botten-Hansen & Petersen (1870 : 209) and Sherborn (1922 : 14) dated the paper 1850. The paper was reprinted as a book differing only in being paginated 1-9 1 , published 'Christiana', 1 850.] 1857. Bidrag til Kundskaben om Middelhavets Littoral-Fauna, Reisebemaerkninger fra Italien. Nyt. Mag. Naturvid. 9 : 1 10-164. (This article formed part of a book of identical title published in Oslo (Christiania) in the same year, paginated 1-55.) Savage, S. 1945. A catalogue of the Linnaean herbarium. London. Schach, H. 1935. Kausal Analyse der Entstehung des polysiphonen Stockes von Campanularia verticillata L. Arch. Entw-Mech. Org. Berlin 132 : 615-647. Schmidt, H.-E. & Benovic, A. 1977. Notes on hydromedusae (Cnidaria) from the Adriatic Sea. J. mar. biol. Ass. U.K. 51 : 635-640. Shaw, G. R. & Nodder, F. P. 1975. The naturalist's miscellany. Vol. 6. London. (Plate 196, cited here under Clytia hemisphaerica, was probably published in 1795 according to Sherborn, C. D. 1895 'On the dates of Shaw and Nodder's "Naturalist's Miscellany".' Ann. Mag. nat. Hist. (6) 15 : 375-376.) Sherborn, C. D. 1914. An attempt at a fixation of the dates of issue of the parts of the publications of the Musee d'Histoire Naturelle of Paris, 1802-1850. Ann. Mag. nat. Hist. (8) 13 : 365-368. 1922. Index animalium . . . Part I. Introduction, bibliography and index A-Aff. London. 1926. Index animalium . . . Part X. Index Eurystomus-funereus. London. 1929. Index animalium . . . Part XVIII. Index o-nigrum-pallens. London. Sigerfoos, C. P. 1893. Note on the formation of blastostyle buds on the reproductive organs of Epenthesis mccradyi. Johns Hopk. Univ. Circ. 12 : 106. Slabber, M. 1769-1778. Natuurkundige verlustigingen, behelzende microscopise waarneemingen van in- en uitlandse water- en land-dieren. Haarlem, Holland. (For bibliographic details see Cornelius, 19750:289.) 1775-1781. Physicalische Belustigungen oder microscopische Wahrnehmungen von drey und vierzigin- und auslandischen Wasser- und Landthierchen. Translated by P. L. St. Miiller. Niirnberg. Splettstosser, W. 1924. Beitrage zur Kenntnis der Gattung Laomedea (sensu Broch). Zool. Jb. (Syst.) 48 : 367^32. Stebbing, A. R. D. 1976. The effects of low metal levels on a clonal hydroid. J. mar. biol. Ass. U.K. 56 : 977-994. 138 P. F. S. CORNELIUS 1979. An experimental approach to the determinants of biological water quality. Phil. Trans. R. Soc. (8)286:465^181. 198 1 . The kinetics of growth control in a colonial hydroid. J. mar. biol. Ass. U.K. 61 : 35-63. Stechow, E. 1913. Hydroidpolypen der japanischen Ostkiiste. II Teil: Campanularidae, Halecidae, Lafoeidae, Campanulinidae und Sertularidae, nebst Erganzungen zu den Athecata und Plumularidae. Abh. bayer. Akad. Wiss. suppl. 3 (2) : 1-162. 1914. Zur Kenntnis neuer oder seltener Hydroidpolypen, meist Campanulariden, aus Amerika und Norwegen. Zoo/. Anz. 45 : 120-136. 1919a. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete, nebst Angaben iibereinige kirchenpauer'sche Typen von Plumulariden. Zoo/. Jb., Syst. 42 : 1-172. 19 190. Neue Ergebnisse auf dem Gebiete der Hydroidenforschung. Mtinchener Med. Wochenschr. (1919)30:852-853. 1919c. Neue Ergebnisse auf dem Gebiete der Hydroidenforschung. Sber. Ges. Morph. Physiol. Munch. 31 : 9-45. — 192 la. Neue Genera und Species von Hydrozoen und anderen Evertebraten. Arch. Naturgesch. 87 (Abt. A. Heft 3) : 248-265. — 192 10. Symbiosen zwichsen Isopoden und Hydroiden. Zoo/. Anz. 53 : 221-223. — 1923a. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete. II Teil. Zoo/. Jb. (Syst.) 47 : 29-270. — 19230. Die Hydroidenfauna der japanischen Region. J. Coll. Sci. imp. Univ. Tokyo, 44 (8) : 1-23. — 1923c. Neue Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen iiber einige andre Formen. Zoo/. Anz. 56 : 1-20. — 1923d. Uber Hydroiden der Deutschen Tiefsee-Expedition, nebst Bermerkungen iiber einige andre Formen. Zoo/. Anz. 56 : 97-1 19. — 1925. Hydroiden der Deutschen Tiefsee-Expedition. Wiss. Ergebn. dt. Tiefsee- Exped. "Valdivia 17 : 383-546. — 1927. Die Hydroidenfauna der Ostsee. Zoo/. Anz. 70 : 304-3 13. 1932. Neue Hydroiden aus dem Mittelmeer und dem Pazifischen Ozean, nebst Bemerkungen iibereinige wenig bekannte Formen. Zoo/. Anz. 100 : 81-92. Stefani, R. 1956. II problema dell'allogonia negli Idrozoi. 1. Osservazioni biologiche ed istologiche in una popolazione di Campanularia caliculata Hincks del Golfo di Cagliari. Archo zoo/, ital. 41 : 337-368. 1959. Sulla variabilita ecologica di un Idrozoo (Campanularia caliculata Hincks). Boll. Zoo/. 26 (2): 115-1 20. Stepanyants, S. D. 1979. Gidroidy vod antarktikh i subantarktikh. Issled. Fauny Morei 22 (30): 1-200. Stephens, J. 1905. A list of Irish Coelenterata, including the Ctenophora Proc. R. Ir. Acad. (B) 25 : 25-92. Teissier, G. 1965. Inventaire de lafaune marine de Roscoff. Cnidaires-ctenaires. Roscoff, France. Thiel, M. E. 1935. Zur Kenntnis der Hydromedusenfauna des Schwarzen Meeres. Zoo/. Anz. Ill : 161-174. Thornely, L. R. 1899. The hydroid zoophytes collected by Dr Willey in the southern seas. In: Willey, A., 1898-1902, Zoological results based on material from New Britain, NewGuinea, Lovalty Islands and elsewhere. Collected during 1895, 1986 and 1897.4 : 45 1-457. Cambridge. 1904. Report on the Hydroida collected by Professor Herdman, at Ceylon, in 1902. Rep. Govt Ceylon Pearl Oyster Fish. GulfManaar, Suppl. Rep. 8 : 107-126. Todd, R. A. 1906. Marine zoology. In Page, W. (editor) The Victoria history of the counties of England: a history of Devonshire in five volumes. Volume 1, pp. 1 3 1-1 58. London. Torrey, H. B. 1904. The hydroids of the San Diego region. Univ. Calif. Publs Zoo/. 2 : 1-43. Toth, S. E. 1969. Aging and regression in the colonial marine hydroid Campanularia flexuosa with special reference to senescence in hydroids. Int. Rev. gen. exp. Zoo/. 4 : 49-79. Totton, A. K. 1930. Coelenterata. Part V. Hydroida. Nat. Hist. Rep. Br. antarct. Terra Nova Exped. 5: 131-252. 1965. A synopsis of the Siphonophora. London. Trebilcock, R. E. 1928. Notes on New Zealand Hydroida. Proc. R. Soc. Viet. 41 : 1-3 1 . Tutin, T. G. 1942. Zostera L. J. Ecol. 30 : 2 1 7-226. Vanhoffen, E. 1910. Die Hydroiden der Deutschen Sudpolar-Expedition 1901-1903. Dt. Sudpol.- Exped. 11 (= Zool. 3) : 269-340. Vannucci, M. 1955. On the newly liberated medusa ofObelia hyalina Clarke, 1 879. Dusenia 6 : 55-60. N.E. ATLANTIC CAMPANULARIID HYDROZOANS 139 Vatova, A. 1928. Compendio della Flora e Fauna del Mare Adriatico presso Rovigno. Memorie R. Com. talassogr. ital. 143 : 1-614. Verrill, A. E. 1873. Results of recent dredging expeditions on the coast of New England. Am. J. Sci. (3) 5: 1-16. Vervoort, W. 19460. Hydrozoa (CI). A. Hydropolypen. Fauna Ned. 14 : 1-336. 19466. Hydroiden uit de Waddenzee. Archs neerl. Zool. 7 : 334-352. 1949. Notes on a small collection of hydroids from Jersey (Channel Islands). Zool. Meded. Leiden, 30: 133-162. 1959. The Hydroida of the tropical west coast of Africa. Atlantide Rep. 5:21 1-325. 1966. Bathyal and abyssal hydroids. Galathea Rep. 8 : 97-174. 1967. The Hydroida and Chondrophora of the Israel South Red Sea Expedition, 1962. Bull. Sea Fish. Res. Stn Israel, 43 : 1 8-54. 1968. Report on a collection of Hydroida from the Caribbean region, including an annotated checklist of Caribbean hydroids. Zool. Verh. Leiden92 : 1-124. 1972. Hydroids from the Theta, Vema and Yelcho cruises of the Lamont-Doherty Geological Observatory. Zool. Verh. Leiden 120 : 1-247. Warburton, F. E. 1953. Antagonism between different species of hydroids on the same shell. Ecology 34: 193-194. Warren, E. 1908. On a collection of hydroids, mostly from the Natal coast. Ann. Natal Mus. 1 : 269-355. Weismann, A. 1 883. Die Entstehung der Sexualzellen bei den Hvdromedusen. Jena. West, D. L. & Renshaw, R. W. 1970. The life cycle of Clytia attenuata (Calyptoblastea : Campanulariidae). Mar. Biol. Berlin 1 : 332-339. Westendorp, G. D. 1843. Recherches sur les polypiers flexibles de la Belgique el particulierement des environs d'Ostende. Bruges, pp. 1-48 + pi. 1. (Published for the Societe Medico-Chirurgicale de Bruges by F. de Pachtere. Reprinted from the Annales of the Societe, vol. 4. I have not seen the Annales version.) 1853. Polypiers flexibles de la Belgique. Collection de bryozoaires, sertulaires, /lustres el spongiaires qu'on rencontre en Belgique et particulierement aux environs d'Ostende. Coutrai, Belgium. [Not seen. These details are from Bedot( 19 10). See note l,p. 123.] Will, J. G. F. 1 844. Horae tergestinae oder Beschreibung und Anatomie der im Herbste 1843 bei Triest beobachteten Akalephen. Leipzig. Winsor, M. P. 1976. Starfish, jellyfish, and the order of life. New Haven, U.S.A. Winther, G. 1879. Fortegnelse over de i Danmark og dels nordlige Bilande fundne Hydroide Zoophyter. Naturh. Tidsskr. (3) 12 : 223-278. Wood, G. W. 1901. The hydroid zoophytes of the Isle of Man, with a notice of species not hitherto reported from the district. Lioar Manninagh 2 : 12-2 1 . Wright, T. S. 1858. Observations on British zoophytes. Proc. R. phys. Soc. Edinb. 1 : 447-455. 1859. Observations on British zoophytes. Edinb. newphil. J. (N.S.) 9 : 106-1 14. 1862. On the reproduction of Thaumantias inconspicua. Q. J. microsc. Sci. (N.S.) 2 : 221-222, 308. 1 863. Observations on British zoophytes. Q. J. microsc. Sci. (N.S.) 3 : 45-52. Wulfert, J. 1902. Die Embryonentwicklung von Gonothyraea loveni Allm. Z. wiss. Zool. 71 : 296-327. Wyttenbach, C. R. 1968. The dynamics of stolon elongation in the hydroid, Campanularia flexuosa. J. exp. Zool. 167:333-351. 1969. Genetic variations in the mode of stolon growth in the hydroid, Campanularia flexuosa. Biol. Bull. mar. biol. Lab., Woods Hole 137 : 547-556. , Crowell, S. & Suddith, R. L. 1973. Variations in the mode of stolon growth among different genera of colonial hydroids, and their evolutionary implications. J. Morph. 139 : 363-375. Yamada, M. 1959. Hydroid fauna of Japanese and its adjacent waters. Publs Akkeshi mar. biol. Stn 9: 1-101. 1965. Marine hydroids from Greece. Publs Seto mar. biol. Lab. 12 : 359-362. Zirpolo, G. 1939. Caso di epibiosi di Obelia geniculata su Hippocampus guttulatus. Annuar. Mus. zool. Univ. Napoli (N.S.) 7 (8) : 1-8. 1940. Nuovo caso di associazione di idroidi e pesci con revisione critica dei casi gia noti. Boll. Soc. nat. Napoli ( 1938-1 939) 50 : 127-139. Manuscript accepted for publication 24 July 1981. Index Valid names are in Roman type, new names in bold and invalid names in italics. New combinations are indicated by an asterisk (*). Main page references are in bold. References to substrates are not included. abietina, Campanularia & Grammaria 1 22 acuminata, Campanulina 74 acuminata, Laomedea 122 acuta, Campanularia 74, 81 adelungi, Obelia 1 1 4 adriatica, Obelia 1 20 Aequorea 122 vitrina 123 affinis, Clytia 7 1 affinis, Eucope 11, 73, 74 africana, Campanularia 60 africana, Orthopyxis 39, 60 agas, Campanularia 54 Agastra 48, 57-58 caliculata 6 1 mira 6 1 , 67 rubra6l,63,61 aha, Campanularia 53, 54 alternata, Clytia 84, 85, 86 alternata, Eucope & Obelia, 1 14 andersoni, Obelia 1 14 angulata, Campanularia & Eulaomedea 98 angulata, Laomedea 41, 42, 43, 44, 98-102, 104, 105; Fig. 17, p. 99 angulosa, Obelia 1 14 antennina, Nemertesia 121 arborescens, Clytia 1 13, 1 14, 117 arcta, Coppinia 122 armata, Oceania 8 1 arruensis, Obelia 1 14 articulata, Eucope 1 14 asymmetrica, Orthopyxis 6 1 , 67 atlantica, Hypanthea 50 atlantica, Silicularia 50 attenuata, Campanularia 73-74, 79, 81 attenuata, Clytia 40, 42, 8 1 attenuata, Obelia 1 14 australis, Obelia 1 14 austrogeorgiae, Obelia 1 13, 117 bicophora, Clytia (Trochopyxis) 73, 80 bicophorum, Phialidium 74, 80 bicuspidata, Obelia 90, 113, 1 14 bidentata, Obelia 40, 43, 44, 85, 96, 97, 1 10, 111,113-117 bifurca, Obelia 11 4, 115 bifurcata, Obelia 1 14 bilabiata, Campanularia 58, 59, 60 bilabiata, Eucopella 58 bilabiata, Silicularia 60 Billardia 123 novaezealandiae 123 biserialis, Obelia 1 14 bistriata, Laomedea 84, 85, 86 borealis, Campanularia 6 1 , 66 borealis, Obelia 1 14 brachycaulis, Campanularia 53, 55-56, 74, 82 brasiliensis, Campanularia 1 14 braziliensis, Obelia 1 14 breviscyphia, Campanularia 60-6 1 , 66 brochi, Laomedea 47, 107, 110 brochi, Paralaomedea 47 buskiana, Thaumantias 74 buskianum, Phialidium 74 calceolifera, Campanularia 102 calceolifera, Eulaomedea 98, 102 calceolifera, Laomedea 41, 42, 43, 44, 47, 98, 100, 101, 102-105; Fig. 18, p. 103 calceolifera, f. of Laomedea angulata 100 calceolifera, Obelia 102 caliculata, Agastra 6 1 caliculata, Campanularia 51, 58, 60-61, 64, 65-66 caliculata, Eucopella 58,61 caliculata, Orthopyxis 48, 6 1 , 63, 65-66 Calycellal22 syringa 55, 57, 78, 122 calyculata, Campanularia 51,61 Camp alar ia 97 conferta 102-105 campanella, Medusa 73 Campanula 51 Campanularia, Eucopella 58, 60, 61 , 63, 67 Campanularia 4 1 , 47, 48, 49, 50, 51-52, 57, 70, 71,97,112 abietina 122 acuta 74, 81 africana 60 agas 54 angulata 98 attenuata!3,14,19,Sl bilabiata 58, 59, 60 borealis 6 1 , 66 brachycaulis 53, 55, 56, 74, 82 brasiliensis 1 1 4 breviscyphia 60, 6 1 , 66 calceolifera 102 caliculata 5 1 , 58, 60, 6 1 , 64, 65-^6 calyculata 51,61 caulini 1 14 cavolinii 1 14 cheloniae\\4, 117, 118,119 INDEX 141 compressa 60, 6 1 , 64, 66-67, 73 confer ta 100, 104 coruscans 1 14 crenata 52-53, 59, 60; Fig. 2, p. 55 crenata f. intermedia 59 decipiens 107 devisa 123 edwardsill, 81 elongata 107,110 everta5\,58,6Q fastigiata 123 flabellatall4 flemingii95,96,\\3 flexuosa 105 fragilis 107, 109,110 fruticosa 121, 122 gegenbauri!3,8Q gelatinosa 95 geniculata sens. Lister 9 1 gigantea 73, 75,81 gracilis sens. Allman 60, 61, 66; sens. Meyen 50;sens. Sars 74, 77,81 gracillima 122 gravieri%4,85 groenlandica 52, 55, 56, 57 hicksoni 54 hincksii 5 1 , 53-55, 80, 82; Fig. 3, p. 55 humilis 123 hummelincki 82 inconspicua 73 Integra 52,60 integriformis 6 1 , 64, 66-67 intermedia 58,60 inter text a 65 , 122 johnstoni 71 , 73, 75, 77, 79-80 kincaidi 9 1 lacerata 122 7flevis54,60,61,65 lennoxensis 58, 60 macrotheca 53,54 magnifica 52,53 maior 1 1 4 morgansi 5 1 mutabilis 123 neglecta 107 noliformis 80 obliqua 84 obtusidens 1 14 parvula 122 paulensis 88 pelagica 73, 78-79 platycarpa 6 1 prolifera 1 14 ptychocyathus 80 pulcratheca 5 1 rara 53 raridentata 73, 74, 75, 76, 80, 8 1 , 90 retroflexa 5 1 60 senulatall,8Q speciosa 52, 53 speciosus 52 spinulosa 1 14 tulipifera 70 tulpifera 70 vermicularis 120 verticellata 67 verticillata67,69 villafrancensis 74, 81 volubiliformis 73, 80 volubilis 48, 5 1 , 54, 55-57, 69, 70, 73, 76, 77, 79; Fig. 4, p. 55 CAMPANULARIIDAE 47^19 CAMPANULARIINAE 48-49, 50-51, 69-70 campanulata, Eucope 7 1 , 73, 74 Campanulata 5 1 verticillata 68 Campanulina acuminata 74 tennis 123 Capsularia 121, 122 Integra 6 1 /aev/5 6 1 , 65 carnea, Podocoryne 43 caulini, Campanularia 1 14 cavolinii, Campanularia 1 14 cheloniae, Campanularia 1 14, 1 17, 1 18, 1 19 chinensis, Obelia 1 14 Clytea 70 vicophora 73 Clythia 57, 70 C/y//z/ajohnstoni 74 poterium 6 1 , 66 Clytia 41 , 42, 47, 48, 49, 57, 69, 70-72 affinis 1 \ alter nata 84,85,86 arborescens 1 13, 1 14, 117 attenuata 40,42,81 bicophora 73,80 compressa 14, 82 cylindrica 42, 73, 80 discoida* 42, 72-73; Fig. 8, p. 72 edwardsi 8 1 ed wards ia 43 elsaeoswaldae 80 eucophora, eucopophora 123 flavidula 73, 74, 76, 77, 80, 81 folleata 9 1 foxi 84, 86 gardineri 9 1 geniculata 84, 86 gigantea 40, 74, 77 gracilis 42, 71,91 gravieriS5,86 hemisphaerica 40, 41, 42, 43, 45, 70-71, 73-82, 86, 87, 89, 90, 91, 92, 94; Fig. 9, p. 75 hendersonae 84 142 hummelincki 4 1 , 42, 44, 69, 82-83, 86; Fig. 10, p. 82 islandica* 42, 84; Fig. 11, p. 84 johnstoni 69, 70, 73, 74, 80 kincaidi 9 1 teem 55, 57, 73 tanguida 8 1 linearis* 42, 84-86; Fig. 12, p. 85 longicyatha 120 longiiheca 1 1 4 mccradyi* 42, 87-88; Fig. 13, p. 87 mo///s55,56,57,80 «o///brw/.s73,80, 81 obeliformis 13,15, &\ obliqua85,86 olivacea 67 Clytia (Orthopyxis) poten'MW 57, 58,60,61,66 Clytia paulensis 42, 43, 44, 76, 85, 86, 88-91, 113, 115, 117; Fig. 14, p. 89 pelagica 73 pentata42,9l Clytia (Platypyxis) 70, 71, 80 Clytia posterior 6 1 poterium61,63 rijckholtii 6 1 , 65 ryckholtii 6 1,65, 123-124 sarsi41,74,78 serrata 85 serrulata 73, 74, 76 simplex 1?>,&\ striata 86 Clytia (Trochopvxis) 70 Clytia ulvae 88, 91 undulata 60, 6 1 , 65 unijlora 74 urnigera 64-65, 73, 78 vicophora 73 villafrancensis 74 viridicans 74 volubilis73,74,76,77,78 volubis 43 CLYTIINAE 38, 48-49, 50, 69-70 clytioides, Orthopyxis 50, 118-119 clytioides, Silicularia 50, 1 18-1 19 clytioides, Tubularia 50, 1 14, 1 17, 118 Cmpanularia 5 1 comata, Cymodocea & Nemertesia 121, 122 commensuralis, Obelia 1 14, 1 17, 118 commissuralis, Obelia 1 14, 1 17, 119 compressa, Campanularia 60, 6 1 , 64, 66-67, 73 compressa, Clytia 74, 82 compressa, Orthopyxis 6 1 , 66 ConcHella 122 intertexta 122 conferta, Campalaria 102-103, 104-105 conferta, Campanularia 100, 104 conferta, Laomedea41, 102-103, 104 congdoni, Laomedea& Obelia 1 14, 119 INDEX convexa, Thaumantias 73 Coppinia arcta \ 22 corona, Obelia 1 14 coruscans, Campanularia 1 14 Coryne65, 122 Couchella\2\ coughtreyi, Obelia 1 14 crenata, Campanularia 52-53, 59, 60; Fig. 2, p. 55 crenata, Eucopella 52, 58 crenata, Orthopyxis 40, 42, 52, 58-60; Fig. 5, p. 59 Cuspidella humilis 123 cylindrica, Clytia 42, 73, 80 cylindrica, Platypyxis 73 cymbaloidea, Epenthesis 73, 74 cymbaloidea. Medusa 73 Cymodocea 1 2 1 comata 122 ramosa 1 2 1 simplex 114, 117,121 decipiens, Campanularia 107 decipiens, Laomedea 107, 109, 110 delicata, Orthopyxis 58, 59, 60 diaphana, Eucope & Thaumantias 1 14 dichotoma, Obelia 40, 41, 43, 45, 73, 96, 100, 101, 102, 104, 112, 113, 114, 117-119, 120,121 dichotoma, Sertularia \ 1 7 dischotoma, Obelia 1 14, 1 17 discoida,* Clytia 42, 72-73; Fig. 8, p. 72 discoida, Oceania 72 discoidum, Phialidium 73 divaricata, Laomedea 1 1 4 divisa, Campanularia 123 dubia, Obelia 114 dubia, Thaumantias 73 dumosa, Lafoea 47, 65, 1 22 echinata, Hydractinia 43 edwardsi, Campanularia 73, 81 edwardsi, Clytia 8 1 edwardsia, Clytia 43 elongata, Campanularia 107, 110 elsaeoswaldae, Clytia 80 Epenthesis 70, 71 cymbaloidea 73, 74 mccradyi 87 equilateralis, Obelia 1 14 Eucalix 48-49, 50-51 morgansi 5 1 retroflexus 5 1 Eucampanularia 47-48, 5 1 , 52, 97 groenlandica 52 integra 52 INDEX 143 speciosa 52 verticillata 52 volubilis52 Eucope 70, 71, 1 12 affinis 71,73,74 alternata 1 1 4 articulata 1 1 4 campanulata 7 1 , diaphana 1 14 exigua 73 fusiform is 1 14 gemmifera 73 pamsitica \ 1 4 pictall, 123 polygena 1 14 polystyla 1 1 primordiale 123 pyriformis 1 1 4 thaumantias 74 thaumahtoides 71 variabilis 73 73, 74 73 bilabiata 58 caliculata 58,61 campanularia 58, 60, 6 1 , 63, 67 crenata52,58 eucophora (& eucopophora), Clytia 123 Eucopium primordiale 123 Eulaomeda 97 flexuosa 105, 117 Eulaomedea 47, 97-98, 107 angulata 98 calceolifera98, 102 flexuosa 98, 107, 1 17 pseudodichotoma 1 1 1 everta, Campanularia 51,58, 60 m>o — r-oo'oooo — OONTT I/I (N CN| l~~ 1 — Q ^™ ^^ ^^ rx|— i«NO(^)TfTt>orr>ON-^--^too^OO oo t~- O 10 m C/5 •* r O ^ ^O /"> r<^ OO Tt >O Tf DA C T *? °f "]>ri4Tri40?0fTrTr?rTl(~-vOfN'<3- ON — ON ^" 00 00 1 U 2 m r- vo ON roOfN«0«0— .fNfNOOOOO Tf OOOO C/3 Q O O m t~- t~-rr)i/~)^1'ONOO^O>O(Nt~~- — — — ON CO en f*"i r — C/5 C/5 *n ^^ ^^ ^^ ^^ rn ^" ON 0*4 ^f ON 0. a t«j >^ >/"> "Q _o ^H w~> m — • s o SO r«-> r-j ON — ^ u. 2 .0 ^ ** i. M — c o *- U _ •a E fe *| •§ x jS SB g "SSg J3J2-2 C'£12 00 O ^^ CJ I- u OU ^~* 5^^ *-» ^_^ UN f^ TO IS * ^ C *7j fj 5S QJ E 3 ^ TO fi C .i- ^ OJQ iu o !>s — -" • O Cd *O r" C/5 ™"* ^^ r" (• ^ ^ C- « i ^^ r? QJ . i— ._- ^^ >*~*^ CO C/5 "_y -^ " "i^ ^^ *""* ^ QJ ?r u W TO ^ c/5 r* O prt u **•• *v O & OJD _*™| ^^ ^* O "" O "^ *-* V E'-s E 2 1 1 1| 1 '5 1| g> | S | g £i >2 g s|.§ ^ £ - Charact ^g 00^ c 1> U i '•= cx-= •?•= — — 'c -S 2 2 p __ 2 l>J=-£-J-'COcO'JQ.flja>w" ^ J— CjD.D,D.OO>Oi--t.- '>? Ouj aCCCggUiUxx^ ZUc/5<< C 'Q, 3 C 03 2>O- 70 mm. A species of Luidia with normally 5 arms; abactinal paxillae with about 4 longitudinal series each side also forming transverse rows with the matching but distinctly smaller superomarginal paxillae, though irregularities may occur where the paxillae of the second and third (sometimes other series) are markedly enlarged and bear a single stout central spine fringed by usually several rings of spinelets, the large spines similar in magnitude to the ambital inferomarginal spines, rarely some of the lateralmost series and many of the median paxillae may have a smaller central spine developed; inferomarginal plates primarily ventral in position, only narrowly visible dorsally, armed at the ambitus with 2 or 3 large spines, usually projecting approximately horizontally, and on the ventral side with 2-4 smaller and somewhat flattened spines among the spinelets: adambulacral plates with 3 large spines in a row at right angles to the furrow, sometimes a more or less enlarged spine proximal to the lateralmost one; elongate, 3- or 4-valved pedicellariae, present on some proximal actinal plates (rarely also on the outer ends of the first few adambulacral plates) the slender valves usually only meeting at the tips; shorter, more often bivalved pedicellariae sometimes present (especially in west african specimens) on some abactinal and/or superomarginal paxillae, no pedicellariae on the furrow face of the oral plates. Colour boldly patterned on the upper side with a dark pentagon on the disc and three to five transverse dark bands, brown, black, greenish or purple, on each arm, the remainder being yellow, cream or white. STATUS OF L. numidica. Madsen (1950) treated Luidia numidica Koehler as a west african variety of L. alternata and noted that the only difference appeared to be the smaller number of spine-bearing paxillae in numidica. A. M. Clark (1953) used subspecific rank for numidica but noted that a specimen from the Cape Verde Islands with relatively numerous spino-paxillae is very similar to some west Atlantic specimens of L. alternata. In 1973, Nataf & Cherbonnier treated numidica as a full species but without comparisons or any comment on its status. Koehler (1911) had only two specimens from West Africa, a poor dry one with R 52 mm and the broken holotype, r 13mm so R probably 80-100 mm. He remarked on the closeness to L. alternata but distinguished that species by the more numerous inferomarginal and paxillar spines, the more attenuated pedicellariae and the 'quite different' paxillae with the peripheral spinelets thin and elongated, not short, thick and granuliform. Study of c. 30 and 15 specimens from west and east Atlantic respectively, mostly from the Gerda and Pillsbury collections, now indicates that, although the armament of the paxillae is somewhat variable in both east and west Atlantic specimens and is also subject to some extent to growth changes, there is a consistent difference in the armament of the smaller paxillae if total size is taken into account and also a significant difference in the number (often also the shape) of the large paxillar spines. Already in west african specimens at R 20-25 mm, over half the median paxillae have more than one (usually 2-4) short coarse similar central spinelets; at R 40-50 mm these number 3-6 and at R c. 60 mm usually 4-8 (see Fig. 3h, i). In L. alternata from the western Atlantic, even at R 100 mm or more, many central paxillae still only have a single more or less coarser central spinelet, the maximum number being about 4. However, these large specimens may have some median paxillae with the peripheral spinelets arranged in two concentric rings of 6-10 each, the outer ones being more slender and cylindrical than the inner ring, the spinelets of which are transitional in shape to the central spinelet (or spine). In general, the shape of the peripheral spinelets can usually be called subclavate but 165 n 01 Fig. 3 (a, b, d-p) Abactinal paxillae, (c, q-v) pedicellariae. (a-g) Luidia alternata alternata, (a) type material of L. quequenensis, Quequen, N Argentina, R 72 mm, median paxillae; (b) holotype of L. bernasconiae, R 100 mm; (c) the same, superomarginal pedicellaria; (d) Gerda st. 1038, R 40 mm, median paxillae; (e) Oregon st. 42 1 5, R c. 60 mm, median paxillae; (f) Pillsbury st. 654, R 100+ mm; (g) Pillsbury st. 767, R c. 90 mm, superomarginal paxilla; (h, i) L. alternata numidica: (h) Pillsbury st. 248, R only c. 22 mm; (i) Cape Verde Is, R 55-60 mm; 0) L. heterozona heterozona, Pillsbury st. 68, Re. 100 mm, disc paxillae; (k) L. ciliaris, Naples, R 140mm, inner lateral paxilla (above) and two median paxillae; (1) L. sagamina aciculata, Atlantide st. 61, R c. 90 mm, superomarginal paxilla (left) and lateral paxillae from above and the side (below); (m) L. sarsi sarsi, W. of Scotland, Re. 100 mm, median paxillae; (n) L. sarsi sarsi, paratype of L. africana, Porcupine st. 36, R c. 85 mm, disc paxillae; (o) L. sarsi elegans, Gerda st 456, R c. 60 mm, median paxillae; (p) L. atlantidea, Pillsbury st. 26, Re. 100 mm, lateral paxillae; (q) L. alternata alternata, Oregon st. 4190, actinal pedicellariae; (r) L. heterozona heterozona, Pillsbury st. 259 (left) and 49 (right), actinal pedicellariae in two views; (s) L. sagamina aciculata, Atlantide st. 6 1 , two actinal plates, one armed only by the pedicellaria, pedicellariae slightly foreshortened; (t) L. ludwigi scotti, paratype, actinal paxillae with 2-, 3- and 4-valved pedicellariae, foreshortened; (u) L. ludwigi scotti, Pillsbury sts 1384 & 842, lateral views of 4- and 3-valved actinal pedicellariae; (v) L. sarsi sarsi, Shetland Is, lateral views of actinal pedicellariae. 166 A.M.CLARK alternate ones may be hardly at all expanded at the tips on the slightly larger paxillae. As for the large paxillar spines on the second and third lateral series of paxillae particularly, even in the Cape Verde Islands specimen mentioned in 1953 only c. 35%, at most, of these lateral paxillae have the large spines. In other specimens from West Africa the percentage is usually c. 20. The shape of the spines is also relatively squat, their length most often 1-2 mm, the longest (in Koehler's large holotype) 3 mm, the length/basal breadth usually less than 3/1. In american specimens such squat spines may also occur but usually the length/breadth ratio is4-(M'5/l and the length may exceed 3 mm even at R only 60 mm. The conclusion reached from these observations is that the specimens from the two sides of the Atlantic are conspecific but the west african form can be recognized as a distinct subspecies numidica of L. alternata, distinguished by the more 'plushy' superficial appearance of the abactinal paxillae, interspersed with shorter and fewer large spines. SYNONYMY. Luidia granulosa Perrier, 1869 was recognized as a synonym of L. alternata by Perrier himself in 1875 and L. variegata Perrier, 1875 by Verrill, 1915. In 1950, Madsen (cited by A. M. C., 1953 : 380) also referred L. bernasconiae A. H. Clark, 1945, to the synonymy but this was refuted by Downey in 1968 & 1973, who maintained that it has only two adambulacral spines, not three or four, and the non-spine-bearing paxillae have 5-10 clavate peripheral spinelets compared with 10-16 slender acute ones in alternata, though on p. 23 of the 1973 paper she describes the peripheral paxillar armament of alternata as subclavate. The dried holotype of L. bernasconiae certainly shows swollen-tipped peripheral paxillar spinelets more clavate than usual for L. alternata but it regularly has three large and a fourth smaller adambulacral spine, as described by A. H. Clark, the common number for L. alternata. In view of the variability in shape of the paxillar armament in L. alternata already mentioned, it is difficult to maintain bernasconiae as distinct. If there are specimens from North Carolina as described by Gray, Downey & Cerame- Vivas (1968) with sporadic large spines but only two adambulacral spines, then these may have to be nomenclatorially distinguished but not as L. bernasconiae. The superficial difference in proportions between L. alternata and L. bernasconiae suggested by the photographs in the 1968 paper is correlated with the much larger size (R c. 135 mm, judging from the scale) of the specimen named alternata, and exaggerated by the greater reduction so that both appear of similar size though R is only c. 77 mm in the specimen named L. bernasconiae. With regard to Luidia quequenensis (and other species of Luidia described by her) Bernasconi, 1942 confusingly modified the terminology of the lateral abactinal paxillae used by Doderlein (1920, see pi. 18, caption) as follows: his 'Randpaxillen' (i.e. the supero- marginal paxillae) become her 'paxilas marginales'; his 'untere Seitenpaxillen' (the two [primary and secondary] outermost series of lateral paxillae) become her 'paxilas infero- marginales'; his 'obere Seitenpaxillen' (the third and fourth series or two outer series of median paxillae) become her 'paxilas superomarginales'. This odd terminology has unfor- tunately been perpetuated by Carrera-Rodriguez & Tommasi (1977). Bernasconi distinguished L. quequenensis from L. alternata in 1943 (pp. 5, 17) by the large number and coarse shape of the spines of the 'paxilas superomarginales', which should be the third and fourth series of paxillae from the true superomarginal ones. However, a specimen from Quequen, kindly presented to the British Museum by Dr Bernasconi, shows numerous spinopaxillae but more of these are in the second from outermost lateral series (what I call the primary row), though the third series also has a considerable number. However, there are many inconspicuous spineless paxillae also in these series so that even the second only has c. 64% of spinopaxillae out of well over 100 paxillae counted on several arms. The largest number of consecutive spinopaxillae found was only 9. Similar frequencies of over 60% of spinopaxillae in the second or third lateral series may occur throughout the range of L. alternata alternata, for instance in some of the Challenger specimens from Bahia, NE Brazil, others from French Guiana (Oregon and Pillsbury collections) and from Georgia (the holotype of L. bernasconiae). NEOTYPE. In the event of future disagreement with the synonymies now proposed and in the NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 167 &::** / - T* ' Fig. 4 Luidia alternata alternata (Say), neotype, BM(NH) reg. no. 1937.5.9.6, Dry Tortugas. Dorsal view, xO'6. 168 A.M.CLARK absence of Say's type material (apparently long-lost), it is very desirable that a neotype from the vicinity of Florida, the type locality, should be nominated. Accordingly I propose a beautiful specimen from the Dry Tortugas now in the British Museum collections, registered number 1937.5.9.6, collected by Colman and Tandy, shown in Figs 4 and 5. The arms vary in length so that R ranges from 150-1 75 mm; r is c. 19mm and R/r = 7-9-9'2/l. Spinopaxillae are fairly numerous on the proximal primary (second) lateral series but become more sporadic distally (c. 55% of 100+ primary paxillae from proximal parts of several arms are spine-bearing). The longest spines measure 4*0 mm in length. Progressively fewer of the third, fourth and fifth series of paxillae are also enlarged with spines but none of the first series or the adjoining superomarginal paxillae and very few midradial paxillae. The median paxillae without enlarged spines have at least one central and about 9 peripheral spinelets, the latter almost cylindrical and blunt-tipped but the slightly larger paxillae have up to 4 central and c. 20 peripheral spinelets, the latter in a double ring of which the inner ones are subclavate and transitional in shape to the shorter, coarser central spinelets. The inferomarginal plates mostly bear two large ambital spines up to 7 mm long, slightly irregular in position on consecutive plates but not regularly alter- nating. There are 2-4, usually 3, more irregular smaller spines on the ventral side of each plate. Most of the primary actinal plates bear a long slender 3- or sometimes 4-valved pedicellaria and proximally also one or two additional pedicellariae based on super- numerary platelets superimposed on the lateral ends of the adambulacral plates. Most adambulacral plates have 3 large spines and a smaller fourth one proximal to the third (lateral-most). On the proximal plates this fourth spine may reach a similar length to the others. DISTRIBUTION. L. alternata alternata from North Carolina to northern Argentina (c. 38° S); 0-160 metres. L. alternata numidica from Senegal to Zaire (Congo), West Africa; 10-100 metres. Luidia atlantidea Madsen Fig. 3p Luidia africana: Koehler, 1911:19; Doderlein, 1920 : 288-289; Mortensen, 1925 (pt) : 178. [Non L. africana Sladen, 1889.] Luidia atlantidea Madsen, 1950 : 192-198, fig. 5, pi. 16, figs 1,2; A. M. Clark, 1953 : 393, 394; 1955 : 22, 32; Nataf& Cherbonnier, 1973 : 76-80, pi. 1, fig. B, pi. 2, figs A, B, pi. 7, figs 1-5, pi. 9, figs E, F. [Non L. atlantidea: Downey, 1973 : 25 = L, sagamina aciculata.] Luidia sarsi: Studer, 1884 : 43. [Non L. sarsi Diiben & Koren, 1845.] DISTRIBUTION. On re-examination, the specimen from North Carolina mentioned under the name of L. atlantidea by Downey (1973) proved to be L. sagamina aciculata. It has a white band along each side of the paxillar area on the arms but this is situated on the two regular rows of lateral paxillae, not on the superomarginal series as in L. atlantidea. It has not been possible to check the Halpern record cited by Downey but the likelihood is that this too would have been a similar misidentification. The species is recorded from Atlantic Morocco to Zaire, including the Cape Verde Islands, 10-80 metres. Luidia barbadensis Perrier Fig. 2b Luidia barbadensis Perrier, 1881 : 29; 1884 : 267 [non pi. 10, figs 7, 8, probabaly = L. sarsi elegans]; Verrill, 1915 : 205-207, pi. 24, fig. 1; Doderlein, 1920 : 216; H. L. Clark, 1941 : 25; John & A. M. Clark, 1954:145; Cherbonnier, 1959:170, fig. 3B, E, fig. 4; Downey, 1973:24-25; Carrera-Rodriguez & Tommasi, 1977 : 67-7 1 , figs 5-9. Luidia convexiuscula Perrier, 1881 : 30; 1884 : 268, pi. 10, fig. 6; Verrill, 1915 : 207. NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 169 Fig. 5 Luidia alternata alternata, neotype. Ventral view of proximal part, x 1 -7. 170 A.M.CLARK REMARKS. Records of five-armed specimens of L. barbadensis are probably based on specimens of L. sarsi elegans, which is partially sympatric with it. Certainly the five-armed specimen captioned as L. barbadensis by Perrier, 1884, pi. 10, figs 7, 8, appears to be elegans, which also differs in the much finer paxillae and the long marginal spines. All 36 Gerda, Pillsbury and Oregon specimens of barbadensis which I have seen have 6 arms. DISTRIBUTION. Southern Florida and the immediately adjacent Gulf of Mexico, southern Bahamas, to SW Caribbean off Nicaragua and east and south to southern Brazil (c. 30° S), 73-430 metres. Luidia ciliaris (Philippi) Fig. 3k Asterias ciliaris Philippi, 1837 : 144. Luidia fragilissima Forbes, 1839 : 123; 1841 (pt) : 135-140, fig. on p. 135 [7-armed specimens]. Hemicnemis ciliaris: Miiller&Troschel, 1840 : 104. Luidia ciliaris: Perrier, 1875 : 342 [1876 : 262]; Sladen, 1889 : 254; Ludwig, 1897 : 61-85, pi. 4, figs 1, 2, pi. 6, figs 25-36; Sussbach & Breckner, 1911 : 209-210; Doderlein, 1920:287, figs 8, 17, 34; Koehler, 1921 : 55, fig. 41; Mortensen, 1927 : 90, fig. 89a; Madsen, 1950 : 205-206, fig. 8; Ursin, 1960 : 30-3 1 ; Tortonese, 1965 : 148-150, fig. 68; Blake, 1973 : 40, pi. 10, figs 1-26. DISTRIBUTION. NE Atlantic, from southern Norway, the Skagerrak, Shetlands, Faeroe Channel, south to the Canary Islands and Azores (?Cape Verde Islands); 1-400 metres. Luidia clathrata (Say) Fig. la Asterias clathrata Say, 1825 : 142. [Non A. clathrata Pennant, 1777.] Luidia clathrata: Liitken, 1859:37-39; Perrier, 1875:332-333 [1876:252]; Sladen, 1889:253; Verrill, 1915 : 200-201, pi. 24, fig. 2; Doderlein, 1920 : 238, 239, 251-252, figs 1, 21; H. L. Clark, 1933 : 19-20; Bernasconi, 1943 : 6-7, pi. 2, fig. 1; A. H. Clark, 1954 : 375; Ummels, 1963 : 95-97, pis 10, 11; Gray, Downey & Cerame-Vivas, 1968 : 139, fig. 8; Downey, 1973 : 22-23, pi. IJigsC, D; Blake, 1973:30-31, pi. 3, figs 1-28; Walenkamp, 1976:19-25, figs 2^t, 19a, pi. 3, fig. 2; 1979: 11-12. NOMENCLATURE. Although A. H. Clark (1954) pointed out that Asterias clathrata Say was preoccupied by Pennant's earlier senior homonym, he commented that 'nothing would be gained by displacing this well established name' and so continued to use it for this common West Indian species. Unfortunately, under the rules of nomenclature (Article 59(a)) such a junior primary homonym 'must be permanently rejected', that is unless the International Commission can be prevailed upon to use its plenary powers to suppress the earlier name (itself a synonym of Asterias rubens Linnaeus, 1758), which action is being requested in a separate proposal. DISTRIBUTION. North Carolina (??New Jersey), Bermuda and the Gulf of Mexico to southern Brazil (c. 3 1° S); 0-175 metres but rarely below 70 metres. Luidia heterozona Fisher Figs2f,i,j,3j, r Luidia heterozona Fisher, 1940:265-268, fig. M4, pi. 23; Madsen, 1950:203-205; John & A. M. Clark, 1954 : 148; A. M. Clark, 1955 : 33; Cherbonnier, 1963 : 182; Nataf & Cherbonnier, 1973 : 74-76, pi. 4, figs A, B, pi. 5, figs 3-6, pi. 9, figs I, J; Sibuet, 1975 : 284, 288. Luidia mortensem Cadenat, 1941 : 53-67, figs 1-3; Cherbonnier, 1963 : 182. Luidia barimae John & A. M. Clark, 1954: 145-148, figs 3, 4, pi. 6, fig. 2; Downey, 1973 : 25; Walenkamp, 1976: 18-1 9, pi. 1, figs 1-3. R up to 255 mm; R/r 7-1 1/1. Of over 150 specimens from both sides of the Atlantic taken by the Gerda, Pillsbury and Oregon, three have 9 arms, all the rest 10, though Madsen (1950) found 2 of the 37 Atlantide specimens to have 9 arms and another only 8. NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 7 1 A species of Luidia with usually 10 arms, relatively slender and attenuated; abactinal paxillae all relatively small and rounded but rather variable in size medially, some slightly larger ones intermingled; laterally two matching longitudinal series each side, outnumbering the adjacent more elongate superomarginal paxillae by 15-18/10; inferomarginal plates mainly lateral in alignment, bearing up to four spines in L. heterozona barimae but rarely more than three in L. heterozona heterozona, the spines erect, long and acute, the uppermost (at least on alternate plates) projecting vertically upwards and the uppermost or the second the longest, the spines alternating fairly regularly in level on successive plates and often also in number 4/3 or 3/2, the plates unusually bare of spinelets between the large spines except at the edges; actinal plates unusually elongate, partially overlying the abradial ends of the adambulacrals; adambulacral plates mostly with three spines in line at right angles to the furrow but in larger specimens, R>70 mm, c. 10-15 proximal plates may have a fourth spine enlarged proximal to the abradial one; pedicellariae bivalved throughout, often very abundant, especially on the paxillae (Fig. 3j) where they are relatively short and rounded, their edges almost fully contiguous, actinal pedicellariae longer, sometimes broadened at the tips, numbering up to three on some proximal actinal plates, similar ones on the infero- marginals and oral plates while one to three more spiniform pedicellariae also occur on the furrow face of each oral plate near the mouth. Colour in life dark chocolate brown above except for a white band along the upper edge of each arm (possibly the superomarginals), the inferomarginal spines white with brown bases. STATUS OF L. barimae. Luidia barimae was established on the basis of two poorly preserved specimens from off Venezuela. Walenkamp (1976) has recorded nine others from Surinam, evidently also in poor condition. About 50 further specimens from the Pillsbury collections off the north coast of South America have now been studied, as well as more than 100 from West Africa of L. heterozona. Though many of these are also poor, some are good enough to show that there are two matching regular longitudinal series of lateral paxillae adjacent to the superomarginal paxillae in the american and well as african specimens. Also the supero- marginal paxillae are similarly markedly larger than the other paxillae (Fig. 2f). This leaves only the number of inferomarginal spines as a possible distinction between the two, L. barimae being described as having four spines proximally, then three, whereas no more than three, more often three and two, are said to be found in L. heterozona by Fisher, Madsen and Cadenat. However, one large Pillsbury specimen from the Gulf of Guinea, West Africa, with R at least 1 50 mm, was found to have four inferomarginal spines on a few proximal alternate plates, while conversely a specimen from Venezuela with Re. 120 mm has no more than three spines, though these reduce to two on the alternate distal plates. Certainly there is no justification for more than a subspecific distinction between specimens from the two sides of the tropical Atlantic and the minor difference in the marginal spines is clearly correlated with different growth rates. DISTRIBUTION. L. heterozona heterozona from Cap Blanc, Mauritania, south to Elephant Bay, Angola; 28-975 metres. As noted above (p. 163), apart from Sibuet's Walda Expedition record (1975), the greatest depth recorded is 400 metres. L. heterozona barimae from off the Gulf of Maracaibo, western Venezuela (c. 12°34'N:71010'W) eastwards to French Guiana (06°07' N : 52°19' W); 38-90 (7100) metres. Luidia ludwigi scotti Bell Figs2a, e, 3t, u Luidia scotti Bell, 1917 : 8-9; A. M. Clark, 1953 : 383-385, fig. 3, pi. 40, fig. 1; John & A. M. Clark, 1954 : 144; Carrera-Rodriguez & Tommasi, 1977 : 62, 65-66. Luidia doello-juradoi Bernasconi, 1941 : 117; 1943 : 8-11, pi. 1, fig. 3, pi. 2, figs 2, 3, pi. 3, figs 4, 5. Luidia rosaurae John & A. M. Clark, 1954: 142-145, figs 1,2, pi. 6, fig. l;Jangoux, 1978 : 95. Luidia ludwigi: Walenkamp, 1976 : 32-37, fig 9, pi. 2, figs 1, 2, 4, pi. 4, fig. 3. R up to 90 mm; R/r 5'3-8'l/l, mean of 9 specimens 7*0/1 . 172 A.M.CLARK A species of Luidia with normally 5 arms: abactinal paxillae with two longitudinal rectangular or squarish lateral series each side, matching each other and also the adjacent slightly smaller superomarginal paxillae, central paxillar spinelets distinctly coarser than the peripheral ones and in smaller specimens, R90 mm. DIAGNOSIS. A species of Luidia with normally 5 arms, becoming long and evenly tapering in large specimens, R>90 mm, but somewhat petaloid in smaller ones; abactinal paxillae relatively small and rounded, with two matching slightly coarser longitudinal lateral series each side but these are inconspicuous, outnumbering the markedly elongate superomarginal paxillae by 17-20/10, paxillar armament of fine spinelets of fairly uniform length, median paxillae mostly with only a single central spinelet, sometimes distinctly coarser in L. 176 A.M.CLARK sarsi sarsi where the peripheral spinelets are also less attenuated than in L. sarsi africana and L. sarsi elegans (see Fig. 3m, n, o); inferomarginal plates mainly lateral in alignment, bearing 2-4 usually 3, large pointed spines, showing some tendency to alternate in position and number on successive plates, the two upper spines similar in length, the uppermost often longer when not at its highest position; actinal plates distinctly broadened and keeled trans- verse to the arm axis; adambulacral plates with three large spines in series transverse to the furrow and sometimes one (or two) enlarged spinelets proximal to the lateralmost; pedicellariae bivalved with fairly broad rounded tips, usually almost circular in cross section but the valves sometimes rather flattened towards the tips, common on the median abactinal paxillae of L. sarsi africana and L. sarsi elegans, occasional in L. sarsi sarsi but longer actinal pedicellariae usually present in all three subspecies proximally, with one on the furrow face of each oral plate (exceptionally represented by two hardly modified spines). Colour in life brownish yellow, reddish or orange above, pale below, arms medially and marginal plates darker, especially in L. sarsi africana. STATUS OF L. africana AND L. elegans. These two taxa of Sladen, 1889 and Perrier, 1875 are here regarded as conspecific with L. sarsi. Confusion in the limits between L. sarsi and L. africana has arisen because the type material of the latter included one specimen from Atlantic Morocco besides the main sample from South Africa, one of the latter conforming to the size and description given by Sladen, as I noted in 1953 (p. 393). The moroccan specimen has fairly numerous globose pedicellariae on the abactinal paxillae, as in the southern specimens. Although such pedicellariae were thought to be rare in north european specimens of L.. sarsi, some larger specimens from west of Scotland collected since 1953 were found to have more or less numerous pedicellariae (Fig. 3m), sometimes two or three on a single paxilla. It is significant that there are no reliable identifications of L. africana from any localities between Cape Verde and Luderitz Bay, Namibia. Despite a critical comparison by Nataf & Cherbonnier (1973) out of 213 five-armed specimens of the ci/zam-group (excluding L. sagamina aciculatd) from this area, every one was found to be referable to L. atlantidea and not to L. africana. Doderlein's record (1920) of L. africana from the Cape Verde Islands (based on two specimens named L. sarsi by Studer in 1884) is clearly in mistake for L. atlantidea since he particularly notes the relative coarseness of the central paxillar spinelets and the occurrence of pedicellariae on the superomarginal but not the abactinal paxillae — the reverse of what is found in L. africana, where both central and peripheral paxillar spinelets are uniformly fine. Madsen (1950) has referred Mortensen's larger moroccan specimens also to L. atlantidea, leaving under the heading of L. africana only some smaller moroccan ones which 'might just as well belong to L. sarsi' and three fragmentary Atlantide specimens which 'are not immediately recognizable as L. africana'. Zoogeographically, it would be expected that the distribution of a northern species also found in the Mediterranean, could well extend south- wards to some extent in north-west Africa, as with Marthasterias glacialis, for instance, which also occurs in South Africa but is unrecorded between Cape Verde and Cape Town. There are in the British Museum collections six specimens from near Cap Blanc (Discovery stations 8005 and 8020, 101 and 261-297 metres) with R 32 to c. 140 mm. These are certainly not L. atlantidea, having uniformly fine paxillar armament, abactinal rather than superomarginal pedicellariae, no white lines emphasizing the positions of the supero- marginals but darker brown midlines to the arms, much as in L. sarsi and L. africana. These specimens are a little unusual in having the pair of lateral spines on the furrow face of each oral plate hardly modified, if at all, into a pedicellaria. One exception is the smallest one, surprisingly since it is usually in larger specimens of species such as L. heterozona that this pedicellaria is more modified. Nearly all the specimens of L. sarsi, africana and elegans seen have quite a well-developed oral furrow pedicellaria, though the two spines from which it is modified may be more or less unequal in size. A second feature of some of these Cap Blanc specimens is that some proximal superomarginal paxillae are unusually broad, almost square, rather than consistently elongated, squarish paxillae being a characteristic of L. NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 77 atlantidea. This is naturally limited to the plates adjacent to those inferomarginals where the highest spine is not at the upper edge of the plate. In describing L. africana, Sladen (1889) cited seven supposed differences from L. sarsi, the larger size (R up to 160 mm in the type material), the more even taper of the arms, the smaller size of the median paxillae, the lesser prominence of the central median paxillar 'granule' (spinelet), the greater length of the uppermost inferomarginal spine relative to the second spine, the greater length of the third (lateralmost) adambulacral spine, achieving equality with the second spine, and the less pointed form of the actinal pedicellariae. Subse- quently Madsen (1950) noted that abactinal pedicellariae are not only much more often found in L. africana than in sarsi, where they are rare, but also that their position, when present in sarsi, is peripheral, while those of africana are central on the paxillar tabulum, so that the pedicellariae of sarsi tend to be seen in profile rather than from the top. He also noted that the inferomarginal spines of L. africana differ in having dark pigmented skin basally. As far as size is concerned, although in Norway, the type locality of L. sarsi, larger specimens usually have R only 50-60 mm, around Scotland it may reach c. 1 10 mm and in the Mediterranean reputedly c. 180 mm (diameter 35 cm). Such larger specimens also have the arms more attenuated and evenly tapering and there is greater disparity in the size of the additional median paxillae. As for the relative lengths of the inferomarginal and adambulacral spines, these are dependent to some extent on growth, the lateralmost adambulacral spine being retarded in growth relative to the second spine, while it would be more correct to say that the two upper inferomarginal spines of L. sarsi are similar in length, the uppermost one often slightly longer, especially on those plates where it is not in its highest position, as is the case in the lectotype now proposed (see below), even though R is less than 40 mm. At R 100+ mm the third spine approximates in length to the other two in specimens from all localities, a characteristic thought by Doderlein (1920) to be diagnostic of L. elegans. With regard to the occurrence of abactinal pedicellariae, in 1953 I noted that in four out of ten northern specimens of L. sarsi studied, some pedicellariae were present. As mentioned above, subsequent Scottish collecting has yielded material with pedicellariae as numerous as in the moroccan paratype of L. africana and some south african specimens. In 1953 I also contended that the pedicellariae are not really central on the paxillae of L. africana (or L. elegans) either but that this may appear so when the paxillae are crowded in preservation and the valves are coarser, pushing the pedicellariae into a more nearly vertical alignment on the tabulum. There is rarely a complete circlet of spinelets, as shown in Mortensen's fig. 3 (1933a) and Madsen's fig. 4c, d (1950). The shape of the pedicellariae, both abactinal and actinal ones, varies to some extent in different specimens from both north and south. Viewed end-on, the outline is usually approximately circular and the shape appears globular but may be slightly oval in one plane or the other if the valves are either thickened or broadened. The very broad shapes of both kinds of pedicellaria shown in Madsen's fig. 4h, i and k (1950), the abactinal ones cockle-shaped and the actinal ones with the individual valves bat-shaped (fan-shaped according to Madsen) seems to be exceptional even among south african specimens, none of the ten now studied having such an exaggerated form. Usually their actinal pedicellariae are either tapered in both side views (like a bishop's mitre) or else the tips are blunter and often thicker, a range also shown by northern specimens. Finally, the shape of the abactinal paxillar spinelets needs to be compared. In many northern specimens of L. sarsi the majority of median paxillae do have the single central spinelet distinctly coarser than the peripheral ones, appearing almost granuliform as foreshortened. This is particularly, but not exclusively, true of smaller specimens, including the lectotype, but in other specimens the thickness of all the paxillar spinelets is similar. A comparable modification of the central paxillar spinelet is noted above in some smaller specimens of L. ludwigi scotti and of course in a much more exaggerated form in L. sagamina aciculata, where the central spinelet is more or less markedly elongated as well as being much thicker than the peripheral spinelets. In general, the paxillar spinelets of south 178 A.M.CLARK african specimens are appreciably more attenuated than those of european specimens and none have been observed to show a coarsening of the central spinelet, though it should be emphasized that no specimens with R< 70 mm are available. To sum up, in comparison between specimens from South Africa and those from Europe and NW Africa, no differences of specific weight can be found. There are slight differences in the armament of the paxillae in some specimens, the southern ones tending to have a finer and more uniform armament but this seems to be shared by some moroccan specimens and is always very subtle. Although the frequency of pedicellariae seems to increase further south, only in occasional south african specimens with extremely broad pedicellariae are these organs distinctive. It is therefore doubtful whether the name africana is worth retaining for a subspecies from South Africa but the distribution gap in west Africa coupled with these minor morphological differences may justify it. With regard to Luidia elegans from the american side of the Atlantic, the only comparison given by Perrier was of the ventral side with that of L. alternata just described, the difference emphasized being the bivalved rather than trivalved pedicellariae. In Doderlein's key (1920), he distinguished L. elegans from the much more closely related L. sarsi and L. africana by the equal length of all three inferomarginal spines being supposedly limited to L. elegans, which was also said to be the only one to have numerous paxillar pedicellariae. This comparison was evidently made primarily on the evidence of a single small specimen of L. sarsi (R 29 mm), one of L. elegans with R 145 mm and two specimens from the Cape Verde Islands which he thought to be L. africana but are more likely L. atlantidea. Inevitably, in smaller specimens the lowest inferomarginal spine, if developed at all, is relatively smaller, while at most stages of growth the two upper spines are similar in length, as discussed above. At Re. 100 mm in European specimens all three spines may be about equal, while the occurrence of pedicellariae is clearly variable and of little taxonomic weight. However, there does appear to be a significant difference in the number of abactinal paxillae across the arm, at least in smaller specimens, R 40-50 mm, on the two sides of the Atlantic. Excluding the two series of superomarginal paxillae, the number in american specimens of about this size is proximally c. 13 compared with c. 17 in those from the East Atlantic. Unfortunately, owing to their irregular arrangement, it is difficult to make a precise estimate of the number of paxillae, especially in larger specimens. Also, the limited amount of material available indicates that the paxillar number may increase at a higher rate in american specimens. There is a small degree of support for this character in the armament of the adambulacral plates. At R c. 40 mm, the third (lateralmost) spine is much smaller than the second in american specimens and even at R 70 mm this spine is still much narrower basally than the second and only about two-thirds as long. Only when R approaches 140 mm does the third spine achieve parity in magnitude with the second on most plates. As for the accessory adambulacral spinelets, in specimens from both sides of the Atlantic one (sometimes two) is usually somewhat enlarged proximal to the third spine (or the space between second and third spines). This spinelet is usually one-third to half, but occasionally as much as three-quarters as long as the third spine. In american specimens this accessory spinelet(s) remains slender but in the eastern Atlantic can be basally stouter and more conical in shape. However, there is so much variation in the adambulacral armament that these differences cannot be of much importance. Finally, the madreporite may remain obscured by the paxillae to a greater extent in specimens from the eastern Atlantic than in american ones but this is probably correlated with size (the plate becoming more prominent in larger specimens) and is affected by preservation. In total, these minor differences do not add up to more than a subspecific distinction between L. elegans and L. sarsi. Finally, Koehler (1923) and Carrera-Rodriguez & Tommasi (1977) have recorded specimens from Uruguay (33° S) and southern Brazil (c. 30^° S) as L. africana or L. elegans. Because of the great geographical discontinuity between them and the rest, comparable to that between L. sarsi sarsi and L. sarsi africana, it is not improbable that a minor morpho- logical difference justifying a taxonomic distinction of eastern south american specimens may exist, most likely again at the subspecific level. NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 179 LECTOTYPE. Because of the complex affinities of Luidia sarsi, it seemed desirable that a lectotype be selected from among Sars' material from Bergen to which Diiben & Koren gave this name in 1845. Thanks to Dr M. E. Christiansen of the Zoologisk Museum, Oslo, it is clear that no type was designated at the time. The specimen chosen (one of seven) was from Manger, near Bergen, Oslo Museum reg. no. E1005a, shown in Fig. 6. R (maximum)/r is 37/8 mm, = 4-6/1. The shorter arms have R 30 and 32mm. (The relatively low R/r ratio is to be expected at this relatively small size.) About 17 lateral paxillae correspond to 10 of the more proximal superomarginal paxillae. The median paxillae nearly all have a single, slightly coarser, central paxillar spinelet and usually 7-1 1 peripheral spinelets. Excluding the superomarginal ones, there are c. 17 paxillae across the arm breadth basally. The paxillae of the outer lateral row have up to three central spinelets. Fig. 6 Luidia sarsi sarsi Diiben & Koren, lectotype, Zoologisk Museum, Oslo reg. no. E1005a, Manger, near Bergen. Dorsal view, x 2-2. 180 A.M.CLARK All the spinelets of the lateral paxillae are distinctly coarser than those of the median ones, while the superomarginal armament is coarser still. The madreporite is not distinguishable. The inferomarginal plates bear 2-4 large spines, rather variable in level on the plate, sometimes alternating, though at the same level on several consecutive plates. The uppermost spine is usually slightly the longest. Most proximal actinal plates bear a single blunt, bivalved pedicellaria but no abactinal pedicellariae were observed. Each adambulacral bears 3 large spines and an enlarged spinelet one-third to half as long as the lateralmost one and proximal to it. Each oral plate bears about four large spines in an abradial row and one other spine near the furrow margin, besides a large bivalved pedicellaria in the furrow. The paralectotypes include four smaller specimens and two larger but broken ones, one with R c. 50 mm and the other R/r 55/9*5. The central spinelet of the median paxillae is also distinctly coarser than the peripheral ones; c. 20 lateral paxillae correspond to 10 superomarginal ones; proximally there are usually four inferomarginal spines. HOLOTYPE OF Luidia elegans. Because of Perrier's rather inadequate description, this specimen was borrowed for reexamination from the Museum of Comparative Zoology, Harvard. The number is M.C.Z. 372. It proved to be in fact the specimen illustrated in Perrier, 1884, pi. 10, fig. 7, under the name of L. barbadensis, though the negative was reversed in printing. One arm is abnormally short, probably regenerated, and curled up at the tip. R is c. 40 mm for the longest arm (Perrier gives 35 mm); r is 7 mm; 1 8 lateral paxillae correspond to 10 superomarginal ones; proximally there are c. 13 paxillae across the arms between (but not counting) the superomarginal ones. The median paxillae mostly have only a single central spinelet, not at all coarser than the peripheral ones. No abactinal pedicellariae were seen. Most inferomarginal plates bear three large spines, sometimes two, showing a tendency for alternation in position. The actinal pedicellariae are bivalved and blunt-tipped. The third adambulacral spine is reduced on some plates. The locality is 'Straits of Florida, 101 fathoms'. Verrill's guess that Perrier's '101 brasses' meant feet being incorrect. DISTRIBUTION. L. sarsi sarsi from Trondheim Fjord, Norway to Cap Blanc, Mauritania, the Azores and the Mediterranean; 9-1 300 metres. L. sarsi africana from Luderitz Bay to the Port Elizabeth area of South Africa; 54-360 metres. L. sarsi elegans from east of New Jersey, U.S.A. (c. 41° N) to the Florida Strait and both east and west Gulf of Mexico; also from southern Brazil (c. 30f S) to Uruguay (c. 33° S); 60-365 metres. There are no reliable records from the Caribbean, Perrier's supposed specimens from Barbados being untraced in the M.C.Z. and it is significant that no specimens appear to have been taken by the intensive collections of the Atlantis around Cuba (H. L. Clark, 1941) or by the Pillsbury in the Caribbean, Lesser Antilles or north of South America. Luidia senegalensis (Lamarck) Asterias senegalensis Lamarck, 1816 : 567. Luidia senegalensis: Miiller & Troschel, 1842 : 78, pi. 5, fig. 4; Perrier, 1875 : 342-343 [1876 : 262]; Doderlein, 1920 : 249-250, figs 9, 20; H. L. Clark, 1933 : 20-22; Bernasconi, 1943 : 5-6; Tommasi, 1958:9-11, pi. 2, fig. 2; Ummels, 1963:94-95, pis 10, 11; Downey, 1973:22, pi. 1, figs A, B; Blake, 1973 : 30, pi. 2, figs 1-31; Walenkamp, 1976 : 25-29, fig. 5, pi. 1, fig. 4; 1979 : 12, pi. 1, figs 1-3. Luidia marcgravii Steenstrup in Lutken, 1859 : 43^6; Verrill, 1915 : 208-209; Boone, 1933 : 76, pis 33-36; Bernasconi, 1958 : 125-127. REMARKS. A very large specimen from Pillsbury st. 750, off Venezuela, 22-26 metres with R up to 252 mm may provide a size record. This locally common Caribbean species with its numerous and particularly attenuated NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 g 1 arms seems even more vulnerable than most Luidias to loss and regeneration of the arm tips, probably due to predation, possibly cannibalism. DISTRIBUTION. Sporadically in southern Florida and from Jamaica eastwards along the Antilles, also Belize and Nicaragua and along the north coast of South America to southern Brazil (Santa Caterina, c. 21° S); the original record from Senegal has never been repeated; the records from Belize and Nicaragua are new and derived from the Pillsbury collections; 1-64 metres, the depth range also being extended by a Pillsbury station (745, off Venezuela, c. 12° N, 67° W); the previous maximum depth was 45 m off French Guiana. Summary of taxonomic changes The number of nominal Atlantic species of Luidia is reduced by the present study from a one-time maximum of 1 7 to 1 1 . Luidia aciculata Mortensen, 19336 is retained as a subspecies of the Indo-West Pacific L. sagamina Doderlein, 1920. L. africana Sladen, 1 889 is reduced to a subspecies of L. sarsi Diiben & Koren, 1 845. L. barimae John & Clark, 1954 is reduced to a subspecies of L. heterozona Fisher, 1940. L. bernasconiae A. H. Clark, 1945 is again synonymized with L. alternata (Say, 1825), having been revived by Gray, Downey & Cerame- Vivas, 1968. L. elegans Perrier, 1 875 is reduced to a subspecies of L. sarsi. L. numidica Koehler, 19 1 1 is treated as a subspecies of L. alternata. L. quequenensis Bernasconi, 1942 is synonymized with L. alternata. L. rosaurae John & Clark, 1954 is synonymized with L. scotti Bell, 1917, which is treated as a subspecies of the East Pacific L. ludwigi Fisher, 1906. Acknowledgements Thanks are due to Miss M. E. Downey of the Smithsonian Institution for access to the Gerda and Pillsbury luidiids; also to Dr M. E. Christiansen of the Zoologisk Museum Oslo, and Professor R. M. Woollacott of the Museum of Comparative Zoology (M.C.Z.), Harvard, for the loan of material. References Bell, F. J. 1893. Catalogue of the British Echinoderms in the British Museum (Natural History). London xvii + 202 pp., 16 pis. 1917. Echinoderma. 1. Actinogonidiata. Brit. Antarct. (Terra Nova) Exped. 1910. Zool. 4 (1): 1-10, 2 pis. Bernasconi, I. 1 94 1 . Dos nuevas especies argentinas de "'Luidia'". Physis, B. Aires 19 : 117-118. 1942. Los Asteroideos sudamericanos de la familia "Luidiidae". Physis, B. Aires 19 : 252-253. 1943. Los Asteroideos sudamericanos de la familia Luidiidae. An. Mus. argent, B. Aires 41 : 1-20, 5 pis. 1958. Equinoideos y asteroideos de la collection del Institute Oceanografico de la Universidad de San Pablo. 2. Bol. Inst. oceanogr. S. Paulo 1 : 1 19-149, 4 pis. 1960. Los equinodermos de la campana 1958 del buque oceanografico Capitdn Cdnepa A.R.A. Adas Trab. I Congr. sudamer. Zool., 1959. 2 : 2 1-32, 3 pis. Blake, D. B. 1973. Ossicle morphology of some recent asteroids and description of some west American fossil asteroids. Univ. Calif. Publs geol. Sci. 104 : 1-59, 3 figs, 19 pis. 1982. Somasteroidea, Asteroidea and the affinities of Luidia (Platasterias) latiradiaia (Gray), 1 840, Palaeontology 25 : 1 67-1 9 1 . Boone, L. 1933. Coelenterata, Echinodermata and Mollusca. Bull Vanderbilt mar. Mus. 4 : 1-217, 133 pis. 182 A.M.CLARK Cadenat, J. 1941. Les Echinodermes de la cote occidentale d'Afrique. Description d'une asterie nouvelle de la region du Cap Blanc. Ann. Soc. Sci nat. Charente Inf. N.S. 3 : 53-67, 3 figs. Carrera-Rodriguez, C. J. & Tommasi, L. R. 1977. Asteroidea de la plataforma continental de Rio Grande do Sul (Brasil). Bolm Inst. oceanogr. S. Paulo 26 (1 ): 5 1-1 30, 33 figs. Cherbonnier, G. 1959. Echinodermes de la Guyane franchise. 2. Bull. Mus. natn. Hist. nat. Paris 31 : 168-1 72, figs 3, 4. 1963. Echinodermes des cotes du Cameroun recoltes par A. Crosnier. Bull. Mus. natn. Hist. nat. Paris 35: 179-193,3 figs. Clark, A. H. 1945. A new starfish of the genus Luidia from the coast of Georgia. J. Wash. Acad. Sci. 35: 19-21. 1954. Echinoderms (other than holothurians) of the Gulf of Mexico. Bull. U.S. Fish. Commn 55 : 373-379. Clark, A. M. 1952. Some echinoderms from South Africa. Trans. R. Soc. S. Afr. 33 : 193-221, 3 figs, Ipl. 1953. Notes on asteroids in the British Museum (Natural History). 3. Luidia. 4. Tosia and Pentagonaster. Bull. Br. Mus. nat. Hist. Zool. 1 : 379-412, 1 5 figs, 8 pis. 1955. Echinodermata of the Gold Coast. Jl W. Afr. Sci. Assn 1 : 16-56, 23 figs, 1 pi. 1977. The South African Museum's Meiring Naude cruises. 4. Echinoderms. Annls S. Afr. Mus. 73: 133-147. Clark, A. M. & Court man-Stock, J. 1976. The echinoderms of southern Africa. Publs Br. Mus. nat. Hist. No. 776, 277 pp., 276 figs. Clark, H. L. 1923. The echinoderm fauna of South Africa. Annls S. Afr. Mus. 13 : 221^35, 4 figs, 6 pis. 1933. A handbook of the littoral echinoderms of Porto Rico and the other West Indian islands. Scient. Surv. P. Rico 16 : 1-147, 7 pis. 1941. Reports on the scientific results of the Atlantis Expeditions to the West Indies. The echinoderms. Mem. Soc. cubana Hist. nat. 15 : 1-54, 10 pis. Doderlein, L. 1920. Die Asteriden der Siboga-Expedition. 2. Die Gattung Luidia und ihre stammesgeschichte. Siboga Exped. 46b : 193-291, 5 figs, 3 pis. Downey, M. E. 1973. Starfishes from the Caribbean and the Gulf of Mexico. Smithson. Contr. Zool. No. 126: 1-1 58, 48 pis. Duben, M. W. von (& Koren, J.) 1845. Norriges Hafs-fauna. Ofvers. K. VetenskAkad Forh. 1844 : 110-116. 1846. Ofversigt af Skandinaviens Echinodermer. K. svenska VetenskAkad. Handl. 1844: 229-328, 6 pis. Engel, H., Croes, A. F. & Schroevers, G. F. 1960. Les Asterides recoltees par 1'expedition du navire-ecole beige Mercator, 1935-36. A. Les Asterides recoltees sur les cotes orientales de 1'Amerique entre la Floride et la Venezuela. (Engel & Schroevers). B. Les Asterides de la cote occidentale d'Afrique. (Engel & Croes). Bull. Inst. roy. Sci. nat. Belg. 36 (54) : 1-10; 10-15, 6 pis. Fell, H. B. 1963. The phylogeny of sea-stars. Phil. Trans. 246B : 38 1^35, 2 pis. Fisher, W. K. 1911. Asteroidea of the North Pacific and adjacent waters. Bull. U.S. natn. Mus. 76 (1) : 1-419, 122 pis. - 1940. Asteroidea. 'Discovery' Rep. 20 : 69-316, 23 pis. Forbes, E. 1839. On the Asteriadae of the Irish Sea. Mem. Wernerian nat. Hist. Soc. 8 : 1 13-130, 2 pis. 1841. A history ofbritish starfishes and other animals of the class Echinodermata. xx + 270 pp., figs. London. Gray, I. E., Downey, M. E. & Cerame-Vivas, M. J. 1968. Seastars of North Carolina. Fishery Bull. Fish Wildl. Serv. U.S. 67 : 127-163, 40 figs. Gray, J. E. 1840. A synopsis of the genera and species of the class Hypostoma (Asterias, Linn.) Ann. Mag. nat. Hist. 6 : 1 75-1 84; 275-290. Jangoux, M. 1975. Note sur le genre Tethyaster Sladen (Echinodermata, Asteroidea). Revue zool. afr. 89 (4): 76 1-768, 4 figs. - 1978. Les etoiles de mer (Echinodermata: Asteroidea) recoltees par le M/V Calamar au large des Guyanes et du Venezuela. Bull. zool. Mus. Univ. Amsterdam 6(13): 93-101 , 3 figs. John, D. D. & Clark, A. M. 1954. The "Rosaura" Expedition. 3. Echinodermata. Bull. Br. Mus. nat. Hist. Zool. 2 (6) : 139-162, 12 figs, 1 pi. Koehler, R. 1895. Notes Echinologiques. Revue biol. N. Fr. 7 : 317-342, 1 pi. NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 183 — 1909. Echinodermes provenant des campagnes du yacht Princesse- Alice. Result. Camp, scient Prince Albert 34 : 1-3 1 7, 32 pis. 1911. Mission Gruvel sur la cote occidentale d'Afrique. Echinodermes. Annls Inst. oceanogr. Monaco 2 (5): 1-25, 3 pis. — 1914. Asteroidea, Ophiuroidea et Echinoidea. Beitr. Kennt. Meeresfauna Westafr. 1 (2) : 129-303, 12 pis. — 1921. Echinodermes. Faune Fr. 1 : 1-210, 153 figs. 1923. Asteries et Ophiures recueillis par 1'expedition antarctique suedoise 1901-03. Further zool. Results Swed. Antarct. Exped. I (1) : 1-145, 1 fig., 15 pis. Lamarck, J. B. P. A. de 1816. Histoire naturelle des animaux sans vertebres. Ed. 1. 2: 522-568: 3 : 1-59; 60-76. Paris. Linnaeus, C. 1758. Systema Naturae. Ed. 10. 1 : 824 pp. Holmiae. Ludwig, H. 1 897. Die Seesterne des Mittelmeeres. Fauna Flora Golf. Neapel 24 : 1-49 1 , 1 2 pis. Liitken, C. 1859. Bidrag til Kundskab om de ved Kysterne af Mellem- og Syd-America levende arter af Sostjerner. Vidensk. Meddrdansk. naturh. Foren. 1859 : 25-96. Mcknight, D. G. 1977. Classification of recent paxillosid sea-stars (Asterozoa: Echinodermata). NZOIRec. 3(12) : 1 13-1 19, figs. Madsen, F. J. 1950. The echinoderms collected by the Atlantide Expedition, 1945-46. 1. Asteroidea. Atlantide Rep. 1 : 167-222, 1 1 figs, 3 pis. Marenzeller, E. von 1893. Veroffentlichungen der Commission fur Enforschung des ostlichen Mittelmeeres. Sber. Akad. Wiss. Wien Math, naturw. Kl. 102 : 1-5. Mortensen, T. 1925. Echinodermes du Maroc et de Mauritanie. Bull. Soc. Sci. nat. Maroc 5 : 178-187, I pi. 1 927. Handbook of the Echinoderms of the British Isles, ix + 47 1 pp., 269 figs. London. 1933a. Echinoderms of South Africa (Asteroidea and Ophiuroidea). Vidensk. Meddr dansk naturh. Foren. 93 : 2 1 5^400, 9 1 figs, 1 2 pis. 19336. The echinoderms of St Helena. Vidensk. Meddr dansk naturh. Foren. 93 : 401-472, 29 figs, 3 pis. Miiller, J. & Troschel, F. H. 1840. Mber. Akad. Wiss. Berlin 1840 : 100-106. 1842. System der Aster iden. 1 . Asteriae. 2. Ophiuridae. xx + 1 34 pp., 12 pis. Braunschweig. Nataf, G. & Cherbonnier, G. 1973. Les asterides d'Afrique occidentale, utilisation du microscope electronique a balayage pour etude systematique des Luidia. Bull. Mus. natn. Hist. nat. Paris (Zool) No. 81 : 69-101, 9 pis. Pennant, T. 1 777. British Zoology. Ed. 4. 4 : 1-1 54, 93 pis. London. Perrier, E. 1 869. Recherches sur les Pedicellaires et les Ambulacres des Asteries et les Oursins. 1 88 pp., 2 pis. Paris [Also in Annls Sci. nat. 12[1869] : 197-304; 13[1870] : 1-81.] 1875. Revision de la collection de Stellerides du Museum d'Histoire Naturelle de Paris. 384 pp. Paris. [Also in Archs Zool. exp. gen. 4[1875] : 265^50; 5[1876] : 1-104; 209-309.] 1881. Description sommaire des especes nouvelles d'Asteries. Bull. Mus. comp. Zool. Harv. 9: 1-31. — 1882. In: Milne Edwards, A. Rapport sur les traveaux de la Commission chargee par M. le ministre de 1'Instruction publique d'etudier la faune sous-marine dans le grandes profundeurs de la Mediterranee et de 1'Atlantique. (Echinodermes). Arch. Missions sci. litt: Rapp. et Instr. ser 3, 9 : 46^9, fig. — 1884. Memoire sur les Etoiles de Mer recueillies dans la Mer des Antilles et le Golfe de Mexique. Nouv. Archs Mus. Hist. nat. Paris (2)6 : 127-276, 10 pis. 1894. Stellerides. Exped. scient. Travailleur- Talisman. 431 pp., 26 pis. Paris. Philippi, R. 1837. Ueber die mit Asterias aurantiaca verwandten und verwechselten Asterian der sicilianischen Kiiste. Arch. Naturgesch. 3 : 193-194. Sars, M. 1835. Radiater. Beskrivelser og Jagttagelser over nogle mdrkelige eller nye i Havel ved den Bergenske Kyst levende Dyr. pp. 39^6. Bergen. Say, T. 1 825. On the species of the Linnaean genus Asterias inhabiting the coast of the United States. J. Acad. nat. Sci. Philad. 5 : 141-154. Sibuet, M. 1975. Asterides abyssales de 1'Atlantique sud. Bull. Mus. natn. Hist. nat. Paris (Zool.) No. 199: 28 1-296, 2 figs, pi. Sladen, W. P. 1889. Asteroidea. Rep. scient. Results Voy. Challenger Zool 30 : 1-935, 1 18 pis. Spencer, W. K. & Wright, C. W. 1966. Asterozoans. In: Moore, R. C. [Ed.] Treatise on invertebrate paleontology. U. Echinodermata 3(1): 4-107, 89 figs. Geological Society of America Inc.: University of Kansas Press. 184 A.M.CLARK Studer, T. 1884. Verzeichniss der wahrend der Reise S.M.S. Gazelle urn die Erde 1874-76 gessammelten Asteriden und Euryaliden. Phys. Abh. K. Akad. Wiss. Berlin [1883, 1884] 3 : 1-64, 5 pis. Sussbach, S. & Breckner, A. 191 1. Die Seeigel, Seesterne und Schlangensterne der Nord- und Ostsee. Wiss. Meeresunters. KielN.S. 12 : 167-300, 3 pis. Tattersall, W. M. & Sheppard, E. M. 1934. Observations on the Bipinnaria of the asteroid genus Luidia. James Johnstone Memorial Volume: Lanes Sea-Fisheries Laboratory, Liverpool, pp. 35-61, 8 figs. Tommasi, L. R. 1958. Os equinodermas do literal de Sao Paulo. 2. Contrcoes avuls. Inst. oceanogr. S. Paulo No. 2: 1-2 7, 6 pis. 1970. Lista dos asteroides recentes do Brasil. Contrcoes Inst. oceanogr. Univ. S. Paulo No. 18 : 1-6 1,60 figs. 1974. Equinodermes do Brasil. 3. P ubl. Inst. oceanogr. Univ. S. Paulo No. 369 : 1-1 5, 12 figs. Tortonese, E. 1 965. Echinodermata. Fauna d'ltalia. 6 : xv + 422, 1 86 figs. Bologna. Ummels, F. 1963. Asteroids from the Netherlands Antilles and other Caribbean localities. Stud. Fauna Curacao 15 : 72-101 , 3 figs, 9 pis. [Also in Natuurwet. Stud. Suriname No. 30.] Ursin, E. 1960. A quantitative investigation of the echinoderm fauna of the central North Sea. Medd. Danmarks Fisk. Havundersog. N.S. 2 (24) : 1-204, 96 figs. Verrill, A. E. 1885. Results of the explorations made by the steamer Albatross off the northern coast of the United States in 1883. Rep. U.S. Fish. Commn 1883 : 503-543, pis 13-19. 1899. Revision of certain genera and species of starfishes with descriptions of new forms. Trans. Conn. Acad. Arts Sci. 1899 : 145-234, pis 24-30. 1915. Report on the starfishes of the West Indies, Florida and Brazil. Bull. Labs nat. Hist. St. Univ. /a 7(1): 1-232, 29 pis. Walenkamp, J. H. C. 1976. The asteroids of the coastal waters of Surinam. Zoo/. Verh. Leiden No. 147: 1-91, 16 figs, 18 pis. 1979. Asteroidea (Echinodermata) from the Guyana shelf. Zoo/. Verh. Leiden No. 170 : 1-97, 31 figs, 20 pis. Zoppi de Roa, E. 1967. Contribucion al estudio de los Equinodermos de Venezuela. Acta biol. venez. 5: 267-333, 29 figs. Manuscript accepted for publication 25 June 1 98 1 New and little known species of Oncaeidae (Cyclopoida) from the Northeastern Atlantic S. J. Malt Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD Introduction A large number of new species and genera belonging to the Oncaeidae have been described in recent years. Many common species have been re-examined and recognized as polytypic, comprising two or more varieties (Tanaka, 1960; Moulton, 1973; Ferrari, 1975; Boxshall, 1977) or two or more species (Heron, 1977). Now that some of the better known species have been redescribed in greater detail it is obvious that many of the newly described species are also in need of more exact description. Taxonomic study of oncaeids has become a very laborious process, demanding the recognition of relatively small morphological differences in these microscopic animals, half the known species of which are less than O6 mm in body length. Most records of oncaeids are from the epipelagic zone, but the more recent discovery of several species extending down into the deep bathypelagic zone (below 2000 m) indicates that previous records reflect the limitations of the sampling programmes rather than any true restrictions in their depth range. A new species, Oncaea heronae, and two little known species belonging to the genus Oncaea are here described. The genera Conaea, Epicalymma and Myctospictosum are synonymized with Oncaea. New locality records are presented for six species and two previously undescribed males and two juveniles of known species are also described. Materials and methods The material described below was donated to the British Museum (Natural History) by the Institute of Oceanographic Sciences (Wormley) and the Marine Biological Association (Plymouth). It was collected in April 1977 at station 9541 (20° N 21° W) in the NE Atlantic during cruise 82 of the R.R.S. Discovery, and in September 1979 at the mouth of the English Channel (48° N 7°30' W) during a cruise of the R.V. Sarsia. The Discovery material was collected by a 80 jn mesh diatom net attached to the RMT 1 + 8 net system. Sarsia material was taken using a pump system with 80 n and 200 // mesh filters. Material was preserved in 10% sea water formalin, stained in chlorazol black, dissected in lactophenol and mounted in polyvinyl lactophenol. Drawings were made with the aid of a camera lucida. Specimens were measured using an eyepiece micrometer, total body length being the distance from the tip of the rostrum to the apex of the caudal rami. Descriptions of species Oncaea tregoubo vi Shmeleva, 1968 O. tregoubovi Shmeleva, 1968 : 1784-1785, figs 1-12. DESCRIPTION Female. Ratio of prosome and urosome lengths 1 -8 : 1 . Prosome elongate, oval in dorsal aspect (Fig. la). Mean body length of 3 specimens 0'34 mm (range 0'31 to Bull. Br. Mus. not. Hist. (Zool.) 42 (3) : 1 85-205 Issued 27 May 1 982 186 S. J. MALT 0*37 mm). Third prosome segment without a dorsal projection. Head (first prosome division) 1*3 times longer than wide. Proportional lengths of ursome and caudal rami 8 : 57 : 5:4: 13 : 13 (Fig. Ib). Genital apparatus located on dorsal surface anterior to midpoint of genital segment; each area armed with a setule. Maximum width of genital segment at level of genital apparatus; length to width ratio 1 '75 : 1 . Caudal rami twice as long as wide. First antenna 6-segmented (Fig. Ic); armature: 1-2, II-7, III-3, IV-3, V-2, VI-5 (some elements may be missing); ratio of segment lengths 10 : 13 : 46 : 13:8: 10. Second antenna 3 -segmented (Fig. Id); first segment bearing 1 long pinnate seta distally; second segment bearing a row of denticles along internal surface; terminal segment longer than second and bearing 5 curved spines and 2 curved setae. Mandible bearing 5 elements (Fig. le): a stout seta on external surface, 2 broad blades, 1 bearing a row of setules along internal edge, and the other unarmed, and 2 setae, 1 long and hirsute, the other small and unarmed. First maxilla bilobed (Fig. 1 f); bearing 1 seta on internal surface; 2 setae on internal lobe and 2 setae, a setose seta and a spine on external lobe. Second maxilla 2-segmented (Fig. Ig); second segment produced distally as an elongate, curved, bilaterally spinulose claw; also having an external pinnate seta and an internal bilaterally spinulose element. Maxilliped 4-segmented (Fig. 1 h); first segment unarmed; internal surface of second segment with 2 spines, distal spine bidentate; third segment reduced; terminal segment produced as a long claw and armed with a row of setules on concave surface, an internal basal spine and an external basal setule. Endopodites and exopodites of natatory legs 3-segmented (Figs 2a-d); external spines of exopodites bilaterally serrate; armature of natatory legs as follows: Coxa Basis Endopodite Exopodite Legl 0-0 1-1 0-1; 0-1; 0,1, 5 I-0;I-1; III, 1,4 Leg2 0-0 1-0 0-1; 0-2; II, 1, 3 1-0; !*-!*; Ill, 1, 5 Leg 3 0-0 1-0 0-1; 0-2; II, 1, 2 1-0; I-1;II,1, 5 Leg 4 0-0 1-0 0-1; 0-2; II, 1,1 I-0;I-1;II,1, 5 * = segment missing from figured specimen (Fig. 2b), present in other material. Terminal spines of exopodites longer than terminal segments bearing them but terminal spines of endopodites shorter than terminal segments; each terminal spine bordered by a serrate membrane unilaterally; endopodites of legs 1-3 terminate in conical projections; all setae on legs plumose. Fifth leg comprising a small cylindrical segment bearing a single terminal seta on body surface near leg (Fig. 1 i). MATERIAL EXAMINED. 399 from sample 9541.24, 2980-3560 m. 20° N 21° W. BM(NH) 1981.114. REMARKS. This species was identified by the following combination of characters: the relative lengths of the segments of the first and second antennae, the armature of the maxilliped and natatory legs, and the relative dimensions of the urosome segments. Some minor differences were found between the present material and the original material figured by Shmeleva (1968). Most notably Shmeleva (1968 : Fig. 9) shows the fourth natatory leg with the internal margin of the third endopodite segment unarmed, whereas the present specimens have a single seta. The absence of this seta is atypical for Oncaea species. It may be significant that Shmeleva does not mention this character in the text of the description, and the original drawing may be inaccurate or based on a specimen with incomplete armature. This species has only been recorded once before, from the southern Adriatic where it was taken between 200 and 300 metres depth. Oncaea ivlevi Shmeleva, 1966 O. ivlevi Shmeleva, 1966 : 932-933, Plate I, figs 1-1 1 . DESCRIPTION. Female. Ratio of prosome and urosome lengths 1-7:1 (Fig. 3a). Body length of 3 specimens 0*33 mm. Third prosome segment without a dorsal projection (Fig. 3b). Head NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 187 1 • 1 times longer than wide. Proportional lengths of urosome segments and caudal rami 8 : 50 : 7:6:18:11 (Fig. 3c). Genital apparatus situated anterior to midpoint of dorsal surface of genital segment; each area armed with a setule. Maximum width of genital segment at the level of genital apparatus; length to width ratio 2:1. Genital segment not markedly swollen. Caudal rami nearly twice as long as wide, curving outwards and each bearing a stout spine at external corner. First antenna armature similar to O. tregoubovi (Fig. 3d); ratio of segment lengths 19 : 19 : 35 : 12 \ 6 : 9. Second antenna armature similar to O. tregoubovi except that first segment armed with a row of setules on internal surface, and second segment with a row of spinules; terminal segment equal in length to second segment (Fig. 3e). Mandible and first maxilla lost during dissection. Second maxilla like that of 0. tregoubovi (Fig. 30- Maxilliped 4-segmented (Fig. 3g); first segment unarmed; internal surface of second segment with 2 spines, longer distal spine spinulose; second segment also bearing a row of denticles along internal surface; third segment reduced; terminal claw bearing a row of spinules along concave surface and an internal basal spine. Armature of natatory legs similar to O. tregoubovi (Figs 3h-k), but terminal segment of second endopodite bearing 1 spine on external surface instead of 2; endopodites 2, 3 and 4 tipped with relatively small conical projections; terminal spines of exopodites 1 and 2 slightly longer than terminal segments; those of exopodites 3 and 4 slightly shorter; all terminal spines of endopodites shorter than terminal segments and bilaterally flanged. Fifth leg comprising a small cylindrical free segment bearing 2 setae, outer double length of inner; and a seta on body surface near the leg. Male. Ratio of prosome and urosome lengths 1-6 : 1 (Fig 4a and b). Mean body length of 6 specimens 0-33 mm (range 0'28 to 0'35 mm). Head I'l times longer than wide. Proportional lengths of urosome segments and caudal rami 7 : 60 : 2 : 3 : 2 : 13 : 13. Genital lappets produced into small postero-lateral processes (Fig. 4c). Genital segment with length to width ratio 2-1 : 1 . Caudal rami twice as long as wide, as in female possessing a stout spine on each ramus. Mouthparts similar to those of female except first antenna and maxilliped. Three distal segments of the first antenna fused. Maxilliped 3 -segmented (Fig. 4d); first segment unarmed; second segment bearing 2 short spines and a row of setules on internal surface; terminal claw bearing spinules on concave surface and a stout external basal spine. Legs 1-5 similar to those of female. MATERIAL EXAMINED. 399 and 6dtf from sample F166-F170, 8(MO m, 48° N 7°30' W. BM(NH) 99 1981. 115-116,^ 1981. 117-1 19 and 1981. 124. REMARKS. This species was identified by the following combination of characters: the armature of the natatory legs, the relative dimensions of the urosome segments, and the presence of the stout caudal spine. The present specimens differ from Shmeleva's descriptions in a few minor details: in the possession of 3 terminal setae in association with the spine on the caudal rami rather than 2, the possession of an extra row of spinules on both the distal spine of the second segment of the female maxilliped and the terminal claw. There are also small differences in the lengths of the terminal exopodite spines. This species has also been recorded from the southern Adriatic where it was taken between 50 and 100 metres depth (Shmeleva, 1966) and from the Atlantic at 15 stations from 9° S 25° W to 9° S 45° W between 10 and 2000 metres depth (Shmeleva, 1969). Oncaea hispida (Heron, 1977) Comb. nov. Conaea hispida Heron, 1977 : 90-95, Figs 33h-j, 34a-k. DESCRIPTION. Female. Ratio of prosome and urosome lengths 1-75 : 1 (Fig. 5a). Body length of 1 specimen 0'56 mm. Third prosome segment without dorsal projection. Proportional lengths of urosome segments and caudal rami 8:48:7:7: 18: 12 (Fig. 5b). Genital apparatus located on dorsal surface anterior to midpoint of genital segment, each area armed 188 S. J. MALT with a setule. Maximum width of genital segment at level of genital apparatus; length to width ratio 1'6 : 1. Caudal rami 1*3 times as long as wide. Appendages similar to those described by Heron with only minor differences apparent (Figs 5c-h and 6a-d). Male. Ratio of prosome and urosome lengths 2 : 1 (Fig. 6e). Body length of 1 specimen 0'55 mm. Head 1-2 times longer than wide. Proportional lengths of urosome segments and caudal rami 10:51 :3:3:3: 18: 12 (Fig.. 6f). Genital lappets produced into small posterior processes. Genital segment with length to width ratio 1'5 : 1. Caudal rami approximately as long as wide. Mouthparts similar to those of female except first antenna and maxilliped. The 3 distal segments of first antenna fused. Maxilliped 3-segmented (Fig. 6g); first segment unarmed; second segment bearing 2 rows of dentiform processes on internal surface; terminal claw unarmed. Legs 1-5 similar to those of female. Fifth Copepodid (female). Ratio of prosome and urosome lengths 2 : 1 (Fig 6h). Body length of 1 specimen O46 mm. Head approximately as long as wide. Urosome 4-segmented (Fig. 6h). Proportional lengths of urosome segments and caudal rami 10 : 45 : 6 : 29 : 10. Genital segment without visible genital apparatus, length to width ratio 1'5 : 1. Caudal rami 1-2 times as long as wide. Appendages similar to those of adult female except that proximal seta on second segment of juvenile maxilliped is pinnate (Fig. 6j). MATERIAL EXAMINED. 19, Irf, 19 juvenile from sample 9541.24, 3980-3960 m, 20° N 21° W. BM(NH)1981.120-122. REMARKS. This species is here transferred from the genus Conaea to Oncaea (for discussion see page 191), and was identified by the following combination of characters: the length of the terminal segment of the second antenna and its armature, the armature of the maxilliped and the length of the third segment of the fourth endopodite. There are minor differences between the present material and Heron's description of the female in the ratio of urosome segment lengths, in the proximal spine of the second segment of the maxilliped which lacks the spinules in the adult female (but not in the juvenile), and the fifth leg has 2, rather than 1, tubercles each bearing at seta. These differences are considered to represent geographical variation within the species, as the only previous records of this species are from the antarctic zone of the southwest Pacific between 1000 and 2000 metres depth. The male and fifth copepodid have not previously been described. Oncaea heronae sp. nov. DESCRIPTION. Female. Ratio of prosome and urosome lengths 1-5 : 1. Prosome elongate and oval in dorsal aspect (Fig. 7a). Mean body length of 4 specimens 0'33 mm (range 0'3 1 to 0'35 mm). Third prosome segment without a dorsal projection. Head I'l times longer than wide. Proportional lengths of urosome segments and caudal rami 9:50:7:7: 13: 14 (Fig. 7b). Genital apparatus located on dorsal surface anterior to midpoint of genital segment. Maximum width of genital segment at level of genital apparatus; length to width ratio 1 -2 : 1 . Caudal rami twice as long as wide. First antenna 6-segmented (Fig. 7c); armature: 1-3, II-6, III-3, IV-1, V-2, VI-5 (some elements may be missing); ratio of segment lengths 12 : 22 : 34 : 13 : 7 : 12. Second antenna 3-segmented (Fig. 7d); first segment bearing 1 long pinnate seta distally; second segment unarmed; terminal segment equal to second segment in length, bearing 1 curved spine and 3 curved setae proximally and 5 curved spines and 1 curved seta distally. Mandible lost during dissection. First maxilla (Fig. 7e) and second maxilla (Fig. 70 with armature similar to O. tregoubovi. Internal surface of second segment of maxilliped bearing 2 overlapping rows of setules and 2 spines; larger distal spine bilaterally dentate (Fig. 7g); terminal claw with smooth concave surface; single internal basal spine. Armature of natatory legs similar to O. tregoubovi except for armature of terminal segments of endopodites 2-4: NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 1 89 Coxa Basis Endopodite Exopodite Legl 0-0 1-1 0-1; 0-1; 0,1, 5 1-0; I-1;III, 1,4 Leg 2 0-0 ?-0 0-1; 0-2; 1, 1, 3 I-0;I-1;III, 1, 5 Leg 3 0-0 1-0 0-1; 0-2; 0,1, 2 1-0; 1-1 ; II, I, 5 Leg 4 0-0 ?-0 0-1; 0-2; 0,1,1 I-0;I-1; II, 1, 5 Bilaterally serrate spines on external margins of exopodites lack pronounced flanges (Figs 7h-k); first endopodite with conical terminal projection; all terminal spines shorter than terminal segments except terminal spines of fourth exopodite and endopodite, these being longer than terminal segments. Fifth leg comprising small free cylindrical segment bearing 2 terminal setae, the longer, internal seta extending to level of genital apertures. Male. Not known. MATERIAL EXAMINED. 499 from sample 9541.24, 3980-3960 m, 20° N 21°W. BM(NH) Holotype 1981.125, paratypes 1981.126. ETYMOLOGY. This species has been named after Gayle A. Heron of the University of Washington, Seattle, in recognition of her work on the taxonomy of the Oncaeidae. REMARKS. The second, third and fourth endopodites of O. heronae, O. brodskii Shmeleva, 1968 and O. longipes Shmeleva, 1968 have the same unusual armature described above, but O. heronae can be distinguished by other small differences in the armature of the natatory legs, by the 2 overlapping rows of setules on the second segment of the maxilliped, and by the structure of the free segment of the fifth leg which is elongate in O. longpipes and reduced to a tubercle in O. brodskii, but small and delimited from the body segment in O. heronae. Oncaea setosa Heron, 1977 O. setosa Heron, 1977 : 73, figs 22a-h. DESCRIPTION. Female. Mean body length of 4 specimens 0'5 mm (range 0'42 to 0*58 mm). Body figured (Figs 8a-c). Appendages of O. setosa similar to those described by Heron with only minor differences apparent (Figs 8d-n). Male. Ratio of prosome and urosome lengths 1-5:1 (Figs 9a and b). Body length of 1 specimen 0'58 mm. Head 1-2 times longer than wide. Proportional lengths of urosome segments and caudal rami 12 : 47 : 3 : 3 : 3 : 18 : 14 (Fig. 9c). Genital lappets extending postero-laterally into acute points. Mouthparts like those of female, except first antenna and maxilliped. First antenna 4-segmented (Fig. 9d). Maxilliped 3 -segmented (Fig. 9e); second segment bearing 2 spines on internal surface and 3 groups of setules; terminal claw bearing a stout internal basal spine. First and second natatory legs like those of female, but terminal spines of third and fourth exopodites longer than those of female, being respectively 1-5 times and twice the length of terminal segments. Two spines, representing the fifth leg, borne on tubercle not clearly delimited from body. Fifth Copepodid (female). Ratio of prosome and urosome lengths 3 : 1 (Figs 9f and g). Body length of 1 specimen 0'51 mm. Head 1-2 times longer than wide. Urosome 4-segmented (Fig. 9h). Proportional lengths of urosome segments and caudal rami 12 : 40 : 8 : 24 : 16. Mouthparts similar to those of adult female except maxilliped, bearing a group of setules on internal surface of second segment (Fig. 9i). Natatory legs bearing terminal spines proportionally longer than those of adult female (Figs 9j and k, lOa and b). (lOj and k, 1 la and b). MATERIAL EXAMINED. 499(1 lost), Irf, 1 9 juvenile from sample 954 1.24, 3980-3960 m, 20° N 21° W.BM(NH) 1981. 127-129. REMARKS. This species was identified by the cluster of spinules on the second segment of the second antenna, the armature of the maxilliped and the relative dimensions of the urosome segments. The female differs from Heron's description in some minor details: the terminal 190 S. J. MALT spines of the third and fourth endopodites are longer in proportion to their terminal segments in the present material. The male and the fifth copepodid have not previously been described. Heron reported O. setosa from 2 stations in the antarctic zone of the southwest Pacific between 1000 and 2000 metres depth. Oncaea rotunda Heron, 1977 O. rotunda Heron, 1977 : 77-79, figs 24o-q, 25a-m. DESCRIPTION. Female, body length of 1 specimen 0-55 mm. Body (Figs lOc-e) and second antenna (Fig. 1 Of) figured. MATERIAL EXAMINED. 19 from sample 9541.24. 3980-3960 m, 20° N 21° W. BM(NH) 1981.130. REMARKS. This specimen was assigned to O. rotunda because of its distinctive second antenna, which has a very short terminal segment relative to the second segment and bears relatively short terminal spines and setae. Also the terminal segment of the fourth endopodite is reduced. Although somewhat smaller than the type material (mean length 0'74 mm), this specimen otherwise corresponds closely to Heron's description. Heron reported O. rotunda from 3 stations in the antarctic zone of the Pacific between 1000 and 2000 metres depth. Oncaea brocha Heron, 1977 O. brocha Heron, 1977 : 60, figs 14f-n, 15a-n. DESCRIPTION. Female. Body length of 2 specimens 0'81 and 0'83 mm. Urosome (Fig. lOg), second antenna (Fig. lOh), maxilliped (Fig. lOi) and second and third natatory legs (Figs lOj and k) figured. MATERIAL EXAMINED. 299 from sample 9541.24, 3980-3960. 20° N 21°W. BM(NH) 1981.131. REMARKS. The specimens were identified by the following combination of characters: the relative lengths of the segments of the first and second antennae, the armature of the maxilliped, of the natatory and of the fifth legs. The second antenna has an additional terminal spine not figured by Heron. This may have been lost from Heron's material or may represent geographical variation. Another minor difference observed was the length of the terminal spines of the second and third endopodites which are relatively longer in the present material. The refractive granules mentioned by Heron are absent from these specimens, but but these may have been artefacts or may represent the positions of cuticular pores. Heron reported O. brocha from 1 station in the antarctic zone of the southwest Pacific between 1000 and 2000 metres depth. Oncaea schmitti (Heron, 1977) Comb. nov. Epicalymma schmitti Heron, 1977 : 82-84, figs 28a-n, 29a-e. DESCRIPTION. Female. Body length 2 specimens 0*33 and 0'34 mm. Body (Figs 1 la and b), second antenna (Fig. 1 1 c) and maxilliped (Fig. 1 1 d) figured. MATERIAL EXAMINED. 299 from sample 9541.24, 3980-3960 m. 20° N 21°W. BM(NH) 1981.132. REMARKS. This species is here transferred from the genus Epicalymma to Oncaea (for discussion see page 191). Specimens are clearly identifiable as O. schmitti by the armature and relative segment lengths of the second antenna, the armature of the maxilliped and the dorsal projections of the caudal rami. However, the specimens are so thinly chitinized that the genital segment had partially collapsed during capture or preservation. Therefore, the drawing of the urosome may not represent the morphology of the living animal. This species NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 1 9 1 has also been recorded from the antarctic zone of the southwest Pacific where it was taken between 1000 and 2000 metres depth. Oncaea umbonata (Heron, 1977) Comb. nov. Epicalymma umbonata Heron, 1977 : 84-86, figs 29f-p, 30a-c. DESCRIPTION. Female. Body length of 1 damaged specimen 0*62 mm, without caudal rami. Body (Figs 1 1 e and 0, second antenna (Fig. 1 1 g) maxilliped (Fig. 1 1 h) figured. MATERIAL EXAMINED. \g from sample 9541.24, 3980-3960 m. 20° N 21° W. BM(NH) 1981.133. REMARKS. This species is here transferred from the genus Epicalymma to Oncaea (for discussion see below). Although the specimen lacks its caudal rami it was assigned to O. umbonata on the basis of its body size and maxilliped armature. There are also differences of the armature of the second antenna between this and the specimens recorded as O. schmitti, the two species are otherwise very close. The natatory legs and other mouthparts correspond closely to Heron's original description. This species has also been recorded from the antarctic zone of the southwest Pacific where it was taken between 1000 and 2000 metres depth. Phylogenetic relationships within the genus Oncaea Giesbrecht (1892) designated the following character states as typical of the genus Conaea: the elongate terminal segment of the second antenna with very large, hook-tipped spines; endopodite of the fourth leg shorter than the exopodite, third segment is reduced (shorter than the first or second segment); the fifth leg reduced to a single plumose seta. However Heron (1977) was able to demonstrate the presence of a minute spinule accompanying the seta of the fifth leg of O. gracilis Dana, 1852 (=syn. C. rapax), the type species. Heron also described 2 new species of Conaea, C. succurva and C. hispida. Both of these have a seta on a small prominence representing leg 5, and a reduced endopodite of the fourth leg, but the third endopodite segment exceeds the first segment in length. These 2 species also lack the hooked tips to the terminal spines of the second antenna. Oncaea expressa Gordejeva, 1973 shares these and other character states but differs in the morphology of the natatory legs (Table 1). No less than 27 other species of Oncaea also show 1 or more of these Conaea-\ike character states (Table 2) The 2 species of Epicalymma which Heron (1977) described also exhibit the following character states: elongate terminal segment of the second antenna with very large hook-tipped spines, endopodite of the fourth leg shorter than the exopodite, third segment reduced (but longer than the first or second segment), fifth leg with a single terminal seta. A continuous gradation of character states within Oncaea, Conaea and Epicalymma is thus becoming apparent as more species of Oncaea are discovered. The armature of the third exopodite segments of the swimming legs also illustrates this. Most Oncaea species have an external spine formula of III, III, II, II but there is-considerable variation within the genus: II, III, III, I (O. ancora Gordejeva, 1973); III, III, III, II (O. minor Shmeleva, 1979 and O. parobscura Shmeleva, 1979); III, II, II, I (O. expressa Gordejeva. 1973); III, II, II, II (O. brodskii Shmeleva, 1968 and O. longipes Shmeleva, 1968); II, III, II, I (O. exigua Farran, 1908); II, II, I, I (O. atlantica Shmeleva, 1967 and O. vodjanitskii Shmeleva and Delalo, 1969; II, III, II, II (O. zernovi Shmeleva, 1966 and O. mollicula Gordejeva, 1975). The typical formula for Conaea is II, III, II, I and for Epicalymma II, III, III, I. Such variation represents a continuum which cannot, in my opinion, justifiably be subdivided into 2 or more genera as it is at present. It is probable that Conaea-\\ke and Epicalymma-like character states have been derived by reduction and loss of armature elements from ancestors within the genus Oncaea. Separation of the Conaea and Epicalymma species would leave 192 S. J. MALT Table 1 Comparison of morphology of natatory legs ofOncaea expressa. O. succurva and O. hispida. Legl Leg 2 Leg 3 Leg 4 O. expressa Exopod. spine Exopod. spine Exopod. spine Exopod. spine Gordejeva, 1973 2 x length 2 x length 2- 5 x length > 2x length term. segm. term. segm. term. segm. term. segm. Endopod. spine Endopod. spine Endopod. spine Endopod. spine shorter than 1-5 x length >4x length > 4 x length term. segm. term. segm. term, segm term. segm. O. succurva Exopod. spine Exopod. spine Exopod. spine Exopod. spine (Heron, 1977) 2 x length >2 x length 2-5 x length >2'5 x length comb. nov. term. segm. term. segm. term. segm. term. segm. (syn. Conaea Endopod. spine Endopod. spine Endopod. spine Endopod. spine succurva) same length 1-5 x length 2- 5 x length 3 x length term. segm. term. segm. term. segm. term. segm. O. hispida Exopod. spine Exopod. spine Exopod. spine Exopod. spine (Heron, 1977) 2 x length >2x length 2-5 x length > 3 x length comb. nov. term. segm. term. segm. term. segm. term. segm. (syn Conaea Endopod. spine Endopod. spine Endopod. spine Endopod. spine hispida) same length 1 -5 x length 2 x length >4 x length term. segm. term. segm. term. segm. term. segm. Oncaea as a paraphyletic group. In order to retain Oncaea as a monophyletic group (sensu Hennig, 1966) it is proposed that Conaea and Epicalymma be synonymized with Oncaea and that the species of Conaea and Epicalymma be transferred to Oncaea, C. gracilis, C. succurva, and C. hispida becoming O. gracilis, O. succurva and O. hispida, also E. schmitti and E. umbonata becoming O. schmitti and O. umbonata respectively. Kazatchenko and Andreev (1977) described Myctospictosum as a genus of uncertain taxonomic position. A single male of M. philippinensis was found on the gills ofMyctophum spinosum in the western Pacific. It is clear from the figures (Kazatchenko and Andreev, 1977 : Figs 9 and 10) that this is an oncaeid. It has the body shape, the 4-segmented first antenna and the 3-segmented second antenna of a typical Oncaea male. The other mouthparts have been interpreted differently by the authors but are clearly those of a male Oncaea. The natatory legs are also typical of the genus. It is proposed, therefore, that Myctospictosum should be synonymized with Oncaea. The armature of the natatory legs and the relative lengths of the posterior segments separate O. philippinensis comb. nov. from any of the known males of the genus Oncaea. Oncaeids are often found clinging by their maxillipeds to other organisms and debris in plankton samples, there is, therefore, no reason to suppose that O. philippinensis is parasitic, it is probably a free-living planktivore like other species of the genus. The generic diagnosis ofOncaea is ammended as follows: ONCAEA Philippi, 1843 Oncaea Philippi, 1843 : 63. AntariaDana, 1846:229. Conaea Giesbrecht, 1891 : 477. Epicalymma Heron, 1977 : 82. Myctospictosum Kazatchenko and Andreev, 1977 : 47. DIAGNOSIS. Cyclopoid shape. Rostral area thickened with rounded posteroventral margin. Female urosome 5-segmented, male 6-segmented. First antenna 6-segmented in female, NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 193 Table 2 Conaea-like character states exhibited by species ofOncaea. c rj u .« o r; CO O SP C « . u Q — C M-5 p ?s ^ U^ -*-^ C QjTO CO ^^ -C J= -« C *> c P.^c So c o o - o « o B - * 2 *><=; ^ -o 3— c£ ^TO> l|s || -g|^ 1| l"8-i§ PCoo mj^t- SCtu £owe § 8 § •§ I ^§^ 'o 8 g B|« a Ha>w20(2B):317-324. Sars, G. O. 1916. Liste systematique des Cyclopoides, Harpacticoides et Monstrilloides recueillis pendant les campagnes des S.A.S. le Prince Albert de Monaco, avec descriptions et figures des especes nouvelles. Bull. Inst. oceanogr. Monaco. 323 : 1-15. Shmeleva, A. A. 1966. New species of the genus Oncaea (Copepoda, Cyclopoida) from the Adriatic Sea. Zool. Zh. 45 (6) : 932-936. (In Russian). 1967. New Oncaea species (Copepoda, Cyclopoida) from south-western part of the Atlantic Ocean. Zool. Zh. 46 (4) : 62 1-622. (In Russian). 1968. New species of planktonic Copepoda : Cyclopoida from the Adriatic Sea. Zool. Zh. 47(12): 1784-1 793. (In Russian). 1969. Especes nouvelles du genre Oncaea (Copepoda. Cyclopoida) de la mer Adriatique. Bull. Inst. oceanogr. Monaco 68, No. 1393 : 1-28. 1979. New species and some previously unknown males of the genus Oncaea from the Mediterranean. Zool. Zh. 58 (4) : 491^98. (In Russian). Tanaka, O. 1960. Pelagic Copepoda. Spec. Publs. Seto mar. Biol. Lab. 10 : 1-177. Manuscript accepted for publication 4 June 1 98 1 NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 195 Fig. 1 Oncaea tregoubovi. a. female, dorsal; b. urosome, dorsal; c. first antenna; d. second antenna; e. mandible; f. first maxilla; g. second maxilla; h. maxilliped; i. fifth leg. 196 S. J. MALT 0-05mm a-d Fig. 2 Oncaea tregoubovi. a. female, leg 1 ; b. leg 2 (damaged); c. leg 3; d. leg 4. NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 197 Fig. 3 Oncaea ivlevi. a. female, dorsal; b. female, lateral; c. urosome, dorso-lateral; d. first antenna; e. second antenna; f. second maxilla; g. maxilliped; h. leg l;i. Ieg2;j. leg 3; k. leg 4. 198 S. J. MALT O-1mm a,b O-O5mm -8P Fig. 4 Oncaea ivlevi. a. male, dorsal; b. male, lateral; c. urosome, lateral; d. maxilliped. NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 199 Fig. 5 Oncaea hispida. a. female, lateral; b. urosome, dorsal; c. first antenna; d. second antenna; e. mandible; f. second maxilla; g. maxilliped; h. labrum (lamella missing). 200 S. J. MALT Fig. 6 Oncaea hispida. a. female, leg 1 ; b. leg 2; c. leg 3; d. leg 4; e. male dorsal; f. urosome ventral (specimen distorted); g. maxilliped; h. female copepodid V, dorsal; i. posterior segments of urosome, dorsal (only posterior half of genital segment figured);], maxilliped. NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 201 b-k Fig. 7. Oncaea heronae sp. nov. a. female, dorsal; b. urosome, dorsal; c. first antenna; d. second antenna; e. first maxilla, f. second maxilla; g. maxilliped; h. leg l;i. leg 2; j. leg 3; k. leg 4. 202 S. J. MALT Fig. 8 Oncaea setosa. a. female, dorsal; b. female, lateral; c. urosome, dorsal; d. first antenna; e. second antenna; f. mandible; g. first maxilla; h. second maxilla; i. maxilliped; j. labrum (lamella missing); k. leg 1 ; 1. leg 2; m. leg 3; n. leg 4. NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 203 Fig. 9 Oncaea setosa. a. male, dorsal; b. male lateral; c. urosome, ventral; d. first antenna; e. maxilliped (distal spine missing); f. female copepodid V, dorsal; g. female copepodid V, lateral; h. urosome, dorso-lateral; i. maxilliped; j. leg l;k. leg 2. 204 S. J. MALT Fig 10 Oncaea setosa. a. female copepodid V, leg 3; b. leg 4; Oncaea rotunda, c. female, dorsal; d. female, lateral; e. urosome, dorsal; f. second antenna; Oncaea brocha. g. female urosome, dorsal; h. second antenna; i. maxilliped;j. leg 2; k. leg 3. NEW AND LITTLE KNOWN SPECIES OF ONCAEIDAE 205 0-1 mm e.f a-d,g,h Fig 11 Oncaea schmitti. a. female, dorsal; b. female, lateral; c. second antenna; d. maxilliped; Oncaea umbonata. e. female, dorsal; f. female, lateral; g. second antenna; h. maxilliped. Larval and post-larval development of the Slender-legged Spider Crab, Macropodia rostrata (Linnaeus) (Oxyrhyncha : Majidae : Inachinae), reared in the laboratory R. W. Ingle Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD Introduction The Slender-legged Spider Crab Macropodia rostrata (Linnaeus) occurs northward to about latitude 65° N and southward to the Moroccan coast and Mediterranean (see Christiansen, 1969; Monod, 1932; Zariquiey Alvarez, 1968). West African and Cape Verde Island specimens have been assigned recently to M. spinulosa (Miers) (see Manning & Holthius, 1981). M. rostrata has been reported also from False Bay, S. Africa (see Barnard, 1950). Previous descriptions (see p. 207) of larval stages of M rostrata are not sufficiently detailed for comparative studies. In 1974 larvae of this species were reared from crabs collected from Tunisian and Plymouth waters and in 1977 from specimens obtained off the Isle of Man. These rearings provided material for the present detailed description and comparisons of the larvae and first crab stages of M. rostrata from three localities within its range of distribution. Materials and methods Ovigerous crabs were trawled from the following localities. (1) Off Port Erin, Isle of Man. 20-24 m., 24-25.3.1977. (2) S.W. of Eddystone Lighthouse, nr. Plymouth, Devon, 90 m., 1 8.6. 1 974. (3) OffCarthage-Salammbo, Tunisia, 4-5 m., 20.2. 1 974. Larvae were reared using methods described by Rice & Ingle (1975 : 104) and Ingle & Clark (1977). Material was fixed in Steedman's fixative (Steedman, 1976 : 148) and later transferred to 70% alcohol. Drawings and measurements were made with the aid of a camera lucida. Measurements given are: (a) distance from base to tip of dorsal spine (D.S.); (b) carapace length, from between eyes to posterio-lateral carapace margin (C.L.). All material has been incorporated into the Collections of the British Museum (Natural History), accession no;- 1981:201-225. Drawings and descriptions (unless otherwise indicated) are of specimens from locality (1). Ten specimens of each stage were dissected and examined from each locality except for ZII of Tunisian material of which 20 specimens were dissected to confirm the absence of a seta on the outer margin of the maxillule basis (see p. 2 10). Whilst in the rearing laboratory females and larvae were maintained at 15°C. Larvae from locality (1) hatched within 10-12 days and those from (2) within 6-7 days of collecting the females. Both hatchings took an average of 30 days to reach first crab stage. Those from locality (3) hatched within 14 and 63 days respectively from time of collecting and an average of 22 days elapsed before the appearance of first crab stage. * Descriptions Larval references. Macropodia phalangium:- Thompson, 1 836 : 37 1 , fig. e (pre-1 st zoea), non fig. 2 = ? Bull. Br. Mus. nal. Hist. (Zool.) 42 (3) : 207-225 Issued 27 May 1982 208 R. W. INGLE Hyas sp. ? Stenorhynchus rostrata:- Stuxberg. 1 874 : 3 ( 1 st zoea). ? Stenorhynchus phalangium:- Cano, 1893:Tav. 35, figs 70, 72, 74, 77-85 (1st zoea, megal. crab). Macropodia rostrata:- Labour, 1928 : 550, PI. Ill, fig. 10 (coloured), PI. XV, fig 7, PI. XVI, figs 3, 5, 8 (prezoea, 1st, 2nd zoeae, megal. 1st crab), non Stenorhynchus phalangium:- Lo Bianco, 1904 : 439; 1904 : Taf. 12, fig 43 (=? Inachus megalopa). FIRST ZOEA Dimensions: D.S. 1-3-1-4 mm.,C.L.O-7-0-8 mm. Carapace (Fig. la). Dorsal spine long and straight, narrowing distally. Rostral and lateral spines not developed. Dorso-median elevation conspicuous above which there is a small seta on either side; ocular eaves developed; a pair of small dorso-lateral setae near base of dorsal spine; posterio-lateral margin of carapace with 'majid' spine and 1-2 additional long thin setae (see inset Fig. la). Surface of carapace with minute spinules. Eyes: Partly fused to carapace. Antennule: (Fig. 2a). Unsegmented and with 2 terminal aesthetascs and 2 setae. Antenna: (Fig. 2a). Spinous process without distal spinules: exopod slightly shorter than spinous process and with one small spinule in proximal half; endopod developed as acute bud. Mandible: (Fig. 2b). Incisor and molar process developed, palp absent. Maxillule: (Fig. 3a). Endopod 2-segmented (incipiently in some specimens), distal segment long and thin with 3 long distal setae; basal endite with 5 spines and 2 setae, coxal endite with 7 setae/spines. Maxilla: (Fig. 4a). Endopod narrow, distally truncate and with 4 long setae; basal endite broad, distal margin concave or incipiently bilobed and with 4 + 5 setae; coxal endite incipiently bilobed, outer lobe with acute outer margin, with 3+4 setae respectively; scaphognathite with 10 long plumose marginal setae, posterior seta very stout. First maxilliped: (Fig. 5a). Basis with 9 setae arranged 2, 2, 2, 3; endopod 5 -segmented with 3,2,1,2,4 + 1 setae; exopod with 4 terminal natatory setae. Second maxilliped: (Fig. 5b). Basis with one proximal seta; endopod 3-segmented with 0, 0, 3-4 + 1 setae; exopod incipiently (in some specimens) segmented with 4 terminal natatory setae. Third maxilliped and pereiopods: Represented as incipient buds. Abdomen: (Figs 6a, b, e). 5-segmented + telson; 2nd segment with a pair of anterio-laterally directed acute dorso-lateral processes; posterio-lateral processes on segments 3-5 long and terminally acute; a pair of minute setae near posterio-dorsal margins of segments 2-5. Telson furcae long and distally with minute spinules, each furca with a small lateral spine; inner medio-lateral margin of telson with 3 long plumed setae, innermost (in some specimens) noticeably short (see Fig. 6e); incipient pleopod buds on segments 2-5. SECOND ZOEA Dimensions: D.S. 1-1-1-2 mm., C.L. 0-8-0-9 mm. Carapace (Figs Ib, c). Dorsal spine curved. Dorso-median elevation well developed; ocular eaves conspicuously expanded (Fig. Ic). Eyes: Moveable. Antennule: (Fig. 2c). With 6 aesthetascs and 1-2 setae. Antenna: (Fig. 2c). Exopod with 2 proximal spinules; endopod slightly more than \ length of exopod. Mandible: (Fig. 2d). Each half dissimilar. Maxillule: (Fig. 3b). Outer margin of basal endite with a seta, distal margin with 5 spines and 3 setae; coxal endite with 7 setae/spines. Maxilla: (Fig. 4b). Endopod setation unchanged; basal endite with 5 + 5 setae; coxal setation unchanged; scaphognathite with 16 setae. First maxilliped: (Fig. 5c). Setation on basis and endopod unchanged; exopod with 6 terminal natatory setae. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 209 Second maxilliped: (Fig. 5d). Basis without setae; endopod with 0, 0, 3 + 1 setae; exopod with 6 terminal natatory setae. Third maxilliped and pereiopods: Represented as conspicuous incipient buds. Abdomen: (Fig. 6c, d). Posterio-lateral processes on segments 3-5 longer than in first stage; pleopod buds long. MEGALOPA Dimensions: C.L. 1 '2-1*3 mm. Carapace: (Figs Id, e). Longer than broad, with a small obtuse rostrum and obtuse but prominent submedian lobes; frontal region with 2-3 median broad longitudinal tubercles, hepatic regions swollen, each protogastric region with a long anteriorly directed process narrowing distally; cardiac region with a posterio-dorsally directed spine. Eyes: Large, with well developed cornea. Antennule: (Fig. 2e). Peduncle 3-segmented, terminal segment with a seta on inner margin; exopod 2-segmented, proximal with 2 and distal segment with 4 aesthetascs. Antenna: (Fig. 2f). Peduncle 3 segmented, proximal segment with a stout ventrally directed process, distal segment with a small ventral seta; flagellum 4-segmented, 2nd segment longest and with 4 setae, 4th segment with 2-3 aesthetascs. Mandible: (Fig. 2g). Molar process slightly reduced, incisor prominent and broad; palp stout and unsegmented, with a small terminal seta. Maxillule: (Fig. 3c). Endopod reduced; distal margin of basal endite with 6-7 spines and 6-7 setae, coxa with 6-7 setae/spines. Maxilla: (Fig. 4c). Endopod reduced and terminally acute, with one long distal seta; basal endite broadly oval, with 6 + 3-4 setae, coxal endite with 3 + 3 setae; scaphognathite with 18-1 9 setae. First maxilliped: (Fig. 7a). Coxa with 4, basis with 9-10 setae; exopod 2-segmented, distal segment with 4 setae; endopod reduced and terminally acute; epipod small. Second maxilliped: (Fig. 7b). Exopod 2-segmented, proximal segment long, distal with 4 setae; endopod 4-segmented, 2nd (carpus) with one, 3rd (propodus) with 3 setae and 4th (dactylus) with one seta and 3 spines; epipod small (not shown in figure). Third maxilliped: (Fig. 8a). Endopod 5-segmented, inner margin of ischium with few minute spinules and 7-9 short setae, merus with 4 setae, carpus with 3, propodus with 6 and dactylus with 4 setae; distal segment of exopod with 4 setae, epipod moderately well developed. Pereiopods: (Figs 8c, 9a, lOa-c). Chelipeds moderately stout and sparsely setose, distal parts of propodus and dactylus curved; merus with small obtuse spinule on inner proximal margin; basis/coxa with small obtuse process on lower margin. Pereiopods 2-5 slender, setose and minutely spinulate, dactylus terminally acute, ischium of 2nd-3rd pereiopods with a prominent curved hook-shaped process. Abdomen: (Figs If, g). 5-segmented + telson; 1st with 4, 2nd-3rd with 2 posterio-dorsal setae, 4th with 4 setae, 5th with 6 setae; telson broader than long, posterior margin subtruncate. A pair of well developed pleopods on segments 2-5, 4th pair smallest, exopod of each with 8 long plumose setae; endopod of each with 2 coupling hooks. FIRST CRAB Dimensions: C.L. 1 -4-1-5 mm. Carapace: (Fig. Ij). Slightly less that l±x longer than broad; frontal region slightly produced, submedian lobes obtuse and with clusters of hook-setae, each protogastric region with a tubercle; orbits long, hepatic regions swollen, epibranchial regions slightly expanded, mesogastric with 2 median cristate tubercles, cardiac region with a prominant tubercle, intestinal with a small median tubercle; dorsal surface and lateral margins of carapace with many hook-setae. Eyes: Large and with a few hook-setae. Antennule: (Fig. 2h). First and 3rd segments of peduncle with a seta; exopod indistinctly segmented, with 7-8 aesthetascs and 2-3 setae; endopod 2-segmented, with 2 terminal and 2 sub-terminal setae. 210 R. W. INGLE Antenna: (Fig. 2h). Peduncular segments with 2, 1-2, and 5 setae respectively; flagellum usually with 3 or 4 demarcated segments, with 3 subterminal setae as shown. Mandible: (Fig. 2i). Molar process acute, incisor expanded as a broad cristate lobe; mandibular palp 2-segmented, distal segment with 2 setae. Maxillule: (Fig. 3d). Endopod very reduced, outer margin of basal endite with a prominent seta, margin with 8 spines and 4 setae; coxal endite with 9 setae/spines. Maxilla: (Fig. 4d). Endopod very reduced and with one apical seta; basal endite with 10 and coxal with 2-3 setae respectively; scaphognathite with 20-2 1 short plumose setae. First maxilliped: (Fig. 7c). Coxa with 10-11 setae, basis with 10 setae; distal segment of exopod with 3 long setae; endopod sub-triangular, with 1-2 small setae on distal margin; epipod well developed and with 6 setae. Second maxilliped: (Fig. 7d). Merus with 2 setae, propodus with 4 setae and a distal spine, dactylus with 1-2 setae and 4 spines; epipod small. Third maxilliped: (Fig. 8b). Ischium with numerous setae (18-20) as shown, inner margin with 3-4 processes; merus with 3 setae, inner margin with an acute process; carpus, propodus and dactylus with 2, 4 and 5 setae respectively on outer surface and margins; distal segment of exopod with 3 short plumose setae; a cluster of short setae near coxal/epipod junction; epipod well developed. Pereiopods: (Figs 8d, 9b, c, lOd, e). Cheliped moderately setose as shown; inner distal propodal margin with 3 acute to sub-acute teeth (see inset to Fig. 8d); distal part of the propodus acute and curved. Pereiopods 2-5 long, thin and with numerous setae; 2nd and 3rd similar in shape, 3rd longest (Fig. 9c); 4th and 5th similar in shape, dactylus curved, inner margin of dactylus of 4th (Fig. lOd) with 4 spines, of 5th with 2-3 spines. Remarks Comparisons of larvae and first crab stages of Mediterranean and British material of M. rostrata (see p. 207) have revealed only two apparent morphological differences. (1) all zoeae II examined from the Tunisian locality are without a seta on the outer margin of the maxillule basal endite; this seta is present in all zoeae II reared from the Plymouth and Isle of Man crabs. The presence of this seta is considered an important larval brachyuran feature (see Rice, 1980 : 299 as 'exopod' seta), and its absence in the Tunisian specimens cannot be explained. Clark (1980) also reported its absence from the maxillule of zoeae II Inachus dorsettensis (Pennant) reared from Isle of Man crabs but found it present in Plymouth material. (2) in the first crab stage of Tunisian specimens the carapace submedian lobes are widely spaced and the outline of the hepatic region is noticeably convex. By comparison, first crab stages from the two British localities are slightly smaller than the Tunisian speci- mens, the submedian lobes are closer together and the hepatic region outline is relatively straight (Fig. 1, cf j & k). These slight morphological differences observed in the first crab stages may express the extremes of geographical variation of M. rostrata since the Mediterranean material probably represents the near southernmost limit of this species (see p. 207). Ingle & Manning (in press) have drawn attention to the noticeable variation of carapace shape among population of pre- and post pubertal crabs of M. rostrata from the N.E. Atlantic and Mediterranean regions. Four species of Macropodia are reported from N.E. Atlantic waters: M. rostrata (Linnaeus), M. deflexa Forest, M. tenuirostris (Leach) and M. linaresi Forest & Zariquiey Alvarez. Larval stages of the first three species mentioned were described briefly by Lebour (1927, 1928); larvae ofM. linaresi are unknown. Lebour (1927, 1928) recognized that the zoeae of M. rostrata and M. deflexa (=egyptia) has a longer and more straight dorsal spine, and longer antennae than M. tenuirostris (= longirostris); she also noted that the antennae of M. deflexa are longer than M. rostrata. Lebour remarked that the megalope of M. rostrata has shorter legs and a less deflected rostrum than M. tenuirostris, and that the 'central lobe' of the rostrum of M. deflexa LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 2 1 1 protrudes more than that of M. rostrata but that otherwise the megalop of both species are 'hardly to be distinguished'. Larval material of M. dejlexa has not been available for examination during this study but it has been possible to compare the larvae of M. rostrata with those of M tenuirostris reared from females collected off the Isle of Man during March 1977. These comparisons are listed in Table 1 . Table 1 Character M. rostrata M. tenuirostris Zoea I Carapace length: Dorsal spine length: shape: Antenna, total length: Second mxpd endop. 3rd sgmt: Zoea II Carapace length: Antenna, total length: Carapace dorso-median elevation: Maxilla, coxal endite: scaphognathite: Abdomen posterio-lateral spines: Second mxpd endop. 3rd sgmt: Megalopa Carapace length: Carapace frontal region: Carapace protogastric processes: Cheliped merus: 0-7-O8 mm 1*3-1-4 mm relatively straight 1-2-1 -2 mm 4 + 1 setae 0-8-0-9 mm 1'3-1'4 mm (exceeding C.L.) pronounced average of 7 setae average of 1 6 setae long 3-4 + 1 setae average 1 -2 mm 2-3 median longit. tubercles forming moderately wide U-shape with a small spine but without secondary pos- terior spinule 0-9-1 -Omm 1-0-1-1 mm noticeably curved 0-9-1 -Omm 3 + 1 setae 1-1-1-2 mm 1-1-1-2 mm (not exceeding C.L.) not pronounced average of 8 setae average of 1 7 setae longer than rostrata 3 + 1 setae average 1*5 mm continuous median carina forming wide U-shape with large spine and secondary posterior spinule Lebour (1928) provided a key to the larvae of five oxyrhynch genera that occur in British waters; Bourdillon-Casanova (1960) extended this number to seven genera in her key to Mediterranean brachyuran larvae. From a study of reared material and of published accounts (see footnote to Table 2), it has been possible to tabulate a sufficient number of comparative larval features from which a provisional key has been constructed to facilitate the identification of larvae of all those oxyrhynch genera represented in the North East Atlantic Ocean north of about 48° 30' N. Detailed studies of larval stages of Achaeus cranchii, Pisa tetraodon, Maja squinado, Macropodia linaresi and M. dejlexa are still required whilst the positive identity (see Rice, 1980 : 308) of the plankton caught zoea II attributed to Dorynchus thomsoni still remains to be established. 212 R. W. INGLE ° - c •s ± It U 00 > 1 o. a — « (N I N = 'r'N •a °~ E ° ~^ _>> u. £> 00 -C ^- 2 C oo O S i o =5 -g & i E o S -t"? S"iS ™" ' O + o c TT a S m «=> g 5 + O i: -i^O X 1^1 + rs u — « g 'S s •S ? 5b — 1 "• Q, I ce ara ral spin pace d I ' ft J J •s ° S. u i: E JS II ^^ gt 00 -^ « ^ TT vO V 00 2 £ 00 § a O. + ~ JS '5. O ' U Oil c oo 00 c + a a *s f> '5.J '5. i 1 i i jd 1 •j? ^ c '* Abdomen posterio-lateral spines: 2 setae on 1st abdominal segment: Setae on 2nd abdominal segment: Dorso-lateral proc. on 3rd abd. sgm Setal formula basis 1st maxilliped: No. setae basis of 2nd maxilliped: Setal formula endopod 2nd maxillip Setal formula endopod maxillule: Telson dorsal spine + or -: megalopa No. abdominal segments (incl. telso No. pairs of pleopods (incl. uropods No. setae on each terminal pleopod: Frontal region of carapace-submedi -rostrum Carapace, each protogastric region i orbital region with: cardiac region with: Spines on abdominal segments: Processes or spine, 1 st ped. sgmt anl c u LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 2 1 3 Provisional key to the larvae of N.E. Atlantic Oxyrhyncha Zoeae Carapace lateral spines present 2 Carapace lateral spines absent 5 Carapace with a group of 5 lateral spines on each side t Dory nchusthomsoni Thomson, (Williamson, 1960). Carapace with one lateral spine on each side 3 Telson medio-lateral margin strongly concave or medially cleft; middle pair of medio-lateral setae longest; each telson furca with one dorsal and one (rarely 2) lateral spines; 3rd abdominal segment with dorso-lateral process 4 Telson medio-lateral margin not concave or cleft; medio-lateral setae of equal length; each telson furca without a dorsal spine, one lateral spine present; 3rd abdominal segment without dorso-lateral process . . . . Rochinia carpenteriThomson, (Ingle, 1979). Carapace dorsal spine much longer than carapace length; abdominal posterio-lateral process very long; 2nd maxilliped basis with 4 setae; each furca of telson with one lateral spine Hyas coarctatus (Leach), H. araneus (Linnaeus), (Christiansen, 1973). Carapace dorsal spine, at the most, slightly longer than carapace length; abdominal posterio-lateral processes short; 2nd maxilliped basis with 3 setae; each furca of telson with 2 lateral spines Majasquinado(Herbst),(Lebour, 1927, 1928). Carapace rostral spine long; each telson furca with a dorsal spine Eurynome aspera (Pennant), (Lebour, 1 928). E. spinosa Hailstone, (see Table 2, footnote). Carapace rostral spine small or absent; each telson furca without a dorsal spine . . 6 Telson medio-lateral margin strongly cleft, marginal setae in 2 groups; maxillule endopod* with 0 + 3-4 setae; abdominal posterio-lateral spines long 7 Telson medio-lateral margin not cleft, marginal setae not in 2 groups; maxillule endopod with 1 + 6 setae; abdominal posterio-lateral spines short tPfaa annaia(Latreille), (Ingle & Clark, 1980). Lateral spine of each telson furca very long (about \ length of furca and exceeding ^ maximum width of telson) Achaeus cranchii Leach, (Bocquet, 1954). Lateral spine of each telson furca short, never reaching | length of furca .... 8 Lateral spine of each telson furca large; middle medio-lateral seta of telson not longest; maxillule endopod with 0 + 4 setae Inachus dorsettensis (Pennant), /. p/ialangium (Fabricius), /. leptochirus Leach, (Clark, 1980). Lateral spine of each telson furca small; middle medio-lateral seta of telson longest; maxillule endopod with 0 + 3 setae Macropodia rostrata (Linnaeus), M . tenuirostris (Leach), (see p. 2 1 1 ), M . deflexa Forest (Lebour, 1928). Megalops Abdomen composed of 5 segments (excluding telson); 4 pairs of pleopods, terminal pair with 8 setae on exopod 2 Abdomen composed of 6 segments (excluding telson); 5 pairs of pleopods, terminal pair with 3-5 setae on exopod 4 Dorsal margins of abdominal segments without spinules; carapace submedian spines not developed Macropodia rostrata (Linnaeus), M. tenuirostris (Leach) (see p. 2 1 1 ), M. deflexa Forest (Lebour, 1928). Dorsal margins of abdominal segments with small spinules; carapace submedian spines well developed 3 Orbital spines present Inachus dorsettensis (Pennant), /. phalangium (Fabricius), /. leptochirus Leach, (Clark, 1 980). Orbital spines absent Achaeus cranchii Leach, (Bocquet, 1954). *Not known for Achaeus cranchii; tHeegaard's (1963) description of zoea I of Pisa tetraodon differs considerably from that of P. armata (see Table 2, footnote) and is excluded from this present key. 214 R. W. INGLE 4 Carapace submedian spines present Hyas coarctatus (Leach), H. araneus (Linnaeus), (Christiansen, 1973). - Carapace submedian spines absent 5 5 First peduncular segment of antenna without a distal process; exopod of each terminal pleopod with 3 setae • Eurynome aspera (Pennant), (Lebour, 1928), E. spinosa Hailstone, (see Table 2, footnote). First pedunular segment of antenna with an obtuse process or spine; exopod of each terminal pleopod with 5 setae 6 6 A stout cardiac spine on carapace . . . Rochinia carpenteri (Thomson), (Ingle, 1979). Without a cardiac spine on carapace 7 7 Rostral spine well developed; each protogastric region with a carina; exopods of lst-4th pereiopods with 10 setae Pisa armata (Latreille), (Ingle & Clark, 1 980). - Rostral spine minute; each protogastric region with (at the most) a tubercle; exopods of 1 st-4th pereiopods with 8 setae . . . A/a/a squinado (Herbst), (Lebour, 1 927, 1 928). Acknowledgements I wish to thank Dr R. B. Manning for sponsoring my visit to Tunisia under the Smithsonian Institution Foreign Currency Program; Dr D. I. Williamson for his assistance in obtaining ovigerous crabs from the Isle of Man region, and Mr A. D. M. Mattacola and crew of the Sepia for their help in collecting material in Plymouth waters. References Barnard, K. H. 1950. Descriptive Catalogue of South African Decapod Crustacea (Crabs and Shrimps). Ann. S. Afr. Mus. 38 : 837 pp. Bocquet, C. 1954. Developpment larvaire d'Achaeus cranchii Leach (Decapode Oxyrhynque). Bull. Soc. zool. Fr. 79 : 50-56. Bourdillon-Casanova, L. 1960. Le meroplancton du Golfe de Marseille: les larves de crustaces decapodes. Red Trav. Stn mar. Endoume3Q : 1-286. Cano, G. 1 893. Sviluppo e Morfologia degli Oxyrhynchi. Mitt. zool. Stn Neapel 10 : 527-583. Christiansen, M. E. 1969. Marine invertebrates of Scandinavia, No. 2. Crustacea Decapoda Brachyura. Universitetsforlaget, Oslo, 1-143 pp. 1973. The complete larval development of Hyas araneus (Linnaeus) and Hyas coarctactus (Leach (Decapoda, Brachyura, Majidae) reared in the laboratory. Norw. J. Zool. 21 : 63-89. Clark, P. F. 1980. British spider crabs of the genus Inaehus; a morphological study of larval development. M.Sc Modern Taxonomy (C.N.A.A.) Thesis. Polytechnic of Central London, 140 pp. London. Heegaard, P 1963. Decapod larvae from the Gulf of Napoli hatched in captivity. Vidensk. Meddr dansk naturh. Foren. 125 : 449^493. Ingle, R. W. 1979. The larval development of the spider crab Rochinia carpenteri (Thomson) [Oxyrhyncha: Majidae] with a review of majid subfamilial larval features. Bull. Br. Mus. nat. Hist. (Zool.) 37: 47-66. & Clark, P. F. 1 977. A laboratory module for rearing crab larvae. Crustaceana 32 : 220-222. 1980. The larval and post-larval development of Gibb's spider crab. Pisa armata (Latreille) [family Majidae: subfamily Pisinae], reared in the laboratory. /. nat. Hist. 14 : 723-735. Ingle, R. W. & Manning, R. B. (in press). Infraspecific variation, synonomy and distribution of the spider crab, Macropodia rostrata (Linnaeus). Colloquium Crustacea Decapoda Mediterranean May 1979 Ancona. Lebour, M. V. 1927. Studies of the Plymouth Brachyura. I. The rearing of crabs in captivity, with a description of the larval stages of Inaehus dorsettensis. Macropodia longirostris and Maia squinado. J. mar. biol. Ass. U.K. 14 : 795-820. 1928. The larval stages of the Plymouth Brachyura. Proc. zool. Soc. Lond. 2 : 473-560. Lo Bianco, S. 1904. Pelagische Tiefseefischeri der "Maja" in der umgebung von Capri. Beitr. Kennt. Meeres und Seiner Bewohner 1 : 1-91 Jena. Manning, R. B. & Holthius, L. B. 1981. West African Brachyuran Crabs (Crustacea: Decapoda). Smithson. Contr. Zool. 306 : 1-379. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 2 1 5 Monod, T. 1932. Brachyuran Maroccana 2e Partie: Dromiidea, Oxystomata, Oxyrhyncha, Brachyrhyncha(excl. Pinnotheridae). Bull. Soc. Sci. nat. Maroc 12 : 7-8. Rice, A. L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Trans. zool. Soc. Lond. 35 : 271^424. & Ingle, R. W. 1975. The larval development of Carcinus maenas (L.) and C. mediterraneus Czerniavsky (Crustacea, Brachyura, Portunidae) reared in the laboratory. Bull. Br. Mus. nat. Hist. (Zool.) 28: 101-119. Steedman, H. F. 1976. General and applied data on formaldehyde fixation and preservation of marine zookplankton (pp. 103-171) In: Steedman, H. F. (Edit.). Zooplankton fixation and preservation. Monographs on oceanographic methodology 4 : 350 pp. UNESCO Press, Paris. Stuxberg, A. 1874. Karcinologiska iaktagelser. Ofvers. K. VetenskAkad. Forh. Stockh. 9 : 3-23. Thompson, J. V. 1836. Of the double metamorphosis in Macropodia phalangium, or spider crab, with proofs of the larvae being zoea in Gegarcinus hydrodomus. Thelpusa erythropus. Eriphia carribaea. and Grapsus pelagicus. Ent. Mag. 3 : 370-375. Williamson, D. I. 1960. A remarkable zoea, attributed to the Majidae (Decapoda, Brachyura). Ann. Mag. nat. Hist. (13)3 : 141-144. Zariquiey Alvarez, R. 1968. Crustaceos Decapodos Ibericos. Investigacion pesq. 32 : i-ix + 5 10 pp. Manuscript accepted for publication 3 July 1981 216 R. W. INGLE Fig. 1 Macropodia rostrata (L.): a, b zoea I & II respectively from lateral aspects; c zoea II carapace and eyes from frontal aspect; d megalopa from dorsal aspect and e lateral aspect of carapace and eye; f dorsal and g lateral aspect of megalopal abdomen; h first and i fourth left megalopal pleopods; carapace of the first crab from j-Isle of man cf. with k-Tunisian waters. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 2 1 7 1 002' Fig. 2 Macropodia rostrata (L.): zoea I, a antennule and antenna, b mandible; zoea II, c antennule and antenna, d mandible; megalopa, e antennule, f antenna, g mandible; first crab, h antennule & antenna, i mandible. 218 R. W. INGLE Fig. 3 Macropodia rostrata (L.): maxillule of- a zoea I; b zoea II; c megalopa; d first crab. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 2 1 9 Fig. 4 Macropodia rostrata (L.): Maxilla of- a zoea I; b zoea II; c megalopa; d first crab. 220 R. W. INGLE Fig. 5 Macropodia rostrata (L.): a & b 1st & 2nd maxillipeds of zoea I; c & d 1st & 2nd maxillipeds of zoea II. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 22 1 Fig. 6 Macropodia rostrata (L.): abdominal segments and telson a & c lateral, b & d dorsal aspects of- a, b zoea I; c, d zoea II; e spines on right medio-lateral margin of telson of zoea I. 222 R. W. INGLE Fig. 7 Macropodia rostrata (L.): 1 st (a, c) & 2nd (b, d) maxillipeds a, b megalopa; c, d first crab. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 223 Fig. 8 Macropodia rostrata (L.): 3rd maxilliped- a megalopa; b first crab; cheliped- c megalopa; d first crab. 224 R. W. INGLE Fig. 9 Macropodia rostrata (L.): a 2nd pereiopod of megalopa; b, c 2nd & 3rd pereiopods of 1st crab. LARVAL AND POST-LARVAL DEVELOPMENT OF THE SLENDER-LEGGED SPIDER CRAB 225 Fig. 10 Macropodia rostrata (L.): a-c 3rd-5th pereiopods of megalopa; d & e 4th & 5th pereiopods of first crab. British Museum (Natural History) British Marine Amphipoda: Gammaridea by R. J. Lincoln 658pp 2,300 figures 4to hard bound ISBN 0 565 00818 £50.00 Amphipods are both numerous and diverse in numbers of genera and species in British coastal waters, but in the absence of any form of modern systematic synopsis or key this group of crustaceans has acquired the reputation of being notoriously difficult to identify. This monograph, which is the first comprehensive and illustrated text on British gammaridean amphipods to be published in more than a century, should go a long way towards solving the problem. The systematic section of the book contains descriptions and figures of all 271 species of marine and brackish water amphipods, in 123 genera and 36 families, recorded from British coasts and the adjacent continental shelf to a depth of 200 metres. Keys are provided at all levels, as well as relevant synonymies and diagnoses of genera and families. The text is illustrated with about 2,300 separate figures which have been drawn by the author from Museum and other material, in many cases with reference to type specimens. The work has been carefully edited to bring corresponding descriptions, keys and figures into close proximity within the text. The systematic section is supported by chapters dealing with morphology, systematics, geographical distribution, biology and ecology, the latter being presented in the form of an annotated subject index of research literature. Finally, there is an extensive bibliography of about 1,200 references that includes most of the British marine amphipod literature published to date. Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. Titles to be published in Volume 42 The tick collection (Acarina: Ixodoidea) of the Hon. Nathaniel Charles Rothschild deposited in the Nuttall and general collections of the British Museum (Natural History). By James E. Kekans Hydroids and medusae of the family Campanulariidae recorded from the eastern North Atlantic, with a world synopsis of genera. By P. F. S. Cornelius Miscellanea Miscellanea Printed by Henry Ling Ltd, Dorchester 902 British Museum (Natural History) Miscellanea Zoology series Vol 42 No 4 24 June 1982 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) © Trustees of the British Museum (Natural History), 1982 The Zoology Series is edited in the Museum's Department of Zoology Keeper of Zoology : Dr J. G. Sheals Editor of Bulletin : Dr C. R. Curds Assistant Editor : Mr"C. G. Ogden ISSN 0007-1498 Zoology series Vol 42 No 4 pp 227-335 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 24 June 1982 .*, LIBRARY ^ a*0; Miscellanea Contents New species of marine nematodes from Loch Ewe, Scotland. By H. M. Platt and Z.N.Zhang .............. The larval development of Crangon crangon (Fabr. 1795) (Crustacea: Decapoda). By A. R.Gurney ............ A revision of the spider genus Cocalodes with a description of a new related genus (Araneae: Salticidae). By F. R. Wanless ........ Anatomy and evolution of the jaws in the semiplotine carps with a review of the Genus Cyprinion Heckel, 1 843 (Teleostei: Cyprinidae). By G. J. Howes 221 247 263 299 A*^ 200. Amphid large (0-7 c.d.) C. caelestis (Gerlach, 1 956) comb. nov. syn. Robbea caelestis Gerlach, 1956 Tail tip rounded, 'a' ratio < 200 4 4 Amphid situated far anterior so that subcephalic and subamphidal setae are on the same level. 7 pairs subventral caudal supplements C. exile (Cobb, 1920) Amphid situated further posterior 5 5 Middle of amphid situated at 10% of length of buccal bulb from anterior . . C. smosp. nov. Middle of amphid situated at least 30% of length of buccal bulb from anterior . . . 6 6 Cephalic setae 8 //m long (0-3 c.d. at base of buccal bulb) C. gallica (Vitiello, 1 974) comb. nov. syn. Robbea gallica Vitiello, 1974 Cephalic setae > 17 ^m long (0-8 c.d. at base of buccal bulb) 7 7 Only 4 subcephalic setae and 4 subamphidal setae between base of amphid and anterior C. porosum Hooper & Cefalu, 1973 Additional subcephalic setae present C. macintyrei sp. nov. Catanema macintyrei sp. nov. Figs 2-3 MATERIAL STUDIED. Holotype: rf, BM(NH) 1981.5.22. Allotype: 9, BM(NH) 1981.5.23. Paratypes: five 1981.5.24-29,31-34. DIMENSIONS. males and five females BM(NH) Holotypecf: Allotype 9,: - 87 M 3710 3770//m;a= 25 25 27 23 - 94 1845 3555 3620 //m; a = 125;b = 39;c = 56; V = 51% 25 25 29 24 = 44//m a=138; b = 37; c = 66; a=137; b = 42; c = 71; a=143; b = 47; c = 72; a=146; b = 45; c = 64; S = 43/zm a=128; b = 41; c = 61; S = 41/an 92:L = 4065/*m; a=145; b = 47; c = 71; V = 50% 93:L = 3720/zm; a =143; b = 40; c = 69; V = 50% 94:L = 4510//m; a=161; b = 46; c = 76; V = 52% 95:L = 4010/zm; a =143; b = 47; c = 78; V = 52% 96:L = 4530//m; a =162; b = 49; c = 79; V = 49% 232 H. M. PLATT & Z. N. ZHANG Fig. 2 Catanema macintyrei: (a) whole body of dy, (b) oesophageal region of cf,; (c) head of cf, (see text for labelling); (d) head of d"7; (e) head of 9,. Bar scales: a = 100 //m; b = 30 ^m; c-e = 10 ^m. Spicule (S) measurement is the arc. Maximum body diameter used is the average of three measurements along the middle portion of the body. DESCRIPTION. Cuticle finely striated from the base of the cephalic capsule, but most conspicuous in the oesophageal and immediate post-oesophageal regions. Typically, the cuticle is covered with a coating of coccoid blue-green algae, but this tends to be lost during preservation. One specimen also had two suctorian ectoparasites attached to the middle part NEW SPECIES OF MARINE NEMATODES 233 Fig. 3 Catanema macintyrei: (a) tail region of c?2; (b) copulatory apparatus of d1,; (c) spicules of three different males; (d) tails of three males and two females. Bar scales: a,c = 30 jum; b = 10 /zm; of the body. Cuticle bears six files of stout setae throughout the body, but they are particularly conspicuous ventrally in the region just posterior to the oesophagus and in the male, anterior to the cloaca. R, sensilla not seen. Six small l'5-2 /zm setose R2 sensilla. Four 17'5-24 //m R3 setae, about 80% of the maximum diameter of the cephalic capsule. The head diameter is usually measured at the level of the R3 sensillae, but they are so far anterior as to make this measurement too inaccurate. Medial and slightly posterior to the R3 setae are four ll-14//m subcephalic setae (labelled SC, in Fig. 2c). Further posterior are four smaller 6-5-8*5 jum setae (labelled SC2 in Fig. 2c). In addition, there is one seta just posterior to the ventral R3 seta (Fig. 2c, V), two level with and dorsal to the middle of the amphid (Fig. 2c, D, & D2) and one ventro-sublateral subamphidal seta (Fig. 2c, A). The disposition of these setae is constant in that each element can be distinguished in the same location in all the specimens, both male and female. Amphid ventrally wound, from outside to centre; 9-10//m wide. Buccal cavity represented by a narrow funnel in the anterior part of the buccal bulb, the latter being about 23 //m long. Tail conical with a characteristic slight ventral inflection at the tip. Spicules paired, curved and proximally cephalate: average length 44 jam (arc) or 32 fj.m (chord). Gubernaculum well cuticularised and dorso-caudally directed. Testis single and outstretched. Ovaries paired and apparently outstretched. DIFFERENTIAL DIAGNOSIS. Catanema macintyrei sp. nov. can be distinguished from the other species in the genus Catanema by the unique disposition of the setae around the head. From 234 H. M.PLATT& Z.N.ZHANG the other sympatric species, C. smo sp. nov., it can also be distinguished by the larger and slightly more posteriorly situated amphid. DISCUSSION. This species will be discussed together with the following species. Catanema smo sp. nov. Fig. 4 Fig. 4 Catanema smo: (a) oesophageal region of cf,; (b) head of cf,; (c) head ofd1,; (d) tail region of rf2; (e) copulatory apparatus of cf,; (0 spicules of three different males; (g) tails of three males and one female. Bar scales: a,d,f = 30 //m; b,c,e = 1 0 //m; g = 50 //m. NEW SPECIES OF MARINE NEMATODES 235 MATERIAL STUDIED. Holotypeicf, BM(NH) 1981.5.38. Allotype: 9, BM(NH) 1981.5.39. Paratypes: four males BM(NH) 1981.5.35-37,30. DIMENSIONS. Holotyperf,: -83 M 2980 3050 /on; a = 122;b = 37;c = 43;S = 38/mi 24 24 25 24 Allotype 9,: - 80 1900 3393 3460/un;a=121;b=44;c=52; V=55% 24 25 29 21 = 3090/mi; a=125; b = 51; c = 54; = 3285/mi; a=131; b = 40; c = 54; a=110; b = 38; c = 50; a=131; b = 38; c = 53; (Spicule and maximum body diameter measured as in C. macintyrei). DESCRIPTION. Cuticle finely striated from the base of the cephalic capsule, but most conspicuous in the oesophageal and immediate post-oesophageal regions. Typically, the cuticle is covered with a coating of coccoid blue-green algae, but this tends to be lost during preservation. The cuticle of the cephalic capsule has a feint punctated or dotted appearance viewed with interference contrast microscopy although this is difficult to depict: it is not obvious with ordinary illumination. There are rows of setae in the oesophageal region but somatic setae are absent from the rest of the body. R, sensilla not seen. Six small (about 1 //m) setose R2 sensilla. Four 17-21 //m R3 setae, about 80% of maximum cephalic capsule diameter and situated at the anterior extremity of the head. Four shorter subcephalic setae and four sublateral subamphidal setae. This disposition of the setae on the anterior part of the head was the same in all specimens. Amphid ventrally wound, situated just posterior to the R? setae and 6-7 //m wide. From a lateral view, the amphid appears oval but when the head is bent slightly the amphid can be seen to be round (Fig. 4c). Buccal cavity a narrow funnel in the anterior part of the buccal bulb. Tail conical with a characteristic ventral inflection in the slightly more cylindrical terminal portion. Spicules paired, curved and proximally cephalate: average length 36 //m (arc) or 27 //m (chord). The proximal cephalation has a characteristic knob which is easy to distinguish even at low magnification (Fig. 4f). Gubernaculum well cuticularised and dorso-caudally ^directed. Testis single and outstretched. Tail has three pairs of subventral setae and a single precloacal ventral seta was present in at least two males. Ovaries paired and apparently outstretched. DIFFERENTIAL DIAGNOSIS. Catanema smo sp. nov. can be distinguished from most other Catanema species with only four subcephalic and four subamphidal setae by the anterior position of the amphid. The only other species with its amphids so far forward is C. exile Cobb, 1920, which has 7 well developed pairs of tubular caudal supplements. DISCUSSION. Superficially, both the species found in the same samples from Loch Ewe appear to be similar. However, on more detailed examination they were shown to be separate species on a number of characters. In practice, the more distinct knob at the proximal end of the spicule could be used to distinguish males of C. smo at relatively low magnification once the existance of two separate species was realized. Other differences include the presence of two conspicuous subamphidal setae in C. smo, presence of somatic setae throughout the body in C. macintyrei and slightly smaller spicules in C. smo. Both species were found mainly 9-12 cm deep in the sand, the epizoic algae being a common feature on deeper-living nematodes. 236 H. M. PLATT&Z. N.ZHANG Ceramonema yunfengi sp. nov. Fig. 5(a-Ki) MATERIAL STUDIED. Holotype: a"«0 2 1 1 +' 1 + r* ** c B c + 1 1 1 O rt CO S C/i •0"3 ag ^M IO CO *^ OO t — • es- e^- — i p— t>- ^" ^o ^ s>" r~» r^ -H — — r^ o CO C oo I V 1 1 . — _j_ 3 o CO .a u * * * * .C 00 ~!r iO v^ ^^ vO *^~> fN C^ ^O CO *^ C?N &&- O 0 U « * * * * s CO "*•? t^ *O ro *O ^D OJ OO ^" *^ f*^ oo ^ "5 -^ ooooooooooo s: 35 0 o |3 j= ^) aj= u «-> GO 0 00 (N C?\ t~~ TT OO ~* ^ <^> 3 *^ 4> ^— ^^ ^^ ^^ c — _ 00 a> s CO m-rj-mt^mfNooO^ooo ~*Tt, *© c V) o jj X O"O O«o"o O"O*O 3 O^D 1 C/3 u- 0 ~ ^- w c I1 Table I Tabular 00 U 'G _ „, (vj r~ ^ Sd2.rr> "" g-^-j in ^ _^ ON ovc"""1 °Na""f'^ S^gON.^U 0 X. — 3-0' — ^ T3 C Q — •t- uT ^ C '5 cS^r'S^ O^ u2S OQ S^> s • S & $ Q £» O 9 U *-i ^ "ts 1> 2 j™ C " S rv ^calculated from I +'vacuoles arrang NEW SPECIES OF MARINE NEMATODES 239 Fig. 6 Pictorial key to the species of Ceramonema: (a) C. attenuatum 9 (after Cobb, 1920); (b) C. carinatum d (after Wieser, 1959); (c) C. chitwoodi 9 (after De Coninck, 1942); (d) C.filipjevi d (after De Coninck, 1942); (e) C. pisanum <3 (after Gerlach, 1952); (0 C. pisanum juvenile 9 (after Gerlach, 1952); (g) C. rectum 9 (after Gerlach, 1957); (h) C. reticulatum d (after Chitwood, 1936); (i) C. salsicum d (after Gerlach, 1956); (j) C. sculpturatum juvenile (after Chitwood, 1936); (k) C. undulatum 9 (after De Coninck, 1 942); (1) C. yunfengi sp. nov. DIMENSIONS. Holotype rf, : Allotype 9, : - 115 M 550 15 23 23 19 - 125 455 680 930/rni; a = 17 25 32 23 = 7-4;c = 3-7; V = 49°/o cT2: L = 860 //m; a = 30; b = 7'5; c = 3'7 a = 35; b = 7'3; c = 3'5 a = 29*; b = 7'3; c = 4-2 a = 25*; b = 7'5; c = 3'5 4 and d"5 were slightly flattened. 240 H. M. PLATT & Z. N. ZHANG Fig. 7 Acantholaimus ewensis: (a) whole body of d1,; (b) head of d1,; (c) head of 9, showing surface structures only; (d) head of 9, showing buccal cavity structure. Bar scales: a=100wm; b-d= 10//m. DESCRIPTION. Cuticle punctated and laterally differentiated. Punctation extends anterior to amphids (Fig. 7c) and in the anterior oesophageal region the lateral punctations are irregular. Further posteriorly, the punctations become smaller and arranged in transverse rows. In the middle of the body, the transverse rows of punctations can be difficult to distinguish. Lateral differentiation consisting of two longitudinal rows of larger punctations begins level with the middle of the posterior oesophageal bulb and ends at about 65% of the tail length. The lateral rows are about 1 -5 /zm apart and there is a space between these rows and the transverse rows of punctations (Fig. 8f). There are four sublateral rows of hypodermal pores, which are larger NEW SPECIES OF MARINE NEMATODES 241 anteriorly (Figs 8d-f). Long somatic setae are present throughout the length of the body and arranged more or less in four sublateral rows. In the oesophageal region, the length of the somatic setae increases from 15-20//m at the anterior to 3 5-40 //m at the posterior. In the middle of the body, the longer setae are about 45 //m long but decrease in length again to about 30-40 //m in the anal region and 10-20 //m in the tail. At the anterior extremity there is a circle of six 2*5-3 /zm stout cephalic sensilla, presumably the R2 sensilla. Just behind are four 30-33 //m R3 cephalic setae which, in some specimens, appear to have a joint at about two-thirds of the length (Fig. 7b). Amphids 8'5-9'5//m wide (45-50% c.d.), circular with a well cuticularised boarder and a ventrally directed posterior inflection. Buccal cavity rather complex but basically it is conical with a distinct dorsal and two subventral teeth and anteriorly bears twelve rugae. Oesophagus has characteristic plasmatic interruptions (Fig. 7d) throughout its length and posteriorly there is a distinct bulb, 28 x 18 //m. Tail elongated (11-12 a.b.d.), gradually tapering but with a slightly bulbous tip and a pointed spinneret (Fig. 8c). Fig. 8 Acantholaimus ewensis: (a) tail of d1,; (b) copulatory apparatus of d,; (c) tail tip of $,; (d) d, cuticle ornamentation about 40 //m from anterior; (e) d, cuticle ornamentation level with the posterior of the oesophagus; (f) d, cuticle ornamentation at middle of body. Bar scales: a = 30 //m; b-f = 1 0 //m. 242 H. M. PLATT & Z. N. ZHANG 7A/ a Fig. 9 /J/zips paraornata: (a) anterior region of d1,; (b) head of d1,; (c) head of another cf; (d) tail of d1,; (e) copulatory apparatus of d1,; (fHO cuticle patterns at positions indicated in Fig. 10. Bar scales: a,d = 30 ^m; b,c,e-l = 10 /am. Spicules paired, equal, 26-28 ^m long and with a characteristic ventral apophysis. Gubernaculum absent. In d1,, a feint cuticularised structure was observed lateral to the spicule (Fig. 8b): it was not detected in the other specimens. There is a single stout ventral precloacal seta, 5-6 //m long. Testis single and mature specimens contain large pear-shaped sperm. Ovary paired, opposed and reflexed. In the female studied, there were sperm present in the oviduct and a large egg in each uterus containing a developing larva. DIFFERENTIAL DIAGNOSIS. Acantholaimus ewensis sp. nov. can be distinguished from Acantholaimus species with two lateral longitudinal rows of larger punctations, A. poly- NEW SPECIES OF MARINE NEMATODES 243 Fig. 10 Rhips paraornata: whole body of rf, showing positions of cuticle pattern drawings in Fig. 9f-l. Bar scale =100 //m. dentatus Gerlach, 195 1 and A. calathus Gerlach, Schrage & Riemann, 1979, on the length of the R3 cephalic setae, somatic setae and tail. A. polydentatus has shorter cephalic setae (20-22 urn, 1-5 h.d. vs. 30-33 /zm; 2'0 h.d.), shorter somatic setae (30 /zm, 0'75 c.d. vs. 45 um, 2-0 c.d.) and a shorter tail (8 a.b.d.vs. 11-12 a.b.d.). A calathus has shorter cephalic setae ( 1 2 um, 1 -0 h.d.), shorter setae (27 um, 1 -0 c.d.) and longer tail ( 1 7 a.b.d.). DISCUSSION. The genus Acantholaimus Allgen, 1933 has been fully revised recently by Gerlach, Schrage & Riemann (1979) who described seven new forms and provided a key to the twelve known species. A. ewensis fits into the first part of their key as follows: 1 (4) Cuticle laterally with two longitudinal rows of larger punctations . . 2 2 (3) Amphid situated close behind the cephalic setae 21 T (2") Tail 8 anal diameters long A. polydentatus Gerlach, 1951. 1" (2') Tail Tl-12 anal diameters long A ewensis sp. nov. 3 (2) Amphid situated half the head diameter behind the cephalic seta. Tail 1 7 anal diameters long. A. calathus Gerlach, Schrage & Riemann, 1979 4 (1) Cuticle with limited or no lateral differentiation, without longitudinal rows of punctations 5 Rhips paraornata sp. nov. Figs 9- 10 MATERIAL STUDIED. Holotype: rf, BM(NH) 1 98 1 .4. 19. Allotype: 9, BM(NH) 1981.4.20. Paratypes: nine males and two females; rf6 BM(NH) 1 98 1 .4.2 1 , remainder in collection of Z.N.Z. 244 H. M. PLATT & Z. N. ZHANG DIMENSIONS. Holotyperf,: - 180 M 990 1 1050m;a = 48;b = 6-l;c = 9-6;S = 7 11 21 23 22 Allotypeg,: - 179 M 938 10800m;a = 37;b = 6-0;c = 7-6; V = 54% 10 26 29 20 rf2:L= 10850m; a = 38; b = 6'l; c=8'7 dy.L=13300m; a = 50; b = 6'5; c= 9*3 rf4:L=11300m; a = 48; b = 6'5; c=ll'9 rf5:L=12600m; a = 44; b = 6'4; c= 8'5 d"6:L=12600m; a = 47; b = 6*9; c=ll'0 d"7:L= 9200m; a = 42; b = 5'3; c= 8-7 d-8:L=12800m; a = 53; b = 6'7; c=10'0 cT9:L=12750m; a = 53; b = 7'4; c=13'0 <*w'.L= 11600m; a = 50; b = 6'9; c=9'0 92:L=10950m; a = 44; b = 6'6; c= 9'0 93:L= 9800m; a = 36; b = 6'5; c= 7-3 DESCRIPTION. The cuticle is conspicuously annulated and heterogeneously ornamented laterally. In the anterior third of the oesophagus, from about the point where the body characteristically narrows (Fig. 9a) each annule bears a row of long and a row of smaller round punctations (Fig. 90, the latter being difficult to distinguish and liable to be over- looked. In the posterior part of the oesophagus, the punctations are smaller and appear to be partly linked diagonally (Fig. 9g), giving a reticulated appearance. The regular nature of the ornamentation breaks down posterior to the oesophagus and lateral differentiation of large dumb-bell-shaped punctations begins (Fig. 9h). Just posterior to the mid-point of the body, the pattern of the lateral differentiation reverses and, at the point of change, the dumb-bell- shaped lateral punctation is represented by a single round dot (Fig. 9j). Lateral differentiation ends just anterior to the anus and transverse rows of discrete punctations are present on the tail (Fig. 91). Short 4-6 0m sublateral somatic setae are present at infrequent intervals throughout the body and at a third of the oesophagus length, there are four longer somatic setae (Fig. 9a). The head bears six setose R, sensilla. The six 2-3 0m R2 sensilla are situated just anterior to the four shorter (1*5-2 0m), R3 sensilla. The elongated first body annule forms a cephalic shield and bears six triangular flap-like extensions anteriorly which alternate with the R2 + R3 sensilla (Fig. 9b, c). The shield is irregularly punctated and bears the large transverse amphids. The amphid has strongly cuticularised borders and is 1 1 0m wide, about 0*85 c.b.d. The buccal cavity is surrounded anteriorly by rugae which protrude beyond the lips. There is a large pointed dorsal tooth and two smaller subventral teeth. The oesophagus widens posteriorly to a weak bulb. The tail is conical and has an unstriated end cone. The male copulatory apparatus is complex, consisting of two long double-jointed spicules, a paired gubernaculum and two lateral pieces. In the holotype, the posterior part of the spicule measures 40 0m as the arc (or 29 0m as the chord) and the anterior part measures 38 0m as the arc (or 35 0m as the chord). In five other males, the total arc length of the spicule is 70-96 0m. The lateral pieces are about 22 0m long and the gubernaculum 18 0m. The cloacal opening is surrounded by irregular cuticular excrescences and anteriorly there is a small ventral spine. The ventral part of the precloacal body rings are more thickly cuticularised: they gradually reduce in thickness anterior to the level of the proximal end of the spicules (Fig. 9d). There is a single testis situated to the right of the gut. The female has two opposed, reflexed ovaries. DIFFERENTIAL DIAGNOSIS. Rhips paraornata sp. nov. can be distinguished from the only other valid species in the genus, R. ornata Cobb, 1920, by having slightly shorter R2 cephalic setae, wider amphids, spicules with each part about the same length and possibly a differently shaped lateral differentiation. NEW SPECIES OF MARINE NEMATODES 245 DISCUSSION. This is only the fifth time that valid specimens of this genus have been reported. Timm( 1961) described a species/?, longicauda from the Bay of Bengal, but the description is very poor and based only on a single immature female: it must be considered a dubious species. The original description of the type and only other species, R. ornata, was by Cobb (1920): the specimens from Loch Ewe are clearly similar to this species from Florida. The species was found again by Wieser & Hopper (1967) who provide a brief redescription and figure the head. Allgen (1932) found what is certainly a male Rhips species from Campbell Island and considered it conspecific with Cobb's species and Gerlach (1957) found R. ornata in Brazil but did not describe it. Cobb (1920) described the unusual triangular scale-like cephalic flaps but only saw the four sublateral ones: Wieser & Hopper (1967) correctly observed all six. The four long cervical setae, located at about one-third the oesophagus length in R. paraornata, were not reported in R. ornata but they are fine and may be lost on handling so that their absence in R. ornata cannot be assumed. The four sublateral cephalic flaps were positioned 'just in front of the ends of the amphids' in R. ornata according to Cobb (1920) and as figured by Wieser & Hopper (1967), whilst the ends of the amphids in R. paraornata extend further beyond the flaps. Both Cobb (1920) and Wieser & Hopper (1967) describe the lateral differentiation in the posterior portion of the body as being 'V-shaped but unfortunately provide no illustrations of the cuticle patterns. If the lateral differentiation is similar in both species, then we would not have chosen to describe the lateral differentiation as 'V'-shaped. It is possible, then, that the cuticle patterns are different. Finally, the spicules ofR. ornata have an anterior part which is 1-5 times the length of the posterior part whilst in R. paraornata the lengths are almost the same, measured as the arc in both cases. Therefore, although there are many points of similarity in overall dimensions and general anatomy, there are sufficient points of difference in relation to the specific sizes of cephalic setae, amphids and spicules and possibly the form of the cuticle lateral differentiation to substantiate the creation of a new species for the specimens from Loch Ewe. Species of the genus Rhips seem to be very closely related to Actinonema, particularly through the excellent redescription of the common species A. pachydermatum Cobb, 1920 by Lorenzen (1972): both have similar amphids, cuticle patterns and the six triangular extensions to the cephalic shield, 'Kopfpanzer', although the flaps in A. pachydermatum are not as conspicuous as those in R. paraornatum and may have been overlooked by earlier workers. However, Actinonema does not have large double-jointed spicules and, according to Lorenzen (1972), between those structures which he terms spicules, but which resemble the lateral pieces of Rhips, there lies a single thin cuticularised tube which Lorenzen (1972) interprets as the cuticularised terminal portion of the vas deferens. We have studied several specimens of A. pachydermatum, sympatric with R. paraornata, and find the cuticularised tube very difficult to distinguish. However, in some specimens it is just possible to make out two tubes: if so, they cannot be vas deferens since A. pachydermatum is monorchic. This problem is of systematic importance since Rhips and Actinomena, together with a number of other genera including Euchromadora, are grouped together in the subfamily Euchro- madorinae. This subfamily was erected by Gerlach & Riemann (1973) without explanation but presumably because of the presence of the 'L'-shaped lateral pieces in addition to spicules and gubernaculum. For Actinonema to fit into this group, which seems reasonable through its similarity to Rhips, the spicules of Actinonema must be considered homologous with the lateral pieces of Rhips and either the cuticularised tube(s?) are vestigial spicules or spicules are absent. For the moment, this problem must remain open for further study. Acknowledgements We would like to thank Dr A. D. Mclntyre of the D.A.F.S. Marine Laboratory, Aberdeen for providing the opportunity for one of us (Z.N.Z.) to spend two years studying benthos at that institute: we named one of our new species in his honour. We would also like to thank the 246 H. M. PLATT & Z. N. ZHANG staff of the benthos section for help, encouragement and the provision of specimens, in particular Dr A. Eleflheriou and Mr D. J. Murison. References Allgen, C. 1932. Weitere Beitrage zur Kenntnis der marinen Nematodenfauna der Campbell-insel. Nyt. Mag. Naturvid. 70 : 97-198. - 1933. Freilebende Nematoden aus dem Trondhjemsfjord. Capita zool. 4 : 1-162. Andrassy, I. 1976. Evolution as a basis for the systematization of nematodes. 288pp. Pitman — London, San Francisco, Melbourne. Boucher, G. 1975. Nematodes des sables fins infralittoraux de la Pierre Noire (Manche occidentale). I. Desmodorida. Bull. Mus. natn. Hist. nat. Paris 285 : 101-128. Chitwood, B. G. 1936. Some marine nematodes from North Carolina. Proc. helminth. Soc. Wash. 3: 1-16. Cobb, N. A. 1920. One hundred new nemas (type species of 100 new genera). Contrib. to a Science of Nematology9 : 217-343. De Coninck, L. A. 1942. Sur quelques especes nouvelles de Nematodes libres (Ceramonematinae Cobb, 1933), avec quelques remarques de systematique. Bull. Mus. r. Hist. nat. Belg. 18: 1-37. Filipjev, I. N. 1918. Free-living marine nematodes of the Sevastopol area. Trudy osob. zool. Lab. Sevastop. biol. Sta. 4 : 1-350 (in Russian). Gerlach, S. A. 1951. Drei bemerkenswerte neue Nematoden aus der Kieler Bucht. Zool. Anz. 147 : 37^3. - 1952. Nematoden aus dem Kiistengrundwasser. Abh. math, -naturw. Kl. Akad. Wiss. Mainz 6:315-372. - 1956. Diagnosen neuer Nematoden aus der Kieler Bucht. Kieler Meeresforsch. 12 : 85-109. 1957. Die Nematodenfauna des Sandstrandes an der Ku'ste von Mittelbrasilien (Brasilianische Meeres-Nematoden IV). Mitt. zool. Mus. berl. 33 : 41 19630. Freilebende Meeresnematoden von den Maldiven II. Kieler Meeresforsch. 14 : 67-103. 19636. Robbea tenax sp. n., ein merkwiirdiger mariner Nematode von den Maldiven. Int. Revue ges. Hydrobiol. Hydrogr. 48 : 1 53-1 58. & Riemann, F. 1973. The Bremerhaven checklist of aquatic nematodes. A catalogue of Nematoda Adenophorea excluding the Dorylaimida. Verojf.Inst. Meeresforsch. Bremerh. Suppl. 4 : 1-404. -, Schrage, M. & Riemann, F. 1979. Die GattungAcantholaimus (Nematoda, Chromadoridae), und Beobachtungen iiber einen mutmasslichen Transportmechanismus fur Spermien bie A. calathus sp. n. Veroff.Inst. Meeresforsch. Bremerh. 18 : 35-67. Haspeslagh, G. 1973. Superfamille des Ceramonematoidea (Cobb, 1933) (Nematoda), evolution et systematique. Annls Soc. r. zool. Belg. 102 : 235-25 1 . Hopper, B. E. & Cefalu, R. C. 1973. Free-living marine nematodes from Biscayne Bay, Florida V. Stilbonematinae: contributions to the taxonomy and morphology of the genus Eubostrichus Greef and related genera. Trans. Am. microsc. Soc. 94 : 578-59 1 . Inglis, W. G. 1968. Interstitial nematodes from St. Vincent's Bay. New-Caledonia. Editions Fondat. Singer- Polignac, Paris, 19672: 29-74. Lorenzen, S. 1972. Die Nematodenfauna im Verklappungsgebiet fur Industrieabwasser nordwestlich von Helgoland II. Desmodorida und Chromadorida. Zool. Anz. 187 : 283-302. 1981. Entwurf eines phylogenetischen Systems der freilebenden Nematoden. Veroff. Inst. Meeresforsch. Bremerh. Suppl. 7 : 1-472. Mclntyre, A. D. & Eleftheriou, A. 1968. The bottom fauna of a flatfish nursery ground. J. mar. biol. Ass. U.K. 48: 113-142. & Murison, D. J. 1973. The meiofauna of a flatfish nursery ground. J. mar. biol. Ass. U.K. 53:93-118. Platt, H. M. 1973. Freeliving marine nematodes from Strangford Lough, Northern Ireland. Cah. Biol. mar. 14:295-321. Steele, J. H. & Baird, I. E. 1968. Production ecology of a sandy beach. Limnol. Oceanogr. 13 : 14-25. Timm, R. W. 1961 . The marine nematodes of the Bay of Bengal. Proc. Pakist. Acad. Sci. 1 : 1-88. Vitiello, P. 1974. Nouvelles especes de Desmodorida (Nematoda) des cotes de Provence. Tethys 5: 137-146. Wieser, W. 1959. Free-living nematodes and other small invertebrates ofPuget Sound beaches. 1 79 pp. University of Washington Press, Seattle. & Hopper, B. Marine nematodes of the east coast of North America. Bull. Mus. comp. Zool. Harv. 135 : 239-244. Manuscript accepted for publication 14 September 1981 The larval development of Crangon cmngon (Fabr. 1795) (Crustacea: Decapoda) A. R. Gurney Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD Introduction The first larval stage of Crangon crangon (Fabr., 1795) was recorded by Du Cane in 1839 and since then descriptions of some of the subsequent larval stages have been added. In 1890 Ehrenbaum described five larval stages but suggested, as they were taken from plankton samples, that his series may not have been complete. Williamson (1901) reared the first three stages in the laboratory, and selected examples from tow net plankton samples concluding that there were probably five larval stages in all. Experimental evidence on the development of larvae under non-circadian light/dark cycles given by Dalley (1980), also suggested there were five larval stages. The present study is based on specimens reared in the laboratory, and describes six larval stages, one more than those previously recorded. Materials and method Ovigerous Crangon crangon (Fabr., 1795) were trawled in the Sound, Plymouth, in March, 1980. Rearing techniques follow those of Fincham, 1977. Representatives of each stage are deposited in the Crustacea collections of the British Museum (Natural History), registration numbers 1981 : 394^00. Description of larval stages Key characters are printed in italic type', many setal counts have been omitted but these are recorded in Table 1 . ZOEA 1 (Fig. 1 ) mean size 2 mm Head (Figs la, b): eyes are sessile. Carapace (Figs la, b): without spines; rostrum short. Antenna 1 (Fig. Ic): single peduncle segment bearing external flagellar segment with three aesthetascs distally plus one seta; terminal setose spine on peduncle segment. Antenna 2 (Fig. Id): expedite broad and flat with ten setae, all plumose except the penulti- mate, subapical medial spine absent; endopodite with terminal spinous seta; spine on distal edge of peduncle segment at base of endopodite. Mandibles (Fig. le): symmetrical. Maxilla 1 (Fig. If): coxa with seven spines; basis with five spines; endopod with five setae plus one small spine. Maxilla 2 (Figs Ig, h): endopod coxa bilobed on inner margin, proximal lobe with six setae, distal lobe with four setae; basis one with four setae; basis two with four setae, endopodite with four lobes on inner margin, fourth proximal lobe with three setae, third sub proximal lobe with two setae, second subdistal lobe with one seta, distal lobe with two setae; exopodite with five setae. Bull. Br. Mus. nat. Hist. (Zool.) 42 (4) : 247-262 Issued 24 June 1 982 248 A. R. GURNEY Maxillipeds 1-3 (Figs li-1): with natatory exopodites. Pereiopod 1 (Fig. 1 m): rudimentary, biramous. Pereiopods 2-4 (Figs 1 m): rudimentary, uniramous. Pereiopod 5: absent. Abdomen (figs la, b): 5 somites, 6th somite continuous with telson. Telson (Fig. In): fans out distally, distal margin with 7+7 terminal plumose spines. ZOEA 2 (Fig. 2) mean size 2 -4 mm Head (Figs 2a, b): eyes 'stalked'. Antenna 1 (Fig. 2c): peduncle two segmented; internal distal margin of second segment extended. Pereiopod 1 (Fig. 2m): 1- segmented endopod with one apical seta; natatory exopodite. Telson (Fig. 2o): distal margin with 8 + 8 terminal plumose spines. ZOEA 3 (Fig. 3) mean size 3'0 mm Antenna 1 (Fig. 3c): external jlagellum with four distal aesthetascs, one narrower than others and one distal seta. Pereiopod 1 (Fig. 3k): 2-segmented endopodite. Telson (Fig. 3n): divided from somite 6 by suture; exopod and endopod of uropod with fringing plumose setae. ZOEA 4 (Figs 4 & 5) mean size 3'5 mm Antenna 1 (Fig. 4c): rudimentary stylocerite. Periopod 1 (Fig. 5d): 3 -segmented endopodite. Abdomen (Fig. 4b): rudimentary buds of pleopods on somites 1-5. ZOEA 5 (Figs 6 & 7) mean size 4- 1 mm Antenna 1 (Fig. 6c): external jlagellum segment with 4 wide distal aesthetascs and one narrow sub-distal aesthetasc plus one setule. Pereiopod 1 (Fig. 7d): 4-segmented endopodite. Pereiopods 2-3 (Fig. 7e): uniramous 1 -segmented. Pereiopod 4 (Fig. 7f): uniramous 4-segmented. Pereiopod 5 (Fig. 7g): uniramous 5-segmented. ZOEA 6 (Figs 8 & 9) mean size 4*2 mm Pereiopod 1 (Fig. 9d): 5-segmented endopodite. Pereiopods 2-5 (Figs 9e-f): uniramous 6-segmented. POST LARVA 1 (Figs 10-12) mean size 4-4 mm Carapace (Figs lOa, b): rostrum short; one dorso-medial tooth. Antenna 1 (Fig. lOc): 3-segmented peduncle; external flagellum 2-segmented with four distal and two sub-distal aesthetascs, internal flagellum 3-segmented; stylocerite with terminal plumose setae. Antenna 2 (Fig. lOd): exopodite with 19 plumose setae; endopodite multisegmented. Mandibles (Fig. 10e): symmetrical. Maxilla 1 (Fig. 10f): coxa with 5 spines; basis with 9 spines plus one plumose seta; endopod with one spine. Maxilla 2 (Fig. lOg): endopod reduced; exopod >22 plumose setae. Maxilliped 1 (Fig. 11 a): endopod reduced with one plumose seta; exopod with 3 short plumose setae distally plus one seta, one plumose seta proximally; epipodite present. Maxilliped 2 (Figs, lib, c): endopod 5-segmented, exopod with reduced setae. Maxilliped 3 (Fig. lid): endopod 3-segmented; exopod with very reduced plumose setae. Pereiopod 1 (Figs lie, 0: exopod reduced; endopod 4-segmented, propodus and dactylus subchelate. Pereiopod 2 (Fig. 1 Ig): 5-segmented, chelate; epipodite present. Pereiopod 3 (Figs 1 1 h, i): 7-segmented. LARVAL DEVELOPMENT OF CRANGON CRANGON 249 Table 1 Morphological comparison of larval stages 1-6 and post larva. (v = visible; d = developed; a = absent; p = present; r = rudimentary; b = biramous; re = reduced) ZOEA STAGE 1 2 3 4 5 6 PL MEAN SIZE 2*0 mm 2*4 mm 3-0 mm 3'5 mm 4'1 mm 4-2 mm 4-4 mm ANTENNA 1 No. of aesthetascs. 3 3 4 4 5 6 6 Stylocerite. a a a V V V d No. of segments. 1 2 2 2 2 2 3 ANTENNA 2 Presence of subapical medial spine. a P P P P P P Exopodite; no. of plumose setae incl. fine setae immediately after spine. 10 11 12-13 13-14 13-14 14-16 19 MAXILLA 1 No. of setae on coxa; 7 7 7 7 8 8 1 basis; 5 7 8 8 9 9 10 endopod. 5 + 1 5 + 1 5 + 1 5 + 1 5+1 5 + 1 4+1 MAXILLA 2 No. of endite setae on endopodite lobes 1,2,3, 4; 2 1.2.3 2.1.2.3 2.1.2.3 2.1.2.3 2.1.2.3 2.1.2.3 re basis 1 ; 4 4 4 4 4 4 re basis 2; 4 4 4 4 4 4 re coxa (prox. & distal). 6 + 4 7+4 7 + 4 7 + 4 7+4 7+4 re No. of plumose setae on exopodite. 5 10 10-13 13-15 15-19 20-22 >22 MAXILLIPED 1 No. of setae on endite of coxa; 4 5 5 5 5 5 re basis. 11 13 14 14 14 14 re No. of plumose setae on exopodite. 4 5 5 5 5 5 re PEREIOPOD 1 No. of segments in endopodite. r/b 1 2 3 4 5 4 No. of setae on endopodite. a 1 2 V V V - Presence of natatory exopodite. a P P P P P re PEREIOPOD 2 No. of segments. r r r r r 6 5 No. of setae. - - - 1 1 - PEREIOPOD 3 No. of segments. r r r r r 6 7 No. of setae. - - - - 1 1 - PEREIOPOD 4 No. of segments. r r r r 4 6 7 PEREIOPOD 5 No. of segments. a r r r 5 6 7 ABDOMEN Somite 6 — distal suture present. a a P P P P P TELSON No. of segments. 14 16 16 16 16 16 10 UROPODS Presence. a a P P P P P 250 A.R. GURNEY Pereiopod 4 (Figs 1 Ij, k): 7-segmented. Pereiopod 5 (Figs 1 11, m): 7-segmented. Pleopod 1 (Fig. 12a): inner margin of exopod with 6 plumose setae, 2 apical setae, external margin with 4 plumose setae. Pleopods 2-4 (Figs 12b-d): exopod inner margin 6 plumose setae, 2 apical setae, external margin with 5 plumose setae. Pleopod 5 (Fig. 12e): exopod inner margin with 5 plumose setae, 2 apical setae, external margin with 4 plumose setae. Telson (Fig. 120: 5 + 5 spines; exopod of uropod with one spine next to subapical medial spine. Discussion The larval stages of Crangon crangon (Fabr., 1795) have not previously been described solely from laboratory reared material. Williamson (1901) figured his larval series from the first three stages reared in the laboratory, supplemented by larval stages four and five and the post larva taken from plankton samples. All earlier records indicate five larval stages. In the present work six larval stages are produced consistently under laboratory conditions. The insertion of an additional larval stage prolonging development is a feature shared by other carideans (Fincham, 1977). In planktonic larvae delaying the onset of metamorphosis until conditions are favourable provides the maximum chance of survival for the mainly benthic juveniles. Minor differences related to the insertion of an additional larval stage become apparent when comparisons to Williamson's (1901) work are made. The numbers of aesthetascs recorded in this paper accord with Williamson up to stage four when he then records 6 (4 distal and 2 subdistal) compared with 5 aesthetascs (4 distal and 1 subdistal) for stage 5; 7 (4 distal and 3 subdistal) in first post larva compared with 6 (4 distal and 2 subdistal) in post larva recorded here. Maxilliped 1 showed variation in the early stages. Williamson records 10 spines on the basis in stage 1, 12 in stage 2, 13 in stage 3 compared with 11 in stage 1, 13 in stage 2 and 14 in stage 3 recorded here. Pereiopods 2-5 in the last two larval stages differed in segmentation. Williamson showed stage 4 pereiopod 2 as 1 -segmented, pereiopod 3 as 3-segmented and pereiopods 4 and 5 as 2-segmented; stage 5 pereiopods 2-5 as 7-segmented. Recorded in this paper stage 5 pereiopod 2 is rudimentary, pereiopod 3 is 1 -segmented, pereiopod 4 is 4-segmented and pereiopod 5 is 5-segmented; stage 6 pereiopods 2-5 are 6-segmented. These differences are almost certainly related to the insertion of the extra larval stage after stage 2 prior to metamorphosis. References Dalley, R. 1980. The survival and development of the shrimp Crangon crangon (L.) reared in the laboratory under non-circadian light-dark cycles. J. exp. mar. Biol. Ecol. 47 : 101-112. Du Cane, C. 1 839. Metamorphosis of Crustacea. Ann. Mag. nat. Hist. 2 : 1 78-1 8 1 . Ehrebaum, E. 1890. Zur naturgeschichte von Crangon vulgaris Fabr. In: Studien tiber Bau, Entwichklung, Lebenweise und Fanguerhdltnisse des Nordsee-Granat im Auftrage der Sektion fur Kusten-und Hochseefischerei des Deutschen Fischerei- Vereins. Moeser, Berlin. 124 pp. Fincham, A. A. 1977. Larval development of British prawns and shrimps (Crustacea: Decapods: Natantia). 1. Laboratory methods and a review of Palaemon (Palaeander) elegans Rathke, 1837. Bull. Br. Mus. nat. Hist. (Zool.)32(l) : 1-28. Williamson, H. C. 1901. On the larval stages of decapod Crustacea — the shrimp (Crangon vulgaris, Fabr.) Rep. Fishery Bd Scotl. 19 : 92-1 19. Manuscript accepted for publication 28 September 1 98 1 LARVAL DEVELOPMENT OF CRANGON CRANGON b /- \ c / I d 251 h Fig. 1 Zoea 1: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (f) maxilla 1 ; (g) maxilla 2; (h) armature of maxilla 2; (i) maxilliped 1 ; (j) armature of maxilliped 1 ; (k) maxilliped 2; (1) maxilliped 3; (m) pereiopods 1-5; (n) telson. Bar scales: a, b = 0'35 mm; c-n = 0' 1 mm. Fig. 2 Zoea 2: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 maxilla 1; (g) maxilla 2; (h) armature of maxilla 2; (i) maxilliped 1; (j) armature of maxilliped 1; (k) maxilliped 2; (1) maxilliped 3; (m) pereiopod 1; (n) pereiopods 2-5; (o) telson; (p) detail of spine, 3rd from right, on telson. Bar scales: a, b = 0'3 mm; c-p = 0' 1 mm. LARVAL DEVELOPMENT OF CRANGON CRANGON n Fig. 3 Zoea 3: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 maxilla 1; (g) maxilla 2; (h) maxilliped 1; (i) armature of maxilliped 1; (j) maxilliped 2; (k) maxilliped 3; (1) pereiopod 1 ; (m) pereiopods 2-5; (n) telson. Bar scales: a, b = 0-2 mm; c-g = 0 1 mm. 254 A. R. GURNEY Fig. 4 Zoea 4: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 maxilla 1 ; (g) maxilla 2. Bar scales: a, b = 0-3 mm; c-g = 0- 1 mm. LARVAL DEVELOPMENT OF CRANGON CRANGON 255 Fig. 5 Zoea 4: (a) maxilliped 1; (b) maxilliped 2; (c) maxilliped 3; (d) pereiopod 1; (e) pereiopods 2 & 3; (f) pereiopods 4 & 5; (g) telson. Bar scales: a-g = 0' 1 mm. 256 A. R. GURNEY Fig. 6 Zoea 5: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 maxilla 1 ; (g) maxilla 2. Bar scales: a, b = 0'3 mm; c-g = 0- 1 mm. LARVAL DEVELOPMENT OF CRANGON CRANGON 257 Fig. 7 Zoea 5: (a) maxilliped 1; (b) maxilliped 2; (c) maxilliped 3; (d) pereiopod 1; (e) pereiopods 2 & 3; (f) pereiopods 4 & 5; (g) telson. Bar scales: a-g = 0- 1 mm. 258 A. R. GURNEY ' Fig. 8 Zoea 6: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (f) maxilla 1 ; (g) maxilla 2. Bar scales: a, b = 0*3 mm; c-g = 0- 1 mm. LARVAL DEVELOPMENT OF CRANGON CRANGON 259 Fig. 9 Zoea 6: (a) maxilliped 1; (b) maxilliped 2; (c) maxilliped 3; (d) pereiopod 1; (e) pereiopods 2 & 3; (0 pereiopods 4 & 5; (g) telson. Bar scales: a-g = 0' 1 mm. 260 A. R. GURNEY Fig. 10 Post Larva 1: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 maxilla 1 ; (g) maxilla 2. Bar scales: a = 0*3 5 mm;b = 0-4 mm;c-g = 0-l mm. LARVAL DEVELOPMENT OF CRANGON CRANGON 261 Fig. 11 Post Larva 1: (a) maxilliped 1; (b) maxilliped 2; (c) detail of endopod of maxilliped 2; (d) maxilliped 3; (e) pereiopod 1; (0 detail of pereiopod 1; (g) pereiopod 2; (h) pereiopod 3; (i) detail of terminal segment pereiopod 3; (j) pereiopod 4; (k) detail of terminal segment pereiopod 4; (1) pereiopod 5; (m) detail of terminal segment pereiopod 5. Bar scales: a-m = 0- 1 mm. 262 a A. R. GURNEY be d Fig. 12 Post Larva 1: (a) pleopod 1; (b) pleopod 2; (c) pleopod 3; (d) pleopod 4; (e) pleopod 5; (0 telson. Bar scales: a-f = 0' 1 mm. A revision of the spider genus Cocalodes with a description of a new related genus (Araneae: Salticidae) F. R. Wanless Department of Zoology, British Museum (Natural History) Cromwell Road, London SW7 5BD. Introduction The genus Cocalodes Pocock, 1897, known from Amboina, Seram, Halmahera and New Guinea, is comprised of 12 species, six of which are described here as new. Petrunkevitch (1928) placed Cocalodes in the subfamily Boethinae, but as far as I am aware it shows no close affinities with the nominate genus Boethus or related genera (sensu Wanless, 1981). It is, however, closely allied to Allococalodes gen. n., proposed here for two new taxa from New Guinea. Both genera are unique in that the majority of males possess a well developed median horn arising from a sclerite between the lower basal margins of the chelicerae (Figs 8 A, 2 1 B). This structure has not been described in Cocalodes by earlier authors, for in the majority of species it does not protrude beyond the edge of the clypeus and is readily overlooked. The horn is not homologous with the paired horns ofPadilla Peckham & Peckham, which arise from the basal region of the anterior surface of the chelicerae. Neither is it homologous with the horn of Thorellia Keyserling which arises from the clypeus. The male palps of both Cocalodes and Allococalodes are also of interest in possessing a functional conductor and median apophysis which seldom occur in other Salticidae. In Allococalodes the functional conductor and median apophysis are lobe-like (Fig. 16E) and relatively simple when compared with the elaborate fan-like functional conductor and bifid median apophysis characteristic of Cocalodes (Fig. 6G). The less complex structures of Allococalodes are possibly vestigial or germinal in respect of those of Cocalodes, but for the present it is a matter for conjecture as the affinities of both genera are uncertain. The standard abbreviations and measurements are those used by Wanless (1978), but for the leg spination the system is that used by Platnick and Shadab (1975). Genus COCALODES Pocock Cocalodes Pocock, 1897:627. Type species Cocalodes leptopus Pocock, by original designation. Simon, 1901:400, 403^405. Waterhouse, 1902:80. Petrunkevitch, 1928:181. Neave, 1939, 1 : 778. Roewer, 1954 : 936. Bonnet, 1956 : 1 172. DEFINITION. Small or large spiders ranging from about 4*5 to 11 -Omm in length. Most species elongate and narrow with long slender legs, the abdomen usually marked with characteristic dark lateral bands; chelicerae robust, in males usually elongate and porrect with a median horn which sometimes protrudes beyond the clypeus; not hirsute, fringes lacking. Carapace: longer than broad, moderately high, widest at about level of coxae II-III; fovea long, weakly sulciform, positioned more or less midway between posterior margins of posterior lateral eyes. Eyes: with black surrounds except anterior medians; posterior medians and posterior laterals on moderately well developed tubercles; Bull. Br. Mus. not. Hist. (Zool.) 42(4): 263-298 Issued 24 June 1 982 264 F. R. WANLESS arranged in three transverse rows, comprised of anterior medians (AM) and anterior laterals (AL), posterior medians (PM), and posterior laterals (PL); anteriors more or less contiguous with apices procurved in frontal view and recurved in dorsal view; anterior medians largest; anterior laterals greater than half diameter of anterior medians; posterior medians relatively large, positioned closer to and on or slightly outside optical axis of anterior lateral eyes; posterior laterals as large or slightly smaller than anterior laterals and set inside lateral margins of carapace when viewed from above; quadrangle formed by posterior median and posterior lateral eyes broader than long and wider behind; entire quadrangle, measured from between bases of anterior medians to posterior margins of posterior laterals, occupying between 47 and 60 per cent of carapace length. Clypeus: between 18 and 42 per cent of diameter of anterior median eyes. Chelicerae: robust, slightly diverging and inclined anteriorly in females; in males usually elongate, diverging and porrect with median horn of variable length which arises from between the lower margins of the cheliceral bases (Fig. 2 1 B); pro- and retromargins usually with three teeth. Maxillae: long and diverging with rounded apices. Labium: oblong, generally less than half maxillae length. Sternum: more or less as in Figs 3B, 14G; slightly elongate scutiform. Coxae: I and II generally larger than HI-IV. Pedicel: short. Abdomen: long slender and tapering; spinnerets moderately long, posteriors longest with long apical articles, anteriors robust, slightly longer than more slender medians; spiracle a transverse slit just in front of anterior spinnerets; tracheal system not examined; position of colulus indicated by scant group of hairs; anal tubercle cone-shaped. Legs: long and slender; spines strong and numerous; claws pectinate; tufts present; scopulae absent. Female palps: long and slender with apical claw. Male palps: moderately complex, intra- and generally inter- specifically distinct. Femora long and bowed; patellae long; tibiae long with broad retro- lateral or ventral apophysis; cymbium with hair tuft protecting embolic tip (Fig. 2H), distal finger-like extension, swelling on retrolateral basal margin and usually a group of peg-like spines on dorsal basal margin (Figs 1 A; ISA); embolus (e) very long and slender, sometimes thread-like, resting distally in well developed membraneous fan-like functional conductoi r (c), the tip of which ends in a sclerotized spur; a bifid median apophysis arises from a trans- lucent pleated membrane on the tegulum (Figs 1 B; 4H); tegulum (t) irregular in form with seminal ducts; median hematodocha in form of a membraneous sac between tegulum and subtegulum only evident in fully expanded palps; subtegulum (st) a sclerotized ring at distal end of basal hematodocha (bh). Epigynes: of various forms; median septum sometimes present and occasionally bearing lateral pouches; introductory ducts long, simply looped or convoluted, of variable width, sometimes narrow (Fig. 8G, H), or sac-like (Figs 4F, G; 6F) or resembling spermathecae (Fig. 141, J); spermathecae of various shapes with leaf-like fertilization ducts; spherical objects frequently present in interstitial spaces (Fig. 2 1C-F). AFFINITIES. The general habitus, presence of a median cheliceral horn, functional conductor and median apophysis suggest that Cocalodes and Allococalodes are closely related, even sister groups. Both genera are also probably allied to Holcolaetis Simon, from Africa and Sonoita Peckham & Peckham from South Africa, as the male papal organs evidently possess an homologous conductor and median apophysis. DIAGNOSIS. Males of Cocalodes are readily distinguished from other Oriental salticids by the distinctive palpal organs. Females are separated with more difficulty by the geographical distribution, relatively large posterior median eyes, by the structure of the epigynes and by having three teeth on the posterior margin of the chelicerae. A more useful diagnosis or assessment of affinities cannot be given until supposedly related genera have been revised. REMARKS. 1. The sclerite from which the median horn arises in Cocalodes and Allococalodes has not been routinely examined in previous revisions and not even in all females of Cocalodes in the present study, as the chelicerae have to be displaced or detatched from the carapace to expose it. A cursory examination of several Salticidae suggests that the sclerite is present in all members of the family. It is usually seen as SPIDER GENUS COCALODES 265 ma B Fig. 1 Cocalodes papuanus Simon. Expanded d1 palp: A, retrolateral; B, prolateral. Abbreviations: bh, basal hematodocha; c, functional conductor; e, embolus; ma, median apophysis; p, peg-like cymbial spines; st, subtegulum; t, tegulum. 266 F. R. WANLESS an indistinct elongate plate embedded in tissue between the bases of the lower margins of the chelicerae. In females of Cocalodes thoracicus Szombathy and in males and females of Holcolaetis it forms a peg-like horn similar to that of C. papuanus Simon (Fig. 2 IB). This does not, however, necessarily support a relation- ship between Holcolaetis and Cocalodes as our knowledge of the development of this sclerite is inadequate. 2. The spherical objects found clustered or irregularly distributed in the interstitial spaces of the epigynes of most female Cocalodes vary in diameter from 8 to 20 //. Sometimes they can be seen through the cuticle of intact epigynes (Fig. 14H), but they are most readily observed in epigynes which have been cleared in lactic acid (Figs 20F; 21C-F). When examined by transmitted light microscopy most spheres appear to have a refractile centre, but with interferance microscopy, the centres are trans- formed into surface depressions which resemble a lunar crater with indistinct lines radiating outwards from the raised margins t(Fig. 2 IF). Some spheres appear to have collapsed and look ragged with irregular and illdefmed craters. These objects, which are not known to occur in other Salticidae, resemble the unidentified spheres described by Forster (1980) from the epigyne of a Gamaso- morpha species (Family Oonopidae). A concensus of opinion (Forster, 1980) favoured a sporozoan infection, but this seems unlikely in the present case as the spheres lack the rod-like structures illustrated by Forster, furthermore their presence in almost every female of a wide range of species suggests they are a natural feature of these epigynes, which could be associated with spermatogenisis, oviposition or even plugging. Clearly, they require further investigation. Lists of species in the genus Cocalodes Pocock, 1897 Cocalodes cygnatus sp. n. C. expers sp. n. C. innotabilis sp. n. C. leptopus Pocock, 1897 C. longicornis sp. n. C. longipes (Thorell, 1881) C. macellus (Thorell, 1878) C. papuanus Simon, 1900 C. platnicki sp. n. C. protervus (Thorell, 1881) C. turgidus sp. n. C. thoracicus Szombathy, 1915 The species of Cocalodes could on the basis of the development of the epigynal septum be divided into two groups. However, these groups have not been proposed as there does not appear to be any correlation with characters of the male palpal organs. This may be a natural phenomenon or the result of incorrectly matching males and females. Matching the sexes has been difficult in the present study and where doubts have existed, these are mentioned in the species descriptions. As is often the case, additional material should resolve many of these problems. Key to species of Cocalodes Males (the males of expers, protervus and turgidus are unknown) 1 Dorsal prong of median apophysis broad and distally truncate (Fig. 2C, F); chelicerae with dorsal cluster of stout bristles (Fig. 2 A) papuanus Simon (p. 267) - Dorsal prong of median apophysis otherwise; chelicerae lacking stout bristles 2 Peg-like cymbial spines present (Figs 4C; ISA) 3 SPIDER GENUS COCALODES 267 Peg-like cymbial spines absent 8 3 Conductor tip cygniform (Fig. 6G) cygnatussp. n. (p. 273) - Conductor tip otherwise 4 4 Dorsal and ventral prongs of median apophysis more or less equal in length (Fig. IOC, D) thoracicus Szombathy (p. 280) - Dorsal prong of median apophysis much longer than ventral prong .... 5 5 Dorsal prong of median apophysis finger-like or scimitar-shaped (Figs 1 3C, E; 1 5D, E) 6 Dorsal prong of median apophysis acuminate (Figs 4C; 7H, I) 7 6 Conductor tip forming a triangular plate (Fig. 13G); dorsal prong of median apophysis scimitar-shaped (Fig. 13 E) longipes (Thorell) (p. 284) Conductor tip otherwise (Fig. 1 5F); dorsal prong of median apophysis finger-like (Fig. 15D, E) innotabilis sp. n. (p. 288) 7 Conductor tip slender with minute barb (Fig. 7G, I) .... macellus (Thorell) (p. 275) Conductor tip robust, barb lacking (Fig. 4H) leptopus Pocock (p. 270) 8 Dorsal prong of median apophysis relatively slender (Fig. 14D, E); cheliceral horn not protruding beyond clypeal margin platnickisp. n. (p. 286) - Dorsal prong of median apophysis relatively robust (Fig. 8B, C); cheliceral horn protruding well beyond clypeal margin (Fig. 8A, D) . . . . longicornis sp. n. (p. 277) Females (the female of innotabilis is unknown) 1 Epigyne with median septum Epigyne lacking median septum 7 2 Epigyne with relatively large openings (Fig. 3D) papuanus Simon (p. 267) - Epigyne with relatively small openings 3 3 Epigynal septum narrow and not extending to posterior margin (Fig. 7C) ... 4 - Epigynal septum broad, extending to posterior margin 6 4 Clypeus white haired; epigynal openings more or less ovoid in outline (Figs 9B, 7C) 5 Clypeus not white haired; epigynal openings indistinct (Fig. 8F) . . longicornis sp. n. (p. 277) 5 Spermathecae large and dark (Fig. 9 B) protervus (Thorell) (p. 279) Spermathecae small and pale, hardly if at all visible through integument (Fig. 7C, D) macellus (Thorell) (p. 275) 6 Epigynal septum with median bulge (Fig. 12 B) turgidus sp. n. (p. 283) Epigynal septum lacking median bulge (Fig. 1 1 B, C) . . . thoracicus Szombathy (p. 280) 7 Epigyne with small median openings and broad posterior ledge (Fig. 1 4H) platnicki sp. n. (p. 286) - Epigyne otherwise 8 8 Epigyne slightly depressed with comma-shaped introductory ducts (Fig. 1 3B) . longipes (Thorell) (p. 284) Epigyne otherwise 9 9 Epigyne with deep median notch on posterior margin (Fig. 5C) . . . expers sp. n. (p. 27 1 ) Epigyne otherwise 10 10 Epigyne with two notches on posterior margin (Fig. 6 D) .... cygnatussp. n. (p. 273) Epigyne without notches on posterior margin (Fig. 4E) . . . . leptopus Pocock (p. 270) Cocalodes papuanus Simon (Figs2A-H;3A-E; 18C,E; 19A,B;21B) Cocalodes papuanus Simon, 1900 : 32, rf. LECTOTYPE rf (here designated) Irian Jaya (MNHN, Paris) [examined]. Simon, 1901:403^04. Roewer, 1954:936. Bonnet, 1956:1173. Proszynski, 1971 :390. Cocalodes armatissimus Strand, 1913:122, d. LECTOTYPE