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.
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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 <?, 5 9 Ixodes loricatus
ex. Lutreolina crassicaudata
Parana Delta (vicinity of 33°43'S, 59°15'W),
Argentina
X.1916
Robin Kemp
K113 1912.6.21.289-292
Ixodes loricatus var. spinosus (4 d1 Ixodes luciae)
ex. Didelphis marsupialis
San Esteban (1 1'28'N, 69'22'W), Venezuela
31.1.1911
Kl 14 None
4 9, 1 5 N, 1 L Ixodes ornithorhynchi
ex. Ornithorhynchus paradoxus (Ornithor-
hynchus anatinus)
Tyalgum Tereed River, perhaps Tyalgum Creek
(28°22'S, 153'13'E), New South Wales,
Australia
IX.1912
Received from W. F. H. Rosenberg
K115 1912.6.21.272-3
Ixodes rasus (2 9 Ixodes cumulatimpunctatus)
ex. Cricetomys gambianus
Bibianiha (05°43'N, 00°27'W) or other populated
places of this name nearby, Ghana
23.XII.1911
H. G. F. Spurrell
K116 1912.6.21.264
3 9 Ixodes ricinus (2 9 Ixodes ricinus)
ex. Canis familiaris
Porlock Weir (5 1'12'N, 03°40'W), England
27.IV.1912
F. J. Cox
Ixodes ricinus is a vector of the viruses Tettnang,
Tribec, Louping ill, tickborne encephalitis, West
Nile, Crimean Congo hemorrhagic fever,
Lipovnic, Uukuniemi, Sumakh, Eyack and
Kharagysh.
30
J. E. KEIRANS
Kl 17 None
Ixodes tasmani! (2 cf, 1 9, 1 N Ixodes tasmani)
ex. Nest of Pseudochirus cooki (Pseudocheirus
peregrinus)
N. W. Coast, Tasmania, Australia
IX.1912
R. N. Atkinson
K118 1912.10.31.7-10
Ixodes tenuirostris (1 cf, 2 9, 3 N Ixodes
trianguliceps)
ex. Microtus glareolus (Clethrionomys glareolus)
Braunton (5 1 °06'N, 04° 1 1 'W), England
21.VI.1912
W. Holland
K119 1912.6.21.267
1 N Ixodes vespertilionis
ex. Rhinolophusferrumequinum
Cave de Meaunes, Southern France (locality not
verified)
20.1.1908
Dr K. Jordan
K120 1912.6.21.268-9
4 N, 1 L Ixodes vespertilionis
ex. Bat?
Monchique (37°20'N, 08°29'W), Portugal
12.V.1910
Dr K. Jordan
K121 1912.10.31.11
1 9 Ixodes vespertilionis
ex. Rhinolophusferrumequinum
Braunton (5 1 °06'N, 04° 1 1 'W), England
22.V.1912
W. Holland
N. C. Rothschild
Note: This 9 is in alcohol but there is a pin
through the idiosoma.
K122 1912.6.21.266
2 N Ixodes vespertilionis
ex. Rhinolophusferrumequinum
St Genes de Malgoires (Saint-Genies-de-
Malgoires) (43°57'N, 04° 13'E), France
IV.1910
Albert Hugues
K123None
1 N Ixodes vespertilionis
ex. bat
Oumasch, Algeria (locality not verified)
5.III.1911
Hon. Walter Rothschild & Ernst Hartert
Note: Theodor (1967: 94) cited a collection of 2 cf ,
3 9 Nycteribia schmidlii schmidlii off Plecotus
auritus christiei, Oumasch near Biskra, Algeria,
5. III. 1911, Hon. W. Rothschild and E. Hartert.
Quite probably this bat was host for both species
of ectoparasite.
K124 1912.6.21.270
1 9 Ixodes vespertilionis
ex. Rhinolophus hipposideros
Uj-Moldova (44°44'N, 2 1 °42'E), Romania
1.V.1910
Lintia Dionisius
K 125 None
9 N, 5 L Rhipicephalus h. haemaphysaloides (4 N,
2 L Rhipicephalus haemaphysaloides)
ex. Lepussp.
Hoshangabad(22°46'N, 77°45'W), India
1912
C. A. Crump
Received from Bombay Nat. Hist. Soc.
K126 1912.6.21.203-204
Rhipicephalus kochi(\ d1 Rhipicephalus jeanneli)
ex. grass in tent
Voi (03°23'S, 38°24'E), Kenya
R. Kemp
19.IV.1910
K127None
Rhipicephalus sp. (3 c? Rhipicephalus kochi; 1 cf
Rhipicephalus longicoxatus; 6 cf, 1 9 Rhipi-
cephalus pravus)
ex. Aged donkey (Equus asinus)
Voi (03°23'S, 38°34'E), Kenya
29. IV.1910
Robin Kemp
Rhipicephalus pravus is a vector of Kadam
virus.
K128 1912.6.21.239
Rhipicephalus sp. (1 9 Rhipicephalus kochi group)
ex. Ox
Curepipe(20°19'S, 57'31'E), Mauritius
3. XI. 1911
C. Brishoe
Note: Tentative identification: Engorged mis-
shapen specimen.
K 129 None
1 cf, 1 9, 10 N Rhipicephalus pravus
Host unknown
Taveta (03°25'S, 37°43'E), Kenya
27.V.1910
Robin Kemp
K130 1953.10.9.17
1 rf Rhipicephalus pravus (1 cf, 1 N Rhipicephalus
pravus)
ex. Jerboa? (Jaculus sp. ?)
Voi (03°23'S, 38°34'E), Kenya
22. IV. 1910
Robin Kemp
K131 1911. 12.9. 1-320 (part)
Rhipicephalus sp. (25 rf, 27 9 Rhipicephalus
pravus)
Host unknown
ROTHSCHILD TICKS IN BM(NH)
31
9 Rhipicephalus
Rombo, Kilimanjaro (03°05'S, 37°20'E),
Tanzania
15. VI. 1910
Robin Kemp
K132 1912.6.21.229
1 cf, 1 9 Rhipicephalus pulchellus
ex. Homo sapiens off ground in tent
Nairobi (0 1 ° 1 7'S, 36°49'E), Kenya
25.IV.1910
Robin Kemp
K133 1919.8. 14.1-70 pt
Rhipicephalus simus (1 c
sanguineus)
ex. Vulpes vulgaris ( Vulpes vulpes)
Toscana(43°25'N, 1 POO'E), Italy
1917
N. Cimballi
Rhipicephalus sanguineus is a vector of Wad
Medani virus and the rickettsia of boutonneuse
fever.
K134 1919.8. 14.1-70 pt
Rhipicephalus simus (1 9 Rhipicephalus
sanguineus)
Host unknown
Firenze (43°46'N, 1 1 ° 1 5'E), Italy
N. Cimballi
K135 1912.12.4.18
Rhipicentor ( 1 9 Rhipicephalus sanguineus)
ex. dog (Canis familiar is)
Livingstone ( 1 2°20'S, 30° 1 8'E), Zambia
1912
A. Douglas
K136 1912.6.21.231-232
2 9 Rhipicephalus sanguineus
ex. Canis familiaris
Dehra Dun (30°19'N, 78'04'W), India
1910
Mrs R. Parson
K137 1912.6.21.234-38
6cf, 149 1 N Rhipicephalus sanguineus
ex. Canis familiaris
Manaus (03° 1 0'S, 60WW), Brazil
1911
Rev. A. Miles Moss
K138None
Rhipicephalus sp. (1 9 Rhipicephalus sanguineus)
ex. Gerbillus indicus (Gerbillus sp.)
Khandala (1 8°55'N, 73°25'E), India
11.V.1912
J. Assmuth
Note: Tentative identification: Teneral runt
female.
K139 1912.6.21.205-220
Rhipicephalus sp. (17
sanguineus)
6 9 Rhipicephalus
ex. Lepus sp.
Mountains near Droh (34°52'N, 05°53'E), Algeria
8.III.1911
Hon. Walter Rothschild & Ernst Hartert
K140 1912.6.21.201
Rhipicephalus sp. (1 9 Rhipicephalus sanguineus)
ex. Erinaceus europaeus
Saint-Genies-de-Malgoires (43°57'N, 04°13'E),
France
21. III. 1909
A. Hugues
K141 19 19.8. 14. 1-70 part
Rhipicephalus sp. (2 9 Rhipicephalus sanguineus)
ex. hedgehog
Hammam Meskoutine (36°27'N, 07°16'E),
Algeria
6.V.1914
Hon. Walter Rothschild & Karl Jordan
K142 1912.6.21.230
Rhipicephalus sp. (1 d1 Rhipicephalus simus)
ex. Vulture
Mduna River, Hlabisa District (28°10'S, 32'10'E),
Zululand Region, Republic of South Africa
27.111.1910
F. Toppin
Received from Natal Museum, Pietermaritzberg
Rhipicephalus simus is a vector of the viruses
Thogoto and Nairobi sheep disease.
K143 None
1 cf, 5 9 Rhipicephalus simus
Host unknown
Tanzania
Dr E. J. Baxter
K 144 None
1 N Rhipicephalus sp.
Host unknown
Biskra (34°50'N, 05°40'E), Algeria
25.111.1909
K145None
2 N Rhipicephalus sp.
ex. Elephantulus rufescens
Voi (03'23'S, 38°34'E), Kenya
18.IV.1910
Robin Kemp
K 146 None
10 N, 2 L Rhipicephalus sp.
ex. Elephantulus rufescens
Voi (03°23'S, 38°34'E), Kenya
19.IV.1910
Robin Kemp
K147None
2 N Rhipicephalus sp.
ex. Elephantulus rufescens
Voi (03°23'S, 38°34'E), Kenya
25.IV.1910
Robin Kemp
32 J. E. KEIRANS
Rothschild specimens missing from British Museum (Natural History) collection
K148 1912.6.21.3-20
Argas persicus
ex. fowls
Larnaca, Cyprus
K149 1911. 12.9.1-320 pt
Amblyomma gemma
Host unknown
Kilimanjaro, Brit. E. Africa
K150 1912.6.21.41 1-415
Aponomma gervaisi var. trimaculatus
ex. Iguana
Kelsey Creek, N. Queensland
K151 1919.8. 14.1-70(pt)
Haemaphysalis leachi
ex. Hedgehog
Meskoutine Hamman, Mespot.
K152 1912.10.31.36-46
Haemaphysalis leachi indica
ex. Wildcat
Harar, Abyssinia
K153 1911. 12.9.1-320 pt.
Hyalomma truncatum
ex. Donkey
British East Africa
K154 1912.6.21.265
1 9 Ixodes ricinus
Host unknown
Hamman Rirha, Algeria
K155 1919.8. 14.1-70 pt.
Ixodes vespertilionis
ex. Vespertilio euryalis
Algeria
K156 1912.6.21.188-200
Rhipicephalus sanguineus
ex. Erinaceus europaeus
Montchique [ = Monchique], Portugal
Locality list for ticks in the Rothschild collection
N = Nuttall, K = British Museum (Natural History) collections. Numbers in parentheses indicate
missing collections.
Abyssinia (Kl 52)
Algeria N561, N3537, N3538, N3539, (N3536), Kl, K2, K87, K89, K91, K92, K123, K139, K141,
K144,(K151),(K154),(K155)
Argentina N588, K19, K20, K21, K22, K34, Kl 12
Australia N546, N547, N550, N552, N553, N556, N567, N569, N633, N642, N643, N645, N646,
N650, N669, Nl 162, N121 1, N1212, N1213, N1392, N1393, N1397, (N551), (N571), (N644), K6,
K33, K40, K41, K42, K43, K44, K45, K46, K47, K49, K57, K58, K59, K60, K103, Kl 14, Kl 17,
(K150)
Belgium N597,(N631)
Bolivia N560, N562, N566
Brazil N641,(N544), K8, K10, K17, K18, K26, K27, K28, K29, K30, K31, K32, Kl 1 1, K137
British East Africa (K 1 49), (K 1 53)
Canada N1105, N1387, N1390, N1399, N1497, N1498, N1499, N1500, N1501, N1502, N1503,
N 1 504, N3502, (N596), (N 1 388)
Chile N 1389
Christmas Island, N360
Colombia K5, K24
Cyprus K84, K85,(K148)
Ecuador K23
Egypt (N539)
England N580, N593, N598, N599, N600, N601, N602, N603, N604, N605, N607, N610, N611,
N618, N621, N622, N623, N652, N1067, N1068, Nl 152, Nl 159, Nl 160, Nl 163, N3160, (N216),
(N217), (N298), (N322), (N324), (N395), (N396), (N509), (N518), (N656), K104, K105, K110,
K116,K118,K121
Ethiopia K36, K37, K38, K63, K64, K65, K66, K82, K93
ROTHSCHILD TICKS IN BM(NH) 33
France N523, K55, K108, Kl 19, K122, K140
Galapagos Islands N397
Germany, Democratic Republic of K3
Germany, Federal Republic of N608
Ghana Kl 1, K48, K80, Kl 15
Guyana K15,K16
India N572, N573, N576, N581, N582, N583, N592, N892, (N3158), (N3159), K76, K81, K125,
K136, K138
Indonesia N72, N73, N74
Ireland N61 2, (N630)
Italy K133,K134
Jamaica K9
Japan N579
Kalao Island (N76)
Kalas Island (N93)
Kenya K12, K13, K61, K75, K90, K94, K95, K96, K98, K126, K127, K128, K129, K130, K132,
K145,K146,K147
Liberia K39, K67
Louisade Archipelago Nl 166
Madagascar, Democratic Republic of N670
Malawi N563
Mauritius K127
Mexico N557, N568, N640, N647, N648, N1386, N1394, N1396, K25
Morocco K88
Papua New Guinea K51, K77, K78, K79, K99, K100, K101, K102
Paraguay N 1391
Philippine Islands N545
Portugal Kl 20, (K 156)
Romania Kl 25
Sardinia Nil 51
Scotland N667, K 107
Senegal N888 (W. Rothschild)
South Africa, Republic of N649, Nl 167, Nl 167a, K50, K142
SriLankaK35,K54
Switzerland N619, N624, N625, N1069, Nl 161
Tanzania K52, K62, K68, K69, K70, K71, K72, K73, K74, K83, K97, K131, K143
Tibet K86
Trinidad-Tobago K53
Uganda-Zaire border N559
Unknown N549, N554, N555, N558a, N558b, N564, N565, N570, N574, N575, N575x, N584, N585,
N589, N590, N591, N613, N617, N634, N637, N638, N639, N826, N1066, Nl 153, Nl 154, Nl 164,
N1395, N1398, N3138, N3330, (N75), (N77), (N78), (N90), (N91), (N92), (N516), (N517), (N548),
(N586), (N587), (N595), (N620), (N628), (N629), (N632), (N674), (N3136), (N3139), (N3140),
(N3141),(N3157),(N3351), K14
U.S.A. N594, N626, N666, Nl 399, K4, K7
U.S.S.R. K56
Venezuela Kl 13
Wales N606, N609, (N746), K109
Yugoslavia K 106
Zambia K 135
Zimbabwe N577,N578
34
J. E. KEIRANS
Host list for ticks in the Rothschild collection
Numbers in parentheses indicate missing collections. All information relating to missing collections is
taken directly from Nuttall's catalogue or British Museum (Nat. Hist.) collection cards.
AlcesalcesN\491,N\499
Anaconda (N3 141)
Anomalurus derbianusjacksoni K98
Anteater K24
Armadillo K34
Arvicola terrestris N662
Ateles geoffroyi geoffroyi N565, N640
BatN591,K120, K123
Bettongia lesueri N646
Bettongia penicillata N 1 393
Bird (N397)
Blastoceros bezoarcticus K2 1
Boa (N548)
Bradypus tridactylus (N544), (N335 1 )
Camel (N586)
Canis familiaris N572, N573, N576, N579,
N645, N892, K9, K38, K56, K60, K83, K86,
K110, K116, K135,K136, K137
Canis, sp. K19
Canis vulpes (N2 1 7)
Cat(K151)
Cat, black N504
Cat, wild K82
Cattle K9
Cercartetus concinnus N 1 1 62
Cervus capreolus (N509)
Chicken K4
Clethrionomys glareolus N6 1 9, K 1 1 8
Coendou bicolor simonsi N560
Coendou prehensilis K 1 8
Columba livia K3
Cricetomys gambianus Kl 1 5
Dactylopsida trivirgata K 1 00
Damaliscus lunatus N 1 1 67, N 1 1 67a
Dasyurus geoffroyi N650
Deer K53
Didelphis marsupialis N64 1 , Kl 1 3
Didelphis marsupialis aurita Kill
Didelphis sp. N638, N647, N648, N1391
Diemenia superciliosa (N57 1 )
Diemenia textilis N569, N642
Diomedea salvinii (N5 1 6)
Dog, 'Badlington' K67
Donkey (Kl 53)
Duck, wild N6 17
Dusicyon gymnocerus K22
Echidnas?. N3138
EirabarbaraKll, K27
Elephantulus rufescens K145, K146, K147
Equusasinus K14, K90, K127
EquuscaballusN582,K51
Erinaceus albiventris (N539)
Erinaceus europaeus N597, N599, N608, (N324),
(N631), K139,(K156)
Erithacm rubecula (N396)
Feliscatus K65
Pel is concolor N637
Felis serval K68
Fe/jssp.K63,K103
Fowls (Kl 48)
Fox (N632)
Fox, chubut K20
Fratercula arctica N634
Genetta stuhlmanni K75
Gerbillus campestris N3537, N3538, N3539
Gerbillus (Dipodillus) sp. K92
Gerbillus gerbillus N 1 398
Giraffa camelopardalis N558a, N558b, N888 (W.
Rothschild)
Grison vittata K28
Hedgehog K141,(K1 51)
Homo sapiens N1498, K9, K39, K132
Hyaena hyaena N575X, N1395, K64
Hydrochoerus hydrochaeris K23
Hydromys sp. K79
Hystrix sp. K50
Ictonyx striatus N577
Iguana (Kl 50)
Isoodon macrourus N669
Isoodon obesulus N 1 2 1 3
Jaculussp. K130
Kangaroo N546, N552, N553,(N551), K77
Lagorchestes conspicillatus N 1 392
Lagostrophusfaciatus K33
Lepussp. K81, K93, K125, K139
Lophuromys flavopunctatus zena K94, K95
Lutra vulgaris (N2 1 6)
Lutreolina crassicaudata Kl 12
Macropus dorsalis K58
Macropus fuliginosus N 1 397
Macropus sp. N633 (N644)
Manissp. Kl 1
Manis tricuspsis N564
Marten K66
Megaderma cor N 1 1 64
Meles meles K 1 06
Microtus agrestis N623
Microtus arvalis N624, N625, N 1069
Monitor (N76),(N93)
ROTHSCHILD TICKS IN BM(NH)
35
Monitor, Australian N550
Monitor strix N554
Morelia spilotes variegata N567
Mus escularis N 1 1 66
Mus minutus (N620)
Mustela erminea N602, N603, N607, N610,
N611, N612, N613, Ml 159, (N656), K105,
K107
Mustela nivalis N604, K104
Mustela putorius N600, N606, N609 (N322),
K109
Mustela vison N1399
Myocastor coypus N826
Myopotomus coypu (N674)
Myotis tricolor N649
Myrmecophaga tridactyla K25
Nasua nasua K26, K29, K3 1 , K32
Neotoma alleni vetula N 1 394
Neotragus pygmaeus K80
Nestlings N594
Notechis scutatus K49
Ochotona princeps Nl 387
Odocoileus virginianus N626, N666, Nl 386
Ophiophagus bungaris (N90)
Ornithorhynchus anatinus Kl 14
Oryctolagus sp. K72
Oryzomys longicaudatus N 1 389
Otomys tropicalis elgonis K96
OxK128
OxenK12, K13
Paradoxurus hermaphroditus K76
Passer montanus N523
Perameles sp. K78, K102
Phalacrocorax aristotelis N652, N667
Phalacrocorax graculus (N395)
Phalanger maculatus K101
Pipistrellus pipistrellus N593, Nil 51, Nil 52,
N1163
'Piquoti' N639
Plecotus auritus christiei Kl 23
Polecat (N746)
Procyon cancrivorus K30
Pseudocheirus sp. Kl 1 7
Pseudocheirus sp. probably K98
Puffin (N5 18)
Putorius energermanis (N 1 388)
Putorius erminea (N298)
Python sp. K5 1
Rangifer tarandus N1500, N1501, N1502,
N1503
Rattus macleari N360
Raven K36
Rhinoceros N563
Rhinolophus ferrumequinum Kl 19, K121, K122
Rhinolophus hipposideros K124
Riparia riparia N605, N1068
Sciurus niger K7
Sciurus sp. N 1 396
Sciurus vulgaris (N630)
Scotophilus temmincki wroughtoni N592
Sheep K35, K54, K84, K85
Sminthopsis murina N 1 2 1 1
Snake, black and white N78
Snake, large N569
Sorex araneus N62 1
Speothos venaticus N562, N566
Squirrel, flying N581
Stag(N629)
Suricata suricata N578
Sus verrucosus celebensis N72
Sus verrucosus subsp. N73, N74
Tachyglossus aculeatus N547, K40, K41, K42,
K43
Talpa sp. N583
Tapirussp. K16
Tapir us terrestris K10
Tayassusp. N557
Tenerec ecaudatus N670
Testudo graeca ibera K87
Testudo pardalis (N77), (N3 140)
Tortoise N549, (N75), (N3 1 39)
Tortoise, Algerian N561
Tortoise, leopard N570
Trachysaurus rugosus K6
Turtle Kl 5
Uria lachrymans (N5 1 7)
Varanus civitatus (N9 1 ), (N92)
Varanus sp. K44, K45, K46, K47, K48
Vesper tilio euryalis (Kl 55)
Vulpes vulpes N1067, Nl 160, K133
Vulture K142
Wallabia bicolor K58
Wallabia sp. K59
Zebra (N3 157)
Zorilla N577
Acknowledgements
I am most grateful to the Misses B. E. Brewster, formerly Arachnida Section, and P. D. Jenkins,
Mammal Section, Zoology Department, British Museum (Natural History), and to Mr D. Macfarlane,
Commonwealth Institute of Entomology, for assisting me in obtaining correct host designations and
locality coordinates. I also thank Dr K. C. Kim, The Pennsylvania State University, for determination
of the Anoplura.
36 J. E. KEIRANS
References
Anonymous (Entomological Research Committee). 1910. Collections Received. Bull. ent. Res. 1 : 87.
Arthur, D. R. 1953. The systematic status of Ixodes percavatus var. rothschildi Nuttall & Warburton,
1911. Parasitology, Cambridge 43 : 222-226.
— 1963. British Ticks. Butterworth & Co. Ltd., London. i-ix, + 213 p.
Cooley, R. A. & Kohls, G. M. 1945. The genus Ixodes in North America. Natn. Inst. Hlth. Bull. 184.
246 p.
Hirst, S. 1914. Report on the Arachnida and Myriopoda collected by the British Ornithologists' Union
Expedition and the Wollaston Expedition in Dutch New Guinea. Trans, zool. Soc. Lond.
20 : 325—334.
1916. On a new variety of European tick (Dermacentor reticulatus, var. aulicus, var. nov.). Ann.
Mag. nat. Hist. (8), 17 : 308.
Hoogstraal, H. 1953. Ticks (Ixodoidea) of the Malagasy faunal region (excepting the Seychelles).
Their origins and host-relationships; with descriptions of five new Haemaphysalis species. Bull. Mus.
comp. Zool. Harv. Ill : 37-1 13.
1955. Notes on African Haemaphysalis ticks. I. The Mediterranean-littoral hedgehog parasite H.
erinacei Pavesi, 1884 (Ixodoidea, Ixodidae). J. Parasit. 41 : 221-233.
1956. African Ixodoidea. I. Ticks of the Sudan. Dep. Navy, Bur. Med. Surg.; Washington, D.C.
1101 p.
1965. Haemaphysalis tibetensis sp. n., and its significance in elucidating phylogenetic patterns in
the genus (Ixodoidea, Ixodidae). J. Parasit. 51 : 452-459.
, Carney, W. P., Kadarson, S. & Van Peenen, P. F. D. 1973. Haemaphysalis (Kaiseriana)
celebensis Hoogstraal, Trapido, and Kohls (Ixodoidea, Ixodidae), a Wallacean member of the
hystricis group: Identity, distribution, hosts, and ecology. /. Parasit. 59 : 556-562.
, Trapido, H. & Kohls, G. M. 1965a. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea,
Ixodidae). The identity, distribution, and hosts of H. (Kaiseriana) hystricis Supino. J. Parasit.
51 : 467^80.
& 19656. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H.
(Kaiseriana) celebensis sp. n., from a wild boar in Celebes. J. Parasit. 51 : 1001-1003.
Hopkins, G. H. E. & Rothschild, M. 1953-1971 . An Illustrated Catalogue of the Rothschild Collection
of Fleas (Siphonaptera) in the British Museum (Natural History). Vols. I-V. British Museum
(Natural History), London.
Kaufman, T. S. 1972. A revision of the genus Aponomma Neumann, 1899 (Acarina: Ixodidae). Ph.D.
thesis University of Maryland, 289 pp.
Neumann, L. G. 1904. Notes sur les Ixodides. II. Archs. Parasit. 8 : 444^64.
Nuttall, G. H. F. 1910. New species of ticks (Ixodes, Amblyomma, Rhipicephalus). Parasitology,
Cambridge 3 : 408^1 16.
& Warburton C. 1911. Ixodidae. Section II. The genus Ixodes. In Nuttall el ai, Ticks. A
monograph of the Ixodoidea. Part II: 1 33-293. Cambridge University Press, Cambridge.
& 1915. Ixodidae. The genus Haemaphysalis. In Nuttall el «/., Ticks. A monograph of the
Ixodoidea. Part III: 349-550. Cambridge University Press, Cambridge.
Roberts, F. H. S. 1960. A systematic study of the Australian species of the genus Ixodes (Acarina:
Ixodidae). Aust. J. Zool. 8 : 392-485.
1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina:
Ixodidae). Aust. J. Zool. 11 : 35-80.
1970. Australian Ticks. Commonwealth Scientific and Industrial Research Organization,
Australia; Melbourne, i-iv, 267 p.
Robinson, L. E. 1926. The genus Amblyomma. In Nuttall el ai, Ticks. A monograph of the Ixodoidea.
Part IV: 1 — 302. Cambridge University Press, Cambridge.
Taylor, F. H. 1946. Spiders, ticks and mites including the species harmful to man in Australia and New
Guinea. Section 1 . Descriptive. Serv. Publs. Sch. publ. Hlth. trop. Med. Univ., Sydney. (6): 7-234.
Theodor, O. 1967. An illustrated Catalogue of the Rothschild Collection of Nycteribiidae (Diptera) in
the British Museum (Natural History). London-British Museum (Natural History), Publication No.
655. pp. viii + 506 + 5 plates.
Warburton, C. & Nuttall, G. H. F. 1909. On new species of Ixodoidea, with a note on abnormalities
observed in ticks. Parasitology, Cambridge! : 57-76.
/ u nipt. F. 1952. The ticks of sea birds. Aust. Nat. Antarctic Res. Rep. Series B. 1 : 12-20.
British Museum (Natural History)
An Illustrated Catalogue of the
Rothschild Collection of Fleas
(Siphonapterd) in the British Museum
(Natural History)
With keys and short descriptions for the
identification of families, genera, species
and subspecies of the Order
The collection of fleas in the British Museum (Natural History) ranks among the most important
in the world. It is based largely on the famous flea collection formed by the Hon. N. C.
Rothschild and presented by him to the Nation in 1913 with the proviso that a catalogue be
prepared and published. Since that time the collection has been augmented by specimens of
fleas from all parts of the world. Although the title refers to the original Rothschild collection
the work in fact deals with the whole of the British Museum (Natural History) collection, in
which some 90% of the known 2,000 or so species are represented. It is a work of
identification seldom equalled in the field of taxonomy.
This series of volumes provides a comprehensive taxonomic monograph on the group, with
keys and descriptions for the identification of families, genera, species and subspecies, with many
excellent drawings.
Volume I
Tungidae and Pulicidae
by G. H. E. Hopkins and M. Rothschild
1953, vii + 362 pp, 45 plates, 466 text figures.
£16.60
Volume II
Coptopsyllidae, Vermipsyllidae,
Stephanociridae, Ischnopsyllidae,
Hypsophthalmidae and Xiphiopsyllidae
by G. H. E. Hopkins and M. Rothschild
1956, xii + 446 pp, 32 plates, 707 text figures.
£20.00
Volume III
Hystrichopsyllidae (Acedestiinae,
Anomiopsyllinae, Histrichopsyllinae,
Neopsyllinae, Rhadinopsyllinae and
Stenoponiinae)
by G. H. E. Hopkins and M. Rothschild
1962, vii + 559 pp, 10 plates, 1,049 text
figures. £23.00
Volume IV
Hystrichopsyllidae (Ctenophthalminae,
Dinopysllinae, Doratopsyllinae and
Listropsyllinae)
by G. H. E. Hopkins and M. Rothschild
1966, vii + 549 pp, 12 plates, 926 text figures.
£33.00
Volume V
Leptopsyllidae and Ancistropsyllidae
by G. H. E. Hopkins and M. Rothschild
1971, viii + 530 pp, 30 plates, 842 text figures.
£40.00
Volume VI
Pygiopsyllidae
by D. K. Mardon
1981, viii + 298 pp, 748 text figures. £50.00
It is expected that the Catalogue will run into some 8 or 9 volumes. Further details : Publications
Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD.
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. Keirans
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
Bulletin of the
British Museum (Natural History)
Hydroids and medusae of the family
Campanulariidae recorded from the
eastern North Atlantic, with a world
synopsis of genera
P. F. S. Cornelius
Zoology series Vol 42 No 2 29 April 1982
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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World List abbreviation: Bull. Br. Mus. not. Hist. (Zool.)
<g) Trustees of the British Museum (Natural History), 1982
The Zoology Series is edited in the Museum's Department of Zoology
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ISSN 0007-1498 Zoology series
Vol 42 No 2 pp 37-148
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 29 April 1982
GENERAL &
I *
Hydroids and medusae of the family \* UBRA«V ^
Campanulariidae recorded from the eastern North ^/
Atlantic, with a world synopsis of genera
P. F. S. Cornelius
Zoology Department, British Museum (Natural History), Cromwell Road, London
SW7 5BD, England
Contents
Synopsis 38
Type designations 38
Introduction 38
Taxonomy and the phenotype 39
Ecology 43
Geographical distribution 44
Key to species 45
Taxonomic section 47
Family Campanulariidae 47
The subfamily divisions and their nomenclature 49
Subfamily Campanulariinae 50
Genus Campanularia 51
Campanularia crenata 52
Campanularia hincksii 53
Campanularia volubilis . . • 55
Genus Orthopyxis 57
Orthopyxis crenata 58
Orthopyxis integra 60
Genus Rhizocaulus 67
Rhizocaulus verticillatus 67
Subfamily Clytiinae nom. nov 69
Genus Clytia 70
Clytia discoida
Clytia hemisphaerica 73
Clytia hummelincki 82
Clytia islandica 84
Clytia linearis 84
Clytia mccradyi 87
Clytia paulensis . 88
Clytia incertae sedis . 91
Subfamily Obeliinae
Genus Gonothyraea 91
Gonothyraea loveni 92
Genus Hartlaubella 94
Hartlaubella gelatinosa
Genus Laomedea
Laomedea angulata. 98
Laomedea calceolifera 102
Laomedea flexuosa 105
Laomedea neglecta 107
Laomedea pseudodichotoma Ill
Bull. Br. Mus. nat. Hist. (Zool.) 42 (2) ; 37-148
Issued 29 April 1982
37
38 P. F. S. CORNELIUS
Genus Obelia
Obelia bidentata . . . . . •
Obelia dichotoma . . . . .
Obelia geniculata .
Obelia spp. indet 120
Taxa now referred to other families
Notes
Acknowledgements
Addendum
References 126
Index 14°
Synopsis
Hydroids and medusae of the family Campanulariidae recorded from the eastern North Atlantic are
revised. Of more than three hundred nominal species just twenty-three are regarded valid. Their
taxonomy, nomenclature, morphology, ecology, reproduction, distribution and relationships are
discussed and a key is provided for their identification. The genera occurring in the eastern North
Atlantic are revised in detail. In addition the genera not represented in this area are briefly reviewed and
notes are included to update a previous revision of the genus Obelia.
Type designations
Type material is designated of the nominal species Campanularia laevis Couch, 1844, to
facilitate comparison with Campanularia laevis Hartlaub, 1905, for which a new name is
proposed (p. 54). Lectotype material is designated ofSertularia uniflora Pallas, 1766 (p. 78),
Laomedea gracilis Sars, 1850 (for which a new name is proposed, p. 78), and Campanularia
intertexta Couch, 1844 (p. 122).
Type species are designated of two genera and a subgenus: Cymodocea Lamouroux, 1816,
family Plumulariidae (p. 121); Eucope Gegenbaur, 1856 (p. 71); subgenus Eucampanularia
Broch, 1910 (p. 52). The type genus of the subfamily Clytiinae nom. nov. is designated as
Clytia Lamouroux, 1812 (p. 69), and that of the subfamily Phialiinae (Family Lovenellidae)
asPhialium Haeckel, 1879 (p. 69).
Introduction
The family Campanulariidae is represented in all oceans. The species are found mainly
within continental shelf depths, and some occur intertidally. Many of the genera and species
are cosmopolitan. For example, many of the species recorded from New Zealand by Ralph
(1957) occur also in British waters (see Geographical distribution, p. 44).
Several European authors have revised the species of Campanulariidae recorded from
their home waters (e.g. Hincks, 1868;Linko, 1911; Broch, 1918;Nobre, 1931;Kramp, 1935;
da Cunha, 1944; Vervoort, 1946a; Leloup, 1947; Russell, 1953; Naumov, 1960, 1969;
Patriti, 1970; Rossi, 1971). But apart from the synoptic lists of Bedot (1901, 1905, 1910,
1912, 1916, 1918, 1925) and a review of the genus Obelia (by Cornelius, 1975a) there has
been no previous attempt to compare the nominal species recorded from the whole of the
eastern North Atlantic. As a result many straightforward taxonomic questions have been left
unresolved. No doubt the wide range of phenotypic variation typical of this family has
caused problems. But it is apparent also that the wide geographical ranges of many of the
species has led to unwitting duplication of work between countries. The resulting literature is
widely scattered, and this too has imposed problems since only a few libraries have all the
relevant publications.
This report attempts to revise the species of the family Campanulariidae, excepting those
in the genus Obelia, recorded from the eastern North Atlantic. Obelia is excluded since it
was revised recently (Cornelius, 1975a) but notes are included to update that revision. The
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
39
area extends southwards to the equator and west to the 30° meridian. It includes the
Mediterranean, Black and Baltic Seas, and extends in one region beyond 30° W to include the
coastal waters of Greenland. In general there have been few records of the family north of
80° N, but this must reflect collecting difficulties since several species have been reported
from these high latitudes (Linko, 1911). One dubious tropical species, Orthopyxis africana, is
not treated fully because it has been recorded only once, near the southern limit of the area.
Most genera of Campanulariidae are nearly cosmopolitan. It happens that the majority are
represented in the eastern North Atlantic. Further, most were described first from European
waters; so that a revision of the NE Atlantic genera comes close to a complete revision. A
comprehensive generic synopsis is not intended here, however, since not all the nominal
species are covered. But at least an interim generic list can be given (Table 1). Most of the
nominal genera are discussed, whether or not they have been recorded in the NE Atlantic,
and the relevant discussion sections can be found through the index. All are at least
mentioned.
Table 1 The subfamilies and accepted genera of the Campanulariidae. Those not recorded from the
NE Atlantic are treated only briefly in this paper, and are marked by an asterisk.
Subfamily/genus
Recent redefinition, if any
Notes
CAMPANULARIINAE
Campanularia Lamarck, 1816
*Eucalix Stechow 192 la
*Orthonia Stechow, 1923a
Orthopyxis Agassiz, 1862
Rhizocaulus Stechow, \9\9b
*Silicularia Meyen, 1834
CLYTIINAE nom. nov.
Clytia Lamouroux, 1812
*Gastroblasta Keller, 1883
*Tulpa Stechow, 192 la
OBELIINAE Haeckel, 1879
Gonothyraea Allman, 1864a
HartlaubellaPoche, 1914
Laomedea Lamouroux, 1812
Obelia Peron & Lesueur, 1 8 1 Oa
p. 50
p. 51
p. 58
p. 67
Ralph, 1957;Stepanyants, 1979
p. 69
p. 71
Kramp, 1961
Ralph, 1957;Stepanyants, 1979
p. 91
p. 92
p. 94
p. 97
p. 112
p. 50
p. 51
p. 50
p. 72
p. 70
Taxonomy and the phenotype
The long synonymies and remarks sections in this paper and in that on Obelia (see
Cornelius, 1975a) reflect the fact that wide phenotypic variation has led to nominal taxa
being based on unimportant characters. Even the usually reliable Hincks (1868) was misled;
but in fairness it should be remembered that Hincks, and his colleagues Alder and Allman,
were breaking new ground when seriously considering the taxonomy of this family. Now,
with a century of hindsight, the taxa they and others proposed can be better assessed. The
early workers did not realize how much these hydroids vary, compared with the usually
much less variable species of Haleciidae, Sertulariidae and Plumulariidae which they knew
already.
Some of the literature on variation in the species of this and other thecate families has been
reviewed recently (Cornelius, \915a, b, 1979). The notes in the \915a paper, on Obelia, are
now summarized and this is followed by some new comments. Authorities for the species
names are mostly omitted here since they are adequately indicated in the Taxonomic Section
(P. 47).
40 P. F. S. CORNELIUS
Colony size in erect species probably increases till mechanical breakage occurs (Crowell &
Wyttenbach, 1957, in Laomedea Jlexuosa; Cornelius, 1975a, in Obelia dichotoma) and is of
little taxonomic value. Obelia geniculata growth was studied by Ralph (1956) and Ralph &
Thomson (1968) in New Zealand. Growth was faster in cool conditions than warm.
Polysiphonic stems occur sometimes in Obelia dichotoma (cf. Millard, 1973) and Laomedea
neglecta, and are usual in Hartlaubella gelatinosa and Rhizocaulus verticillatus. The species
Obelia plicata Hincks, 1868, was based on O. dichotoma material of this kind (p. 119).
Occasional overgrowth of one colony by another of the same species is a rather different
phenomenon now called auto-epizoism (p. 1 19).
Branching was most frequent in low water temperatures in Obelia geniculata in New
Zealand (Ralph & Thompson, 1968). Similar results were obtained from 'Clytia attenuata1
by West & Renshaw (1970) who discussed the taxonomic implications (see also notes on
Growth, p. 42).
Internode length, extent of annulation, curvature, amount of asymmetric thickening and
angle of flexure have all be used to define species limits in the Campanulariidae; but only
asymmetric thickening (in Obelia geniculata} and sometimes curvature (in Laomedea
Jlexuosa) seem good characters (Cornelius, 1 975a; below, p. 1 1 3).
Perisarc tanning has been included in some species descriptions, for example in that of the
now discredited Obelia longissima (discussion in Cornelius, 1975a), but has not been taken
as a unique species character. The intensity of tanning increases with age in many species. In
some the perisarc does not seem to darken appreciably, but this might be due simply to its
thinness. Knight (1965, 1970, 1971) studied the tanning process in Laomedea Jlexuosa.
Hydranth characters are not useful at species level but the shape of the hypostome is
consistent throughout some genera. Tentacle number is usually too variable to be useful,
notably in Clytia hemisphaerica and Obelia dichotoma, but it has been used in defining
Orthopyxis crenata.
Hydrothecal characters are useful in this family. But despite their confusing intraspecific
variation (Broch, 1910, and later workers) we hardly know how the characters develop (e.g.
Berrill, 1949, in Obelia; Berrill, 1950, in Clytia hemisphaerica, Laomedea Jlexuosa &
Orthopyxis integra; Knight, 1965, in Laomedea Jlexuosa; Beloussov, 1973, in Gonothyraea
loveni). Several authors have reported chitinous structures in the hydrothecae of
Gonothyraea loveni and Obelia bidentata which, however, have proved merely to be
regularly arranged folds in the delicate hydrothecal walls. Minor variations in the pattern of
cusps on the hydrothecal rims of several of the species have been given undue weight by
some authors (see Remarks under O. bidentata). Presence or absence of the hydrothecal
diaphragm is a subfamily character but is not useful at genus or species level.
Mammen (1965) noted that an oblique hydrothecal diaphragm viewed from the 'front'
appears transverse in optical section — a point not realized by some taxonomists. It follows
that rotation of a sloping diaphragm produces a whole series of angles, from horizontal to the
true maximum slope. Further, even when correctly viewed, a transverse diaphragm is not a
consistent character (Cornelius, 1 975a).
Naumov (1969 : 123) stated that many hydroids have larger hydrothecae in cool waters
than warm, but offered data in only one species (Orthopyxis integra). Although the relation
might well be valid in many species, detailed proof is needed. Possibly it has already caused
taxonomic confusion since large, northern specimens of Clytia hemisphaerica have been
referred to the invalid C. gigantea by several authors (p. 8 1 ) — but not all of the large
specimens were from cool areas.
Hydrothecal pedicels are variable in length, in amount of annulation and in the presence or
absence of a smooth central portion. Most species vary widely in these characters. Pedicels of
reptant species are usually longer than those of upright colonies, perhaps in response to
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 41
greater feeding opportunities away from the substrate than close to it. A peculiar case is the
long hydrothecal pedicels of floating colonies of the usually reptant Clytia hemisphaerica,
which have often been regarded a distinct species (C. sarsi nom. nov., p. 78). Implicitly,
minor variations in pedicel length are phenotypic and overall length ranges are genotypic;
but evidence is lacking.
A more useful pedicel character than length is the occurrence of spiral grooving in some
species in place of the more usual annuli. Spiral grooving seems constant in those species
having it, and has not been found even exceptionally in annulated species; but annulated
pedicels have been recorded in the usually smooth to spiral Rhizocaulus verticillatus
(p. 68). When present a spirally ornamented pedicel is a safe character.
Another reliable character is the formation in some species of a sub-hydrothecal
'spherule', first described by Ellis (1755) and noted by many subsequent authors. It is formed
between two annuli just below the hydrotheca and spaced apart on the pedicel by a distance
about equal to its width (Fig. 6). Some authors have regarded possession of spherules by
distinct species as indicating affinity, and this seems usually justified. But spherules are
simple modifications of existing structures and might have evolved more than once. Thus
Clytia hummelincki is alone in its genus in having a spherule.
The spherule structure is otherwise found in the genera Campanularia, Orthopyxis and
Rhizocaulus. It recalls the basic arthropod joint in having structures analogous to arthrodial
membranes, but in other ways it resembles the vertebrate ball-and-socket joint. Considerable
passive flexibility is achieved with little materials, enabling the hydrotheca to be rapidly
orientated downstream in response to local water movement. It would seem that fewer
materials are needed in this arrangement than would be needed to construct a rigid,
unbending pedicel which could maintain the hydrotheca broadside on in strong currents.
The spherule joints seem to be an evolutionary advance on the simple annulations seen in
many species of Campanulariidae. Uniformly annulated pedicels bend a little at each
annulation and have tissue-attachment problems associated with repeated asymmetric
compression along their whole length. A hydrotheca supported by a spherule can simply
flip-flop from side to side in response to local current surges, and bends just at one point. In
addition, spherules would appear more resistant to vertical compression than annuli.
Murdock (1976) considered very briefly the role of annuli in Obelia sp. main stems. He
observed that they help bending, an obvious conclusion not often repeated. Hughes (1980)
studied Laomedea flexuosa and Obelia dichotoma at a few sites on British coasts and found
higher numbers of pedicel annuli in sheltered situations than exposed.
Gonothecal shape provides good taxonomic characters in Laomedea but in Clytia, Obelia
and Orthopyxis it does not. Some nominal species have been based on immature gonothecae,
for example the invalid 'Laomedea conferta' (p. 104).
Most species have monomorphic gonothecae but strongly marked sexual dimorphism does
occur. It has caused taxonomic confusion in Laomedea calceolifera, which was formerly
regarded as two species. The two kinds of gonothecae proved to be male and female of the
same species. Gonothecae of L. angulata, L. flexuosa and Gonothyraea loveni tend towards
sexual dimorphism, and the few L. pseudodichotoma specimens available indicate it occurs
in that species too. In the other species described herein the gonothecae are monomorphic,
so far as is known.
Nematocysts have not yet proved useful in delimiting species in Obelia (Cornelius, 19750)
but little is known of their potential value in the rest of the family (review in Ostman, 1979).
As in most hydroids, they are among the smallest of nematocysts and their study requires
refined techniques. Although Ostman reported slight differences in basal armature between
the nematocyst threads of some pairs of species here regarded conspecific [Clytia
hemisphaerica (Linnaeus, 1767) and C. sarsi nom. nov. ( = Laomedea gracilis Sars, 1850);
Obelia dichotoma (Linnaeus, 1758) and '0. longissima (Pallas, 1766)'], rather few
populations have yet been studied. She commented that microbasic mastigophores have
42 P. F. S. CORNELIUS
hitherto been separated on characters which are optical artefacts, a conclusion borne out by
some unpublished electron micrographs she has kindly shown me (Ostman, pers. comm.). If
the populations having distinct, although very similar, nematocyst types can be
distinguished on other characters also then Ostman's conclusions will be corroborated. But
the undischarged capsules are only 6 urn to 8 um long, and the fine details of thread
structures are unlikely to prove useful in routine identification. Some correlation with gross
morphological characters would seem desirable. See also page 78 concerning 'Clytia gracilis
(Sars, 1850)'.
Habitat preferences and substrate associations are discussed under ecology. Few members of
the family have a regular habitat association and most seem able to live on a wide variety of
substrates. The prime exception is Laomedea angulata, living exclusively on eel grasses; but
since other hydroids live there too this does not help in identification.
Medusa generation Russell (1953) showed best the extent to which intraspecific variation in
hydromedusae has led to many invalid species being described; but the factors controlling
this apparently phenotypic variation are almost unknown.
The problems surrounding the two nominal species of Obelia recognized from the medusa
stage in British waters still remain (summarized in Cornelius, 19750, but based largely on
Russell, 1953 and pers. comm.). In addition there is no clue as to why Obelia medusae on
release should not always be at the same stage of development. Some authors have based
nominal species of Obelia partly on tentacle number at the time of release, and although this
seems unwise the cause of the variation is still unexplained.
In contrast, four tentacles on release is characteristic of the medusae of most of the Clytia
species (see generic diagnosis, p. 71). Adult medusae of Clytia linearis, C. hummelincki, C.
paulensis and Orthopyxis crenata (but see p. 59) are undescribed, as are the hydroid stages
of C. discoida, C. pentata, C. islandica and arguably C. mccradyi. When all stages in the life
cycles of these species are known some of the nominal taxa may fall.
It is remarkable that only a little taxonomic confusion has resulted from the unusual habit
in Orthopyxis Integra of releasing medusae on some occasions and not on others (p. 63).
Growth is affected by many factors and has been widely studied in this family. Hammett
(1943) is now thought to have studied growth in Laomedea Jlexuosa, not Obelia geniculata
as he stated (Crowell, 1957; Cornelius, 19750). Toth (1969) studied colony 'senescence' in L.
Jlexuosa. Wyttenbach, Crowell & Suddith (1973) reviewed their own work on stolon
elongation in thecate and athecate hydroids, treating Laomedea calceolifera, L. Jlexuosa and
Gonothyraea loveni among the present family. They reported similar results in the two
Laomedea species but found generic differences in the growth cycles of the stolon tips.
Cyclic increases in length had been demonstrated earlier in L. Jlexuosa by Wyttenbach
(1968, 1969) alone; and Beloussov (1961) and Hale (1964) had still earlier found the same
peculiar phenomenon in stolons of Laomedea Jlexuosa and Clytia hemisphaerica
respectively. Hale (19730, b) later reported further morphogenetic work on C. hemi-
sphaerica stolons and reviewed the literature. Beloussov's (1973) important paper described
more work on the stolons of G. loveni but he did not take environmental factors into account.
Nevertheless, phenotypic responses to changes in temperature and feeding opportunity are
known to occur in the stolon of L. Jlexuosa (e.g. Crowell, 1957, 1961), mainly in 'alterations
largely due to the sensitivity of zones of prospective growth'. These observations are
interesting in themselves, but their experimental requirements make them unsuitable for
regular taxonomic use.
Phenotypic response to temperatue change was noted in the hydroid stage of 'Clytia
attenuata" by West & Renshaw (1970) who incidentally regarded that species as valid (but
see p. 40). In vitro colonies at 13°-15°C were unbranched and could not be distinguished
from ''Clytia cylindrica Agassiz'; but at 17°-19°C a kind of branching occurred which these
authors considered characteristic of C. attenuata. Whatever the validity of the two nominal
species involved, West & Renshaw drew attention to a taxonomic difficulty resulting from
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 43
phenotypic response. Their extensive review and discussion mainly concerns western North
Atlantic species ofClytia hydroids and medusae, and further comment would be out of place
here.
Finally Stebbing (1976, 1979, 1981; see also p. 107) has studied the influence of toxic
inorganic ions on the growth and death of Laomedea Jlexuosa. He found that growth was
actually enhanced at sub-inhibitory concentrations of the toxic ions. It seems that care
should be taken when assessing the morphological characters of material collected from
slightly polluted places.
Ecology
Associations. Only one of the species included here has an apparently obligatory substrate
association: Laomedea angulata, which has been reliably recorded growing only on eel
grasses. Obelia geniculata tends to occur on brown algae whereas the very similar O.
dichotoma grows usually on other substrates; but each occurs occasionally on the substrate
more usual for the other. The other North Atlantic species of the family show no marked
substrate associations. They repeat the general hydroid pattern of a few species having some
substrate specificity with the majority having only loose associations or none at all; but the
association of L. angulata with eel grasses is unusually close. Nishihira (1968) reported
'Clytia edwardsia' to be 'characteristic of Zostera marina in Japanese waters. However,
both Nishihira and Picard (1955, in Algeria) reported many hydroid species growing on
Zostera but not confined to it.
Brackish water and estuarine species. Many species of Campanulariidae are tolerant of
reduced salinity. But while many of the species included here occur either occasionally or
habitually in low salinity areas, none has been reported from fresh water. The species
comprise more than a third of the present faunal list: Laomedea angulata, L. calceolifera, L.
neglecta, Clytia paulensis, Gonothyraea loveni, Hartlaubella gelatinosa, Obelia bidentata,
O. dichotoma and O. geniculata. All records refer to the hydroid stage and none to the
medusa. Further details are given in the Habitat sections of these species.
A similar impression of the family was given by Calder (1976). He found as many as 40
brackish water hydroids in South Carolina, and of these no fewer than ten were from the
Campanulariidae.
Interactions between species. Although the phenomenon of overgrowth has been studied in a
variety of coelenterates and other colonial invertebrates (review in Larwood & Rosen, 1979),
among hydroids it has been recorded infrequently. Sustained overgrowth of one hydroid
colony by another is unusual, and in the present family there are a few instances only. For
example, occasional colonies of Obelia dichotoma with erect stems comprising more than
one hydrocaulus were once regarded a distinct species (O. plicata, p. 1 19); and the regularly
polysiphonic stems of some other species treated here may be derived in the same way
(Rhizocaulus verticillatus, p. 67; Hartlaubella gelatinosa, p. 95; Obelia bidentata, p. 113).
Overgrowth has been recorded occasionally in Laomedea neglecta (p. 107) but the species is
not well enough known for this to be assessed. Millard (1973) listed several species of
thecates from other families showing growth of one colony on another, and introduced the
descriptive term auto-epizoism.
Antagonism between colonies is widely known among other coelenterates and in many of
the invertebrate phyla (Larwood & Rosen), but like overgrowth has seldom been reported
among hydroids. Hughes (1975) reviewed work on a few species of Campanulariidae ('Clytia
volubis Packard', C. hemisphaerica, Obelia dichotoma); while in another family Warburton
(1953) recorded aggression between a colony of Hydractinia echinata (Fleming, 1828) and
one ofPodocoryne ?carnea Sars, 1 846, on a gastropod shell inhabited by Pagurus sp.
44 P. F. S. CORNELIUS
Geographical distribution
Most species of Campanulariidae are widely distributed, some occurring nearly throughout
the World in shallow waters. For example, of the 18 species recorded from New Zealand no
fewer than seven occur also in British waters (Ralph, 1957). The corresponding figures for
southern African seas are 21 species and 8 (after Millard, 1975; the immigrant Gonothyraea
loveni would be additional), and for the antarctic area 13 and five (after Stepanyants, 1979,
with some taxonomic revision). But few species of the family appear uniformly distributed,
and many have a patchy local distribution. The local variations are best documented in
European waters, towards which the following notes on the eastern North Atlantic species
are unavoidably biased. Further details are given in the Taxonomic Section under each
species.
Orthopyxis Integra. Although one of the most nearly cosmopolitan of all shallow-water
hydroids this species has not been found in the Kattegat, Skagerrak, Baltic Sea and Dutch
waters; and has only occasionally been recorded from Belgium, western Scotland and the
Irish Sea.
Clytia hummelincki. So far this species has been reported only from the West Indies, Florida,
Massachusetts, South Africa and Ghana (p. 83).
Clytia paulensis. Known for some years from parts of NW France but only recently added to
the British faunal list (p. 89). However, a specimen collected in S Devon in 1899 has now
been correctly identified. In the 1970s the species was found in Devon and Suffolk.
Gonothyraea loveni. In South Africa this species is known from Cape Town docks only, and
Millard (1975) considered it had spread from Europe to the Southern Hemisphere on ships.
Hartlaubella gelatinosa. The several nineteenth century Scottish records contrast with a
single Scottish record this century, in 1932 (p. 95). But the species is still common at least as
far north as NW England (J. Clare, pers. comm.) and the lack of recent Scottish records may
be misleading.
Laomedea angulata. There are few reliable records from the British Isles this century, in
contrast to an abundance of nineteenth century records (p. 100). Apparently the species has
yet to regain its former distribution after the temporary decline of the Zostera beds in the
1930s (described by Tutin, 1942). Although L. angulata was recorded from the Scillies in
1967 (Robins, 1969) there is apparently no other reliable British record since those from S
Devon before 1910 (Marine Biological Association, 1957). (But see Addendum.)
Laomedea calceolifera. Although widespread in North Atlantic waters, relatively con-
spicuous, and distinctive when fertile, this species has been reliably recorded only twice from
British waters (S Devon, c. 1 87 1 , by Hincks, 1871; Norfolk, in 1 95 1 , by Hamond, 1957), with
a third dubious record (Norfolk, in 1899, quoted by Hamond, 1957). Probably the species
reaches its northern limit in southern England but the paucity of British records is still
remarkable as the species is well known from NW France (p. 104).
Laomedea pseudodichotoma. This species has yet to be recorded away from the coastal
waters of tropical W Africa but it would be remarkable if this indicated the true geographical
range (p. 1 12).
Obelia bidentata. This species was first reported from British waters only some 25 years ago;
and the first record from the south coast of England is reported here (p. 115). It seems
unlikely that the nineteenth century British collectors would have overlooked so distinctive
a species, and the absence of earlier records may be genuine. The species was not known in
Europe until the 1900s. Indeed, it may have been an immigrant into E Atlantic waters from
the American coast, but this is not certain and early confusion with Hartlaubella gelatinosa
is not excluded as a reason for the absence of earlier records. See also the next species.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 45
Obelia dichotoma & O. geniculata. There are records of these widespread species on many
swimming vertebrates, including a turtle, a shark, the blueback herring and a sea-horse; and
also on drifting kelp (pp. 117, 118, 1 20). Taken together the records suggest that these species
might be transported over vast distances. The potential was realized long ago in other
hydroid families (e.g. Alcock, 1892; Lloyd, 1907; Heath, 1910; review in Gudger, 1928,
1937). Transport might explain the appearance of the previous species in European waters
around the 1900s, carried perhaps by ships rather than by vertebrates. However, turtles
regularly cross the Atlantic (Parker, 1939; E. N. Arnold, pers. comm.) so that natural means
are not excluded. Clytia hemisphaerica, another widespread species, has similarly been
found on fish, attached to their crustacean ectoparasites (p. 77). It seems plausible that
continuous transport across deep ocean basins will promote exchanges between the gene
pools of these species on different continental shelves; and might explain why many hydroid
species are virtually cosmopolitan at shelf depths. Paradoxically, it may be that the hydroid
stages of such species sometimes travel further than their medusae which live for just a few
weeks.
Key to species (hydroid stages)
Many of the species of Campanulariidae are so variable that overlap in characters occurs,
and identification by a dichotomous key is not always possible. Young and infertile
specimens are particularly difficult and even with the help of long Museum series some
specimens cannot be identified. Close study of a single, undamaged hydrotheca is often
useful and the outline of the unabraded rim can be diagnostic. Hydranth characters are
seldom useful for identification in this family.
Provisional identification can frequently be made from the illustrations of the gonothecae.
Characters based on reproductive structures are mostly omitted from the key, however, as
many specimens are infertile. When identifying fertile material it can be helpful to determine
whether the ova develop within the gonotheca or in an external acroyst, and if the gonotheca
contains developing medusae; but these characters too are mostly avoided in the key.
1 Colony with erect stems each supporting several to many hydrothecae .... 2
Colony mainly stolonal, each stem or pedicel supporting one or just a few hydrothecae . 1 8
2 Rim of hydrotheca even to sinuous 3
Rim of hydrotheca definitely cusped [Rims often abrade smooth in Gonothyraea loveni,
Hartlaubella gelatinosa and Laomedea neglecta] 10
3 Terminal region of hydrotheca flared 4
Terminal region of hydrotheca not flared 5
4 Gonothecal aperture narrow; recurved in mature 9 . Laomedea calceolifera (p. 102; Fig. 18)
- Gonothecal aperture broad, never recurved [tropical]
Laomedea pseudodichotoma (p. Ill; Fig. 2 1 )
5 Internodes curved 6
Internodes straight 8
6 Hydrotheca thickened, sometimes much so Obelia geniculata (p. 119)
Hydrotheca with little or no thickening 7
7 Hydrotheca usually 1^ times long as broad, or longer; gonothecal aperature usually raised;
releases medusa Obelia dichotoma (p. 1 1 7)
- Hydrotheca not much longer than broad; gonothecal aperture not raised; no medusa
stage Laomedea flexuosa (p. 105; Fig. 19)
8 With sub-hydrothecal spherule [a locally distributed species] Clytia hummelincki (p. 82; Fig. 10)
Lacking sub-hydrothecal spherule 9
9 Internodes rigidly straight; terminal tendrils present in autumn; gonotheca borne on stolon;
no medusa [on eel grass] Laomedea angulata (p. 98; Fig. 17)
46 P. F.S.CORNELIUS
- Internodes slightly curved; terminal tendrils unusual; gonotheca nearly always axillary;
medusa released [on many substrates, but including eel grass] . Obelia dichotoma (p. 1 1 7)
1 0 Hydrothecal cusps sharp 11
- Hydrothecal cusps blunt, square, notched or rounded 15
1 1 Hydrothecal cusps usually bimucronate (hard to see; four species difficult to separate when
immature) 12
Hydrothecal cusps all same length
Clytia hemisphaerica (p. 73; Fig. 9; see also C. mccradyi p. 87, Fig. 13)
12 Mature colony tall and bushy, with polysiphonic stem and second-order branching;
gonothecal aperture usually raised, slightly narrower than gonotheca
Obelia bidentata (p. 1 13)
Mature colony small, with up to c. 20 hydranths; stem monosiphonic (except occasionally
in L. neglecta); gonothecal aperture not raised, as broad as gonotheca .... 13
13 Hydrothecal pedicels longer than hydrotheca . . . . Clytia paulensis (p. 88; Fig. 14)
Hydrothecal pedicels roughly same length as hydrotheca or shorter 14
14 Each internode of constant diameter; hydrothecal cusps strengthened by substantial chitinous
strip; medusa released [Mediterranean southwards] . . Clytia linearis (p. 84; Fig. 12)
- Internodes slightly bulging; hydrothecal cusps without strengthening strip (folds in
hydrothecal wall can be confusing); no medusa [Mediterranean northwards] (see also young
Obelia bidentata) Laomedea neglecta (p. 107; Fig. 20)
1 5 Hydrothecal margin with rounded cusps 16
Hydrothecal margin with square cusps, often notched (abrade easily) 17
16 Mature colony large, polysiphonic; most hydrothecae with subhydrothecal spherule; no
medusa stage Rhizocaulus verticillatus(p. 67; Fig. 7)
Mature colony not usually large, always monosiphonic; no spherule; medusa released
Clytia hemisphaerica (p. 73; Fig. 9) (also C. mccradyi, ?S France only; p. 87; Fig. 13)
17 Small slender colony, stem monosiphonic; primary branching only; medusa retained as
gonomedusa external to gonotheca .... Gonothyraea loveni (p. 92; Fig. 15)
Large bushy colony, stem polysiphonic; with secondary branching; large ova, developing
into planulae within gonotheca; no medusa. . Hartlaubella gelatinosa (p. 95; Fig. 1 6)
18 Sub-hydrothecal spherule present 19
Sub-hydrothecal spherule absent 24
19 Rim of hydrotheca even 20
Rim of hydrotheca cusped or undulating 21
20 Hydrotheca usually much thickenend [common]
Orthopyxis integra (p. 60; Fig. 6) (also O. crenata with even hydrothecal rims; see text)
- Hydrotheca unthickened [scarce] Clytia hummelincki(p. 82; Fig. 10)
21 Hydrotheca > 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, <S = 9, on short pedicels; flask-shaped, smooth, aperture at end of neck of
indefinite length; neck forms after body of gonotheca; on stolon or (Hincks, 1868) on
hydrothecal pedicels.
DISPERSIVE STAGE. Planulae, brooded in the 9 gonotheca. There is no medusa stage.
REPRODUCTIVE SEASON. Apparently the only published information is of a fertile specimen
off Norfolk, 16 June, 1951 (Hamond, 1957). None of the dated specimens in the BMNH is
fertile. Possibly reproduction in this species is usually vegetative. Hamond found fertile
material just once, and only a few of the BMNH specimens have gonothecae.
DISTRIBUTION. Common from southern England northwards, but probably present in
scattered localities further south. Not recorded from NW France or Belgium, nor reliably
from the Netherlands (Teissier, 1965; Leloup, 1952; Vervoort, 1946a). However, the species
is well known from the south coast of England (Ellis, 1755; Marine Biological Association,
1957). Vervoort (1949) recorded a single specimen from the Channel Isles but was 'unable to
trace records along the NW coast of France'. There are some records from the Mediterranean
Sea (Stechow, 1923a; Riedl, 1959; Naumov, 1969) but Picard (19586) excluded the species
from his faunal list. There is a record from N Spain (Santander; Rioja y Martin, 1906) and
another from Mauritania (Billard, 193 la); but in general there are few records further south
than the British Isles.
Northerly records include: Iceland, numerous examples (Kramp, 1938); Greenland,
widespread up to 72° N (Kramp, 1943); N Norway and N coast of Russia (Mathiesen, 1928;
and Naumov, 1969, as C. groenlandica).
HABITAT. Both Mathiesen (1928, N Norway) and Naumov (1969, Russian seas) reported a
usual depth range of 25-100 m, Naumov giving extreme limits of 5-250 m. Kramp (1943,
Greenland) gave a range of 25-650 m.
Hincks (1868) stated the substrate to be other hydroids. All the BMNH material is on
hydroids, especially Tubularia larynx (sens, auct., e.g. Hincks, 1868), Hydrallmaniafalcata
(Linnaeus, 1758) and Abietinaria abietina (Linnaeus, 1758), and other sertulariids.
Although Couch (1844) reported material on the antennae of crabs and on a bivalve (Pinna
fragilis, as '.P. ingens'), his description of the gonotheca suggests his material was Clytia
hemisphaerica. He evidently confused the two species.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 57
REMARKS. Some nomenclatural confusion between the present species and Sertularia uni-
flora Pallas, 1766, is discussed below (pp. 77-78).
Rees & Thursfield (1965) suggested that C. volubilis might prove conspecific with
Rhizocaulus verticillatus (p. 67). Their evidence was some similarity in the hydrothecae
and gonothecae of the two species. However, the long BMNH series confirms the several
constant differences. The linear dimensions of hydrothecae, hydrothecal pedicels, gono-
thecae and stolon diameters in C. volubilis are about half the same dimensions in R.
verticillatus. And while none of the C. volubilis specimens has polysiphonic, erect stems
these are present in all the R. verticillatus specimens. There is no intermediate material.
Further, the perisarc tubes in R. verticillatus are nearly all parallel. Had the two forms been
conspecific, colonies of C. volubilis s. str. with some aggregation of the perisarc tubes might
have been found; but there are no such specimens in the BMNH series.
As noted by Hincks (1868) the distinction between C. volubilis and the hydroid stage of
Clytia hemisphaerica was overlooked by Johnston (1847), but recognized soon afterwards by
Alder (1 857). Couch ( 1 844) also confused the two.
Confusion between C. volubilis and the species now called Calycella syringa (Linnaeus,
1767) occurred in the mid-eighteenth century but was resolved by Linnaeus (1767) himself.
Essential details are given in the above synonymy, and further discussion in Cornelius
(1978). The species is currently referred to the family Campanulinidae.
Campanularia groenlandica Levinsen, 1893, although widely recognized, was apparently
founded on C. volubilis material from the Davis Strait. Syntype material in the BMNH shows
features present in the original illustrations of groenlandica, for example spirally sculptured
hydrothecal pedicels and blunt cusps on the hydrothecal rim; and the gonotheca shown in
the original illustration is identical with that normal in C. volubilis. Thus the two taxa
appear conspecific. Material has been recorded as C. groenlandica from Trondheim Fjord
(Mathiesen, 1928) north to 68° 20' N (Kramp, 1943; 50-525 m depth); and Naumov (1969)
recorded 'C. groenlandica' from the N coast of Russia. '
The type material of Clytia mollis Stechow, 19190, examined here, comprises immature
colonies of C. volubilis. Stechow's original illustration incorrectly shows a truncate, wide
mouthed gonotheca. It is simply a young one in which the long neck has yet to form. The
hydrothecal pedicels illustrated are topped by sub-hydrothecal spherules not present in
Clytia. Stechow tentatively included in the synonymy of 'C mollis' the nominal species
Clytia laevis Weismann, 1883, based on Naples material. However it is clear from
Weismann's description that C. laevis was founded on normal Clytia hemisphaerica
material, and it is here referred to that species.
Genus ORTHOPYXIS Agassiz, 1862
Clytia: Westendorp, 1843 : 23 (part; see Remarks under Orthopyxis Integra).
ISilicularia Meyen, 1834 : 206 (?part; see Remarks and p. 50).
Campanularia: Macgillivray, 1842:465 (part); Couch, 1844:40 (part); Hincks, 1868: 160 (part);
Millard, 1975 : 203 (part); (see Remarks below, and under O. Integra).
Clytia (Orthopyxis) Agassiz, 1862 : 297.
Clythia Agassiz, 1862 : pi. 28 (lapsus for Clytia).
Orthopyxis Agassiz, 1862 : 355; Ralph, 1957 : 834; Arai & Brinckmann-Voss, 1980 : 101.
Hincksia Agassiz, 1862 : 355 (sic).
Eucopella von Lendenfeld, 1883a : 188.
Agastra Hartlaub, 1897 : 452 (nom. mid.), 504; Kramp, 1961 : 160.
Leptomedusa Browne, 1900 : 7 14 (see notes on Nomenclature under O. Integra).
TYPE SPECIES. Clytia (Orthopyxis) poterium Agassiz, 1862; by monotypy; may be conspecific
with O. integra. Orthopyxis was introduced by Agassiz as a subgenus of Clytia on page 297 of
his work, comprising the 'new' species poterium alone; but on page 355 he upgraded it to
genus. On that page he implicitly used the combinations 'Orthopyxis (Orthopyxis)
poterium\ 'Orthopyxis (Campanularia) volubiliformis' and 'Orthopyxis (Laomedea) Integra'
58 P. F. S. CORNELIUS
(of various authors). Thus poterium should rightly be taken as type species by monotypy
of the subgenus Orthopyxis Agassiz, 1862. Nutting's (1915 : 63) designation of Campanu-
laria caliculata Hincks, 1853, as type species must be disregarded since caliculata was not
originally included. It was unfortunately repeated by Arai & Brinckmann-Voss (1980).
DIAGNOSIS. Campanulariidae forming stoloniferous or short unbranched upright colonies;
stolon anastomosing; true diaphragm absent; hydrotheca fundamentally radially symmetri-
cal but often asymmetrically thickened; medusa reduced, lacking manubrium and tentacles,
not feeding, believed facultatively retained in at least one species.
REMARKS. The genera Silicularia Meyen, 1834, and Hypanthea Allman, 18760, are
discussed on page 50.
Ralph (1957) listed some works in which Orthopyxis was discussed in relation to
Eucopella von Lendenfeld, 1883a (based partly on Campanularia bilabiata Coughtrey,
1875); and other discussion was provided for example by Bale (1914), Nutting (1915) and
Fraser (1918). The species Eucopella campanularia was described in greater detail in
another paper (von Lendenfeld, 1883/?). Bale, Ralph and others referred Eucopella to
Orthopyxis; but Hirohito (1969) held the two genera distinct on the presence or absence
respectively of marginal vesicles in the (retained) 'eumedusoid'. He referred caliculata
Hincks, 1853, to Eucopella, stating an intention to discuss the generic question later.
However, caliculata is here referred to O. Integra; and generic separation seems unjustified. I
provisionally follow several previous authors in regarding Eucopella congeneric. However,
Kramp (1961) accepted the genus and included in it the two species bilabiata Coughtrey,
1875, and crenata Hartlaub, 1901. He designated bilabiata type species. The two species
Kramp included may not be distinct: they are treated here under O. crenata (p. 60).
Kramp's generic separation, from the 'medusa' genus Agastra, was based on minor differ-
ences in the medusa and is not upheld here.
The genus Hincksia Agassiz, 1862, was proposed to accommodate solely the well
described species Campanularia tincta Hincks, 186 la; but Bedot (1910:311), Stechow
(1923a : 94) and Rees & Thursfield (1965 : 93) referred the genus to Campanularia auct. In
proposing the genus Agassiz stated merely 'The genus Hincksia is characterized by its one-
sided, ringled, fertile hydra', hardly an acceptable diagnosis. Nevertheless, type material of
C. tincta is available (BMNH reg. no. 1899.5.1.219-220), as noted by Rees & Thursfield.
Although dry the material shows the characters of the genus Orthopyxis Agassiz, 1862, and
Hincksia and Orthopyxis (not Campanularia) can be regarded congeneric. Under the first
reviser principle I retain Orthopyxis, which has been widely used, and suppress Hincksia,
which has not. O. tincta is an Australian species and so is outside the present scope, but it
may be noted in passing that the type material has an anastomosing stolon like Orthopyxis s.
str. and a highly distinctive, closely ringed gonotheca. Useful synonymies of the species were
provided by Nutting (1915), Stechow (19230) and Rees & Thursfield (1965), and a redescrip-
tion by Stepanyants (1979). Campanularia tincta sensu Warren, 1908, is mentioned here
under C. africana Stechow, \923d, a junior synonym of O. crenata (p. 60).
The genus Agastra Hartlaub, 1897, was based on Helgoland material of the medusa stage
of Orthopyxis integra (see also p. 67). Kramp (1961) resurrected the genus but there seems
no doubt that its synonymy with Orthopyxis is justified.
Orthopyxis crenata (Hartlaub, 1 90 1 )
(Fig. 5)
^.Campanularia bilabiata Coughtrey, 1875 : 291-292, pi. 20, figs 46-49.
^.Campanularia everta Clarke, 1876:251, 253-254, pi. 39, fig. 4; Garcia Corrales el at.,
1978 : 24-25, fig. 9 (syn. C. lennoxensis Jaderholm).
Eucopella crenata Hartlaub, 1901 : 364-366, pi. 22, figs 27-31, 33-35; Hirohito, 1969 : 7, fig. 7.
^.Campanularia lennoxensis Jaderholm, 1904/7 : 268-269, pi. 12, figs 4-5.
Campanularia ?inlermedia Stechow, 19 \9a : 66-68, fig. V.
^Orthopyxis delicata Trebilcock, 1928 : 3, pi. 2, fig. 1 ; Garcia Corrales el ai, 1978 : 22-23, fig. 8.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 59
Campanularia crenata forma intermedia: Picard, 195 la : 345.
Campanularia crenata: Picard, 1955: 186; Millard & Bouillon, 1973:47^18, fig. 6B-F; Millard,
1975 : 204-206, fig. 68A-F (?syn. Orthopyxis delicata Trebilcock, 1928); Garcia Corrales et al.,
1978:1 9-22, fig. 7; (non Allman, 1 8766).
Orthopyxis crenata: Trebilcock, 1928:3; Ralph, 1957:838-840, fig. 6g-v (syn. O. formosa
Trebilcock, 1928); Rees & Thursfield, 1965 : 104.
TYPE MATERIAL AND LOCALITY. The species was based partly on material from French Pass,
Bare Island, New Zealand, and partly on the original description of Campanularia bilabiata
Coughtrey, 1875. I have located none of the type material. Ralph (1957) restricted the type
locality to French Pass.
MATERIAL EXAMINED. I have seen no Atlantic material of this species.
DESCRIPTION AND IDENTIFICATION OF HYDROID STAGE. The lack of available material of this
species and the taxonomic confusion surrounding the whole genus together make redescrip-
tion difficult. The following identification notes are adapted from Ralph (1957), Millard &
Bouillon (1973) and Millard (1975). Differing from O. Integra as follows: hydrothecal rims
smooth through gently wavy to crenate, with 8-12 short rounded cusps, commonly varying
within a colony (always smooth in O. Integra); hydranth with c. 14 tentacles (> 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<a, 1938;
Stechow, 1925; Picard, 195 la; Vervoort, 1967; Millard & Bouillon, 1973; Millard, 1975;
Hirohito, 1977). Colonies both stolonal and erect. When erect branching sympodially, up to
c. 10 hydrothecae in extent, to 20 mm. Hydrothecal pedicels finely ringed throughout (6-37,
usually c. 20) or with smooth central portions; distal pedicels shorter than proximal.
Internodes of erect stems arcuate, narrowly ringed basally. Hydrothecae long, sides parallel
to slightly divergent, often with slight asymmetric bulge; with 8-16 long narrow marginal
cusps recalling those of Clytia paulensis and Obelia bidentata but each with internal
Fig. 12 Clytia linearis. The specimen illustrated is a syntype of C. alternata Hargitt, 1924, here
regarded conspecific. (a) part of colony, (b-c) hydrotheca and part of rim. (d) $ gonotheca.
Scales: (a) 500 /mi; (b) 250 /mi; (c-d) 500 //m.
86 P. F. S. CORNELIUS
stiffening strip reaching to tip and extending downwards sometimes to middle of hydrotheca;
intervening bays rather wide and square, bulging out; diaphragm usually oblique but some-
times transverse. Hydranth with 12-15 tentacles (in holotype specimen of C. alternata
Hargitt). Mature gonotheca (?rf = 9) elongate, on 2-3 ringed pedicel, either on stolon or on
erect shoot in axil; widest in centre, tapering below and usually above; one-flapped circular
deciduous operculum; young gonotheca shorter, more sharply truncated. Gonophore with
one or more rows of developing medusae; pre-release medusae with hemispherical umbrella
and four marginal tentacles.
Variation Hydrothecal length varies greatly, at least from 400 /^m to 1100//m (Millard,
1975). Internodes and pedicels are shorter in upper parts of the colony (Hirohito, 1977).
Hirohito found little difference between Red Sea and Japanese specimens, indicating a rather
constant phenotype. Hydrothecal pedicels were on the whole shorter in the Japanese
specimens, but gonothecae were similar in size. Billard (1938) reported that the embayments
between the hydrothecal cusps are often distorted in preserved material due to lateral
folding.
DISPERSIVE STAGE. A medusa, with hemispherical umbrella and four marginal tentacles on
release; earliest stage identical with newly liberated Clytia hemisphaerica medusa, even in
cnidom according to Picard (195 la). Adult medusa not described (Millard, 1975). Russell
(1953) noted that the Mediterranean medusa Phialidium viridicans Leuckart, 1856, might
prove valid. It might follow that Clytia linearis is its hydroid, but evidence is only
circumstantial and the hydroid stages of C. hummelincki and C. paulensis must also be
considered. 'P. viridicans' is here provisionally referred to C. hemisphaerica.
DISTRIBUTION. Circumglobal in tropical to warm shallow seas (Millard & Bouillon, 1973;
Hirohito, 1977) extending northwards in the Atlantic Ocean at least to the south coasts of
France ('extremely common' intertidally at Banyuls, Picard, 195 la) and Spain (10-20 m off
Alicante, Garcia Corrales et al., 1978). Billard (1907) recorded the species south of Madeira;
Picard (1955) from Algeria; Rees & Thursfield (1965, as C. striata) from the Cape Verde Is
and Rees & White (1966, as C. striata) from the Azores. Vervoort (1967) gave a list of known
Indo-Pacific localities; and some possible records under other specific names are discussed
below.
HABITAT. On cirripede and pteropod shells and on other hydroids (Billard, 1904a; Vervoort,
1967; Millard & Bouillon, 1973; Millard, 1975; Hirohito, 1977); also on intertidal rocks
(Picard, 195 la). Intertidal (Picard) to 110m (Billard, 1907; Millard).
REMARKS. Ritchie (1907) recorded material from the Azores as Clytia geniculata Thornely,
1904, a name originally applied to specimens from Sri Lanka. Rees & Thursfield (1965)
tentatively thought C. geniculata conspecific with C. striata (Clarke, 1907), which Millard &
Bouillon (1973) and Millard (1975) referred to the present species. But Hirohito (1977)
described what he considered undoubted C. striata material (on a pteropod) and considered
the species valid.
Picard (1950) assigned material from near Marseille to Clytia ?obliqua Clarke, 1907) but
that species seems identical with Clytia linearis. Clarke based the distinction merely on the
angle of slope of the hydrothecal cusps. The type material of C. obliqua, examined here, does
not otherwise differ from the original description of C. gravieri.
The type material of Clytia alternata Hargitt, 1924, was examined also by Hirohito
(1977). As he stated, it resembles the description of C. linearis so closely that the two species
can be regarded conspecific. C. foxi Billard, 1926, was based on slight differences and may
also be the same species. Vervoort (1967) drew attention to the close similarity between C.
alternata and C. gravieri Billard, 1904, which also seems conspecific. Billard (1938) had
earlier thought them conspecific along with Laomedea bistriata Leloup, 193 la. He saw four
marginal tentacle bulbs in the pre-release medusa and hence assigned the species to Clytia.
Detailed discussion of several Pacific forms was provided by Hirohito.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 87
Clytia mccradyi (Brooks, 18886)
(Fig. 13)
Oceania sp. Brooks, 1888a : 29-30.
Epenthesis mccradyi Brooks, 18886 : 147-1 62, pis 13-15; Sigerfoos, 1893 : 106.
Oceania mccradyi: Mayer, 1900 : 50, pi. 2 1 , figs 56-59.
Phialidium mccradyi: Mayer, 1910 : 271-272, pi. 34, figs 2-3, pi. 35, figs 1-3; Kramp, 1959 : 149;
Kramp, 1961 : 170; Bougis, 1963 : 2,2-2,3, pi. 9, fig. 2.
Further references were given by Mayer (1910) and Kramp ( 1 96 1 ).
TYPE MATERIAL AND LOCALITY. Green Turtle, 1886, and Nassau, 1887, Bahamas (medusa
stage only); material not located.
MATERIAL EXAMINED. None available.
DESCRIPTION OF HYDROID STAGE. The identity of the hydroid stage is unclear. The species
was originally based on mature medusae collected from the plankton. Brooks (18886) later
saw young medusae released from a hydroid colony and, identifying the young medusae as
the same species as the adults earlier described, concluded he had found the hydroid. Mayer
(1910) was sceptical. Brooks' description of the hydroid did not differ from the hydroid of C.
hemisphaerica. The description included stolonal gonothecae with lateral constrictions
characteristic of the better known hemisphaerica. Only Brooks has reported a conventional
hydroid stage. Others (Sigerfoos, 1893; Mayer, 1910; Bougis, 1963) have described a much
reduced hydroid stage comprising only hydroid blastostyles within gonothecae, attached to
the gonad of the medusa.
Fig. 13 Clytia mccradyi. Redrawn after Mayer (1910 : pi. 35, fig. 1 ). Diameter c. 15mm.
DISPERSIVE STAGE. A medusa. The following description is after Brooks (18886) and Mayer
(1910). Bell shallow, c. 15 mm diameter, less than half as high as broad, flexible; stomach
short, less than l/8th as long as diameter of bell, quadrate, with four simple, slightly recurved
lips; gonads short, oval, about mid-way between axis and bell margin; velum well developed;
1 6-24 long, contractile marginal tentacles (type material had 1 6 only), 'many times as long
as diameter of bell', each with statocyst at base; 8-16 other marginal statocysts; 1-4 (?4+)
gonothecae borne on one to all of the gonads in many individuals. Gonads of medusa
develop either eggs or sperm, or into hydroid blastostyles which produce medusae direct
(details in Sigerfoos, 1893 and Mayer, 1910). 4-8 tentacles on release (Brooks, 18886).
REPRODUCTIVE SEASON. Brooks (18886) recorded fertile hydroid material in June, 1887, at
Nassau, Bahamas; Mayer (1910) in July at Tortugas, Florida. Medusae from April to July
(Mayer).
DISTRIBUTION. In the eastern North Atlantic I know of only one record: Villefranche, S
France, found 'from time to time' (Bougis, 1963). Reported elsewhere in the World from
the Bahamas and Florida (Kramp, 1959, 1961).
88 P. F. S. CORNELIUS
HABITAT. Brooks (18886) reported the 'free' hydroid stage on algae in the harbours of Nassau
and Green Turtle, Bahamas.
The hydroid stage is at least sometimes partially suppressed and attaches to or grows on
the medusa. Thus the species seems adapted to an oceanic existence and might be found
away from coastal waters. Possibly the isolated record from the Mediterranean is evidence of
a natural trans-atlantic dispersal.
REMARKS. The unusual life cycle of this species has been checked by at least three
independent workers (Sigerfoos, 1893; Mayer, 1910; Bougis, 1963) since the original descrip-
tion by Brooks (18886). Brooks alone claimed to have found a benthic hydroid stage, but
Mayer evidently doubted his observation. Although Mayer studied live medusae of the
species closely, Brooks' account is explicit and suggests strongly that the benthic hydroids he
described were of this species. At the time Mayer wrote such plasticity in the life cycle of a
single hydromedusa species would have been regarded unusual and requiring more rigorous
proof than today, when several such examples are known (summary in Naumov, 1969).
Brooks stated that the medusa has either 4 or 8 tentacles on release, a departure from the
unvarying four usually considered normal in this genus (see p. 42).
Clytia paulensis (Vanhoffen, 1910)
(Fig. 14)
Campa nularia pa ulensis Vanhoffen, 1910 : 298, fig. 19.
Clytia ulvae Stechow, 19190 : 47-48, fig. N.
Clytia paulensis: Stechow, 19230: 110, fig. N; Philbert, 19356:25-26, fig. 4; Picard, 1955: 186;
Millard, 1966 : 481-483, fig. 15 (?syn. C. ulvae Stechow, 1919a); Millard, 1975 : 221, fig. 73a-d.
Obelia paulensis: Naumov & Stepanyants, 1972 : 37, fig. 2a-b.
TYPE LOCALITY AND MATERIAL. Shallow water in crater lagoon of St Paul Island, S Indian
Ocean (38° 40' S, 77° 34' E), 26 Apr 1903, on Sertularella polyzonias (Linneaus, 1758), coll.
Deutsche Siidpolar-Expedition 1901-1903; material not located.
TYPE MATERIAL OF OTHER SPECIES EXAMINED. Clytia ulvae Stechow, 1919<2, infertile colony
on Ulva (green alga), Marseille, microslide preparation; Munich Zoological Museum. Also 1
non-type specimen, 'Valdivia' sta. 100 (S Africa), det. E. Stechow as C. ulvae; MZM.
OTHER MATERIAL EXAMINED (All BMNH material is listed. None is fertile.). Breakwater at S
end of Landguard Pt, Felixstowe, Suffolk, SE England, ELWS, Sep-Oct 1976, infertile
colony on Crisia sp. (Bryozoa) itself on Tubularia indivisa Linnaeus, spirit + 1 microslide
preparation, coll. & det. R. G. Hughes; 1977.5.7.2. Off Berry Head, Tor Bay, Devon, SW
England, c. 15m, Mar-Jun 1973, infertile colony on Nemertesia sp., spirit* 1 microslide
preparation, coll. R. G. Hughes; 1973.8.13.1 (see Remarks; mentioned, Hughes,
1975 : 291). Off Berry Head, c. 15m, summer 1974, infertile colony on Nemertesia sp., co-
epizoic with C. hemisphaerica, spirit + 1 microslide preparation, coll. & det. R. G. Hughes;
1977.5.7.1 (Fig. 14). Mewstone Ground, near Plymouth, Devon, Oct 1899, infertile colony
on Laomedea flexuosa, 1 microslide preparation, coll. Marine Biological Association of
U.K., ex E. T. Browne colln; 1961.11.14.16 (see Remarks). R. Ranee, nr St Malo, NW
France, infertile colony on Hydrallmania falcata (Linnaeus, 1758), spirit + 2 microslide
preparations, coll. M. Philbert; 1935.7.10.1 (?mentioned, Philbert, 19356). Sta. SCD258 W,
Univ. Cape Town Ecol. Survey, 14 Jul 1961, infertile colonies on Obelia dichotoma, spirit +
1 microslide preparation, pres. N. A. H. Millard; 1962.6.18.9.
DESCRIPTION OF HYDROID STAGE. Colony stoloniferous, comprising a tortuous, branched,
mostly unringed stolon from which usually unbranched pedicels arise at intervals. Pedicels
long, occasionally branching as in C. hemisphaerica with similar upward-curved bases
bringing branches approximately parallel with main pedicel; annulated basally, below
hydrotheca and sometimes centrally, forming 1-2 smooth central portions. Hydrotheca
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
89
Fig. 14 Clytia paulensis. (a) single hydrotheca on pedicel (right) adjacent to same of C.
hemisphaerica (left), SW England; 1977.5.7.1. (b) part of (a), right hand specimen, enlarged, (c)
gonotheca (after Millard, 1975 : fig. 73D). Scales: (a, c) 500 /mi; (b) 100/zm.
rather longer than in C. hemisphaerica, length : breadth ratio usually 3-4, exceptionally 1^;
7-1 1 rounded bimucronate cusps; flared appearance immediately below rim resulting from
outward bulging of larger embayments; diaphragm oblique; longitudinal folds in hydrothecal
wall in microslide preparations can look like striations (Millard, 1975) but are artefacts.
Hydranths in present material with 16-22 tentacles. Gonotheca (not seen) ?d" = 9, cylindrical,
tapering gradually below and slightly above, smooth (after Stechow, 19230; Philbert, 19356;
Millard, 1975); borne on stolon on short annulated pedicel. 1-3 medusa buds per blastostyle,
pre-release medusa with 4 tentacle buds (Millard, 1975); free medusa not yet described.
Variation. Cusps on hydrothecal rim variable in both length and breadth; always rounded in
present material. Embayments between cusps irregular in depth so that adjacent ones are
sometimes similar, obscuring bimucronate condition. Hydrotheca length : breadth ratio
usually 3^t but Millard (1966) gave 1|-3|, once 1$ (BMNH 1935.7.10.1). Angle of slope of
diaphragm variable.
MEASUREMENTS. See Table 3.
DISPERSIVE STAGE. A medusa. Advanced embryos still in the gonotheca have been widely
reported as having 4 tentacle buds, as they have in other Clytia species, but the free medusa
has yet to be described. See also Dispersive Stage under Clytia linearis.
REPRODUCTIVE SEASON. No information.
DISTRIBUTION. A widespread, warm water species recorded in the N Atlantic north to
Suffolk, SE England. However, C. paulensis has been found at only three English localities
(present material). Fertile material was reported as 'very common' near St Malo, NW France
(Philbert, 19356) but all the English material has been infertile.
90 P. F. S. CORNELIUS
Table 3 Clytia paulensis Measurements in //m.
NW France NW France South Africa
(1937.7.10.1) (Philbert, 19356) (Millard, 1966,
1975)
S Indian Ocean
(holotype; Vanhoften,
1910)
Hydrotheca
Length
220-350
300-720
360^00
Breadth (max)
130-150
130-330
200
Pedicel length
550-900
500-1800
Gonotheca (?rf = 9)
Length
Breadth (max)
Aperture diameter
Pedicel length
900-950
330^10
290
50-90
Other European records are 'the Mediterranean' (Picard, 19586), Naples (Riedl, 1959) and
the Glenan Isles, NW France (Fey, 1969); but not the Roscoff area (Teissier, 1965; L.
Cabioch, pers. comm.).
HABITAT. Lower shore (present material) to 200m (Stechow, 19230; Riedl, 1959;
Mediterranean); 0-138 m, once 384 m (Millard, 1975, 1977; S Africa). Tolerant of reduced
salinity at least as low as 17%o (Calder, 1976). Usually recorded epizoic on hydroids and
other inert animal substrates. The following have been reported: Laomedea sp., Clytia
hemisphaerica (as C. gracilis), Dynamena sp., Halecium beanii Johnston, 1838, Nemertesia
antennina (Linnaeus, 1758), Pennaria disticha sensu Brinckmann-Voss, 1970, Sertu-
larella sp. and spines of an echinoid, Cidaris sp. (as Dorocidaris sp.) (all by Stechow,
19230); Sertularia cupressina Linnaeus, 1758, and Halecium beanii (by Philbert, 19356);
Hydrallmania falcata (Linnaeus, 1758), Obelia dichotoma and a bryozoan, Crisia sp.
(present material); and Nemertesia sp., Tubularia sp. and Scrupocellaria scruposa (Bryozoa)
in Suffolk and Essex, SE England (by R. G. Hughes, pers. comm.).
REMARKS. The four tentacle buds of the pre-release medusa and the close similarity to C.
hemisphaerica indicate that paulensis is correctly referred to Clytia. C. paulensis was placed
in Obelia by Naumov & Stepanyants (1972) because the material they saw was 'strongly
branched' but this seems slim reason. Stepanyants (1979) later referred C. paulensis to
'Obelia bicuspidatd* Clarke, 1875, a conclusion with which I cannot agree (see notes under
O. bidentata, p. 1 1 7).
The earlier Torbay material listed above, collected in 1973, was the first to be recorded
from the British Isles. E. T. Browne's Plymouth material had been collected in 1899 but was
overlooked. Browne first identified it as Campanularia raridentata Alder, here referred to
Clytia hemisphaerica; but in 1927 added a note doubting his determination (E. T. Browne
ms notebooks 13 : 148-149; Zoology Library, BMNH). The material came to the BMNH as
part of the E. T. Browne bequest and was re-identified by the then curator as 'Clytia gracilis
Sars'. The hydrothecal rims, with their rounded bimucronate cusps, clearly differ from those
of C. gracilis s. str. auct. and the specimen resembles C. paulensis in all respects. Although
Browne regarded the material as something unusual in 1899 C. paulensis was not described
until 1910, and he may well have been the first in the World to collect this species.
The type material of Clytia ulvae Stechow, 19190, from Marseille, has a bimucronate
hydrothecal rim and is undoubtedly C. paulensis. Nevertheless, Stechow was among the first
to collect C. paulensis from Europe and to recognize it as different from C. hemisphaerica.
Clytia paulensis is perhaps difficult to identify. It differs from C. hemisphaerica in its
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 91
bimucronate hydrothecal rim, its slender hydrotheca and pedicel and, so far as is known, its
consistently smooth gonotheca. Both species are widespread in the World. But in European
waters C. paulensis has yet to be found north of Suffolk, SE England, whereas C.
hemisphaerica ranges much further north. Other west European Campanulariidae having
bimucronate hydrothecal rims are Obelia bidentata and Laomedea neglecta which produce
taller colonies and branch differently.
Clytia incertae sedis
Two Clytia medusae, with 5 and 6 radial canals and diameters of 8 mm and 1 3 mm respec-
tively, were taken in the southern Adriatic Trough on 9 August, 1969. They were referred to
Phialidium pentata Mayer, 1900, by Schmidt & Benovic (1977), making the first NE
Atlantic record of that species. Kramp (1961) had referred "P. pentata' to Phialidium
folleatum McCrady, 1857 (= Clytia folleata) but there remains a possibility that both are
abnormal variants of C. hemisphaerica. Indeed, Schmidt & Benovic considered: 'It might be
possible that all [recorded] specimens [of pentata, folleata and also P. gardineri Browne,
1904] are abnormal forms of C. hemisphaerica'. I concur with their conclusion that more
material is needed to resolve these problems.
Leloup (1940 : 21, as Laomedea) recorded Campanularia kincaidi Nutting, 1899, from
the Azores, at 1 187 m depth, without description or comment. The record was repeated by
Rees & White (1966:277, as Obelia). The only previous record from the Atlantic
was of two colonies from the Caribbean, also by Leloup (1935 : 20). In the absence of more
definite indication it seems best to omit the species from the present survey. The nominal
species was provisionally referred to Clytia by Cornelius (19750 : 280).
Subfamily OBELIINAE Haeckel, 1879
Obelidae Haeckel, 1879 : 163 (part).
Obelinae: Mayer, 1910 : 231 (part); Russell, 1953 : 296.
NOMENCLATURE. The root of the subfamily name is Obelia, and the spelling Obelinae is
incorrect.
DIAGNOSIS. Campanulariidae with erect hydrocaulus and true hydrothecal diaphragm; no
sub-hydrothecal spherule; stolon not anastomosing; medusa liberated but reduced (Obelia),
or vestigial and retained (the rest).
TYPE GENUS. Obelia Peron & Lesueur, 1 8 100 (by present designation).
SCOPE. The genera Gonothyraea Allman, 18640, Hartlaubella Poche, 1914, Laomedea
Lamouroux, 1812 and Obelia Peron & Lesueur, 18100.
REMARKS. The subfamily name is the oldest available. The Obeliinae was recognized also by
Mayer (1910) and Russell (1953), who like Haeckel based their classifications on the medusa
stage alone. All the included genera occur in the eastern North Atlantic and are defined
below.
Genus GONOTHYRAEA Allman, 18640
Gonothyraea Allman, 1864a : 374.
Gonothyrea auct. (laspus pro Gonothyraea).
Campanularia, Laomedea and Obelia part, auct. (see Remarks).
TYPE SPECIES. Laomedea loveni Allman, 1 8590; designated by Millard (1975). The originally
included species were L. loveni, Campanularia geniculata sensu Lister (= G. loveni; see
Cornelius, 19770 : 47) and L. gracilis Sars. The last named was based on a mixed series
comprising what was probably Clytia hemisphaerica (hydroid) and G. loveni, but following
designation of lectotype material (p. 94) it is now subjectively referred solely to C.
hemisphaerica.
92
P. F. S. CORNELIUS
DIAGNOSIS. Campanulariidae forming upright, branched colonies; stem flexuose; hydro-
thecae tubular, pedicellate, alternate; diaphragm present; no sub-hydrothecal spherule;
gonophore a gonomedusa.
REMARKS. I have discussed elsewhere the past confusion between G. loveni and nominal
species of Campanularia, Laomedea and Obelia (in Cornelius, \911a). This confusion
probably delayed by some decades the discovery of the alternation of generations in
medusoid coelenterates; and, it can be claimed, in other animal groups.
Gonothyraea loveni (Allman, 1 859a)
(Fig. 15)
Laomedea gracilis Sars, 1857: pi. 2, fig. 4 only (not pp. 51-54, nor figs 1-3, 5, = Clytia
hemisphaerica, q.v.).
Laomedea loveni Allman, 1859a : 138-140.
Gonothyraea (Laomedea) loveni: Allman, 18646 : 376.
Gonothyraea hyalina Hincks, 1866 : 297-298; Hincks, 1868 : 184-185, pi. 35, fig. 2.
Gonothyraea loveni: Hincks, 1868 : 181-183, pi. 25, fig. 2.
Obelia loveni: Naumov, 1960 : 264-265, fig. 152 (syn. G. hyalina Hincks); Naumov, 1969 : 285-287,
fig. 1 52 (syn. G. hyalina Hincks).
NOMENCLATURE. Bedot (1912 : 294; 1916: 107) listed but two uses of the combination
Obelia hyalina, and none of O. loveni, in his synoptic works (1901-1925). Obelia hyalina
Clarke, 1879, and O. hyaliana Vannucci, 1955, are different nominal species.
16d
Figs 15-16 Fig. 15 Gonothyraea loveni. (a) tip of colony, Bay of Biscay; 1959.9.17.59. (b) same,
hydrotheca. (c) 9 gonotheca and gonomedusa, Vadso, E Finmark, Norway, intertidal;
1912.12.21.184. Scales (a-c) 500 urn. Fig. 16 Hartlaubella gelatinosa. (a) part of 9 colony. Note
large embryos. SW England; 1959.9.17.57 (microslide preparation), (b) same, two blastomeres.
Note conspicious chromosomes (see Remarks), (c) hydrothecal rims. Israel; 1932.8.13.1. (d)
unopened rf gonotheca, NE England; 1 969. 1 1 .28.2. Scales: (a, d) 500 //m; (b-c) 50 /^m.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 93
TYPE LOCALITY AND MATERIAL. Firth of Forth, Scotland (Allman, \S59a : 137); material not
located.
TYPE MATERIAL OF OTHER SPECIES EXAMINED. Gonothymea hyalina Hincks, 1866, syntypes,
Shetland Is, Scotland, several colonies in spirit, coll. J. G. Jeffreys, on 'Tubularia,
Halecium,etc.\ex Hincks colln; 1899.5.1.157.
OTHER MATERIAL EXAMINED. BMNH collection, about 80 specimens.
DESCRIPTION. Colony erect, stem monosiphonic, delicate, variably flexuose, internodes
usually slightly curved, irregularly branched, up to c. 100mm. Annuli above origins of
branches. Hydrothecal pedicels usually annulated throughout but smooth central portion
frequent; usually tapering distally to roughly half proximal diameter. Hydrotheca
cylindrical, campanulate, length 1^-3 times maximum breadth; rim delicate, slightly out-
turned, castellate, raised portions often notched (Fig. 15); often longitudinal folds in
hydrotheca. Hydranth with 20-25 tentacles; branched tentacles recorded but rare
(Hamond, 1957); hypostome spherical. Gonotheca (rf narrower than 9, otherwise similar;
Miller, 1973) cylindrical, truncated above, tapering below. Conspicuous sporosacs or ova
develop internally and later extrude up to four together as tentaculate, sub-spherical reduced
medusae termed gonomedusae (formerly called meconidia; see Dispersive stage).
Variation. Internode length and curvature, length of hydrotheca and the degree of tanning
and translucency of the perisarc are all variable.
DISPERSIVE STAGE. A planula larva. The reduced medusa is retained until after the planulae
have escaped. It does not swim but simply drops off. Ellis (17560, b, c, 1767, but not 1755;
see Cornelius, \911a) recorded that the 'released' gonomedusae exhibited strong tentacle
movements and adopted a 'worm-like' shape, but his observations seem not to have been
repeated.
The gonomedusae were interpreted as highly developed sporosacs by Allman (1 859a) who
coined for them the long standing term meconidia. But Goette (1907) and more recently
Miller (1973) have shown that they are actually reduced medusae. Miller introduced the
preferable term gonomedusae. Wulfert (1902) provided a summary of early reproductive
studies on this species.
REPRODUCTIVE SEASON. Published records suggest that in the English Channel and southern
North Sea the species breeds almost throughout the year (Mar-Sep, Hamond, 1957; Jan-Apr
& Sep-Nov, Marine Biological Association, 1957; May-Aug & Oct-Nov, Teissier, 1965);
but an intertidal population which I studied in 1974 in Sussex, SE England, bore
gonomedusae only during the first two weeks of April. Some of the published records might
refer to colonies with developing gonothecae, or with empty ones.
Fertile gonothecae were reported at Naples from January to May and in September by Lo
Bianco (1909).
DISTRIBUTION. Widespread in suitable habitats and often common. Occurs throughout
western Europe north to W Greenland (but not E), Iceland, Faeroes, Spitzbergen, Barents
Sea, Murman coast and White Sea (Mathiesen, 1928; Kramp, 1929, 1938; Calder, 1970). In
the Baltic G. loveni is said to penetrate as far as Helsinki in the Gulf of Finland and the Aland
Isles in the Gulf of Bothnia (Linko, 1911; Stechow, 1927; Naumov, 1969). It has been
recorded from the Mediterranean Sea (Picard, 19586; Riedl, 1959) and Black Sea (Naumov,
1969); and on the Atlantic coast from W France and Morocco (Billard, 1927; Patriti, 1970).
Millard (1975) recorded the species in South Africa only from Cape Town docks and
considered this and other Southern Hemisphere records to result from transport by ships.
Hence G. loveni might be found further south than Morocco. Rees & White (1966) listed a
dubious old Azores record from the unusual depth of 845 m.
HABITAT. Intertidal, usually in pools, and offshore. Naumov (1960, 1969) gave a normal
depth range of 0-30 m, with an extreme lower limit of 200 m; and Mathiesen (1928)
94 P. F. S. CORNELIUS
similarly gave 6-200 m. The BMNH material falls within these limits. The record at 845 m
off the Azores listed by Rees & White (1966) is much deeper, and needs confirmation. G.
loveni is tolerant of reduced salinity at least to 12%o (Calder, 1976). It has been recorded on a
variety of animal, plant and inorganic substrates (Hincks, 1868; Hamond, 1957; Marine
Biological Association, 1957; Teissier, 1965), and there is no regular association.
REMARKS. Hummelinck (1936), Naumov (1960, 1969) and others listed by Calder (1970)
regarded Gonothyraea loveni and G. hyalina as conspecific and I concur.
The distinctive method of reproduction has been described in part by many authors from
Ellis onwards. Nevertheless G. loveni was widely confused with Obelia spp. and with
Laomedea Jlexuosa until Wright (1858, 1859) realized it was distinct (Cornelius, 19770;
Cornelius & Garfath, 1980). But Wright did not provide a name. The species was soon
named by Allman (18590) who took Joshua Alder's suggestion (in litt.) that the species was
valid. Alder was perhaps unaware of Wright's publications. From the literature it would
appear that Wright and Alder worked independently; but since both communicated freely
with Hincks (e.g. 1 868 : preface) they could have been in touch directly. It may be difficult to
find out who among these British workers really was first to recognize G. loveni.
Most of the life cycle is well understood and has been redescribed in detail by Miller (1973;
see also Bergh, 1879, Wulfert, 1902, and Goette, 1907). Aspects of stolon growth were
described by Wyttenbach, Crowell & Suddith (1973).
Laomedea gracilis Sars, 1850, 1857, was based on a mixed type series and this has caused
confusion. The material illustrated in 1857 was partly G. loveni (see synonmy) but
predominantly Clytia hemisphaerica (see Stechow, 19230 : 111). The material resembling C.
hemisphaerica in this series is here designated lectotype. Hence the species name loveni
Allman, 18590, remains available. In any case, the name gracilis is preoccupied. This and
other problems relating to "L. gracilis'' are considered under C. hemisphaerica, where a new
name is introduced in place of L. gracilis (p. 78).
Obelia hyalina Clarke, 1879, has been referred to Obelia dichotoma (by Cornelius,
19750 : 266) and is not G. hyalina Hincks. Hence Billard's (19310) record of 'Obelia hyalina
Clarke' from Mauritania refers to O. dichotoma and not G. loveni. This homonymy is
discussed further under O. dichotoma (p. 1 19).
The affinities of G. loveni are discussed above (pp. 47-49).
Genus HARTLAUBELLA Poche, 1914
Sertularia, Campanularia, Obelia and Laomedea auct., part.
Obelaria Haeckel, 1879 : 172 (part).
Obelaria Hartlaub, 1897 : 488 (homonym).
Hartlaubella Poche, 1914 : 76.
TYPE SPECIES. Sertularia gelatinosa Pallas, 1 766; by monotypy.
DIAGNOSIS. Erect, colonial Campanulariidae with polysiphonic stems and second order
branching; hydrotheca pedicellate, with diaphragm and castellated rim, without spehrule; no
medusa stage (see Remarks under H. gelatinosa).
REMARKS. The genus name Obelaria Haeckel, 1879, was proposed as a nom. gen. nov. for
the hydroid stage of Obelia, of which name it is a junior synonym (Cornelius, 19750 : 254).
Haeckel included Sertularia gelatinosa Pallas, 1 766, in its scope. Confusion was caused later
when Hartlaub (1897) independently coined the generic name Obelaria to apply to a gen.
nov. comprising Sertularia gelatinosa alone. Although Hartlaub's name is a homonym of
Haeckel's and cannot be used, Hartlaub's generic concept is accepted here. The next avail-
able name for the genus is Hartlaubella Poche, 1914, of which gelatinosa has always been
the only member.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 95
Hartlaubella gelatinosa (Pallas, 1 766)
(Fig. 16)
Sertularia gelatinosa Pallas, 1 766 : 1 16-1 1 7.
Campanulariaflemingii Deshayes & Edwards, in Lamarck, 1836 : 133-134.
Laomedea gelatinosa: Couch, 1884 : 4-5, 39-40; Hincks, 1852 : 85-86; Da Cunha, 1944 : 65-66;
Vervoort, \946a : 300-303, fig. 133 (syn. Campanulariaflemingii Deshayes & Edwards).
Obelia gelatinosa: Hincks, 1868 : 151-154, pi. 26, fig. 1; Naumov, 1960 : 268-269, figs 157-158;
Naumov, 1969 : 290-29 1 , figs 1 57-1 58.
Obelaria gelatinosa: Haeckel, 1879 : 172, 173, 176; Hartlaub, 1897 : 488-495 (non Haeckel); Nutting,
1915: 88-90, pi. 24, figs 1-5.
Hartlaubella gelatinosa: Poche, 1914:76; Stechow, 1925:522; Stechow, 1927:309; Teissier,
1965: 17;Calder, 1970: 1 543; Cornelius &Garfath, 1980 : 283.
Campanularia gelatinosa: Ralph, 1957 : 820, fig. Ib-f.
TYPE LOCALITY AND MATERIAL. Belgian coast (Pallas, 1766; Ralph, 1957); specimen not
located.
MATERIAL EXAMINED. BMNH collection, about 50 specimens.
DESCRIPTION. Colony elongate; small colonies loosely conical, large colonies bushy, up to c.
200 mm; main stems straight or branched, polysiphonic. Final branches monosiphonic,
arranged irregularly all round stem, ± dichotomous, flexuose; internodes usually curved but
sometimes straight, length varied, 5-9 rings basally. Hydrothecae on ringed, slightly tapering
pedicels, long-campanulate to cylindrical; rim often abraded even but initially castellate
with notch of varied depth in centre of each blunt cusp; embayments rounded; diaphragm
transverse. Hydranth with 22-28 tentacles, hypostome spherical. Propagation by stolons in
spring. Gonotheca ^ = 9, axillary, inverted-conical, sides smooth to sinuous; aperture wide,
distal, on short collar; ova (4^6) and embryos larger than usual in family; nuclei of
blastomeres large, with unusually conspicuous chromosomes.
Variation.lntQrnode length and curvature, angle of flexure of stem and length : breadth ratio
of hydrotheca are all variable. The hydrothecal rim often abrades smooth.
DISPERSIVE STAGE. Planulae, developing within the gonotheca. Some authors have
mistakenly reported a medusa stage.
REPRODUCTIVE SEASON. May-August in NW France (Teissier, 1965); July, 1934,
Northumbria (H. O. Bull, in Evans, 1978).
DISTRIBUTION. Recorded from southern Scotland, Oslo Fjord, Danish waters and part of the
Baltic south to the Mediterranean and Black Seas. The species is common in Dutch, Belgian,
Irish, Welsh, English and N & W French waters (Vervoort, 19460; Leloup, 1952; BMNH
collection; Billard, 1927; Teissier, 1965).
Trustworthy Scottish records are few and there seems only one this century, although the
species is still common in NW England: Tay Estuary (Fleming, 1820; Alexander, 1932);
Shetlands, Berwick Bay and Solway Firth (Johnston, 1847); ?Dundee (BMNH
1851.7.25.227, specimen not located). Hincks (1868) repeated several of these records and
Norman (1869) gave another Shetlands locality. Recent English records are numerous,
northerly ones including Northumberland (Alexander, 1932) and Morecambe Bay (J. Clare,
pers. comm. & BMNH 1970.8.28.6-10).
The species was recorded from 13 localities in Oslo Fjord by Christiansen (1972) but his
statement that the species occurs north to Finmark is questionable, and Linko (1911) and
Broch (1918) listed no records so far north. Neither also did Kramp (1929, 1938), who
likewise considered the species absent from Iceland and the Faeroes. Naumov (1969) gave an
Iceland record without further detail; but with greater precision stated the northern limit in
the Baltic to be Tort Liepaja = Libava', Latvia. Kramp (1935) listed several Danish records
north to Frederikshavn, NE Denmark, and repeated Stechow's (1927) record from Trave-
miinde, near the southern limit of the Baltic Sea. Rasmussen (1973) reported the species
from Siaelland Island, southern Denmark.
96 P. F. S. CORNELIUS
Mediterranean records are few. Naumov (1969) gave 'Black Sea, Mediterranean and
Gibraltar'. Although Picard (19586:197) expressly excluded the species from the
Mediterranean list Rossi (1950 : 205) had earlier recorded it from NW Italy, the Adriatic and
Strait of Gibraltar. Linko (1911) listed Black Sea material.
The species is known from Portugal (Nobre, 1931; Da Cunha, 1944) but has yet to be
recorded so far south as Morocco (Patriti, 1 970).
HABITAT. Intertidal, particularly in pools and run-off from saltings, and shallow depths off-
shore. Often in estuaries and tolerant of reduced salinity at least to 6-2%o(Vervoort, 1946a).
Also tolerant of silt.
The lower depth limit is probably c. 15 m. A record from 30 m off the Scilly Isles was
based on Obelia bidentata material (BMNH 1969.9.9.6; mentioned, Robins, 1969). Couch
(1844) recorded material from beneath intertidal stones and on algae, but that under stones
might have been Laomedea neglecta (cf. p. 107).
REMARKS. The nomenclatural history of this species is involved. The name Sertularia
gelatinosa was first applied by Pallas (1 766) to the 'Corallina confervoides, gelatinosa alba,
geniculis crassiusculis, pellucidis' of Ray (1724 : 34, para. 7). However, Ellis (1755 : 20,
p. 1 1 , figs B, b) assigned bryozoan material to Ray's species, providing clear illustrations.
Linnaeus (1758 : 812) included the descriptions of both Ray and Ellis in a single species to
which he gave the new name Sertularia spinosa. This name is currently applied to a
bryozoan species in the combination Vesicularia spinosa (Linnaeus, 1758), for example by
Prenant & Bobin (1956). The bryozoan name Sertularia sericea Pallas, 1776 : 1 14, was a
nom. nov. for S. spinsoa and is its junior objective synonym. Pallas elsewhere
(1 766 : 1 16-1 1 7) adequately described gelatinosa.
Fleming (1820) noted that Ray and Ellis each described a different species but referred
Ray's description to Obelia geniculata, not to the present species. However, Pallas' account
is clear and he contrasted gelatinosa with both O. geniculata and O. dichotoma. Also he
noted the polysiphonic stem and cusped hydrothecal rim — both unusual in Obelia. Even so,
it could be argued that Pallas had material of Obelia bidentata before him since that species
superficially resembles H. gelatinosa. There is some evidence that O. bidentata did not then
occur in European waters. Even if it did, Pallas might have overlooked the fine cusps on the
hydrothecal rim which are a main distinction. But despite these small doubts it seems highly
likely that Pallas' description indeed refers to the present species. Hincks (1868: 152)
himself commented that Pallas' description was 'admirable, and is the only one we possess
which is not positively incorrect'; so the case is strong. Hincks summarized some of the
additional taxonomic confusion surrounding the species between 1 820 and 1 868.
Campanularia flemingii Deshayes & Edwards, in Lamarck, 1836, was based on material
of the present species described by Fleming (1820) from Scotland — although Deshayes &
Edwards gave the type locality as coasts of England! Fleming had referred his material to
gelatinosa but noted that it disagreed with Pallas' description in having even hydrothecal
rims. Fleming thought Pallas might have mistaken tentacle tips for castellations on the
rim, and assumed that gelatinosa always had an even rim. Deshayes & Edwards thought
Pallas too careful to make this mistake and concluded that two species were involved, one
with castellations and one without. Probably Fleming's material simply had hydrothecae
in which the rims were worn smooth! Johnston (1838, 1847) realized the confusion and
referred flemingii back to gelatinosa. He was followed by Bedot (1905) and Vervoort
(1946a), and I concur. Gray (1848), however, gave C. flemingii specific rank, but did not cite
material. Gray seems usually to have relied heavily on Johnston's work but on this occasion
clearly did not. Possibly Edwards, who sometimes worked on the British Museum collec-
tions, persuaded him to accept the species.
Thaumantias leucostyla Will, 1844 : 73, pi. 2, figs 16-17, based on an Obelia medusa, was
referred to the present species by Bedot (1912 : 328) without comment; but as gelatinosa has
no medusa stage this must be wrong (see next paragraph). It would be difficult to identify the
medusa beyond Obelia sp. from Will's description.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 97
Several authors, including Hincks (1852, 1868), have stated that H. gelatinosa releases a
medusa, which it does not (Cornelius, \915a : 279). It could be that Hincks and the others
saw medusa release from Obelia bidentata. But the first European records of that species date
from the early 1900s; and it seems improbable that Hincks would have made such a mistake
when identifying a hydroid. His comment that the branches of gelatinosa 'droop slightly' fits
bidentata, but this is not conclusive. Another possible explanation of his apparent mistake is
that he took the unusually large ova of//, gelatinosa for developing medusae.
Couch (1844) had much earlier given a correct (and posthumous) description of the life
cycle, reporting planula development and release, and early development of the young
colony. In addition he noted that the planulae were propelled by cilia. But his contempor-
aries were still muddled, and had Couch lived a little longer he might have corrected some of
the ensuing confusion. Van Beneden (1843, 1844) reported medusa release in H. gelatinosa
but his material was actually Obelia dichotoma (see Cornelius, 19750; and 19770 for other
references). Hincks (1852) also attributed a medusa to gelatinosa. Van Beneden's error of
identification was later appreciated by Hincks, who referred Van Beneden's material to
a synonym of O. dichotoma, namely O. longissima (Pallas, 1766). Similarly, the much later
report by Godeaux (1941) that gelatinosa had a medusa was also based on O. dichotoma
material. It might be relevant that Godeaux worked at the Van Beneden Institute! Then
Leloup (1947), paralleling Hincks, referred Godeaux' material to O. longissima.
A convenient distinction between O. bidentata and H. gelatinosa is that in side view the
branches of bidentata show a graceful sigmoid curvature lacking in gelatinosa.
The large chromosomes illustrated here recall in shape and size those of Obelia medusae,
shown by Faulkner ( 1 929).
Genus LAOMEDEA Lamouroux, 1812
Laomedea Lamouroux, 1812 : 184.
Campanularia Lamarck, 1816 : 112 (part).
Lomedea Pickering, in Dana, 1846 : 689 (lapsus pro Laomedea, see p. 78).
Campalaria Hartlaub, 1897 : 449.
Eucampanularia Broch, 1910: 1 84 (part; see p. 52).
Eulaomedea Broch, 1910 : 189; Millard, 1975 : 223.
Cmpanularia Mulder & Trebilcock, 1914: 11 (part; lapsus pro Campanularia).
Laomedea (Paralaomedea) Hummelinck, 1936 : 57; Vervoort, 19460 : 285.
Eulaomeda Rees & Thursfield, 1965 : 102 (lapsus pro Eulaomedea).
TYPE SPECIES. Laomedea flexuosa Alder, 1857 (proposed designation by Cornelius, 1981).
Broch (1905 : 10) proposed 'Laomedea loveni Allman, 18590' as type species but loveni was
not among the species originally included in the genus and is not eligible.
DIAGNOSIS. Colonial Campanulariidae with: polyp generation forming upright colonies;
stolon branching but not anastomosing; hydrotheca pedicellate, lacking spherule; true
diaphragm present; annular perisarc thickening inside base of hydrotheca; gonotheca
stolonal or axillary, aperture typically circular, wide; gonophores sessile, interpreted as
vestigial medusae in many species.
REMARKS. I have previously pointed out (Cornelius, 19750) that Laomedea Lamouroux,
1812, is a junior synonym of Obelia Peron & Lesueur, 18100. But the name Laomedea is so
well known that I have submitted a case to the International Commission on Zoological
Nomenclature recommending that it be conserved by application of the Plenary Powers. I
have proposed that Laomedea flexuosa Alder, 1857, be designated type species although it
was not originally included. The alternative, of applying the Rules of Nomenclature, would
result in the virtually unused name Campalaria Hartlaub, 1897, being employed for the
present genus. Further details of the case, and another concerning the genus name
Campanularia (p. 5 1 ), have been presented elsewhere (Cornelius, 1981).
The subgenus name Eulaomedea Broch, 1910, type species Laomedea flexuosa Alder,
98 P. F. S. CORNELIUS
1857, by monotypy, is a junior synonym. Stechow (1923a : 95) referred Eulaomedea to
Laomedea Lamouroux, 1812, and apart from subgeneric use by Splettstosser (1924),
Hummelinck (1936) and Vervoort (1959) there seems to have been no further appearance of
Eulaomedea in the literature until Rees & Thursfield (1965). These authors upgraded
Eulaomedea to genus status, including within its scope "E. angulata\ 'E. flexuosa"1 and '£.
calceolifera\ Finally Millard (1975) employed Eulaomedea to embrace flexuosa and
calceolifera alone. Thus the name has not been widely used and Broch (e.g. 1918) himself
came to drop it, without comment. For further details see Cornelius (1981).
The subgenus Paralaomedea was apparently introduced by Hummelinck (1936). The type
species is Laomedea neglecta (Alder, 18566; by monotypy). Vervoort (19460) followed
Hummelinck's usage. But the unusual acrocyst of neglecta was shown by Splettstosser
(1924) to be medusoid in origin, so there seems inadequate reason for subgeneric separation.
Laomedea angulata Hincks, 18616
(Fig. 17)
Laomedea angulata Hincks, 18616 : 261, pi. 8; Hummelinck, 1936 : 51-52, fig. 5; Picard, 19586 : 191
(syn. L. sphaeroidea Stechow); (non Da Cunha, 1944 : 63, fig. 36; = Laomedea calceolifera).
Campanularia angulata: Hincks, 1868: 170-171, pi. 34, fig. 1, woodcut 14 (p. 136); Fraipont,
1880 : 433-466, pis 32-34; Billard, 19046 : 46, 47, 53, 55, 57, 65, 67, 72-82, 97, 144, 173, pi. 3, figs
1-7, pi. 5, figs 1-2, 7, 10; Faure, 1965 : 419-426, figs Ib, d, 2a, b, 3a, b.
Laomedea sphaeroidea Stechow, 1932 : 85-86.
Eulaomedea angulata: Rees & Thursfield, 1965 : 101-102.
TYPE LOCALITY AND MATERIAL. Hincks (18616) based the original description on specimens
from South Devon and the Isle of Man. Some of this material is preserved as follows: (i)
Hancock Museum, Newcastle upon Tyne, infertile colony on single blade of Zostera L. (eel
grass), in spirit, labelled 'Laomedea angulata, Ramsey, Isle of Man. Revd T. Hincks' and, on
a second label, 'Campanularia angulata Hincks' (mentioned, Cornelius & Garfath, 1980);
(ii) BMNH, several colonies on blades of Zostera, in spirit, in two tubes. One contains a
single, wide blade of Zostera supporting an infertile colony of C. angulata; the other tube
five Zostera blades, all much narrower than that in the first, each supporting one or more
fertile colonies of C. angulata. The two tubes, in one jar, jointly bear the registered number
1899.5.1.149. In the bottom of the jar, detached from the tubes, were two faded labels written
by Hincks: 'Campanularia angulata Hincks, Isle of Man', and ' Campanularia angulata
Hincks, Britain'. There seems no indication which label originally belonged to which tube.
Although the narrow leaved Zostera resembles that in the Hancock Museum specimen,
labelled Isle of Man, there is no further indication that the BMNH thin leaved specimen
came from there too.
Probably all this material was identified by Hincks at some time, but whether he saw it
before or after the date of publication of the first description is not clear. Hence, it is not
possible to decide whether the specimens should be treated as syntypes or neotypes.
TYPE MATERIAL OF OTHER SPECIES EXAMINED. Laomedea sphaeroidea Stechow, 1932, fertile
colonies taken from Zostera, Sete, S France, microslide preparations; Munich Zoological
Museum.
OTHER MATERIAL EXAMINED. BMNH collection, c. 30 specimens from Britain and the
Channel Islands and 13 microslide preparations from the Lagoon of Venice. Amsterdam
Zoological Museum and Leiden Natural History Museum, Netherlands, c. 20 specimens.
DESCRIPTION. Mature colony comprising several erect, monosiphonic stems inserted at
approximately regular intervals on an almost straight, little branched or unbranched, smooth
hydrorhiza; recorded reliably only on eel grasses; stolons usually (Fraipont, 1880; Billard,
19046) growing parallel with the leaf margins. Stems flexuose, internodes markedly straight
(rarely, faintly curved), angle between them 90°-120°, length : breadth ratio variable; \-c. 10
annuli basally in BMNH material. Late in season stems terminate in long, often curling
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
99
Fig. 17 Laomedea angulata. (a) part of colony, including hydrocaulus and stolonal 9 gonotheca
with embryos at different stages of development, Jersey; 1959.9.17.11. (b-c) cf gonothecae,
Plymouth, SW England; 1959.9.17.3, 12. (d) vegetative terminal stolon; as (c). Scale (a-d)
500 /on.
tendrils approximately same width as internodes except at often recurved tip region which is
wider. Tendrils probably represent modified hydranths: those ofObelia dichotoma differ (see
Remarks). Hydrothecae campanulate, delicate, usually slightly flared distally, length c. \{
times maximum breadth, thin walled and not thickened; diaphragm usually transverse but
occasionally slightly oblique; pedicels 3-1 5 ringed, slightly tapering distally, sometimes with
smooth central portion (e.g. BMNH 1959.9.17.11). Hydranth with 24-30 long tentacles
alternately elevated and depressed; hypostome large, spherical when dead but (Fraipont,
1880) when alive constantly changing in shape; tissues colourless. Gonothecae apparently
always borne on stolon. 9 elongate-ovate, aperture distal, wide, on slightly demarcated neck,
usually with sub-terminal internal strengthening ring; on short, 3-6 ringed pedicel; contain-
ing several eggs (see Remarks); planulae brooded internally. <5 similar, aperture narrower;
100 P. F. S. CORNELIUS
containing several cf gonophores; sometimes said to be more tapered distally than 9, but this
not evident in BMNH series. No medusa stage.
Variation. Apparently minor. Broch (1933) regarded L. calceolifera a variety of the present
species, proposing a forma typica for L. angulata s. str., but his opinion is no longer followed.
DISPERSIVE STAGE. Planulae, which develop within the gonotheca (e.g. BMNH
1 959.9. 17.11). Also vegetative tendrils which break away to form new colonies.
REPRODUCTIVE SEASON. Sexual reproduction June-August in NW France (Teissier, 1965),
but fertile material recorded early as April in Isle of Man (Bruce et al., 1963, material not
examined). Vegetative reproduction involving tendrils typically August to November in NW
France (Teissier, 1965).
DISTRIBUTION. A southerly species which probably occurs no further north than the British
Isles. Unchecked published records and data with BMNH specimens indicate the following
localities in NW Europe: SW Scotland (Rankin, 1 90 1 ; Ritchie, 1911; Chumley , 1 9 1 8); N & S
Ireland (Hincks, 1868); Isle of Man (part of type series; also Bruce et al., 1963); Scilly Isles,
1967 (Robins, 1969); S Devon, 1898 & 1906 (type series; also Marine Biological Associ-
ation, 1957; a 1936 record on a barnacle seems unlikely); Dorset, BMNH; Roscoff(Fraipont,
1880; Faure, 1965; Teissier, 1965); Channel Islands (Vervoort, 1949; Leiden NHM &
BMNH); Netherlands (Vervoort, 1946#; Leiden NHM). There seems only a single,
unchecked record from the east coast of Britain, from St Andrews Bay (Crawford, 1895,
repeated in Laverack & Blackler, 1974); and the species has not often been recorded from the
many Zostera beds of southern England (see Addendum).
Published records from Spain southwards include the following: Santander, N Spain,
(Rioja y Martin, 1906); NW Italy (Rossi, 1950; also Naples, Riedl, 1959, and lagoon of
Venice, BMNH material); 'Mediterranean' (Picard, 19586).
At Castiglione, Algeria, Picard (1955) found no less than 1 8 species of hydroids growing on
the eel grass Posidonia, but did not report L. angulata. A single colony of the species was
erroneously reported from Portugal by Da Cunha ( 1 944), his illustration showing a mature 9
gonotheca of L. calceolifera. The record from the Falkland Islands by James Ritchie,
repeated by Rees & Thursfield (1965), was based on Obelia dichotoma material.
The northernmost material I have seen came from the Isle of Man (type series; also Bruce
et al., 1963). More northerly material was reported from the Clyde Sea by both Rankin
( 1 90 1 ) and Ritchie ( 1 9 1 1 ), the latter repeating some of Rankin's records and giving some new
ones. However, the depth range given by the two authors, 30-1 30 m, exceeds the depths from
which the species has otherwise been recorded. There are records of one of the substrate
plants, Zostera, from the Clyde Sea area so it is conceivable Rankin and Ritchie at their
deeper localities dredged up loose plants which had sunk; but dead Zostera often floats! Den
Hartog (1970) gave 7 m as the deepest British record of Zostera marina L. (in the Scilly Isles)
but cited Danish and Mediterranean records down to 1 1 m and one from the Pacific coast of
the U.S.A. at 30 m, adding credibility to at least some of the Clyde Sea records. He gave the
NE Atlantic distribution of Z. marina as Algeria (Castiglione), S France (rare) and N
Mediterranean north to the arctic coast of the U.S.S.R., so it can be assumed that the
northern limit of L. angulata is not determined by availability of eel grass.
However, the close association of this species with the eel grasses certainly affects its
distribution. Thus Zostera almost died out in England in the 1930s (Tutin, 1942), and there
is a virtual lack of L. angulata records since the 1900s. See also Addendum.
L. angulata was not recorded in several surveys of North Sea coasts, as follows:
Northumberland and Durham (Robson, 1914), Norfolk (Hamond, 1957; Hamond &
Williams, 1977), Belguim (Leloup, 1952), Denmark (Kramp, 1935; Rasmussen 1973) and W
Sweden (Rees & Rowe, 1969). The record from Danish waters by Vervoort (19460) probably
referred to Broch's (1928) record as Campanularia conferta and should be rejected (W.
Vervoort, pers. comm.). The species has still to be recorded from Denmark (K. W.
Petersen, pers. comm.); but a record from the Great Belt, Kattegat, by Winther (1879,
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 101
repeated in Stechow, 1927) if confirmed would be the most northerly of the species. A record
dated 1960 from 20 m depth off SW Wales, quoted by Crothers (1966), seems dubious since
both depth and area would be unusual. However, Zostera does occur there (K. Hiscock, pers.
comm.).
HABITAT. Intertidal and shallow sublittoral, the deepest reliable record being 6-8 m
(Studland Bay, Dorset, during or before 1 890, coll. R. Kirkpatrick, pres. F. Beckford; BMNH
1 899.7.22. 1 ). Deeper records, from the Clyde Sea and off SW Wales, cannot be substantiated.
Faure (1965) reported that L. angulata grew in a zone on the shore at Roscoff above that
occupied by L. calceolifera; but the numerous records from shallow offshore localities show
that L. angulata is not confined to the intertidal zone. A record from 64 fathoms (130 m)
near the Falkland Is repeated by Rees & Thursfield (1965) was based on Obelia dichotoma
material.
L. angulata has been widely recorded on the eel grasses, Posidonia and Zostera (Hincks,
186 16; Fraipont, 1880; Philbert, 19356; Hummelinck, 1936; Vervoort, 19460; Rossi, 1950;
Riedl, 1959; Faure, 1965) and only seldom on other substances (sertularian hydroids by
Betencourt, 1888; Laminaria by R. Oppenheim, in Vervoort, 1949; Dictyota dichotoma by
Philbert, 19356; Balanus improvisus by W. J. Rees, in Marine Biological Association, 1957;
Fucus by several recorders listed in Teissier, 1965). The overwhelming majority of published
reports, and all the BMNH, Amsterdam Zoological Museum and Leiden Natural History
Museum material are from Zostera, however, and it is likely that records on other substrates
are wrong. Possibly some at least refer to L. calceolifera, since the two species were confused
for several decades. Although Betencourt noted terminal tendrils on his material it might
have been Obelia dichotoma which also has tendrils and can look remarkably similar.
Nishihira (1968) made a detailed study of the hydroids epizoic on Zostera in northern
Japan, but did not report L. angulata which seems (Stechow, 19236) not to occur there.
Picard (1955) made a similar study in Algeria, where he found L. angulata absent from the
Posidonia beds of Castiglione.
The species is tolerant of brackish conditions. Hummelinck (1936) noted a tolerance of
9-8%o Cl, a figure repeated by Vervoort 19460) and, incorrectly as salinity, by Naumov
(1960, 1969). The corresponding salinity figure is 17'7%o. Morri (19790) found L. angulata
in a range of salinities down to 27%oin Tuscany, Italy.
REMARKS. Athough Hincks (1868, 1871) and several subsequent authors distinguished
correctly between the present species and L. calceolifera some later authors (e.g. Babic,
1912; Broch, 1928, 1933; Vervoort, 19460; Naumov, 1960, 1969) regarded them
conspecific. A summary of this confusion and a taxonomic assessment of the two species was
provided by Faure (1965). The main differences are as follows, in approximate order from
most to least useful. The gonothecae differ both in structure and position (hydrorhizal in L.
angulata, on the stem and axillary in L. calceolifera). The internodes in angulata are usually
straight, and curve slightly in most calceolifera specimens. Terminal tendrils are often
present, particularly in autumn, in angulata but are unrecorded in calceolifera (Obelia
dichotoma sometimes has them also). Eel grasses are probably the only substrate for angulata
but calceolifera occurs on a variety of inert, solid substrates. Faure reported angulata growth
at its peak in summer, and that of calceolifera in spring, at the same place. He reported erect
stems spaced at 2-5 mm intervals in angulata and often clustered in calceolifera; but the
extent to which this character in angulata is phenotypic, induced by the elongate shape of
the eel grass leaves, is not known. Finally, Faure reported that colony height was 5-10 mm in
angulata, 13-1 6 mm in calceolifera, each stem bearing respectively at maturity 4-6 and
10-15 hydrothecae. Despite this list of differences it remains difficult to identify some
infertile material, even with the aid of long reference series; and some young colonies are
probably impossible to identify.
A straight hydrorhiza occurs in Obelia geniculata also, in which it can be interpreted as a
genotypic character adapted to keeping neighbouring, parallel-running hydrohizae ad-
equately spaced along their whole lengths. O. geniculata occurs on broad thalloid algae, and
102 P. F.S.CORNELIUS
spacing is probably an advantage. L. angulata is unusual among thecate hydroids in being
closely adapted to its substrate in a number of obvious ways, and may have taken to living
on eel grasses in relatively recent geological time. It seems likely that since eel grasses are
themselves quite recent, being derived from more conventional terrestrial angiospermes,
other campanularian and laomedean hydroids were living on algal substrates long before L.
angulata or its ancestors took to eel grasses. Maybe hydroids of this group, having straight
hydrorhizae, were pre-adapted to growing on the long narrow leaves of eel grasses.
However, at least some orientation of stolon growth occurs in L. angulata in a direction
parallel with the eel grass leaf since diagonal or transverse stolons do not occur. O. geniculata
stolons, although straight, are not orientated along the lamina and it may be that angulata
stolon tips have a geotropic response which keeps them growing vertically (upwards or
downwards) along the leaf, which floats upright when the tide is in. The fact that stolons of
angulata grow sometimes along the narrow edges of the substrate leaves suggests that the
planula does not seek a central position; or that once a tendril attaches and forms a new
colony the hydrorhizal tip cannot locate the centre of the leaf; but there is no experimental
evidence.
The terminal tendrils of L. angulata were found to be most common from August to
November at Roscoff by Faure (1965). Billard (19046) suggested that their function was to
attach to adjacent eel grass leaves and hence enable the species to colonize fresh plants
vegetatively. The tendrils of BMNH specimens differ from those of occasional Obelia
dichotoma colonies in having dilated recurved end regions, the ends of O. dichotoma tendrils
being uniform in width and approximately straight. Study of the BMNH material suggests
that tendrils in L. angulata are modified hydranth/hydrotheca complexes.
The most detailed histological and general biological account of L. angulata is that of
Fraipont (1 880). However, he stated that the female gonophore contains a single egg; but one
of his illustrations (pi. 34, fig. 3) showed a female gonotheca containing several planulae, as
illustrated also here, and more than one egg seems normal.
The nominal species Laomedea sphaeroidea Stechow, 1932, was based on material from
near Sete (Cette), S France, which Stechow (\9\9a) had earlier referred to L. angulata. The
type material is referrable to L. angulata as defined here and it seems unnecessary to
recognize the second species. Picard (19586 : 191) also suggested this synonymy, without
comment.
Laomedea calceolifera (Hincks, 1871)
(Fig. 18)
? ] Laomedea exigua Sars, 1857: 50-5 1 .
Campanularia calceolifera Hincks, 1871 : 78-79, pi. 6; Faure, 1965 : 419^26, figs la, c, 2c, d, 3c, d;
Miller, 1973 : 377-386, figs 6c, d, 10 a-d.
Campalaria conferta Hartlaub, 1897 : 495-496, pi. 19, figs 2-12.
Laomedea angulata: Babic, 1912 : 45 7^60, figs l-5;DaCunha, 1944 : 63, fig. 36.
Laomedea conferta: Splettstosser, 1924 : 403-420, figs X'-Z1, A2-T2; Da Cunha, 1944 : 63-64.
Laomedea calceolifera: Billard, 19316 : 390; Picard, 1955 : 187.
Obelia calceolifera: Picard, 1955 : 187.
Laomedea (Campalaria) conferta: Hamond, 1957 : 3 1 5, fig. 23.
Eulaomedea calceolifera: Rees & Thursfield, 1965 : 102; Millard, 1975 : 223-224, fig. 73g-k.
TYPE LOCALITY AND MATERIAL. Salcombe, Devon, England, 'on stones &c', offshore (Hincks,
1871:73-74, 79), several colonies in spirit and one microslide preparation; BMNH
1899.5.1.155. Some of the spirit material is attached to a worm tube, possibly a sabellariid
(det. J. D. George).
TYPE MATERIAL OF OTHER SPECIES. Prof. Dr M. Dzwillo tells me that there is no material
labelled "Campalaria conferta Hartlaub' in the Zoologisches Institut und Zoologisches
Museum, University of Hamburg, where Hartlaub worked; and the type material is probably
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
103
therefore lost. The original illustrations were detailed, however, and some appraisal of
Laomedea conferta is possible.
I could not locate type material of Laomedea exigua Sars.
OTHER MATERIAL EXAMINED. All BMNH non-type material is listed. Camara do Lobos,
Madeira, 146m ('80 fms'), coll. & pres. R. Kirkpatrick, 9 colonies in spirit; 1922.3.4.6.
'Probably Woods Hole', Massachusetts, U.S.A., ex James Ritchie collection, parts of two
colonies on microslide; 1964.8.7.83 (mentioned, Rees & Thursfield, 1965 : 102). Cape
Town, Republic of South Africa, on moored raft, 15 Dec 1949, rf & 9 colonies on two
microslides; South African Museum coll. SH 423 (Fig. 18).
/D\
Fig. 18 Laomedea calceolifera. (a) part of 9 colony, showing two mature gonothecae and,
uppermost, an immature one. 'Probably Woods Hole', ex James Ritchie colln (mentioned, Rees
& Thursfield, 1965 : 102); 1964.8.7.83. (b-c) optical sagittal section and sketch of part of
lowermost gonotheca in (a), (d) cf gonotheca, Gulf of Pago, NW Yugoslavia; after Babic (1912).
(e) cf, Cape Town docks, ex South African Museum, microslide preparation SH423. Scale (a-c,
e, and probably d) 500 /urn.
DESCRIPTION. Mature colony comprising one to several erect, monosiphonic stems inserted
at short, irregular intervals on a smooth, tortuous hydrorhiza. Stems probably usually
unbranched, flexuose; internodes slightly curved to almost straight, long, 3-10 annulations
basally. Terminal tendrils not recorded. Hydrothecae campanulate, delicate, flared distally,
length c. \{ x breadth, thin walled, rim even; diaphragm transverse to oblique; pedicels 3-20
ringed, sometimes with smooth central portion (Fig. 18). Hydranth (BMNH 1964.8.7.83)
104 P. F. S. CORNELIUS
with 15-20 tentacles; hypostome large, probably sub-spherical in life. Gonothecae in axils,
single or in clusters of up to 3; 9 club-shaped with sub-terminal introverted curving tubular
aperture on one side, several embryos developing internally; d1 roughly cylindrical, elongate,
tapering gradually below, more sharply above, aperture terminal, central, at end of
introverted tube (as 9 but straight). Young 9 gonotheca truncate (Miller, 1973), lacking
distinctive aperture until late in development (? young d1 similar). Reproduction described in
detail by Miller. No medusa stage.
Variation. Little information. Male gonothecae vary in width (Fig. 18).
DISPERSIVE STAGE. Planulae, which develop within the gonotheca. Splettstosser (1924, as C.
conferta) described the early stages in sporasac development and more recently Miller (1973)
gave a detailed account of the whole reproductive cycle. Miller interpreted the sporosacs as
retained medusae ('gonomedusae').
REPRODUCTIVE SEASON. Fertile material recorded April-August in NW France (Teissier,
1965), June off Norfolk (Hamond, 1957), December in the austral seasons of South Africa
(present material).
DISTRIBUTION. Widely distributed in warmer parts of the Atlantic Ocean, from South Africa
(Millard, 1975) north to the English Channel and southern North Sea on the European coast
and to Maine (Fraser, 1946) on the American side. However, published localities are few.
For example the only definite British record since Hincks' description of the type material
is from off Norfolk (Hamond, 1957, as L. conferta). Possibly the several British records of L.
exigua Sars refer to this species: Swanage, Dorset (Hincks, 1868; possibly BMNH
1899.5.1.153 is this material); KJlve, N Somerset (Bassindale, 1941); and Great Yarmouth,
Norfolk (Harmer, in Hamond, 1957). Other European records are as follows: R Ranee and
Roscoff, NW France (Philbert, 19356; Faure, 1965; Teissier, 1965), Helgoland (Hartlaub,
1897, as Campalaria conferta and Portugal (Da Cunha, 1944, as L. angulata and L.
calceoliferd). The species has been recorded from the 'Mediterranean Sea' by several authors
(Babic, 1912, as L. angulata; Billard, 19316; Philbert, 19356; Picard, 19586) and, more
precisely, from Castiglione, Algeria (Picard, 1955, as Campanularia conferta). Lastly Linko
(1911) reported material from Sebastapol, Black Sea. Naumov (1969) stated that the species
occurred along the 'Atlantic coast of England' and in the North Sea but he confused
calceolifera with angulata, to which his notes may partly refer (see also Remarks
concerning 'C. exigua').
HABITAT. Extreme lower shore (Faure, 1965) and shallow offshore (Hincks, 1871),
sometimes deeper. Millard (1975) listed a record of an infertile colony from 70 m off South
Africa; and one of the specimens listed above was said to come from 146 m ('80 fins') off
Madeira.
Faure (1965) and Teissier (1965) recorded a usual substrate of rocks and pebbles.
However, some authors (Hartlaub, 1897; Splettstosser, 1924; Vervoort, \946a; all as
'conferta') have noted an affinity for Sabellaria tubes (see also notes under Type material)
while other 'living' substrates recorded include a spider crab (Hyas araneus) carapace and an
ascidian (Styela coriacea) (both by Hamond, 1957, as Laomedea conferta), and intertidal
Lithothamnion pools (Teissier, 1965). Morri (1979<a, 6) recorded L. calceolifera in a range of
salinities down to 27%oin brackish lagoons in Italy.
REMARKS. For distinguishing features from Laomedea angulata see that species. Infertile
specimens can be difficult to distinguish from some young, infertile L. angulata and Obelia
dichotoma colonies.
Type material of the nominal species Campalaria conferta Hartlaub, 1897, could not be
located but the original description and detailed illustrations agree with the type material of
the present species in vegetative features. A difference, however, is that the material
described by Hartlaub had gonothecae with truncated ends. Miller ( 1 973) has shown that the
peculiar asymmetric terminal region with its introverted tubular aperture forms late in 9
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
105
gonotheca development, indicating that Hartlaub's account was probably of immature L.
calceolifera material. Splettstosser (1924) and Hamond (1957) probably also had immature
material which, following Hartlaub, they assigned to C. conferta.
The identity of Laomedea exigua Sars, 1857, remains obscure. The species was based on
Gegenbaur's (1854) material from Sicily briefly described as Campanularia sp. nov. It might
prove conspecific with calceolifera, which it predates. Van Beneden (1866), Hincks (1868)
and Leloup (1947, 1952) assigned English Channel material to exigua but it seems likely
their material was male calceolifera. Naumov (1960, 1969) provided a redescription of L.
exigua, copying Hincks' figure of a colony and a gonotheca, already copied from Gegenbaur.
He recorded the nominal species from the Black, Adriatic and Mediterranean Seas and the
eastern North Atlantic from 'Gibraltar to the North Sea inclusive', but did not cite material.
Probably his records refer to calceolifera. Bassindale's (1941) record from SW England might
also have been young L. calceolifera.
Dimorphic gonothecae were probably first recognized in L. calceolifera by Babic (19 12, as
L. angulatd), whose illustration of the male gonotheca is redrawn here.
Stolon growth was studied by Wyttenbach et al. (1973).
Laomedea flexuosa Alder, 1857
(Fig. 19)
Laomedea flexuosa Hincks, in Alder, 18566 : 440 (nom. nud.); Alder, 1857 : 122-123.
Campanularia flexuosa: Hincks, 1868 : 168-170, pi. 33.
non Eulaomeda flexuosa: Rees & Thursfield, 1965 : 102-103 (lapsus pro Eulaomedea; redet. Obelia
dichotoma).
Fig. 19 Laomedea flexuosa. (a) New England, USA; 1915.3.6.7. (b) same, rf gonotheca. (c) 9
gonotheca, ?British Isles: 1912.12.21.290a. (d) part of colony with internodes longer than
normal, S Greenland; 1 938.3. 1 .297. Scale (a-d) 500 urn.
106 P. F.S.CORNELIUS
NOMENCLATURE. The original publication of the name flexuosa (Hincks, in Alder, 18566)
lacked description, figures or other definitive indication but a description was published a
year later (Alder, 1857). The species has been widely recognized and a lengthy synonymy is
unnecessary.
TYPE MATERIAL AND LOCALITY. The type locality has been restricted to the British Isles by
Cornelius & Garfath (1980). We listed the syntype material, which is preserved in the
Hancock Museum, Newcastle upon Tyne.
OTHER MATERIAL EXAMINED. BMNH series, about 50 microslide preparations.
DESCRIPTION. Colony comprising often branched, erect stems up to c. 30 mm arising at
irregular intervals from a straight, branching stolon. Stems and branches flexuose; internodes
characteristically curved, length varied between colonies but constant within, each having
1-10 annuli basally. Hydrothecae campanulate, robust, sometimes asymmetrically thick-
ened, length equal to or slightly longer than greatest width; rim even, diaphragm transverse;
pedicels 3-20 ringed, slightly tapering distally, occasionally with smooth central portion.
Hydranth with 17-23 tentacles, hypostome spherical. 9 gonotheca subcylindrical, tapering
basally, truncate distally; sides smooth to slightly sinuous; opening wide, terminal; pedicel
short, ringed, rf shorter, tapering distally to narrower aperture than in 9. Nematocysts
described by Ostman (1979). No medusa stage.
Variation.The following features are variable in the BMNH series: length : breadth ratio and
radius of curvature of internodes (between, not within, colonies); length of hydrothecal
pedicel and number of annuli; and gonothecal shape.
DISPERSIVE STAGE. Planulae, which develop within the gonotheca. Miller (1973) interpreted
the gonophores as retained medusae ('gonomedusae'). Another description of their develop-
ment was given by Goette ( 1 907).
REPRODUCTIVE SEASON. Fertile material recorded April-July, Isle of Man (Bruce, 1948);
June-July, 1934, Northumbria (H. O. Bull, in Evans, 1978); May-December, NW France
(Teissier, 1965); February-March, May, September, S Spain (Chas Brinquez & Rodriguez
Babio, 1977); November, Naples (Lo Bianco, 1909).
DISTRIBUTION. Recorded widely in the North Atlantic but status in some areas unclear.
There are records from the Murman and White Sea coasts and Norway (intertidal pools,
Mathiesen, 1928), the Faeroes and Iceland (Kramp, 1929, 1938), Hardanger Fjord, Norway
(Brattegard, 1966) and the Shetlands southwards (Hincks, 1868; many other British
authors). Records from warmer areas are numerous: Santander, N Spain (Rioja y Martin,
1906), S Spain (Chas Brinquez & Rodriguez Babio, 1977), Portugal (Da Cunha, 1950),
Mauritania (Billard, 1906), Morocco (Patriti, 1970), Ghana (Buchanan, 1957); but not
southern Africa (Millard, 1975). In the Mediterranean Sea the species has been recorded
from Naples (Lo Bianco, 1 909; Riedl, 1 959) and 'E Adriatic' (Pieper, 1 884).
The northern and southern limits on the North American coastline are also unclear.
Although Kramp (1938) reported the species from W Greenland, Calder (1970) did not
record it from Canadian waters and Eraser's ( 1 944) most northerly record was from the Gulf
of St Lawrence. The species has been reported along most of the United States coast, but the
status in the Caribbean is just a single record which was doubted by both Fraser ( 1 944) and
Vervoort(1967).
HABITAT. Recorded on a wide variety of inert and living substrates (Hincks, 1868; Vervoort,
1946a; Marine Biological Association, 1957; Teissier, 1975; Chas Brinquez & Rodriguez
Babio, 1977), probably lacking a strong substrate preference although often found on fucoid
algae. Occurs intertidally, and probably not often deeper than the 37 m recorded by Miller
( 1 96 1 ) and the 40 m from which a BMNH specimen was collected (reg. no. 1971.5.11 .24, W
Scotland). However, a deep record from '12 positions 20^48 miles (32-77 km) SW of
Eddystone', SW England, 73-92 m (Marine Biological Association, 1957), suggests the
species is not unusual at such depths; and Crawshay (1912) reported the species from
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 107
80- 100m at several stations in the western English Channel. The four microslide
preparations listed under this species by Rees & Thursfield (1965), from depths down to
160 m, are ofObelia dichotoma material.
REMARKS. Laomedea flexuosa is type species of the unacceptable genus Eulaomedea Broch,
1910, and of the genus Laomedea (discussion of both on pp. 97-98).
Alder (1856/7) introduced the name flexuosa in passing when describing Laomedea
neglecta in the phrase 'L. flexuosa Hincks, MS'; but the introduction was invalid as it lacked
an acceptable indication. It is clear that in or shortly before 1856 Alder and Thomas Hincks
concurred thai flexuosa was a valid species, and were the earliest to recognize it. When first
mentioning the name Alder (18566) cited as indication 'the young of Johnston's small
variety of L. gelatinosa'; but Johnston (1847 : 472) said that he had not seen material. He
stated simply: 'variety a — with even margins and simple stem (not seen by me)'. Johnston
apparently referred to hints by earlier authors that there was a species awaiting
recognition. But Johnston did not see flexuosa material himself, nor did he cite this earlier
literature. Hence Alder's (18566) indication of Johnston's brief account does not satisfy the
requirements of nomenclature, and the name flexuosa must date from the later paper (Alder,
1857).
The earlier mentions of the species which Johnston (1847) apparently had in mind are
not easily traced. He may have thought that Couch (1844 : 39-40) had seen some flexuosa
material when, in describing gelatinosa, he stated that species often to be just one inch
(2 5 '4 mm) high compared with the greater height usual in gelatinosa s. str. It can be noted
that Couch might have included neglecta also within gelatinosa, since he recorded that
'gelatinosa' sometimes grew under stones. Following Couch's early death it was left to Alder
and Hincks to define correctly the three species involved. Thus, Alder (18566 : 440) stated
that 'if observed [by earlier workers — possibly Couch] neglecta has been passed over as ... L.
flexuosa'.
L. flexuosa has been used by several experimental biologists. Stolon growth has been much
studied and was mentioned above (p. 42). Phenotypic response to simple environmental
changes was described by Crowell (1957; 1961; see p. 42). Embryonic rupture of the
hydrothecal bud was studied by Knight (1965, 1970, 1971). Stebbing (1976, 1979) investi-
gated the influence of inorganic toxins on growth, noting that sub-inhibitory doses actually
enhanced growth rates. Factors such as straightness of stolon, growth rate and frequency of
gonophore production were all affected by water quality. In a later paper (Stebbing, 1981) he
studied several factors affecting growth.
The material assigned to this species by Rees & Thursfield (1965) I refer to Obelia
dichotoma.
Laomedea neglecta Alder, 18566
(Fig. 20)
Laomedea neglecta Alder, 18566 : 440, pi. 16, figs 1-2; Vervoort, \946a : 308-310, fig. 316 (syn. L.
brochi Splettstosser; L. lauta Hummelinck); (non DaCunha, 1950 : 142-143, fig. 9).
Campanularia fragilis Hincks, 1863 : 46-47; Hincks, 1868 : 175-176, pi. 32, fig. 3 (syn. C. elongata
Van Beneden).
Laomedea decipiens Wright, 1863 : 49, pi. 5, fig. 9.
Campanularia elongata Van Beneden, 1866 : 164-165, fig. 6 (p. 150); Leloup, 1947 : 24-25.
Campanularia neglecta: Hincks, 1868: 171-172, pi. 30, fig. 2; Hincks, 1872 : 390-391, pi. 20, fig. 4.
Campanularia decipiens: Hincks, 1868: 173-174.
Laomedea brochi Splettstosser, 1924 : 376-403, figs A-Z, A -W.'
Laomedea ?lauta Hummelinck, 1930 : 35-37, fig. 5.
Laomedea (Paralaomedea) neglecta: Hummelinck, 1936 : 57 (syn. ?L. brochi Splettstosser; L. lauta
Hummelinck).
TYPE MATERIAL AND LOCALITY. Single infertile colony preserved as two microslide
preparations, Hancock Museum, Newcastle upon Tyne, Northumberland; designated
108 P. F. S. CORNELIUS
neotype by Cornelius & Garfath (1980). Although the neotype was once part of Alder's
collections and was probably identified by him it almost certainly came from an Essex
locality, whereas most of Alder's collections came from NE England. The type locality
remains 'Cullercoats and Tynemouth, Northumberland, on undersides of stones' (designated
by Nutting, 1915; repeated by Cornelius & Garfath).
TYPE MATERIAL OF OTHER SPECIES EXAMINED. Laomedeo. lauta Hummelinck, 1930, holotype;
Rijksmuseum van Natuurlijke Historic, Leiden, reg. no. 259. Hummelinck's illustration of
this material is very accurate.
OTHER MATERIAL EXAMINED (L. neglecta is only poorly represented in the BMNH collec-
tion). Forth Cressa, St Mary's, Isles of Scilly, on underside of block of granite, c. LWM, 17
Apr 1903, fertile colonies, 'ova in an external capsule' (ms note by E. T. Browne, Zoological
Notebook 10 : 15, preserved in BMNH), spirit + 2 microslide preparations, coll. E. T.
Browne; 1948.9.8.1 19 (Fig. 20). Mewstone Echinoderm Ground, near Plymouth, Devon, c.
50 m, October 1899, infertile colonies, coll. Marine Biological Association of the U.K., det.
E. T. Browne (ms note in Zoological Notebook 13 : 149), spirit material (1948.10.1.126)+ 1
microslide preparation (1959.9.17.19). Gaso Ranna, Gullmarsfjord, Sweden, 20-30 m, 27
Aug 1962, infertile colonies, spirit material + 1 microslide preparation, coll. W. J. Rees;
1962.11.8.9. Zooligisch Museum, Amsterdam: Brehorn, Zuider Zee, Netherlands, 14 Jun
1927, infertile and <5 fragments in spirit (Zuider Zee Onderzoek sta. 1; det. & mentioned
Hummelinck, 1936 : 57, fig. 9 a-g) (Fig. 20). Dollard, NE Netherlands, fertile colony in
spirit, coll. A. P. C. de Vos, 9 Aug 1954. Kornwenderland, NE Zuider Zee, 4-6 m, coll.
Zoological Station, den Helder, 29 Sep 1938, several infertile fragments in spirit, det. W.
Vervoort. Rijksmuseum van Natuurlijke Historie, Leiden: Aberystwyth Bay, Wales, 1 7 Jun
1939, intertidal, several infertile fragments, coll. P. W. Hummelinck; RMNH reg. no. 968.
Strand Renesse, Schouwen, Netherlands, 20 Dec 1941, infertile fragments on Tubularia
?indivisa stems, coll. J. Viergever; RMNH reg. no. 702 (some of these specimens are
polysiphonic, having two hydrocauli fused basally). Kornwenderland, 6m, 29 Sep 1938,
four small colonies on one microslide, coll. Zoological Station, den Helder, det. W.
Vervoort; RMNH reg. no. 1012.
DESCRIPTION. A small species. Hydrorhiza smooth, tortuous, sometimes branched;
short, delicate monosiphonic to bisiphonic stems arising at irregular intervals. Internodes
long, narrow, often wider in middle than at ends, 3-10 rings basally, curved (recalling L.
flexuosa) to almost straight (e.g. Hummelinck, 1936 : fig. 9a); each sharply inturned basally;
hydrothecal pedicel on short distal process in line with axis of internode. Hydrotheca long,
delicate, unthickened, cylindrical; length = 3 x width; diaphragm oblique to transverse; rim
usually bimucronate but often abrades smooth; pedicel long, tapering distally, up to c. 20
annuli, sometimes with 1-3 smooth central portions. Hydranth with c. 20 tentacles (Van
Beneden, 1866, as C. elongata; present material, 1971.5.11.11). Gonotheca (after
Splettstosser, 1924, as L. brochi) rf = 9, cylindrical to inverted-conical, truncated above.
Acrocyst in 9, eggs possibly extruded singly (Hincks, 1868; Splettstosser, 1924).
Variation. The delicate hydrothecal rim easily abrades smooth. Some published descriptions
suggest that there are minute spines associated with the cusps on the hydrothecal rim but
these seem simply to be folds. Splettstosser (1924 : fig. B, as L. brochi) illustrated a
hydrotheca in which the bimucronate condition was obscure, and quite apart from damage
by abrasion it seems that not all specimens are perfectly bimucronate. Other bimucronate
species vary in the same way (Obelia bidentata, Clytia paulensis; see also p. 40).
DISPERSIVE STAGE. Planulae, brooded in 9 acrocyst derived from vestigial retained medusa
(Splettstosser, 1924). Developing eggs have been reported in the endoderm tissues of the stem
(Splettstosser).
Hincks (1872) described what was apparently a vegetative spore developing in place of a
hydranth, on a specimen collected from British waters in June.
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
109
REPRODUCTIVE SEASON. Probably April-October, but infertile colonies not unusual during
this period. All available information is given: fertile material recorded Netherlands, 14
June, 1927 (Hummelinck, 1936; also present material) and 9 August, 1954 (present
material); Isle of Man, August (Bruce et al., 1963); Norfolk, England, 18 September, 1954
(Hamond, 1957); Northumbria, June (J. H. Robson, in Evans, 1978); SE Scotland, October
(Laverack & Blackler, 1974); Scilly Isles, 27 April, 1903 (present material).
Infertile material recorded Netherlands, 27 August, 1929 (Hummelinck, 1936), 29
September, 1938 & 20 December, 1941 (present material); Aberystwyth, Wales, 17 June,
1939 (present material); Norfolk, 2 July, 1952 (Hamond, 1957); W Sweden, 27 August,
1962; SW England, October, 1899 (present material).
Fig. 20 Laomedea neglecta. (a) part of colony, Zuiderzee, Cl = 14'8%o; Amsterdam Zoological
Museum collection, (b) same, one hydrotheca. (c) same, part of hydrothecal rim. (d) gonotheca
with acrocyst, Is of Scilly, SW England; 1948.9.8. 1 19. Scales: (a, c-d) 500 ^m; (b) 100 //m.
DISTRIBUTION. Although infrequently reported L. neglecta seems widely distributed in the
North Atlantic. The most northerly and southerly records are from Iceland (Kramp, 1938)
and the Adriatic Sea (Vatova, 1928). All other published records are listed (see also Material
list): England (Alder, 1 856/7; Hincks, 1868, 1872; Hamond, 1957); Isle of Man (Herdman, in
Wood, 1901 : 20; Bruce et a/., 1963, as Campanularia fragilis); Scilly Isles (Vallentin, in
Browne & Vallentin, 1904; repeated in Robins, 1969); SE Scotland (Wright, 1859, as
Laomedea decipiens, see Remarks; Laverack & Blackler, 1974); Aberystwyth, Wales
(present material); Netherlands (Vervoort, 1946a); Belguim (as C. fragilis, Van Beneden,
1866, repeated in Leloup, 1947, 1952); Kattegat (Jagerskiold, 1971; Rasmussen, 1973); W
France (Billard, 1927).
Picard (19586) did not include the species in the mediterranean faunal list but Vatova's
(1928) adriatic record seems soundly based. The records by Broch (1933), from the Adriatic,
110 P. F. S.CORNELIUS
and Da Cunha (1950), from Portugal, were probably based on Obelia bidentata material.
L. neglecta has also been recorded from the eastern coast of North America (Fraser, 1944).
HABITAT. Intertidal to at least 50 m. All available information is given: intertidal, under-
neath stones and on other hydroids (in Britain, Alder, 18566; Hincks, 1868); under stones,
on Tubularia stems and at 50 m depth (present material); 1 5-50 m on sea-beds of mud, clay
and shell gravel (Kattegat, Jagerskiold, 1971); 'on stone embedded in meshes of whelk pot',
offshore, and under rocks in tidal creek (SE England, Hamond, 1957); on pebbles & oysters
(W France, Billard, 1927); LWM, underside of rock (Scilly Isles, present material, Browne &
Vallentin, 1904; repeated in Robins, 1969).
Collected in salinity of 26'7%ofrom Zuider Zee (Hummelinck, 1936 : 57; data with speci-
men in Zoologisch Museum, Amsterdam).
REMARKS (see also Remarks under Laomedea flexuosa). Laomedea neglecta is a small
species and although widespread has been little recorded. It was first described from English
waters but there are still only a few records from Britain and Europe.
Infertile colonies resemble Obelia bidentata, and there is some overlap in hydrothecal
length. Although the stems of L. neglecta are probably always narrower than those of O.
bidentata, identification of small specimens can still be difficult.
The spines associated with the hydrothecal rim by several authors are merely folds in the
hydrothecal wall appearing as artefacts during preservation.
Campanularia fragilis Hincks, 1863, was probably founded on a colony of the present
species. The original illustration was actually published, without a binominal, a year earlier
than the species name and description, in vol. 10 of the same journal, forming plate 9, figure
3. The illustration seems to represent L. neglecta. Hincks distinguished the two species on
the 'markedly flexuose character of the stem', the long hydrotheca with plain rim and the
small size of the colony. Probably the hydrothecal rims had abraded smooth. The holotype
(not found) was collected from under stones in a rock pool, a likely habitat for L. neglecta.
Hincks later (1868) referred to the species as C. ?fragilis, and significantly regarded C.
elongata Van Beneden, 1866, type locality Ostend, Belgium, as conspecific. Vervoort
(1946a :310) quoted Maitland's (1897) opinion that elongata and fragilis were conspecific;
an opinion Vervoort considered to have been a pers. comm. from Van Beneden to Maitland.
Leloup (1947) concurred with this synonymy. There seems to have been no material
recorded as elongata since the original description. Van Beneden illustrated a long
hydrotheca with even rim and described the (infertile) colony as small. The hydranth had c.
20 tentacles. His description, like that of Hincks, probably represents L. neglecta with
abraded hydrothecal rims.
Laomedea decipiens Wright, 1863, although described with some precision, was
illustrated only by a small woodcut of a hydrotheca. Wright likened the species to L. neglecta
except that the hydrothecal rim was 'even, and had the appearance of being double for about
half its length from the rim', arguably a misinterpretation; and that 'the reproduction' of L.
decipiens resembled 'exactly' the process he had described (Wright, 1859) in Opercularella
lacerata (Johnston, 1847). That is, there was an external acrocyst. But Wright said his
'decipiens' material differed in that each acrocyst contained only three ova, while those of 0.
lacerata had seven or eight. L. neglecta is now well known to have an acrocyst containing
rather few ova. Further, its hydrothecal rims often wear smooth. Hence it is plausible that
decipiens was based on fertile material of neglecta in when the hydrothecal rims had worn.
Hincks (1868) repeated Wright's description and indicated that Wright had obtained his
material from the Firth of Forth — Wright had given no locality. Wright and Hincks were in
close contact (Hincks, 1868 : preface) and Wright probably told Hincks the locality. Hincks
did not know the gonosome of neglecta and was rightly cautious when treating decipiens.
The record from the Firth of Forth was repeated uncritically by Leslie & Herdman (1881)
and Pennington ( 1 885), but the species seems not to have been recognized since.
Laomedea brochi Splettstosser, 1924, and Laomedea lauta Hummelinck, 1930, were
N.E. ATLANTIC CAMPANULARIID HYDROZOANS
11
referred to the present species by Hummelinck (1936) and Vervoort (1946a) with good
reason. See also the notes under Variation.
Da Cunha (1950) recorded L. neglecta from Portugal growing on a species of Sertularella.
However, his illustration shows a medusa developing within a gonotheca and his material
was probably Obelia bidentata.
The affinities of L. neglecta are discussed above (p. 47-49).
Laomedea pseudodichotoma Vervoort, 1959
(Fig. 21)
Laomedea (Eulaomedea) pseudodichotoma Vervoort, 1959:316-318, figs 56-57; Vervoort,
1966: 104.
TYPE MATERIAL AND LOCALITY. Lectotype designated from original syntype series by
Vervoort (1966), 50mm d1 colony with developing and mature gonothecae, 13°43'N,
17° 23' W (off Senegal); mostly preserved in University Zoological Museum, Copenhagen,
with part on microslide in Rijksmuseum van Natuurlijke Historic, Leiden. 'Paralectotypes',
9 colonies, from 5° 37' N, 0° 38' E (off Ghana); also preserved in Copenhagen.
OTHER MATERIAL EXAMINED. Off Abidjan, Ivory Coast, 35m, 2 Mar 1966; Rijksmuseum
van Natuurlijke Historic, Leiden, reg. no. 10410 (Fig. 21).
DESCRIPTION (partly after Vervoort, 1959, 1966). Colonies so far recorded up to 50mm,
polysiphonic basally; stem almost straight; roughly pinnate; some secondary branching;
branches approximately alternate, in one plane, in the only available colony. Internodes
slender, 2-3 ringed basally; some tanning. Hydrothecal pedicels short, 5-10 ringed; some
axillary. Hydrothecae delicate, conical, some slightly swollen below; diaphragm oblique in
side view, basal chamber large; rim even, circular. Gonothecae dimorphic, d1 long,
cylindrical to slightly tapering basally, widest about ± from aperture; truncate, aperture
simple, not raised, as wide as gonotheca. Immature cf gonotheca much shorter. 9 gonotheca
long, widest in terminal |, tapering gradually below and abruptly above; truncate; aperture
raised, half width of gonotheca; gonophores thought to be heteromedusoid, sessile.
Fig. 21 Laomedea pseudodichotoma. (a-c) part of colony, single hydrotheca and 9 gonotheca,
Abidjan, Ivory Coast, 35 m. (d) rf gonotheca, off Senegal; syntype series. Scales: (a) 500 //m;
(b-d)500/an.
112 P. F. S. CORNELIUS
DISPERSIVE STAGE. Almost certainly no medusa (Vervoort, 1959). Presumably planulae are
released in the normal way.
REPRODUCTIVE SEASON. Fertile material recorded 30 January, 1946, 50 m, off Ghana, 9; 2
March, 1966, 35 m, off Ivory Coast, 9; 25 April, 1946, 65-89 m, off Senegal, rf; (Vervoort,
1959; present material).
DISTRIBUTION. Recorded only from the coastal waters of Ghana, Ivory Coast and Senegal.
HABITAT. So far found only on Sertularella cylindritheca (Allman, 1888), at depths from
35 m to 89 m (Vervoort, 1959; present material).
REMARKS. This species is known only from tropical west Africa.
Genus OBELI A Peron & Lesueur,
Obelia Peron & Lesueur, 18100 : 355; Peron & Lesueur, 718106 : 43.
Monopyxis Ehrenberg, 18340 : 297; Gray, 1848 : 84.
Obelomma Haeckel, 1879 : 176.
Obeliopsis Le Danois, 1913:110.
For other synonymy see Remarks, Cornelius (\915a) and page 114. See also note 3 (p. 124)
concerning the date of introduction of the genus name Obelia.
TYPE SPECIES. Obelia sphaerulina Peron & Lesueur, 1810# (nom. nov. pro Medusa marina
Slabber, 1769); by monotypy. For taxonomic purposes the type species was taken to be
conspecific with hydroid O. dichotoma (Linnaeus, 1 758) by van der Hoeven ( 1 862 : 280) and
Russell (1953 : 297), but this link is subjective. For the time being at least there is great
difficulty in relating medusae collected from the plankton to their hydroids (e.g. Cornelius,
1975#). Peron & Lesueur cited as indication a specimen taken in Dutch waters, as illustrated
in the German edition of Slabber's (1775-1781 : pi. 9, figs 5-8) work. But Goy (1980 : 72)
links also to the published description an unpublished illustration by Lesueur. The specimen
illustrated therein was taken near Le Havre, in the Museum of which town the illustration is
preserved (see also note 3, p. 124).
Naumov (1960, 1969) and Stepanyants (1979) designated Sertularia geniculata Linnaeus,
1758, type species of Obelia; but geniculata was not an originally included species. For
discussion see Cornelius ( 1 91 5a, 1981).
DIAGNOSIS. Colonial Campanulariidae with: polyp generation forming upright colonies,
branched or unbranched, variably flexuose; stolon not anastomosing; internodes annulated
proximally, supporting hydrotheca on distal lateral process; hydrotheca bell shaped,
hydranth with sub-spherical hypostome; gonotheca inverted cone-shaped, usually with
raised terminal aperture but sometimes simply truncate; medusa umbrella flat, eversible,
mesoglea thin; velum reduced to absent; manubrium long; about 16 marginal tentacles on
release, numerous in adult, short.
REMARKS. An exhaustive synonymy and restriction of this genus have already been
published (Cornelius, 1975a) but the most recent redefinitions of Obelia are by Stepanyants
(1979) and Arai & Brinkmann-Voss (1980). The genera referred to Obelia by Cornelius
included: Slabberia Oken, 1815 (a 'rejected work' for nomenclatural purposes);
Campanularia Lamarck, 1816 (part; but see p. 51); Thaumantias Forbes, 1848 (part; see
also p. 71); Eucope Gegenbaur, 1856 (part; here referred to Clytia, see p. 71);
Schizocladium Allman, 1871; Obelaria Haeckel, 1879 (but see p. 94); Obeletta Haeckel,
1879; Obelissa Haeckel, 1879; and Monosklera von Lendenfeld, 1885. The nomenclatural
problems involving Obelia with Laomedea are discussed above (p. 97).
Three species of Obelia are recognized from the hydroid stage in the eastern North
Atlantic but their medusae cannot be told apart (Russell, 1953; Cornelius, 1975a; Arai &
Brinckmann-Voss, 1980). Two of the species were described from the hydroid stage before
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 1 1 3
any of the medusae, and their 'hydroid names' can be regarded as having safe seniority. The
description of the third valid hydroid stage, O. bidentata Clarke, 1875, post-dates several
binominals applied to the medusa stage and this name cannot yet be regarded safe. But there
are difficulties in identifying to species level the early descriptions of the medusa and linking
them with the correct hydroid stages. Thus it seems probable that the name O. bidentata
will be unchallenged for some time, and that a working stability has been reached.
The subgenus Monopyxis was introduced in the combination Sertularia (Monopyxis)
geniculata Linnaeus, 1758, by Ehrenberg (1834a : 297). S. geniculata is type species of the
subgenus, by monotypy. Since there are only 3-5 species of Obelia recognized or
provisionally accepted from the hydroid stage (e.g. Cornelius, \915a) it seems superfluous to
subdivide the genus, and Monopyxis can be referred to Obelia. Ehrenberg's account was
based on material from Norway, Denmark and Germany. Gray (1848 : 84) used Monopyxis
as a supergenus, including in it the species Obelia geniculata, Hartlaubella gelatinosa,
Campanularia flemingii (now referred to H. gelatinosa, p. 96) and Monotheca obliqua (e.g.
Hincks, 1868, as Plumularia); but this supergenus name was not employed again. Finally
Hincks (1868) included Monopyxis in his synonymy of Obelia.
The genus Obelomma Haeckel, 1879, was established to include Obelia medusae with 48
tentacles on release. It included three nominal species, each incorporating several others
within the synonymies Haeckel presented. Most of those incorporated are now referred to
Obelia, of which Obelomma should be regarded a synonym.
The generic names Obelaria Haeckel and Obelaria Hartlaub are considered under
Hartlaubella (p. 94); and Eucope Gegenbaur is treated under Clytia (p. 7 1 ).
The genus Obeliopsis Le Danois, 1913, was erected to include material referred to a single
species (see p. 120) and can be confidently referred to Obelia.
General notes on the medusa stage and indications to other literature are given under O.
dichotoma (p. 118) and in Cornelius (1975<a). Applications of the genus name Obelia to
bryozoan species were also listed in that paper.
The three species of Obelia (hydroid) occurring in the NE Atlantic can usually be
identified from the characters given by Cornelius (1975a : table 1); but occasional specimens
of O. dichotoma approach O. geniculata in having a slight thickening of the internodal
perisarc. Some of these specimens cannot be confidently identified to species.
Obelia bidentata Clarke, 1875
(For illustrations see Cornelius, \915a)
Obelia bicuspidata Clarke, 1875 : 58, pi. 9, fig. 1; Stepanyants, 1979 : 37-38, pi. 7, fig. 1 (syn. O.
austrogeorgiae Jaderholm, 1904a; Clvtiapaulensis Vanhoffen, 1910).
Obelia bidentata Clarke, 1875 : 58-59^ pi. 9, fig. 2; Jaderholm, 19046 : 270-271 (syn. O. bicuspidata
Clarke; first reviser); Cornelius, 1975a : 260-265, fig. 2 (syn. O. bicuspidata Clarke).
Clytia arborescens: Billard, 1 907 : 1 67 (non Pictet, 1 893).
For further synonymy see Cornelius ( 1 975a) and Table 4.
NOMENCLATURE. Jaderholm (19046) acted as first reviser when using the specific name
bidentata for this species, and several contemporary authors followed him (references in
Cornelius, \975a). Although the superseded name bicuspidata has been widely used, O.
bidentata is the available name under the current conventions of zoological nomenclature.
DISPERSIVE STAGE. The medusa of this species has probably still to be reared to maturity but
when young resembles those of the other two Obelia species (Cornelius, 1975<2, \911a). I did
not mention in the 1975# paper that Professor K. Ramunni 'reared the medusa of this
species' from Bengal material (mentioned in Annandale, 1915, as O. spinulosd). The age to
which he reared it was not recorded. Ramunni was by far the earliest to realize that the
medusa resembled that of the other two Obelia species, and to see it released.
DISTRIBUTION. Recorded from the English Channel and southern North Sea southwards
through most European and African coastal waters but not from the Black Sea or the Baltic.
114
P. F.S.CORNELIUS
Table 4 Synonymies among the nominal species of Obelia described from the hydroid stage (after
Cornelius 1975a, with additions). Although the lists are World-wide most of the nominal species have
been recorded from the eastern North Atlantic. Many were first described in genera other than Obelia.
Discussion of species treated in the 1975a paper is not duplicated here. The references cited in the
table are mainly as in that paper, only the few additional ones being included in the present reference
list.
(a) Synonyms of Obelia bidentata Clarke, 1875
O. bicuspidata Clarke, 1 875
Campanularia spinulosa Bale, 1888
O. andersoni Hincks, 1889
O. bifurca Hincks, 1889
Gonothyraea longicyatha Thornely, 1899 (non
O. longicyatha Allman, 1877)
O. corona Torrey, 1 904
Obelia sp. Clarke, 1907
O. bifurcata Thornely, 1908
O. multidentata Fraser, 1914
O. oxydentata Stechow, 1914
O. longa Stechow, 1921
Clytia longitheca Hargitt, 1924
O. longitheca Hargitt, 1 924 (sic)
O. attenuata Hargitt, 1924
Laomedea bicuspidata var. picteti Leloup, 1932
L. spinulosa var. minor Leloup, 1932
L. bicuspidata var. tenuis Vervoort, 1946
IClytia arborescens: Billard, 1893 (see present
paper, p. 1 17)
(b) Synonyms of Obelia dichotoma (Linnaeus,
1758)
Sertularia longissima Pallas, 1 766
Sertolare genicolata Cavolini, 1785 (lapsus
pro Sertularia geniculata)
ICymodocea simplex Lamouroux, 1816
ITubularia clytioides Lamouroux, in Freycinet,
1824
Campanularia maior Meyen, 1 834
C. brasiliensis Meyen, 1834
C. cavolinii Deshayes & Edwards, 1 836
C. caw//m Chiaje, 1841
Laomedea gracilis Dana, 1 846
O. commissuralis McCrady, 1857
L. divaricata McCrady, 1857
Eucope parasitica Agassiz, 1865
E. pyriformis Agassiz, 1 865
E. articulata Agassiz, 1865
Campanularia flabellata Hincks, 1 866
O. plicata Hincks, 1 868
Schizocladium ramosum Allman, 187 1
O. pygmaea Coughtrey, 1 876
O. hyalina Clarke, 1879
O. adelungi Hartlaub, 1884
O. helgolandica Hartlaub, 1884
O. australis von Lendenfeld, 1885
C.cheloniae Allman, 1888
O. angulosa Bale, 1888
O. chinensis Marktanner-Turneretscher, 1890
O. arruensis Marktanner-Turneretscher, 1 890
O. nigrocaulus Hilgendorf, 1898
O. gracilis Calkins, 1899
(b) Synonyms of Obelia dichotoma (Linnaeus,
1758) (continued)
O. surcularis Calkins, 1899
O. fragilis Calkins, 1 899
O.griffini Calkins, 1899
O. rhunicola Billard, 1901
O. borealis Nutting, 1901
O. dubia Nutting, 1901
O. solowetzkiana Schydlowsky, 1902
C. obtusidens Jaderholm, 1905a
L. congdoni Hargitt, 1909
O. piriformis Bedot, 1910
O. pyriformis: Mayer, 1910
L. sargassi Broch, 1913
O. undotheca Stechow, 1923
O. nodosa Bale, 1924
O. coughtreyi Bale, 1 924
O. obtusidentata Bedot, 1925
O. dischotoma Billard, 1927 (lapsus pro
dichotoma)
O. everta Hargitt, 1927
1O. commensuralis Gudger, 1937 (?lapsus pro
commissuralis)
O. alternata Fraser, 1938
O. equilateralis Fraser, 1938
O. microtheca Fraser, 1938
O. tenuis Fraser, 1938
O. racemosa Fraser, 1 94 1
O. irregularis Fraser, 1943
O. braziliensisVannucci Mendes, 1946
O. biserialis Fraser, 1 948
O. hyaliana Vannucci, 1955 (see present paper,
p. 119)
(c) Synonyms of Obelia geniculata (Linnaeus,
1758)
Laomedea lairii Lamouroux, 1 82 1
Campanularia prolifera Meyen, 1834-
Eucope diaphana L. Agassiz, 1 862
E. alternata A. Agassiz, 1865 (nom. nov. pro E.
diaphana L. Agassiz; non Thaumantias
diaphana A. Agassiz)
E. polygena A. Agassiz, 1 865
E. fusiformis A. Agassiz, 1865
O. gymnopthalma Spagnolini, 1 87 1
Monosklerapusilla\on Lendenfeld, 1885
O. geniculata vars /, // & /// Marktanner-
Turneretscher, 1890
Campanularia coruscans Schneider, 1 897
O. geniculata f. subsessilis Jaderholm, 19506
O. geniculata f. gaussi Vanhoffen, 1910
O. geniculata ff. intermedia, subantarctica &
subtropica Ralph, 1956
N.E. ATLANTIC CAMPANULARIID HYDROZOANS 1 1 5
There are few English records but from the north coast of France southwards there are many:
NW France, Belgium, Netherlands (earliest records in Table 5; also Vervoort, \946a;
Leloup, 1947; Teissier, 1965); Cadiz, Gibraltar, Morocco, Mauritania, S of Madeira
(Stechow, 1925); Algeria, Senegal (Picard, 19516, 1955); Ghana (Buchanan, 1957); Gambia
(Vervoort, 1959); Morocco (Patriti, 1970); Azores, Nigeria, Sierra Leone (Cornelius, \915a);
South Africa (Millard, 1975). The recorded northern limits are north of the Wash (Table 5)
and Schiermonnikoog in the Frisian Islands (53° 30' N, 6°15'E; RMNH Leiden, W.
Vervoort, pers. comm.). By coincidence the most southerly records, from South Georgia and
Tierra del Fuego (Cornelius, \915a), are from a similar latitude, 54° S. A record from Adelie
Land, Antarctica, may be based on Clytia paulensis material (see Remarks).
My statement (Cornelius, 1975#) that Kramp (1961) recorded the species from Helgoland
was incorrect and there are no records from that island (W. Werner, pers. comm.) or from
any other German waters. The Kent material which I listed was wrongly identified.
The Dutch record of Leloup (1933 : 1 1) listed in Table 5 was referred by Hummelinck
(1936 : 56) and Vervoort (19460 : 300) to Hartlaubella gelatinosa; but Leloup also recorded
gelatinosa in his paper. I have not located Leloup's material but if O. bidentata it would be
among the earliest Dutch records.
Some aspects of the European distribution deserve comment. Although the species is
distinctive and occurs widely in warmer parts of the Atlantic and Indo-Pacific oceans, it was
not recorded from the eastern side of the Atlantic until the 1900s. It might previously have
been confused with H. gelatinosa, as suggested by both Billard and Leloup in various
publications. Confusion certainly occurred in some of the published records of the species
from Belgium and England (possibly Leloup, 1933; Robins, 1969; both as H. gelatinosa; see
previous paragraph and Table 5). Further, the species was poorly documented before 1900
and the descriptions available then were in obscure literature. It is understandable that when
Billard (1902) recorded the first French material he referred it to Obelia bifurca Hincks,
1889, described from the Indian Ocean. (O. bifurca is now regarded conspecific.) Much later
Stechow (1925) too referred material to O. bifurca, 50 years after O. bidentata had first been
described; and it is evident that news of bidentata had spread slowly.
Although the European records (Table 5) appear to show the arrival of the species in
Cherbourg and Ostend in 1902, the Netherlands in 1926, Norfolk in 1951, the Scilly Isles in
1966 and Hampshire in 1975 (an unusually hot summer), there is no satisfactory proof of a
gradual extension of range. Indeed, the apparent spread of the species along the coasts of
France, Belgium and the Netherlands follows quite closely the coming of hydroid experts in
those countries, by coincidence from west to east (Billard, Leloup, Hummelinck, Vervoort).
South-east England has seldom been given close attention by hydroid workers. Hamond
(1957) recorded O. bidentata off Norfolk in 1951 as 'rather common' over a wide area,
suggesting that it was already long established — but for how long cannot be decided. Parallel
evidence comes from another overlooked species, similarly told from its near relatives by a
bimucronate hydrothecal rim: Clytia paulensis. Although some BMNH material of
paulensis was collected from near Plymouth in 1899 (p. 90) it was not identified for 80
years; and the species was first recognized from British waters independently in 1973. It
seems that for a long time it too was overlooked and thought to occur no further north than
NW France; but now it has been found again in southern England and, like O. bidentata, in
East Anglia.
It remains puzzling that the nineteenth century English and Belgian experts active around
the English Channel (Couch, Van Beneden, Gosse, Hincks) did not find O. bidentata. Hincks
(1889) in fact described the species from Indian Ocean material, as O. bifurca, and would
surely have recognized British examples. So it might really have been absent at that time; and
it may be relevant that in 1937-1938 Vervoort (19466) did not record the species from the
Waddenzee, N Netherlands, where it now occurs. Finally, although O. bidentata has been
recorded from the well worked offshore region around Roscoff (Billard, 1912; Teissier, 1965)
it is scarce and has been found only in deep water (L. Cabioch, pers. comm.). See also page
44.
116
P. F.S.CORNELIUS
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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
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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
<?v<?/7a, Obelia 1 14
, Orthonia 5 1
, Orthopyxis 60
exigua, Eucope 7 3
exigua, Laomedea 102, 103, 104, 105
fabredomergui, Obeliopsis 120
fastigiata(um), Campanularia & Stegopoma 123
flabellata, Campanularia 1 14
flabellata, Obelia 1 14, 120
flavidula, Clytia 73, 74, 76, 77, 80, 81
.flavidula, Oceania 73, 81
flemingii, Campanularia 95, 96, 1 1 3
flexuosa, Campanularia 105
flexuosa, Eulaomeda 105, 1 1 7
flexuosa, Eulaomedea 98, 107, 1 1 7
flexuosa, Laomedea 40, 41 42 43 47 94
105-107; Fig. 19, p. 105
folleata, Clytia 91
folleatum, Phialidium 9 1
forbesi, Thaumantias 74
./&*/. Clytia 84, 86
fragilis, Campanularia 107, 109, 110
fragilis, Obelia 1 14
fruticosa, Campanularia 121, 122
fruticosa, Lafoea 122
fruticosa, Laomedea 1 24
fruticosa, Sertularia 121, 124
fruticosus, Thyroscyphus 121
fusiformis, Eucope 1 14
gardineri, Clytia 91
gardineri, Phialidium 9 1
Gastroblasta 48-49, 70, 72
raffaelei 74, 81
gaussi, f. of Obelia geniculata 1 14
gegenbauri, Campanularia 73, 80
gelatinosa, Campanularia 95
gelatinosa, Hartlaubella 40, 43, 44 48 95-97
113, 115, 118,119; Fig.16, p. 92
gelatinosa, Laomedea 47, 95, 107
gelatinosa, Obelaria 95
gelatinosa, Obelia 95
gelatinosa, Sertularia 48, 94, 95
gemmifera, Eucope 73
genicolata, Sertolare 1 1 4
geniculata (sens. Lister), Campanularia 9 1
geniculata, Clytia 84, 86
geniculata, Monopyxis 1 1 3
geniculata, Obelia 40, 42, 43, 45, 96 101-102
113,114,118,119-120
geniculata, Sertularia 1 12, 1 13, 1 14, 1 19
gigantea, Campanularia 73, 75, 81
gigantea, Clytia 40, 74, 77
gigantea, Laomedea 74
Gonothyrea 9 1
Gonothyraea 47, 48, 49, 91-92
/mz//mz92,93,94, 119
longicyatha 1 1 4
loveni 40, 41, 42, 43, 44 49 73 79 91
92-94; Fig. 15, p. 92
gracilis, Campanularia sens. Allman 60, 61 , 66;
sens. Meyen 50; sens. Sars 74, 77, 8 1
gracilis, Clytia 42, 7 1,91
gracilis, Laomedea 4 1 , 7 1 , 73, 74, 75, 76 78-79
91,92,94, 114
gracilis, Lomedea 78, 117
gracilis, Obelia sens. Calkins 1 14
gracilis, Silicularia 50, 118-119
gracillima, Campanularia & Lafoea 122
Grammaria abietina 122
grandis, var. of Rhizocaulus verticillatus 69
grandis, Stegella 69
gravieri, Campanularia 84, 85
gravieri, Clytia 42, 85, 86
griffini, Obelia 1 14
groenlandica, Campanularia 52, 55, 56, 57
groenlandica, Eucampanularia 52
gymnopthalma, Obelia 1 14
Hartlaubella 48-49, 9 1,94
gelatinosa 40, 43, 44, 48, 95-97, 113, 115,
118, 11 9; Fig. 16, p. 92
144
INDEX
helgolandica, Obelia 1 14
hemisphaerica, Clytia 40, 4 1 , 42, 43, 45, 70-7 1 ,
73-82, 86, 87, 89, 90, 9 1 , 92, 94; Fig. 9,
p. 75
hemisphaerica, Medusa 70, 71, 73, 74
hemisphaerica, Oceania 73, 78
hemisphaerica, Thaumantias 73, 74
hemisphaericum, Phialidium 7 1 , 74, 77
hendersonae, Clytia 84
hexaradiata, f. of Pseudoclytia pentata 74
hicksoni, Campanularia 54
Hincksella 123
Hincksia 57, 58
tincta 58
hincksii, Campanularia 5 1 , 53-55, 80, 82;
Fig. 3, p. 55
hincksii, Paracalix 5 1
humilis, Campanularia & Cuspidella 123
hummelincki, Campanularia 82
hummelincki, Clytia 41, 42, 44, 69, 82-83, 86;
Fig. 10, p. 82
hummelincki, Laomedea 82
hyaliana, Obelia 92, 1 14, 1 17, 119
hyalina, Gonothyraea 92, 93, 94, 1 19
hyalina, Obelia 92, 94, 1 14, 119
hybridum, var. of Staurostoma laciniatum 84
Hydractinia echinata 43
Hypanthea48,5Q
atlantica 50
repens 50, 66
Hypanthia 50
inconspicua, Campanularia 73
inconspicua, Thaumantias 73, 74, 76
indivisa, Tubularia 123
integra, Campanularia 52, 60
integra, Capsularia 6 1
integra, Eucampanularia 52
integra, Orthopyxis 40, 42, 44, 48, 57, 60-67,
78, 122, 123; Fig. 6, p. 62
integriformis, Campanularia 6 1 , 64, 66-67
intermedia, Campanularia 58, 60
intermedia, f. of Obelia geniculata 1 14
intermedia, f. of Orthopyxis ( = Campanularia)
crenata 59
intertexta, Campanularia 65, 122
intertexta, Conchella 122
irregular is, Obelia 1 14
islandica, Clytia* 42 , 84; Fig. 1 1 , p. 84
islandicum, Phialidium 84
johnstoni, Campanularia 7 1 , 73, 75, 77, 79-80
johnstoni, Clvthia 74
johnstoni, Clytia 69, 70, 73, 74, 80
kincaidi, Campanularia, Clytia, Laomedea &
Obelia 91
Kirchenpaueria pinnata 124
lacerata, Calycella, Campanularia, Capsularia
& Laomedea, 122
lacerata, Opercularella 1 10, 122
lacerata, Wrightia 122
laciniatum, Staurostoma 84
laevis, Campanularia 54, 60, 6 1 , 65
laevis, Capsularia 6 1 , 65
/aev/s,dytia55,57, 73
Lafoea 122
dumosa47,65, 122
fruticosa 122
gracillima 122
parvula 122
LAFOEIDAE47
lairii, Laomedea 1 44
languida, Clytia 8 1
languida, Oceania 74
languidum, Phialidium 74, 81
Laomedea 41,47,48,49,91, 97-98, 1 22
acuminata 122
angulata 41, 42, 43, 44, 98-102, 104, 105,
125; Fig. 17, p. 99
bicuspidata var. picteti 1 1 4
bicuspidata var. tennis 1 14
brochi 47, 107,110
calceolifera 41, 42, 43, 44, 47, 98, 100, 101,
102-105; Fig. 18, p. 103
conferta4\, 102-103, 104
congdoni 1 14, 119
decipiens 107 ', 109,110
divaricata 1 1 4
exigua\02, 103, 104, 105
flexuosa 40, 41, 42, 43, 47, 94, 97, 105-107;
Fig. 19, p. 105
fruticosa 124
gelatinosa47,95, 107
gigantea 74
gracilis 41, 71, 73, 74, 75, 76, 78-79, 91, 92,
94, 114
hummelincki 82
kincaidi 9 1
lairii 1 14
lautalQl, 108,110
lovem'91,92,97
neglecta 40, 43, 47, 48, 96, 107-111;
Fig. 20, p. 109
obliqua 122
plicata \\1.\\9
pseudodichotoma41,44, 111-112; Fig. 21,
p. Ill
repens 60, 6 1 , 62, 66
sargassi 1 14, 119
sauvagii 121
sphaeroidea 98, 102
spinulosa 1 14
tenuis 122-123
lauta, Laomedea 107, 108, 110
INDEX
145
laxa, Sertularia 121
lennoxensis, Campanularia 58, 60
Leptomedusa 57,61
Leptoscyphus tennis 122-123
leucostyla, Obelia 96
leucostvla, Thaumantias 74, 96
linearis,* Clytia 84-86; Fig. 12, p. 85
linearis, Obelia 84
lineata, Thaumantias 74
Lomedea 97
gracilis 78, 1 1 7
longa, Obelia 1 1 4
longicyatha, Clytia 120
longicyatha, Gonothyraea 1 1 4
longicyatha, Obelia 1 14, 120
longissima, Obelia 40, 4 1 , 96, 1 1 4, 1 1 8
longissima, Sertularia 1 1 4
longitheca, Clytia & Obelia 1 14
LOVENELLIDAE38,69
loveni, Gonothyraea 40, 41, 42, 43, 44, 49, 73,
79, 9 1,92-94; Fig. 15, p. 92
loveni, Laomedea 9 1 , 92, 97
loveni, Obelia 92
lucida. Medusa 72, 73
lucida, Thaumantias 73
madeodi, Plumularia 124
macrogona, Orthopyxis 6 1
macrotheca, Campanularia 53, 54
magnifica, Campanularia 52, 53
maior, Campanularia 1 14
marina, Medusa 1 1 2
mccradyi, Clytia* 42, 87-88; Fig. 13, p. 87
mccradyi, Epenthesis, Oceania & Phialidium 87
Medusa 4%, 10,11,124
campanella 73
cymbaloidea 73
hemisphaerica 70, 71, 73, 74
lucida 72, 73
marina 1 12
microtheca, Obelia 1 14
minor, var. of Laomedea spinulosa 1 14
mira, Agastra 6 1 , 67
Modeeria rotunda 123
/H0//w,Clytia55,56,57,80
Monopyxis 1 12, 113
geniculata 1 1 3
Monosklera4&, 112
pus ilia 1 14
Monotheca obliqua 113, 122
morgansi, Campanularia & Eucalix 5 1
multidentata, Obelia 1 14
Multioralis 70, 72
mutabilis, Campanularia 123
mutabilis, Scandia 123
neglecta, Campanularia 107
neglecta, Laomedea 40, 43, 47, 48, 96, 107-111;
Fig. 20, p. 109
neglecta, Paralaomedea 47, 98, 107
Nemertesia antennina 1 2 1
comata 121
ramosa 121, 122
nigrocaulus, Obelia 1 14
nodosa, Obelia 1 14
noliformis, Campanularia 80
noliformis, Clytia 73, 80, 81
novaezealandiae, Billardia 123
Obelaria4S,94,l\2
gelatinosa95
Obeletta4*,\\2
Obelia 39, 41, 42, 47, 48, 49, 77, 91, 94, 96, 97,
112-113, 124
adelungi 1 14
adriatica 120
alternata 1 1 4
andersoni 1 14
angulosa 1 14
arruensis 1 1 4
attenuata 1 14
aus tralis 1 14
austrogeorgiae 113, 117
bicuspidata9Q,113, 114
bidentata 40, 43, 44, 85, 96, 97, 110, 111,
113-117
bifurcal\4,\l5
bifurcata 1 14
biserialis 114
borealis 1 1 4
braziliensis 1 1 4
calceolifera 102
chinensis 1 14
commensuralis 114, 1 17, 118
commissuralis 114, 1 17, 119
congdoni 1 14, 119
corona 114
coughtreyi 114
dichotoma 40, 41, 43, 45, 73, 96, 100, 101,
102, 104, 112, 113, 114, 117-119, 120,
121
dischotoma 1 14, 117
dubia 1 1 4
equilateralis 1 1 4
everta 1 1 4
flabellata\\4, 120
fragilis 1 1 4
gelatinosa 95
geniculata 40, 42, 43, 45, 96, 101-102, 113,
114,118,119-120
gracilis 1 14
griffini 1 14
gymnopthalma 1 14
helgolandica 1 1 4
hyaliana92, 114, 117, 119
hyalina 92, 94, 114,119
irregularis 1 1 4
kincaidi 9 1
146
INDEX
leucostyla 96
linearis 84
longa \ \ 4
longicyatha 1 14, 120
longissima 40, 4 1 , 96, 1 1 4, 1 1 8
longitheca 1 1 4
loveni 92
microtheca 1 1 4
multidentata \ 14
nigrocaulus 1 14
nodosa 1 1 4
obtusidentata 1 1 4
oxydentata \ 1 4
paulensis 88, 90
piriformis 1 1 4
plana 120
p//cflta40,43, 114, 117, 119
polystyla 1 1
pygmaea 1 1 4
pyriformis 1 14
racemosa \ \ 4
rhunicola 1 14
solowetzkiana 1 14
sphaerulina 1 12
spinulosa 113, 114
surcularis 1 1 4
tennis 1 14
undotheca 1 1 4
OBELIDAE50,91
obeliformis, Clytia 73, 75, 8 1
OBELIINAE48,49,91
OBELINAE50,91
Obeliopsis\l2, 113
fabredomergui 1 20
ObelissaW, 112
Obelomma 112,113
obliqua, Campanularia 84
obliqua, Clytia 85, 86
obliqua, Laomedea 1 22
obliqua, Monotheca 113, 122
obliqua, Plumularia 113, 122
obtusidens, Campanularia 1 14
obtusidentata, Obelia 1 14
Oceania 70, 71,87
armata 8 1
discoida 72
hemisphaerica 73, 78
languida 74
mccradyi 87
olivacea, Clytia 67
Opercularella lacerata 110, 122
Orthonia48,49,50,51
everta 5 1
ORTHOPYXINAE50
Orthopyxis 4 1 , 48, 49, 50, 5 1 , 57-58
africana 39 , 60
asymmetrica 6 1 , 67
caliculata 48, 6 1 , 63, 65-66
clytioides 50, 118-119
compressa 6 1 , 66
crenata 40, 42, 52, 58-60; Fig. 5, p. 59
delicata 58-59, 60
everta 60
//iteffra 40, 42, 44, 48, 57, 60-67, 78, 122,
123; Fig. 6, p. 62
macrogona 6 1
poterium 57-58, 60
tincta* 58
volubiliformis 57, 74
oxydentata, Obelia 1 14
ParacalixSl
hincksii 51
pulcratheca 5 1
volubilis 51
Paralaomedea 47, 97, 98
brochii 47
neglecta47,98, 107
parasitica, Eucope \ 14
parvula, Campanularia 122
parvula, Lafoea 122
paulensis, Campanularia 88
paulensis, Clytia 42, 43, 44, 76, 85, 86, 88-91,
113, 11 5, 117; Fig. 14, p. 89
paulensis, Obelia 88, 90
pelagica, Campanularia 73, 78-79
pelagica, Clytia 73
pelagica, Laomedea 74, 75-76, 8 1
pentata, Clytia 42, 91
pentata, Phialidium 9 1
pentata, Pseudoclytia 74
Phialella quadrata 123
PHIALIDAE69
Phialidium 48, 69, 70, 71, 72
bicophorum 74, 80
buskianum 74
discoidum 73
folleatum 9 1
gardineri 9 1
hemisphaericum 1 1 , 74, 77
islandicum 84
languidum 74, 81
mccradyi 87
pentata 9 1
temporarium 74
variabile69, 74
v/riVftcarts71,73,74,86
PHIALIINAE (& PHIAL1NAE) 38, 69
Phialium 69
picta, Eucope \ 1 4
picteti, var. of 'Laomedea bicuspidata '114
pileata, Thaumantias 74
pinnata, Kirchenpaueria 124
piriformis, Obelia 1 14
plana, Obelia & Thaumantias 120
platycarpa, Campanularia 61
INDEX
147
Platypyxis 70,71,80
cylindrica 73
plicata, Laomedea 117, 119
plicata, Obelia 40, 43, 1 14, 1 1 7, 119
Plumularia macleodi 124
obliqual!3, 122
Podocoryne carnea 43
polygena, Eucope 1 1 4
polystyla, Eucope 1 1
polys tyla, Obelia 71
posterior, Clytia 6 1
poterium, Clythia, Clytia & Orthopyxis 57-58,
60,61,63,66
primordiale, Eucope 123
prolifera, Campanularia 1 14
Pseudoclytia 70, 72
pentata f. hexaradiata 74
pseudodichotoma, Eulaomedea 1 1 1
pseudodichotoma, Laomedea 41, 44, 111-112;
Fig. 21, p. Ill
ptychocyathus, Campanularia 80
pulcratheca, Campanularia 5 1
pulcratheca, Paracalix 5 1
punctata, Thaumantias 74
pusilla, Monosklera 1 14
pygmaea, Obelia 1 14
pyriformis, Eucope 1 14
pyriformis, Obelia 1 14
quadrata, Phialella 123
racemosa, Obelia 1 14
raffaelei, Gastroblasta 74, 81
ramosa, Cymodocea 121
ramosa, Nemertesia 121, 122
ramosum, Schizocladium 1 1 4
rara, Campanularia 53
raridentata, Campanularia 73, 74, 75, 76, 80,
81,90
raridentata, Thaumantias 74
repens, Hypanthea 50
repens, Laomedea 60, 6 1 , 62, 66
repens, Silicularia 50
retroflexa, Campanularia 5 1
retroflexus, Eucalix 5 1
Rhizocaulus 41 , 48, 49, 50, 52, 67
verticillatus 40, 4 1 , 43, 48, 52, 57, 67-69;
Fig. 7, p. 68
verticillatus var. grandis 69
rhunicola, Obelia 1 14
rijckholtii, Clytia 6 1 , 65
ritteri, Campanularia 60
rosea, Silicularia 50
rotunda, Modeeria 123
rubra, Agastra6l,63,61
ryckholtii, Clytia 6 1 , 65, 1 23-124
sargassi, Laomedea 114, 119
sarnica, Thaumantias 74
sarsi, Clytia 4 1,74, 78
sauvagii, Laomedea 121
Scandia mutabilis 123
Schizocladium 48, 112
ramosum 1 14
sericea, Sertularia [Bryozoa] 96
serrata, Clytia 85
serrulata, Campanularia 73, 80
serrulata, Clytia 73, 74, 76
Sertolare genicolata 1 14
Sertularia 70-7 1
dichotoma 1 1 7
fruticosa!21, 124
gelatinosa48,94, 95
geniculata 1 12, 1 13, 1 14, 1 19
laxa 121
longissima 1 14
(Monopyxis) geniculata 1 1 3
sericea [Bryozoa] 96
spinosa [Bryozoa] 96
syringa 70, 78
uniflora 55, 57, 73, 74, 77-78
verticillata48,51-52,67,70
volubilis 48, 5 1 , 52, 55, 70, 73, 77-78
Silicularia 48, 49, 50, 57,70
atlantica 50
bilabiata 60
clytioides 50, 118-119
gracilis 50, 118-119
repens 50
rosea 50
simplex, Clytia 73, 8 1
simplex, Cymodocea 114, 1 17, 121
Slabberia 112
solowetzkiana, Obelia 1 14
speciosa, Campanularia 52, 53
speciosa, Eucampanularia 52
speciosus, Campanularia 52
sphaeroidea, Laomedea 98, 102
sphaerulina, Obelia 1 12
spinosa, Sertularia & Vesicularia [Bryozoa] 96
spinulosa, Campanularia 1 14
spinulosa, Laomedea 1 14
spinulosa, Obelia 1 13, 1 14
Staurophora 84
Staurostoma 84
laciniatum 84
Stegella grandis 69
Stegopomafastigiatum 123
striata, Clytia 86
striata, Obelia 85
subantarctica, f. of Obelia geniculata 1 14
subsessilis, f. of Obelia geniculata 1 14
subtropica, f. of Obelia geniculata 1 14
surcularis, Obelia 1 14
syringa, Calycella 55, 57, 78, 122
syringa, Sertularia 70, 78
temporarium, Phialidium 74
148
INDEX
tenuis, Campanulina, Laomedea & Lepto-
scyphus 122-123
tenuis, var. of 'Laomedea bicuspidala' 1 14
tenuis, Leptoscyphus 122-123
tenuis, Obelia 1 14
thaumantias, Eucope 74
Thaumantias 4%, 1Q,1\, 112
buskiana 74
convexa 73
diaphana 1 14
dubia 73
forbesi 74
hemisphaerica 73, 74
inconspicua 73, 74, 76
leucostyla 74, 96
lineata 74
lucida 73
pileata 74
plana 120
punctata 74
raridentata 74
sarnica 74
thompsonil3,14
typica 74
thaumantoides, Eucope 71,73
thompsoni, Thaumantias 73, 74
Thyroscyphus fruticosus 1 2 1
tincta, Campanularia 58, 60
tincta, Hincksia & Orthopyxis* 58
Trochopyxis 70
bicophora 73, 80 (as Clytia)
Tubularia clytioides 50, 1 14, 1 17, 118
indivisa 123
tulipifera, Campanularia & Tulpa 70
Tulpa48,49,70
tulipifera & tulpifera 70
tulpifera, Tulpa 70
typica, f. of Laomedea angulata 100
typica, Thaumantias 74
. Clytia 88, 91
undotheca, Obelia 1 14
undulata, Clytia 60-6 1 , 65
uniflora, Clytia 74
uniflora, Sertularia 55, 57, 73-74, 77-78
urnigera, Clytia 64-65, 73, 78
var iabile, Phialidium 69, 74
\ariabilis, Eucope 73
vermicularis, Campanularia 120
verticellata, Campanularia 67
verticillata, Campanularia 67, 69
verticillata, Campanulata 68
verticillata, Eucampanularia 52
verticillata, Sertularia 48, 5 1-52, 67, 70
verticillata, Verticillina 68
verticillatus, Rhizocaulus 40, 41 , 43, 48, 52, 57,
67-69; Fig. 7, p. 68
Verticillina 67
verticillata 68
Vesicularia spinosa [Bryozoa] 96
vicophora, Clytea & Clytia 73
villafrancensis, Campanularia 74, 8 1
villafrancensis, Clytia 74
viridicans, Clytia 74
viridicans, Phialidium 71 , 73, 74, 86
vitrina, Aequorea 123
volubiliformis, Campanularia 73, 80
volubiliformis, Orthopyxis 57, 74
volubilis, Campanularia 48, 51, 54, 55-57, 69,
70, 73, 76, 77, 79; Fig. 4, p. 55
volubilis, Clytia 73, 74, 76, 77, 78
volubilis, Eucampanularia 52
volubilis, Paracalix 5 1
volubilis, Sertularia 48, 51, 52, 55, 70, 73,
77-78
volubis, Clytia 43
Wrightia 122
British Museum (Natural History)
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The Evolving Earth: 276x219 mm, 280pp, 138 line illustrations, 42 halftones
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The tick collection (Acarina: Ixodoidea) of
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Hydroids and medusae of the family Campanulariidae recorded
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Miscellanea
Miscellanea
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ISSN 0007-1498 Zoology series
Vol 42 No 3 pp 149-225
British Museum (Natural History)
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London SW7 5BD Issued 27 May 1982
GENERAL
Miscellanea
Contents
Page
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
A new species of free-living nematode from the
Firth of Clyde, Scotland
P. J. D. Lambshead
Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Introduction
A new species of free-living marine nematode, Odontophoroides paramonhystera is
described from a fine sandy beach in the Firth of Clyde, Scotland. It is similar to
Odontophoroides monhystera (Gerlach, 1953) from which it differs primarily in having a
precloacal supplement. Odontophoroides monhystera sensu Hopper, 1968, is renamed
Odontophoroides hopperi nom. nov.
Odontophoroides paramonhystera sp. nov.
Figs 1,2, 3 &4
MATERIAL STUDIED. Eleven males and nine females, deposited at the British Museum
(Natural History): Holotype- d1,, BM(NH)198 1.6.42; Paratypes- d1, BM(NH)
1981.6.43-50 & 1981.6.59-60,$,^ BM(NH)198 1.6.5 1-57 & 1981.6.61-62. Also an 'en face'
view of a juvenile was prepared, BM(NH)1981.6.58. In addition specimens of O.
monhystera, collected by Dr S. Lorenzen (1973) and Drs G. Boucher & M. N. Helleouet
(1977), were examined.
LOCALITY. Low water spring mark, fine sandy beach, Scalpsie Bay, Isle of Bute, Scotland.
MEASUREMENTS. See Table 1 .
DESCRIPTION. Slender, colourless, worms. No pigment spots or ocelli seen in preserved
specimens. Cuticle finely, transversely striated (Fig. 3a, b). Six papillae visible in 'en face'
view (Fig. Ic, juvenile): whether R, or R2 not clear. Four R3 cephalic setae, about 1J head
diameters long. Two fine subcephalic setae associated with the subventral R3 cephalic setae
in females; not observed in males. Six cervical setae present; two subdorsal and four sub-
ventral, posterior subventral cervical setae more posterior in males than females. Stout
lateral setae at base of amphids; about 15 //m long, but difficult to measure accurately due to
the angle at which they project. Somatic setae in four irregular rows; length variable, but
often longer in oesophagus and anterior gut regions. Amphids prominent, shaped like the
head of a looped Shepherd's crook, with cuticularized borders; wider in males. Corpus
gelatum displaced in about half of the male specimens but none of the females. Buccal cavity
large, conical, cuticularized, with a complex arrangement of teeth (Fig. Ic, d). Six teeth
present, bifurcate with a minor projection growing out of the main tooth. The four sublateral
teeth are large and heavily cuticularized. Lateral teeth reduced to flat plates. Teeth attached
at base to heavily cuticularized anterior section of buccal cavity (Fig. Id). Two subdorsal,
cuticularized tooth-like projections present; no equivalent subventral structures. In some
preserved specimens the mouth was open, partially everting the teeth. Oesophagus
muscular, especially in region which surrounds posterior section of buccal cavity; bulbs
absent. Excretory pore prominent, located opposite posterior half of buccal cavity. Cellular
sac in close association with posterior of excretory cell (Fig. 2b); function unknown. Caudal
glands present lying entirely posterior to the anus. Tail conical.
Bull. Br. Mus. nat Hist. (Tool.) 42 (3): 149-155 Issued 27 May 1982
150
P. J. D. LAMBSHEAD
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A NEW SPECIES OF FREE-LIVING NEMATODE
151
Fig. 1 (a) Head of male; (b) head of female; (c) 'en face' view of juvenile (dorsal is to the right of
illustration); (d) teeth of female (dorsal is to the right of illustration); (e) cloacal region. Bar
scale = 1 5 yum.
Male. Spicules paired, equal, arcuate, cephalate at proximal end slightly hooked at distal
end; about 4 a.b.d. long. Gubernaculum appears to surround spicules and has narrow,
paired, dorso-caudally directed apophyses. One supplement, with pore, 1 a.b.d. anterior to
cloaca. Two testes, not clearly visible, outstretched, in tandem, ventral and apparently to left
of gut; posterior testis about 250 //m behind anterior testis. Vas deferens widens into sac-like
tube, of granular appearance, 4 a.b.d. anterior to cloaca.
Female. Vulva in anterior position. Vagina with cuticularized walls, surrounded by band
of muscle, which is attached to ventral cuticle, the uterus, and to a tube-like structure. This
tubed structure can give the false impression of connecting the sac associated with the
excretory cell to the vagina (Fig. 2b). Muscle band presumably controls the opening of the
152
P. J. D. LAMBSHEAD
Fig. 2 (a) Whole female; (b) vulval region; (c) tail of male. Bar scale = 1 5 //m.
vulva. Single posterior ovary, outstretched, extending for most of the length of the body,
ventral and to left of gut.
DIFFERENTIAL DIAGNOSIS. Odontophoroides paramonhystera sp. nov. is very similar to O.
monhystera (Gerlach, 1953): the females and juveniles appear morphologically
indistinguishable. However, males of the new species possess a precloacal supplement
(lacking in O. monhystera) and the spicules are distally less slender.
A NEW SPECIES OF FREE-LIVING NEMATODE
153
Fig. 3 Anterior of female, left lateral views: (a) showing amphid, cephalic and subcephalic setae,
and lateral seta; (b) showing fine subcephalic seta associated with subventral R3 seta.
Discussion
The previously monotypic genus Odontophoroides was erected by Boucher & Helleouet
(1977) for Synodontium monhystera Gerlach, 1953. Gerlach (1953) originally described S.
monhystera from Italy, and the types have been lost (Riemann, pers. comm.). However
Gerlach (1953) noted 'Praanal-organe fehlen'. Subsequently a female was described by
Hopper (1968) from Prince Edward Island, Canada, and males, females and juveniles
described by Lorenzen (1973) from Helgoland, North Sea. Boucher & Helleouet (1977) also
described a male (which has unfortunately been lost, Boucher pers. comm.), females and a
juvenile from the English Channel. The type material of S. monhystera by implication,
remains the specimens described by Gerlach (1953). Gerlach's (1953) and Boucher &
Helleouet's (1977) descriptions show the distal ends of the spicules as less slender than
Lorenzen's (1973) specimens. This character may, therefore, prove to be unreliable.
Odontophoroides monhystera sensu Hopper, 1968, is depicted without lateral subcephalic
setae. The text refers to two lateral subcephalic setae, but notes them as positioned '8 // and
23 //, respectively, from anterior extremity' which would not seem to place them in their
typical situation, i.e. at the base of the amphids. Lorenzen (1973) demonstrates that such
setae are only present in adults and fourth stage larvae and Boucher (1974) shows the range
of variation which may occur in the position of subcephalic setae in the related genus
Odontophora, where species may or may not have lateral subcephalic setae. The presence or
absence of these setae may therefore not be a stable generic character. In other respects
Hopper's (1968) specimen is clearly Odontophoroides, being monodelphic (V= 19%) and
154
P. J. D. LAMBSHEAD
Fig. 4 (a) Left lateral view of female head showing subventral and subdorsal teeth, and subdorsal
tooth like projection; (b) right lateral view of female head showing lateral tooth.
having 'six prominent bifid odontia'. As females of this genus appear indistinguishable it
cannot safely be referred to either species and so to avoid zoogeographic problems it is here
renamed Odontophoroides hopperi nom. nov. (type Synodontium monhystera sensu
Hopper, 1968). It is characterized by the lack of lateral subcephalic setae at the base of the
amphid and the absence of fine subcephalic setae associated with the subventral R3
subcephalic setae. The discovery of further specimens, particularly males, should elucidate
the relationship between O. hopperi and the European species but until such time the status
of this species remains equivocal. As a consequence of this analysis the original generic
diagnosis of Odontophoroides given by Boucher & Helleouet (1977) should be modified to
include forms with or without lateral subcephalic setae.
Acknowledgements
I would like to express my thanks to Drs Riemann, Lorenzen and Boucher for kindly lending
specimens of O. monhystera, to Dr Jensen for helpful discussion, to Mr Coles for technical
advice and to the BM(NH) Photographic Unit for the photomicrographs.
References
Boucher, G. 1974. Nematodes libres marins des lies Hautes de Polynesie. 1-Comesomatidae et
Axonolaimidae. Cah. Pacif. 17 : 205-232.
A NEW SPECIES OF FREE-LIVING NEMATODE 1 55
Boucher, G. & Helleouet, M. N. 1977. Nematodes des sables fins infralittoroux de la Pierre Noire
(Manche occidentale). III. Araeolaimida et Monhysterida. Bull. Mus. natn. Hist. nat. Paris 427,
Zoo/. 297: 85-122.
Gerlach, S. A. 1953. Die Nematodenbesiedlung des Sandstrandes und des Kustengrandwassers an der
Italienischen Kiiste. I. Systematischer Teil. Arch. Zoo/. Ital. 37 : 517-640.
Hopper, B. E. 1968. Marine nematodes of Canada. I. Prince Edward Island. Can. J. Zoo/.
46: 1103-1111.
Lorenzen, S. 1973. Freilebende Meeresnematoden aus dem Sublitoral der Nordsee und der Kieler
Bucht. Veroff.Inst. Meeresforsch. Bremerh. 13 : 307-316.
Manuscript accepted for publication 8 June 1981
Notes on Atlantic Asteroidea. 2. Luidiidae
Ailsa M. Clark
Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Introduction
In reviewing the Atlantic species of Asteroidea, recent proposals for inclusion of the family
Luidiidae in the order Paxillosida rather than the Platyasterida (as in the 'Treatise of
Invertebrate Paleontology') need to be discussed. Also, several taxonomic complications to
do with the large genus Luidia have become apparent, particularly in reassessing the ranks of
similar taxa from the continental shelves on the two sides of the ocean. These are better
disposed of in a preliminary paper.
The large collections of the Pillsbury and Gerda of the University of Miami from both
sides of the tropical Atlantic have supplemented the material already in the British Museum
(Natural History). Only L. patriae Bernasconi from Argentina is not available, so comments
on its affinity with L. scotti Bell are based on the published description. The number of
nominal species occurring in the Atlantic is reduced by synonymy or lowering to infra-
specific rank from 17 to 11. Coverage ofL. atlantidea Madsen, L. barbadensis Perrier, L.
ciliaris (Philippi) and L. senegalensis (Lamarck), all of which are limited to either east or
west, is restricted to inclusion in a tabular key of principal characters and an outline of the
distribution. Also L. dathrata (Say) raises only a nomenclatural problem, being another
species of limited range. A neotype for L. alternata alternata (Say) and a lectotype for L.
sarsi sarsi (Diiben & Koren) are designated and discussed.
The good size range of specimens from the Miami collections has allowed for some
comments on post-metamorphosal growth changes.
Systematic account
LUIDIIDAE Sladen, 1889
Astropectinidae: Luidiinae Sladen, 1889 : 244.
Luidiidae: Verrill, 1899 : 201; Fisher, 1911 : 105; Bernasconi, 1943 : 2-3; Fell, 1963 : 433; Spencer &
Wright, 1966 : U42; Downey, 1973 : 21; Blake, 1973 : 22-24 (pt); A. M. Clark & Courtman-Stock,
1976 : 43; McKnight, 1977 : 1 16; Blake, 1982 : 186.
AUTHORITY. Under Article 36 of the International Code of Nomenclature (coordinate
categories), the authority and date of a name in the family group can originate from its use as
a subfamily name, so the citation of Verrill, 1899 as author and date for Luidiidae (e.g. by
Spencer & Wright in the 'Treatise on Invertebrate Paleontology', 1966) is incorrect.
ORDINAL POSITION. In 1963 Fell, followed by Spencer & Wright (1966) split off the
Luidiidae to the order Platyasterida (otherwise including only the lower Palaeozoic family
Palasteriscidae) from its longstanding position close to the Astropectinidae because of its
transversely matching longitudinal series of adambulacral, actinal and inferomarginal plates.
This has never seemed to me satisfactory in view of the close resemblance of Luidia and
Astropecten, especially in transverse section of the arms, and I fully agree with Jangoux
(1975) and McKnight (1977), who dismissed Fell's disposition because of close anatomical
resemblances between Luidia and certain astropectinids, and the different alignment of the
ambulacral and adambulacral plates (the Platasteriscidae having only a very shallow
Bull. Br. Mus. nat. Hist. (Zool.) 42 (3) : 1 57-1 84 Issued 27 May 1 982
158 A.M.CLARK
ambulacral groove, unlike any recent asteroids). The Luidiidae should therefore be included
in the order Paxillosida Perrier, 1884, formerly lumped with Perrier's Valvatida in Sladen's
larger order Phanerozonia (1889). Blake (1982) has independently reached the same
conclusion on the basis of his more detailed morphological studies of the various series of
ossicles, coupled with the absence of any fossil luidiids in pre-Miocene strata. The Luidiidae
shares with the Astropectinidae not only the paxilliform abactinal plates but also the
presence of superambulacral plates, the knobbed tube feet with double ampullae and the
horizontal framework to the body of enlarged marginal plates (though this is not exclusive to
the Paxillosida). The main differences are the lack of transverse matching of the actinal series
with the adambulacrals and inferomarginals in astropectinids — a character probably not of
more than familial weight — and the reduction and paxilliform nature in luidiids of the
superomarginals, so that only their position matching the inferomarginals indicates their
true identity. (In the Clathrata and Alternata groups of species of Luidia, the two lateralmost
series of abactinal paxillae each side also match the marginals in length and are indis-
tinguishable from the superomarginal paxillae unless the latter are broadened — which
reaches an extreme development in Luidia (Platasterias) latiradiata (Gray), referred to the
Clathrata group by Blake (1973); other groups of Luidia species have smaller lateral
abactinal paxillae.)
DIAGNOSIS. A family of Paxillosida with 5-1 1 fairly long, gently tapering arms
(usually described as 'strap-shaped'), not broadening basally into the disc; lower surface
almost flat (and in preserved specimens often also the upper, owing to collapse of the fine
paxillar reticulum); median paxillae irregular and smaller than the lateral ones, of which at
least the two outermost series are arranged in longitudinal series and matched transversely;
papulae distributed all over the upper side between the abactinal plates, compound at their
outer ends; superomarginal plates paxilliform, similar in structure to the adjacent abactinal
plates, though up to twice as long in species where the abactinal paxillae are particularly
small (only in the subgenus Platasterias of Luidia are the superomarginals considerably
broader than the lateral paxillae, though still essentially paxilliform); inferomarginal plates
massive, lateral or lateroventral in position but never conspicuous in dorsal view, each raised
into a flat-topped keel bearing some large spines, especially near the upper end (the ambitus
or broadest part of the arm), interspersed with spinelets of varying size, the sides of the keel
armed with many fine geometrically-arranged fasciolar spinules; a small actinal plate (rarely
two proximally) intercalated between each inferomarginal and the matching adambulacral
plate, often acting as the base for a spiniform pedicellaria (in a few species of the Alternata
group of Luidia there are supernumerary reduced, pedicellaria-bearing actinal plates
superimposed on the abradial ends of the adambulacral plates); interradial areas compact,
rarely with more than a few series of actinal plates; adambulacral plates broad and raised into
a shallow keel, bearing 2-4 large spines along the crest (the fourth, if present, proximal to the
third), the innermost (furrow spine) compressed, curved and sabrelike; pedicellariae well
developed, sessile, with 2-, 3- or even 4-valves, absent in a few species or individuals, those
of the upper side usually nutcracker-like with broad blunt jaws, the marginal and ventral
ones more elongated and spiniform; internally, superambulacral plates linking the
ambulacrals and inferomarginals; tube feet pointed or ending in knobs, their internal
ampullae double.
REMARKS. Doderlein (1920) monographed Luidia Forbes, 1839, distinguishing the following
groups and subgenera:
Clathrata group: subgenera Senegaster and Petalaster Gray, 1 840;
Alternata group: subgenera Maculaster, Alternaster and Armaster;
Quinaria group: subgenera Integraster, Denudaster, Penangaster and Quinaster,
Ciliaris group: subgenus Hemicnemis Miiller & Troschel, 1840 (other names being
Doderlein's).
Fell (1963) pointed out that the type species of both Luidia and Hemicnemis is L. ciliaris
(Philippi) so that Hemicnemis is a synonym. Except for Armaster, which he lumped with
NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 59
Alternaster, he perfunctorily raised all the others to generic rank, a move which has failed
to gain acceptance, notably from Blake (1973) in an exhaustive study of the ossicles of
luidiids and related asteroids. At the same time, Blake reduced Platasterias Gray, 1.87 1 to the
rank of a subgenus of Luidia (whereas Fell had included it in the subclass Somasteroidea),
since P. latiradiata is undeniably more closely related to Luidia clathrata (Say) than
clathrata is to L. ciliaris. The family is accordingly considered to be monogeneric.
Where Atlantic taxa of Luidia are concerned, the limits between Doderlein 's main groups
are particularly blurred. For instance, L. scotti Bell, 1917 (omitted by Doderlein), now
treated as a subspecies of L. ludwigi Fisher which Doderlein put under the Alternata group,
loses its bold colour pattern characteristic of that group soon after preservation and must
have a different pigment to L. alternata. Since L. ludwigi scotti, together with L. armata, also
lacks the enlarged abactinal spines found in the majority of species of the Alternata group, it
bridges the gap to the similarly smooth species of the Clathrata group, both groups having
the two outermost series of abactinal plates matching up with the marginal plates. L.
barbadensis Perrier has the lateral paxillae only slightly outnumbering the marginals by c.
12/10, which serves to differentiate it from the two above groups as defined by Doderlein,
who listed it as 'incertae sedis' under the Alternata group. Its coarse and squarish lateral
paxillae certainly link it more closely with the Alternata group than with the Quinaria and
Ciliaris groups with their finer and more numerous paxillae.
One of the main characters by which at least the Atlantic species of Luidia can be split up
is the alignment of the inferomarginal plates, whether or not they extend above the ambitus.
In the Clathrata and Alternata groups they are ventro-lateral (see Fig. la, b) but in the
Quinaria and Ciliaris groups they extend higher. The difference in alignment is accompanied
by some difference in armament. With more ventrally-aligned inferomarginals, the largest
spines are at the ambitus and project horizontally, or nearly so (though if there is more than
one ambital spine then the uppermost is more or less reduced); the spines below the ambitus
are appreciably smaller also and are somewhat appressed. When the inferomarginals extend
on to the upper surface all their spines are erect, the uppermost one (or two) being largest, the
top one projecting almost vertically upwards in its natural position, at least on those (often
alternate) plates where it arises from its highest locus.
Secondly, the number of lateral paxillae corresponding to the superomarginal ones —
whether matching exactly or in excess — is another easily determined character, again
dividing the Clathrata and Alternata groups from the Quinaria and Ciliaris ones, though L.
barbadensis is an exceptional intermediate, as already mentioned.
These two characters are therefore the most important in the tabular key (table 1) to the
Atlantic species, besides the obvious difference in arm number for some species. The
occurrence of pedicellariae is variable and not reliable for more than local differences at the
infraspecific level, though their location and the number of valves may be useful.
ONTOGENY. The reproduction and larval stages of Luidia sarsi were studied by Tattersall &
Shepherd (1934). An early post-metamorphosal stage of this species is exemplified by the
diminutive holotype, R 8 mm, of Astrella simplex Perrier, 1882, synonymized with L. sarsi
byLudwig(1897).
Fully metamorphosed specimens of other species taken by the Pillsbury in the tropical
Atlantic show that all the primary plates, even the small actinal ones, occur from an early
stage, though the actinals at first lack any armament.
At R c. 6 mm, the upperside paxillae of each arm consist of a band of somewhat irregular
median paxillae flanked on either side by two complete regular longitudinal series, the
outer of which is the superomarginal series and the other the primary lateral series. Almost
immediately a secondary lateral series begins to develop proximally between the two but
probably does not become fully developed to the arm tip until R is c. 25 mm, at least in L.
alternata numidica Koehler. In L. heterozona Fisher, where the lateral paxillae are relatively
smaller, the secondary lateral series is not initiated until R c. 25 mm and is still not quite
complete at R 50 mm. At least in species with fairly coarse paxillae, the primary lateral
160
A. M. CLARK
Fig. 1 Half arm sections, viewed from the proximal side, of: (a) Luidia clathrata, Pillsbury st.
652, Re. 120 mm; (b) L. alternata alternata, Pillsbury st. 767, R c. 90 mm; (c) L. sarsi elegans.
Albatross st. 2 1 77, R c. 115 mm; (d) L. sagamina aciculata, Atlantide st. 6 1 , R c. 90 mm.
series can usually be recognised by the fact that both its abradial and adradial basal lobes are
overlain by the opposing lobes of the secondary lateral and outermost median series of
paxillae respectively (see Fig. la, b, d) but this does not always hold good when the skeleton
is more delicate. The median paxillae also increase in number to some extent, less so in
species where they are stouter, as in L. alternata (Say) where the total number of paxillae
across the arm proximally between the two opposite superomarginal series increases from 9
or 10 at R 6 mm to only c. 12 or 13 at R 50 mm. There are still only c. 12 in the largest
specimen of L. alternata alternata studied, R 175 mm, so the tabula of the larger paxillae in
such species may be markedly broadened.
With regard to the armament, as usual in newly metamorphosed asteroids, the spinelets
and future spines are all attenuated, often trifid, with the terminal point prolonged. This is
still true up to R c. 9 mm, except for the furrow and second adambulacral spines, which are
simple and unbranched, though still uniform in length with the other spinelets. At R c.
10 mm, an ambital inferomarginal spine begins to enlarge in L. alternata and the second
adambulacral spine also increases disproportionately in size, while a few of the more lateral
paxillae begin to develop a single large central spine. In species with a third adambulacral
NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE
161
Fig. 2 (a to d) Dorsal views of a proximal part of an arm, showing the upper ends of several
inferomarginals, the adjacent superomarginals and some abactinal plates, denuded: (a) Luidia
ludwigi scotti, paratype, Terra Nova st. 42, R 62 mm; (b) L. barbadensis, Gerda st. 239, R c.
130 mm; (c) L. sarsi elegans (as in Fig. Ic); (d) L. alternata alternata, Pillsbury st. 767, R c.
15 mm; (e to g) the same views with armament entire (inferomarginal omitted in (f)) of: (e) L.
ludwigi scotti (as in Fig. 2a); (f) L. heterozona barimae, Pillsbury st. 652, R 200+ mm; (g) L. sarsi
africana, Namibia, Re. 100 mm; (h) one adambulacral, actinal and inferomarginal, in ventral
view of L. alternata alternata (as in Fig. Ib); (i) L. heterozona heterozona, Pillsbury st. 24, R
100+ mm, lateral view of two successive inferomarginals and superomarginals with a few lateral
paxillae; (j) apex of jaw of L. heterozona barimae (as in Fig. 20; (k) L. sagamina aciculata, (as in
Fig. Id), two lines of adambulacral, actinal and inferomarginal plates, the one on the left
denuded. The proximal side is to the right in (a-g), to the left in (h, i & k).
162 A.M.CLARK
Table 1 Tabular key to the Atlantic species of Luidia. Entries in brackets signify occasional
occurrence.
12 34567 8 9 10 11
senegalensis
9(8)
_
10
c
_
vl
2
4
_
_
clathrata
5
-
10
c
-
vl
2(3)
4
-
-
ludwigi scotti
5
+
10
c(s)
-
vl
2(1)
4
3
-
patriae
5
_7
10
u
-
vl
2
4
—
-
alternata (2 subspp.)
5
+
10
S
-
vl
2,3
3(^)
3
-
barbadensis
6
-
12-13
c
-
vl
3(2)
4
3
-
sagamina aciculata
5
-
16-19
s
+
1
+ 2/3
2(3)
3
+
heterozona (2 subspp.)
10
-
15-18
u
+
1
+ 3/2,4/3
(4)3
2
+
sarsi (3 subspp.)
5
-
17-20
u
+
1
+/- 4/3,3/2
3(^)
2
-K-)
atlantidea
5
-
15-19
c
-
1
+/- 3/2(4)
3(^)
2
-K-)
clitoris 7(8) 15-18 c (+) 1 + 3-52 -/3 -
1 . Arm number.
2. Colour pattern of upper side:
+ — patchy (may fade in long-preserved L. ludwigi scotti).
uniform or darker along midradial area or lighter along midradial area or lighter along
superomarginal or adjoining lateral paxillae.
3. Number of consecutive lateral paxillae corresponding to 10 superomarginal ones.
4. Paxillar armament:
c — central spinelets distinctly coarser than peripheral ones,
u — central and peripheral spinelets uniform.
s — one central spinelet both longer and stouter on most paxillae.
S — one central spine (1-5 mm long) abruptly enlarged on some scattered (mostly lateral)
paxillae.
5. Shape of superomarginal paxillae:
+ — elongate.
rounded or squarish.
6. Alignment of inferomarginal plates:
1 — lateral.
vl — ventro-lateral.
7. Inferomarginal spines:
+ — alternating on consecutive plates.
all on the same levels.
8. Number of large erect inferomarginal spines.
9. Number of adambulacral spines.
10. Number of valves on actinal pedicellariae (when present, - = absent).
1 1 . Oral furrow pedicellariae:
+ — present.
absent.
spine in the adult, this is probably evident proximally by R 12-15 mm and the other spines
and spinelets grow and coarsen into their adult proportions at this size or soon after. Luidia
sagamina aciculata Mortensen is exceptional in the development of the third adambulacral
spine, which may be lacking altogether, especially in american specimens, judging from the
few so far studied. Increase in numbers of spines also shows a growth gradient along the arm.
In L. heterozona Fisher, for instance, only about four proximal adambulacral plates have the
third spine enlarged at R 20 mm but there is even a fourth spine on the first 10-1 5 plates, and
almost equal to the fully developed third one, by R c. 80 mm.
Actinal and abactinal pedicellariae, when developed, probably first appear between R 1 5
and 20 mm but oral furrow pedicellariae, for example in L. heterozona, are not fully
modified from pairs of spines until R exceeds c. 50 mm.
NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 63
PREVIOUS WORKS. Useful relevant publications on the Luidiidae include the general
monograph of Doderlein (1920), Nataf & Cherbonnier (1973) on the tropical West African
area and Downey (1973) and Walenkamp (1976 and 1979) on the West Indian area.
DISTRIBUTION. The species of Luidia are mostly restricted to the shelf, though several
Atlantic taxa extend to the upper bathyal, including L. barbadensis to 430 metres, L.
ciliaris to 400 metres, L. heterozona and L. sagamina aciculata to 975 metres and L. sarsi
exceptionally to 1 300 metres. (The maximum depth for both L. heterozona and L. sagamina
aciculata derives from Jean Charcot (Walda cruise) st. 40, c. 04° 07' N, 01° 35' W, off Ghana,
975 metres, and is surprising since previous maxima for the two respectively were only 400
and 150 metres, cited by Nataf & Cherbonnier (1973), the vast majority of records for both
species being from 50-100 metres.)
Geographically, five Atlantic taxa appear to be restricted to the american side of the
ocean, namely L. barbadensis, L. clathrata, L. ludwigi scotti, L. patriae and L. senegalensis
(the last despite its name), while L. atlantidea and L. ciliaris are entirely eastern (american
records for L. atlantidea proving incorrect, see p. 168. The remainder are distributed as
follows:
WEST ATLANTIC EAST ATLANTIC
alternata — N Carolina to N Argentina numidica — Senegal to Zaire (Congo)
barimae — Venezuela to French Guiana heterozona — Mauritania to Angola
sagamina aciculataC?) — N Carolina to sagamina aciculata — Mauritania to
Florida Strait Zaire & St Helena
elegans — E U.S.A. & Gulf of Mexico sarsi — Norway to Mauritania & Azores
eleganstf] — S Brazil africana — SW & S of southern Africa
The relationships of these are discussed below to assess their respective ranks.
LUIDIA Fofbes
Luidia Forbes, 1839:123; Sladen, 1889:244-248; Fisher, 1911:105; Sussbach & Breckner,
191 1 : 198; Doderlein, 1920 : 193-246; Bernasconi, 1943 : 3-5; A. M. Clark, 1953 : 379-380; Fell,
1963 : 433; Downey, 1973 : 21-22; Tommasi, 1974 : 7; A. M. Clark & Courtman-Stock, 1976 : 43.
Type species: Luidia fragilissi ma Forbes, 1839 (a synonym of Asterias ciliaris Philippi, 1837), by
monotypy.
Hemicnemis Miiller & Troschel, 1840:105; Doderlein, 1920:217 (as subgenus). Type species:
Asterias ciliaris Philippi, 1837.
Petalaster Gray, 1840: 183: Doderlein, 1920:215 (as subgenus). Type species: P. hardwicki Gray,
1 840, cited by Fisher, 1911 (though with Petalaster in synonymy), non P. Columbia Gray, designated
by Fell, 1963.
Luydia (lapsus) Diiben & Koren, 1846 : 254.
Astrella Perrier in Milne-Edwards, 1882 :21. Type species: A. simplex Perrier, 1882 (a synonym of
Luidia sarsi Diiben & Koren, 1 845), by monotypy.
Senegaster, Maculaster, Alternaster, Armaster, Integraster, Denudaster, Penangaster & Quinaster-
(subgenera) Doderlein, 1920:215-217, raised to generic rank by Fell, 433-434 (except for
Armaster).
Luidia alternata (Say)
Figs Ib, 2d, h, 3a-i,4, 5
Asterias alternata Say, 1825 : 144-145.
Luidia alternata: Liitken, 1859:42^13; Perrier, 1875:334-336 [1876:254-256]; Sladen, 1889:
250-251, 740; Verrill, 1915 : 201-203 [non L. alternata bicolor}; Doderlein, 1920 : 241, 267-268,
figs 7, 1 1; H. L. Clark, 1933 : 20, pi. 1 ; Bernasconi, 1943 : 14-15, pi. 3, figs 1,6, pi. 4, fig. 1; Engel &
Schroevers, 1960 : 6; Ummels, 1963 : 97-98, pis 8, 10; Gray, Downey & Cerame-Vivas, 1968 : 138,
fig. 6; Tommasi, 1970 : 8, pi. 8, fig. 24; Downey, 1973 : 23-24, pi. 2, figs A, B; Blake, 1973 : 33-34,
pi. 6, figs 25-42; Walenkamp, 1976 : 29-32, figs 6, 7, pi. 3, fig. 3, pi. 4, fig. 4.
Luidia granulosa Perrier, 1869 : 109-1 10, pi. 2, fig. 18.
Luidia variegata Perrier, 1875:337[1876:257].
164 A.M.CLARK
Luidia numidica Koehler, 1911 : 3, pi. 1, figs 8-11; 1914 : 167, pi. 4, fig. 7; Doderlein, 1920 : 235, 242;
Nataf & Cherbonnier, 1 973 : 7 1-74, pi. 1 , figs A, C, pi. 5, figs 1 , 2, pi. 7, fig. 6, pi. 9, figs A-D.
Luidia quequenensis Bernasconi, 1942 : 253; 1943 : 15-18, pi. 2, fig. 4, pi. 3, figs 2, 3, 7, pi. 5, figs 1, 2;
1960 : 21-22, pi. 3, fig. 1 ; Carrera-Rodriguez &Tommasi, 1977 : 65.
Luidia bernasconiae A. H. Clark, 1945 : 19-21; Gray, Downey & Cerame-Vivas, 1968 [?pt] : 138, fig.
7; Downey, 1973:25.
Luidia alternata var. numidica: Madsen, 1950 : 206-209, fig. 9.
Luidia alternata numidica: A. M. Clark, 1953 : 388-389, pi. 41 , fig. 1 ; 1955 : 22; Engel & Croes, 1960 :
11-1 2, pi. 6, figs. 1,2.
R up to 175 mm. R/r ranging from c. 5-6/1 at R 3(MO mm to 7-9/1 at R> 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, R<c. 40 mm, where the median paxillae often
have only a single central spinelet, this may stand out from the surface slightly but not as
much as in L. sagamina; larger specimens with multiple central spinelets have them
relatively shorter and making a uniform surface; inferomarginal plates mainly ventral in
alignment, bearing one or two (in the largest specimens, R c. 90 mm, sometimes three) stout
ambital spines, the upper one more or less shorter than the lower when there are two, on the
ventral side 4 or 5 much smaller, almost cylindrical, but slightly tapering spines;
adambulacral plates with 4 large spines, the abradial two in line parallel to the furrow;
pedicellariae 3- or 4-valved on at least the more proximal actinal plates, the valves varying in
shape from broadly spatulate to evenly tapering and triangular (see Fig. 3t, u), sometimes
bivalved pedicellariae present on the paxillae and inferomarginal plates, absent from the
furrow face of the oral plates. Colour (recently dried) patterned above with dark rose-
coloured patches on the centre of the disc and bands across the arms, sometimes linked by
continuous colour midradially.
SYNONYMY. The poorly-preserved holotype and two paratypes of Luidia rosaurae John &
Clark, 1954, from off Venezuela, were thought to differ from L. scotti Bell, 1917, of southern
Brazil in having only a single large ambital inferomarginal spine, finer ventral armament and
more numerous pedicellariae. Walenkamp's material from Surinam (1976) and a number of
Pillsbury and Gerda specimens from northern South America and the Florida Strait now
studied, show that there is considerable variation in the development of the second (upper)
inferomarginal spine, the thickness of the ventral spines and the frequency of the
pedicellariae; consequently L. rosaurae cannot be maintained as specifically distinct.
Walenkamp deferred a decision on this point in default of material from the type locality of
L. scotti. Simultaneously, he rejected the synonymizing of Luidia doellojuradoi Bernasconi,
1941 with L. scotti, proposed by A. M. Clark (1953), on the grounds that its marginal
paxillae are larger than the lateral ones and the R/r ratio is only 7/1 . However, he was misled
in this by Bernasconi's peculiar terminology of the lateral and superomarginal paxillae (see
p. 166), the true superomarginal paxillae of L. doellojuradoi being in fact smaller than the
lateral ones. Also in the 1 5 Terra Nova specimens of L. scotti the R/r ratio ranges upwards
from 7/1; in the lectotype (incorrectly called holotype in 1953) it is 7*5/1. Thanks to Dr
Bernasconi, there are two specimens which she named as L. doellojuradoi in the British
Museum collections. Apart from the greater extent of the actinal pedicellariae on the arms
and the occurrence of some inferomarginal pedicellariae — characters not considered to be of
specific weight — no significant differences from L. scotti are apparent. The ventral
armament of the inferomarginal plates has become appressed during preservation, which
lends a rather coarser appearance than is shown by the well preserved type material of L.
scotti.
It is also possible that L. patriae Bernasconi will prove to be synonymous with scotti.
Walenkamp (1976) percipiently noted the close affinity between L. ludwigi Fisher, 1906,
from California and specimens from the north coast of South America conspecific with L.
rosaurae, referring the latter to the synonymy of L. ludwigi. Fisher's holotype has R
1 10-1 1 5 mm and superficially resembles the larger Atlantic specimens seen except that it is
rather coarser. A paratype of L. ludwigi with R 72 mm compares closely with a paralecto-
type of L. scotti of almost the same size but there appears to be a significant difference in the
number of paxillar spinelets, the median paxillae of ludwigi rarely having less than 6 coarse
ones whereas in L. scotti the number is most often only about 3. Possibly there is also a
colour difference, Fisher having described L. ludwigi (at least when freshly preserved) as
being reddish above 'sometimes mottled with lighter', whereas Atlantic specimens seem to
be more boldly patterned dark (?red) and light. In conjunction with the geographical
difference, these small differences are now considered to warrant a subspecific distinction.
NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 73
One further nomenclatural threat still remains. Doderlein (1920), in listing L. ludwigi,
noted that it is possibly (vielleicht) a synonym of L. armata Ludwig, 1905, from the Gulf of
Panama. Accordingly, one of the two intact syntypes of L. armata from the U.S. National
Museum has been studied; it has R 47 mm and is slightly decalcified and flaccid. The
armament of the paxillae is more or less appressed, emphasizing the spiniform shape of the
central spinelets. Although the larger Pacific specimens of L. ludwigi have markedly coarser
and shorter central spinelets, tending to appear granuliform, smaller Atlantic specimens
from the vicinity of Florida with R c. 40 mm, have a very similar relatively elongate
armament and also agree in the shape and arrangement of the paxillae, plates and spines.
However, the syntype of L. armata differs in having a very large, highly modified bivalved
pedicellaria on each oral plate — a feature somehow overlooked by Ludwig and not found in
the type material of L. ludwigi seen or any Atlantic specimens. It also has relatively small
actinal interradial areas with no sign of a second row of actinal plates. At R 40 mm, L. scotti
already has a single median interradial actinal plate distal to the main series and in larger
specimens, R 50-60 mm, there are one or two such plates each side of the interradius. These
two characters, but particularly the first, indicate that there are two distinct species in the
East Pacific.
DISTRIBUTION. Eastern Gulf of Mexico west of Florida, (c. 21 {° N : 84° W) and Florida Strait,
also the northern and eastern coasts of South America from Venezuela to northern
Argentina (c. 39° S); 33-126 (7135) metres. The northernmost record from 85 miles west of
St. Petersburg, Florida, is thanks to Dr. K. Serafy and with seven Gerda stations in the Florida
Strait provides a considerable extension of the range from South America. About 35
specimens from 12 Pillsbury stations came from Venezuela to French Guiana and also
served to extend the bathymetric range both up and down, the three shallowest — 33, 36 and
42 metres — being from Surinam and French Guiana. There is also a specimen labelled as
being from Pillsbury st. 1384, which is in the Puerto Rico Trench (c. 19°45' N : 67W W) at
7956 metres! Although it has relatively long arms (R/r 65/8 mm = 8'l/l), about the
maximum found in scotti, the paxillar structure, form of the pedicellariae, general armament
and banded colour pattern leave no doubt of the identification, but the locality is evidently a
mistake.
Ludiapatriae Bernasconi
Luidia patriae Bernasconi, 1941 : 1 17-1 18; 1943 : 11-13, pi. 1, figs 1,2.
R 72 mm (holotype); R/r 7-2/1 .
A species of Luidia with normally 5 arms; abactinal paxillae with the two lateralmost
series forming transverse rows with the matching superomarginal paxillae and all rectangular
or squarish in shape: inferomarginal plates mainly ventral in alignment, bearing two slightly
flattened and slightly curved large spines near their upper ends at the ambitus, the lower one
larger, and on the ventral side about six pointed spines, cylindrical or somewhat flattened;
adambulacral plates with four large spines, the two abradial ones in line parallel to the
furrow: pedicellariae absent throughout. Colour (dried) dark rose above with darker areas in
the centre of the disc and on the median parts of the arms.
AFFINITIES. L. patriae is very similar to L. ludwigi scotti, of which Bernasconi had as many
as 80 specimens (as L. doello-juradoi), though only six or seven of L. patriae, which she
supposed to differ in the absence of pedicellariae (not now thought to be a character of
specific weight), the form, number and arrangement of the ventral inferomarginal spines and
in lacking the two or three enlarged spinelets abradial to the four main adambulacral spines
described for L. doello-juradoi. Possibly L. patriae will also prove to be a synonym of L.
ludwigi scotti.
DISTRIBUTION. Northern Argentina. 34±-37^° S [?also from Uruguay, 33°S]; 100-126 metres.
174 A.M.CLARK
Luidia sagamina aciculata Mortensen
Fig. Id, 2k, 31, s
?Luidia alternate, bicolor Verrill, 1 9 1 5 : 203 , pi. 1 2, fig. 1 .
Luidia sagamina: Downey, 1973 : 24.
Luidia aciculata Mortensen, 19336 : 425-426, fig 7, pi. 20, figs 7-12; Fisher, 1940 : 268-269, fig. M5;
Nataf & Cherbonnier, 1973 : 80-82, pi. 3, figs A, B, pi. 6, figs 1-6, pi. 9, figs G, H: Sibuet, 1975 : 284,
288.
Luidia sagamina var. aciculata: Madsen, 1950 : 199-203, figs 6, 7.
Luidia sagamina aciculata: A. M.Clark, 1955 : 33; A. M. Clark & Courtman-Stock, 1976 : 23,32,45.
Luidia atlantidea: Downey, 1973 : 25 [Non L. atlantidea Madsen, 1950.]
Rupto 140 mm; R/r 7-0-10-5/1.
A species of Luidia with normally 5 arms; abactinal paxillae all with an enlarged median
spinelet (sometimes replaced by a pedicellaria) both stouter and longer than the remaining
spinelets; two (or in the largest specimens three) longitudinal series of lateral paxillae also
matching transversely but outnumbering the much longer superomarginal paxillae by
16-19/10; inferomarginal plates mainly lateral in alignment, bearing two or three,
sometimes four large spines, tending to alternate in number and position on consecutive
plates, the uppermost spine the longest and projecting vertically upwards in its natural
position, at least on every second plate; adambulacral plates with two or three large spines in
a line at right angles to the furrow, when only two, the abradial one is relatively large;
pedicellariae usually present on at least the proximal actinal plates, where they are more or
less broadened basally and tapering so that the individual valves are triangular, though not
closely contiguous except terminally, abactinal pedicellariae bivalved and shorter, some-
times absent but one or more elongate bivalved pedicellariae always present on the furrow
face of each oral plate near the mouth, though not fully developed from spinelets in small
specimens. Colour in life purple to purple-brown above, white below, marginal spines dark
brown basally.
SYNONYMY. Madsen (1950) decided that Luidia sagamina Doderlein, 1920 from Japan and
L. aciculata Mortensen, 1933 from St Helena are conspecific, following comparison of
Mortensen's type material with a specimen from Sagami Bay and others collected off West
Africa by the Atlantide. Although he retained aciculata as a subspecies, he could not find any
real difference between it and sagamina. This prompted Downey (1973) to drop aciculata
and use L. sagamina for some american specimens, though conversely Nataf and
Cherbonnier in the same year retained aciculata at the specific level, without explanation.
Madsen had also referred to L. sagamina aciculata a specimen from Durban, Natal and in
1976 (Clark & Courtman-Stock) I retained this terminology, though without having seen any
specimens from either Japan or Natal. Since then, further echinoderm material from off
Natal (A. M. Clark, 1977) has shown that the fauna of SE Africa has much in common with
that of southern Japan and the East Indian area. This would imply that the Durban specimen
should be named L. sagamina sagamina if two subspecies are to be recognized, as seems
desirable to me in the absence of precedent from related species of moderate depths common
to similar areas of the Atlantic and Indo-West Pacific.
A further complication is raised by the possibility that Luidia sagamina aciculata is
conspecific with the (two?) specimens from the Florida Strait which Verrill (1915) called
Luidia alternata variety bicolor. VerrilFs drawings show relatively small single spines (or
enlarged spinelets) on all the abactinal paxillae, quite different from the sporadic and much
larger spines of L. alternata. Also the lateral paxillae outnumber the inferomarginal plates
(the superomarginals were not distinguished by the artist) and the inferomarginal spines are
regularly alternating on consecutive plates, as in L. sagamina and aciculata. Downey (1973)
recorded a small specimen (R in fact 2 1 mm not 8 mm as given in her paper) from South
Carolina as L. sagamina and mentioned another (p. 25) which she referred to L. atlantidea
because of light lateral bands along the arms but which also proves to be conspecific with the
first. A further specimen, R 24mm, from Gerda st. 1039 in the Florida Strait may be
NOTES ON ATLANTIC ASTEROIDEA 2. LUIDIIDAE 1 75
Halpern's supposed L. atlantidea, mentioned by Downey. Unfortunately the Bahama
Expedition's specimens of L. alternata var. bicolor cannot now be found at the University of
Iowa. Since the revival of bicolor as a specific name would mean that both L. sagamina
sagamina and aciculata became junior synonyms and there is an element of doubt about the
identity, it is desirable that the name bicolor should be formally set aside.
Verrill's specimens evidently had only two adambulacral spines, which is also the case in
the small Silver Bay and Gerda specimens. Mortensen's holotype of L. aciculata from St.
Helena, Fisher's Discovery specimen from off the Congo (Zaire), a Pillsbury specimen from
the Gulf of Guinea (Fig. 2k) and an Atlantide west african specimen all usually have three
adambulacral spines. Unfortunately, Madsen recorded four other Atlantide specimens with
only two such spines and this is also true of two other Pillsbury Guinea specimens. Clearly,
this character of adambulacral armament cannot be used to distinguish yet another
american, as opposed to african, subspecies, comparable to the subspecies of L. heterozona
and L. alternata, which would make for consistency of taxonomic treatment. The present
american specimens are too small and too few to suggest other possible differential
characters.
DISTRIBUTION. North Carolina to south and west Florida and from Cap Blanc, Mauritania
south to Zaire and from St. Helena, 20-975 metres. As noted above (p. 163), apart from
Sibuet's Walda Expedition record (1975), the greatest depth recorded is 1 50 metres.
Luidia sarsi Diiben & Koren
Figs Ic, 2c, g, 3m-o, v, 6
Asteriassp. aff. A. aranciaca: M. Sars, 1835 : 39.
Luidia fragilissima (pt, five-armed specimens) Forbes, 1841 : 135-140.
Luidia Sarsii Dttben & Koren, 1845:113; Perrier, 1875:342 [1876:262]; Sladen, 1889: 258;
Sussbach & Breckner, 1911:210.
Luydia Savignyi (pt) Diiben & Koren, 1 846 : 254, pi. 8, figs 23, 24.
Luidia elegans Perrier, 1875 : 336-337 [1876 : 256]; 1884 : 269[?], pi. 10, fig. 7 [and probably also fig.
8], captioned 'L. barbadensis'; Verrill, 1885 : 543, pi. 13, fig. 39; 1915 : 203-205, pi. 16, figs4,4a, pi.
19, fig. 1; Doderlein, 1920:289-290, fig. 36; Gray, Downey & Cerame-Vivas, 1968 : 140, fig 9;
Downey, 1973 : 25, pi. 3, figs C, D; Blake, 1973 : 40-41, pi. 10, figs 27-52; Carrera-Rodriguez &
Tommasi, 1977 : 71-75, figs 10-12.
Astrella simplex Perrier, 1882 : 21, fig. 25; 1894 : 193-194, pi. 14, fig. 3.
Luidia africana Sladen, 1889 : 256-258, pi. 44, figs 1, 2, pi. 45, figs 1, 2; Koehler, 1923 : 132; H. L.
Clark. 1923 : 252; 1925(pt) : 8; Mortensen, 1933<z : 239-240, figs 3, 4; Madsen, 1950(pt) : 188-192,
fig. 4, pi. 16, figs 3, 4; A. M. Clark, 1952 : 195; 1953 : 393-394, figs 10, 1 1; Nataf & Cherbonnier,
1973:79, pi. 8; A. M. Clark & Courtman-Stock, 1976:23, 30, 44, fig. 26. [Non L. africana:
Koehler, 191 1 : 19; Doderlein, 1920 : 228, nee Mortensen, 1925 : 178 = L. atlantidea.}
Luidia sarsi: Bell, 1893 : 72; Koehler, 1895 : 320, pi. 9. figs 6, 7; Ludwig, 1897 : 85-104, pi. 4, figs 2, 3,
pi. 7, figs 1-12; Koehler, 1909 : 59; Doderlein, 1920 : 288-289, fig. 35; Koehler, 1921 : 57, fig.;
Mortensen, 1925 : 178 [Isarsi ace. Madsen, 1950]; 1927 : 69, fig. 39b; Tattersall & Sheppard, 1934 :
36-55 (larvae); Madsen, 1950: 187; Tortonese, 1965: 150-152, fig. 69. [Non L. sarsi: Studer,
1 884, = L. africana ace. Doderlein, 1920 but much more likely = L. atlantidea.]
Luidia paucispina von Marenzeller, 1 893 : 3-4, pi. 1 , fig. 1 .
SIZE. The maximum size varies in different parts of the range. In NE Atlantic specimens (L.
sarsi sarsi) R reaches only c. 1 10 mm (BM coll., W of Scotland) but the same subspecies from
the Mediterranean, L. sarsi africana from South Africa and L. sarsi elegans from America
may reach 180-190 mm R; R/r 5'0-10'5/1, usually 7-5-10-0/1 at R>90 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.
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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^ -*-^ <U I— < ,._
6CO tJ -4— > 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><u W*j Jxi5 m2^'Z^ H ' o «G c/5
O. graci/w (Dana, 1852)
O. schmitti (Heron, 1977)
O. umbonata (Heron, 1977)
O. ancora Gordejeva, 1973
O. atlantica Shmeleva, 1967
O. exigua Farran, 1908
O. expressa Gordjeva, 1973
O. hispida (Heron, 1977)
O. mollicula Gordejeva, 1975
O. succurva (Heron, 1977)
O. alboranica Shmeleva, 1979
O. prendeli Shmeleva, 1966
O. shmelevi Gordejeva, 1972
O.tenellaSars, 1916
O. tregoubovi, Shmeleva, 1968
O. brodskiiShme\e\&, 1968
O. heronae sp. nov.
O. longpipes Shmeleva, 1968
O. memorata Gordejeva, 1973
O. minima Shmeleva, 1968
O. rotunda Heron, 1977
O. rotundata Boxshall, 1977
O. subtilis Giesbrecht, 1 892
O. vodjanitskii Shmeleva & Delalo, 1969
O. zernovi Shmeleva, 1966
O. ornata Giesbrecht, 1 89 1
O. infantula Gordejeva, 1972
O. africana Shmeleva, 1979
O. curvata Giesbrecht, 1902
O. englishi Heron, 1977
O. ivlevi Shmeleva, 1966
4-segmented in male, both with long third segment. Second antenna 3-segmented, terminal
segment bearing proximal group of 3 or 4 setae and a distal group of 6 to 8 setae. Mandible
bearing 2 blades and 2 or 3 setae. First maxilla bilobed, inner lobe with 3 elements. Second
maxilla with setose claw. Maxilliped 4-segmented in female and 3-segmented in male,
second segment in female bearing 2 setae on the internal surface. Terminal segment
produced as a long, curved claw. Legs 1-4 3-segmented, endopodites slender. Exopodites
armed with serrate spines *on external margin. Leg 5 reduced to small rod or knob-like
segment fused to urosome somite and bearing 1 or 2 apical setae. Gender feminine.
TYPE SPECIES: Oncaea vensusta Philippi, 1843. Sixty-eight species of Oncaea are now
known, over half of which have been described within the past 15 years (Shmeleva, 1966,
1967, 1968, 1969, 1979; Razouls, 1969; Gordejeva, 1972, 1973, 1975; Boxshall, 1977;
Heron, 1977; Kazatchenko and Andreev, 1977).
194 S. J. MALT
Acknowledgements
I wish to thank Dr H. S. J. Roe (I.O.S.) and Dr R. P. Harris (M.B.A.) for arranging for
material to be donated to the BM(NH), also Dr G. A. Boxshall and Dr R. J. Lincoln for
reading and commenting on the manuscript. This work was undertaken while the author was
in receipt of a BM(NH) postgraduate studentship.
References
Boxshall, G. A. 1977. The planktonic copepods of the northeastern Atlantic Ocean: some taxonomic
observations on the Oncaeidae (Cyclopoida). Bull. Br. Mus. nat. Hist. (Zool.) 31 (3) : 101-1 55.
Dana, J. D. 1 846. Notice of some genera of Cyclopacea. Am. J. Sci. ser. 2, 1 : 225-230.
1852. Crustacea. U.S. Explor. Exped. 13 : 1-1618.
Ferrari, F. D. 1975. Taxonomic notes of the genus Oncaea (Copepoda : Cyclopoida) from the Gulf of
Mexico and northern Caribbean Sea. Proc. biol. Soc. Wash. 88 (2 1 ): 2 1 7-232.
Giesbrecht, W. 1891. Elenco dei Copepodi pelagici raccolti dal tenente di vascello Gaetano Chierchia
durante il viaggio della R. Corvetta 'Vettor Pisani' negli anni 1882-1885, e dal tenente di vascello
Francesco Orsini nel Mar Rosso, nel 1 884. Alt. Acad. naz. Lincei Re. 7 (10) : 474^8 1 .
1892. Systemik und Faunistik des pelagischen Copepoden des Golfes von Neapel und der
angrenzenden Meeres-Abschnitte. Fauna Flora Golf. Neapel 19 : 1-83 1 .
1902. Copepoden. Result. Voyage S. Y. Belgica, Zool. 1-49.
Gordejeva, K. T. 1972. New species of Oncaea (Copepoda, Cyclopoida) from the tropical zone of the
Atlantic Ocean. Zool. Zh. 51 (7) : 963-968. (In Russian).
1973. New species of the genus Oncaea (Cyclopoida) from the tropical Atlantic. Zool. Zh.
52(10): 1572-1 576. (In Russian).
1975. Pelagic Cyclopoida (Copepoda) from the tropic Atlantic and South Seas. Zool. Zh.
54 (5): 776-779. (In Russian).
Hennig, W. 1966. Phylogenetic systematics. Univ. of Illinois Press, Urbana. 263 pp.
Heron, G. A. 1977. Twenty-six species of Oncaeidae (Copepoda : Cyclopoida) from the southwest
Pacific-Antarctic area. Biology of the Antarctic Seas. VI. Antarctic Res. Ser. Washington 26 : 37-96.
Kazatchenko, V. N. & Andreev, G. V. 1977. Parasitic copepods (Crustacea) collected during 57th
cruise of 'Vityaz' in the western tropical Pacific and seas of the Indo-Malayan archipelago. Trudy
Inst. Okeanol. 107 : 30^8. (In Russian).
Moulton, T. 1973. Principal component analysis of variation in form within Oncaea conifera
Giesbrecht 1 89 1 , a species of copepod (Crustacea). Syst. Zool. 22(2) : 141-156.
Philippi, A. 1843. Fernere Beobachtungen iiber die Copepoden des Mittelmeeres. Arch. Naturgesch.
9(1): 54-71.
Razouls, C. 1969. Description d'une espece nouvelle du genre Oncaea (Copepoda,, Cyclopoida). Vie
M///>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
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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.
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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
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By James E. Kekans
Hydroids and medusae of the family Campanulariidae recorded
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Miscellanea
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Miscellanea
Zoology series Vol 42 No 4 24 June 1982
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.*, 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*^<?ittl ir.AT/nM^"O^
New species of marine nematodes from Loch Ewe,
Scotland
H. M. Platt
Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Z. N. Zhang
Shandong College of Oceanography, Qingdao, Shandong Province, People's Republic of
China1
Introduction
During an investigation of the shallow sublittoral nematode fauna off a small sandy bay in
Scotland one of us (Z.N.Z.) encountered a number of previously undescribed species. These
were studied in detail at the British Museum (Natural History) and we here present
descriptions of six of the more interesting species: Gerlachius novusetosus, Catanema
macintyrei, Catanema smo, Ceramonema yunfengi, Acantholaimus ewensis and Rhips
paraornata. A new diagnosis is given for the subfamily Gerlachinae Andrassy, 1976. The
genus Robbea Gerlach, 1956 is synonymized with Catanema Cobb, 1920; a new generic
diagnosis and a key to the species is given. Keys are also provided for the genera
Ceramonema Cobb, 1920 and Acantholaimus Allgen, 1933. Several other minor
nomenclatorial changes are suggested. Species belonging to these five genera have not been
found previously in British waters and Rhips Cobb, 1920 has not previously been recorded
from Europe.
Material and methods
All specimens came from the same place; sublittoral sand in 3 m of water off 'north beach',
Firemore Bay, Loch Ewe, Wester Ross, Scotland. Detailed descriptions of the bay and its
fauna can be found in Steele & Baird (1968), Mclntyre & Eleftheriou (1968) and Mclntyre &
Murison (1973). The glycerine mounted specimens were studied using a Leitz Ortholux II
microscope equipped with differential interference contrast facility. All drawings were made
using a drawing tube. The formulae used for the dimensions are a modification of Filipjev's
(1918) formula and the de Man ratios, as described in Platt (1973). The abbreviations 'S' and
'V are the spicule length and the relative position of the vulva respectively. The
abbreviations 'a.b.d.' and 'c.d.' are the anal (or cloacal) body diameter and corresponding
(body) diameter. Type material has been deposited at the British Museum (Natural History).
Systematic descriptions
The classification follows that given in Gerlach & Riemann (1973).
Subfamily GERLACHINAE Andrassy, 1976
DIAGNOSIS. Meyliidae. Cuticle smooth or feintly striated. Amphids circular, lightly
cuticularised or invisible. Testes paired, opposed. Ovaries reflexed.
'also, D.A.F.S. Marine Laboratory, Victoria Road, Aberdeen AB9 8DB
Bull. Br. Mus. nat. Hist. (Zool.) 42 (4) : 227-246 Issued 24 June 1 982
228
H. M. PLATT & Z. N. ZHANG
DISCUSSION. Andrassy (1976) erected the genus Gerlachius to distinguish Meylia lissa
Gerlach, 1956 from the other two species of the genus, M. alata Gerlach, 1956 and
M. spinosa Gerlach, 1956. According to Andrassy, Gerlachius is characterized by its
smooth cuticle and invisible amphids: the specimens described below are similar to
G. lissus in many respects apart from these two main generic characters. However,
the cuticle in G. novusetosus sp. nov. was observed under interference contrast and
can otherwise be seen only as feintly striated so that this feature may have been over-
looked in G. lissus. Likewise, the amphid may have been overlooked, since in the
male G. novusetosus sp. nov. it was difficult to distinguish. Therefore, at this stage we
prefer simply to widen the subfamily and genus diagnosis to permit forms with a smooth
cuticle without amphids until these animals become better known.
The subfamily Gerlachinae can now be distinguished from Meyliinae since the ovaries are
outstretched in the latter (Lorenzen, 198 1) but reflexed in the former.
Fig. 1 Gerlachius novusetosus: (a) whole body of d; (b) head of rf; (c) head of 9; (d) copulatory
apparatus of rf; (e) 9 reproductive system. Bar scales: a,e = 40 um; b,c,d = 10 um.
NEW SPECIES OF MARINE NEMATODES 229
Gerlachius novusetosus sp. nov.
Fig. 1
MATERIAL STUDIED. Holotype: d1 BM(NH) 1981.4.10.
Allotype: 9 BM(NH) 1 98 1 .4. 1 1 .
DIMENSIONS.
Holotype rf: - 34 M 433 505 /zm;a = 23;b= 14-9; c = 7-0; S = 31 um
11 18 22 17
Allotype 9: - 34 280 463 515/zm;a = 19;b= 15-l;c = 9-9; V = 54%
12 22 27 17
DESCRIPTION. Short, relatively stout body. Cuticle feintly striated. Short 2'5-3-5//m
sublateral somatic setae present (Fig. la): they are not bilaterally symmetrical. Four 5*5 um
submedian cephalic setae (R3?) seated on short l-5 um peduncles. A short stout additional
seta, 2-2-5 um long, is associated with each cephalic seta positioned as shown in Fig. Ib, c.
Amphid round, not strongly cuticularised, 7 um long and 6*5 um wide in female (male
amphid less distinct but about 5-5 jum long). Posterior to the amphid there is a subcuticular
reticulate structure (Fig. Ic) of unknown function. Buccal cavity absent. Oesophagus short,
without a bulb. Tail conical, 3-4 a.b.d. Three conspicuous caudal glands lying entirely in the
tail.
Spicules curved and slightly cephalate proximally: chord length 23 um, arc length 3 1 um.
Gubernaculum has a dorso-caudally directed apophysis. Anterior to cloaca the cuticle
striations are thickened ventrally to produce a longitudinal row of contiguous pegs which
extends about 190 um from the cloaca (Fig la, d). Testes paired, opposed.
Vulva cuticularised and prominent. Ovaries paired, opposed, reflexed.
DIFFERENTIAL DIAGNOSIS. Gerlachius novusetosus sp. nov. differs from the only other
species, G. lissus (Gerlach, 1956) in having four short additional cephalic setae, shorter
cephalic setae (5*5 um vs 15 um), shorter oesophagus (b = 15 vs 7) and precloacal cuticular
differentiation.
DISCUSSION. The reticulate organ posterior to the amphid and the four additional pegs
alongside the cephalic setae have not been reported previously. However, this whole group
of what appear to be primitive desmoscolids are poorly known, the only other records being
G. lissus (Gerlach, 1956), Meylia alata Gerlach, 1956 and M. spinosa Gerlach, 1956; all
from Kiel Bay.
CATANEMA Cobb, 1920
Robbea Gerlach, 1956 syn. nov.
The genus Catanema was originally described by Cobb, 1920 to accommodate C. exile from
Jamaica, a male specimen with fine cuticle striations, narrow buccal cavity 'enclosed in a
swelling', seven pairs of subventral postcloacal tubular organs, proximally cephalate spicules
and a large dorso-caudally directed gubernacular apophysis. Three further species have
subsequently been assigned to the genus: C. cobbi Inglis, 1968, with ten pairs of subventral
postcloacal organs, cephalic cuticle 'modified into blocks' and a dorsally directed
gubernacular apophysis; C. porosum Hopper & Cefalu, 1973, with five pairs of subventral
postcloacal organs; C. gerlachi Hopper & Cefalu, 1973. This last species was a new name
for a specimen from the Maldive Islands originally described by Gerlach (1963#) under the
name Eubostrichus exilis (Cobb, 1920). However, as Hopper & Cefalu (1973) point out,
Gerlach (19630) makes no mention of the buccal bulb so characteristic of this genus and the
gubernaculum lacks the prominent apophysis, which in Gerlach's specimen is thin and lies
almost parallel to the spicules.
230 H. M.PLATT& Z.N.ZHANG
Hopper & Cefalu (1973) decided to keep this record within the genus Catanema.
However, we feel that it is more probable that Gerlach would not have overlooked a buccal
bulb if it were present (having already described nematodes with this character, e.g. Robbea
caelestis Gerlach, 1956) so we transfer this species back to Eubostrichus, becoming
Eubostrichus gerlachi (Hopper & Cephalu, 1973) comb. nov. Eubostrichus is characterized
as having modified 'ponds' on the tail (Hopper & Cefalu, 1973: porids = tubular setae serving
as outlets for glands): Gerlach (1963 p. 95) depicts similar structures in both E. parasitiferus
Chitwood, 1936 and his E. exists ( = E. gerlachi) from the Maldive Islands and both species
have similarly shaped guberaacula.
In 1956, Gerlach erected the genus Robbea for a male specimen from Brazil, considering
that the muscular buccal bulb distinguished the taxon from all other related genera.
Although the type, R. caelestis, is certainly depicted as having a very prominent buccal bulb,
some species described subsequently seem to have somewhat less prominent bulbs, depicted
as similar to those described for Catanema. This being so, we propose to synonomize
Robbea with Catanema so that Catanema now also includes the following species: C.
caelestis (Gerlach, 1956), C. gallica (Vitiello, 1974) and C. tenax (Gerlach, 19636). C.
gerlachi (Boucher, 1975) was described from a female only: since male characters are of
importance in this taxon, and despite the fact that C. gerlachi seems to be unique in the
length of its subcephalic setae, we prefer to treat it as a species inquirenda.
In proposing this synonomy, there remains two main points to discuss: the amphid of the
type and the presence of subventral tubular supplements on the male tail. In Cobb's (1920)
description of the type, C. exile, he describes the amphids as 'minute labial tubes'. However,
he also states that they were 'forward-pointing . . . difficult to see'. A similar anterior position
was found in C. smo sp. nov. described here and we feel confident that Cobb misinterpreted
what were spiral, albeit minute, amphids. Cobb (1920) also depicted the type with seven
prominent pairs of subventral tubular supplements on the tail, similar to the modified porids
found in Eubostrichus. We cannot be sure whether these supplements, found in C. exile, C.
cobbi and C. porosum are homologous with the stout caudal setae described in C. caelestis,
C. gallica, C. tenax and the two new species described here. However, their presence or
absence seems to be no more significant a reason for suggesting a generic split than say the
presence of huge cervical suckers in C. tenax or the reticulate head of C. cobbi. Therefore, we
consider the presence or absence of caudal supplements as a variable infrageneric character.
Finally, we would comment on the arrangement of setae on the head. Inglis (1968)
described 6 + 4 cephalic setae, the lateral components of the R2 circle being 'markedly dorsal'
in position (although, according to the direction of the amphid spiral, he mistook ventral for
dorsal). However, in most species, there are four prominent setae, often longer than the
others; we judge these to be the four R3 sensilla. The six R2 sensilla are small in size, often not
depicted but can be clearly seen in the two species described here. If so, the R, sensilla must
be reduced to papillae and may require SEM to resolve. In all species, each R3 sensillum
seems to be accompanied by a more medially situated subcephalic seta, which may be at
about the same level as the R3 sensilla as in C. tenax or more posterior as in C. caelestis. It is
these which Inglis (1968) called the four cephalic setae. In addition to these subcephalic
setae, there are others on the cephalic capsule which, according to the several specimens of
each species described here, seem to be quite stable in position. Most species, including C.
smo sp. nov., have setae located either side of the base of the amphid. In at least C. cobbi, C.
macintyrei sp. nov. and C. tenax there are additional setae in specific locations. For example,
in C. cobbi there is a pair of setae either side of the base of the amphid and an additional seta
associated with each ventral subcephalic seta. In C. macintyrei sp. nov. there is a seta close
behind each subcephalic seta, a small seta associated with each ventral R3 seta and a seta
lying dorsal to the amphid.
We propose the following generic diagnosis: Catanema. Desmodoridae. Stilbonematinae.
Elongated body. Cuticle finely striated posterior to cephalic capsule and in life covered with
coccoid blue-green algae. Cephalic capsule may appear reticulate or feintly punctated. Four
anteriorly directed R3 cephalic setae and four subcephalic setae. Additional subcephalic
NEW SPECIES OF MARINE NEMATODES
231
setae may be present on the cephalic capsule. Spiral amphid. Buccal cavity minute and
funnel-shaped leading to a muscular buccal bulb. Oesophagus narrow, terminating in a
rounded posterior bulb. Spicules curved and proximally cephalate. Gubernaculum with a
solid dorsally or dorso-caudally directed apophysis. Single outstretched testis. Subventral
pairs of tubular organs may be present on the tail. Tail conical.
TYPE SPECIES. Catanema exile Cobb, 1920
Key to the species of Catanema
1 Ventral suckers present in oesophageal and post-oesophageal region
C. tenax (Gerlach, 1963) comb. nov.
syn. Robbea tenax Gerlach, 1963
Ventral suckers absent 2
2 Cephalic cuticle in conspicuous blocks. 1 0 pairs subventral caudal supplements
C.coAA/Inglis, 1968
- Cephalic cuticle smooth or with feint punctations 3
3 Tail tip pointed, 'a' ratio > 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: <? BM(NH) 1 98 1 .4. 1 2.
Allotype: 9, BM(NH) 1981.4.13. Paratype 92 BM(NH) 1981.4.14.
DIMENSIONS.
Holotype cf : - 181 M 1328 1 560 //m; a = 78; b = 8-6; c = 1 1 -7; S = 24 ^m
16 18 20 18
Allotype 9,: -125. M ,791 927/rni;a = 54;b = 7-4;c = 6-8; V = 44%
15 17 17-5. 12
92: - 141 M ' 874 1020/nn; a = 58;b = 7-2;c = 7-0; V = 46%
16 17-5 17-5 12-5
DESCRIPTION. Body colourless, elongated and conspicuously annulated. Cuticle annules in
male 5'5//m, 7/^m and 5 /*m long in the anterior oesophagus, posterior oesophagus and
cloacal regions respectively. Total number of body annules 277 in cf, 186 in 9, and 184 in 92.
Each annule divided into eight plates, so that there are eight longitudinal crests running the
Fig. 5 Ceramonema yunfengi: (a) whole body of cf,; (b) cf, head; (c) cf, tail; (d) 9, head. Bar scales:
NEW SPECIES OF MARINE NEMATODES 237
length of the body. Vacuoles are present beneath the cuticle which are not confined to the
longitudinal crests. In the male, but not the female, the annule in the cloacal region is
elongated (about 12//m) relative to the rest (Fig. 5c). Somatic setae absent except for two
caudal setae situated on the distal cone of the male only. Cephalic capsule elongated: <5
33-5 x 1 6'5 //m; 9, 29 x 1 5 /zm; 92 3 1 -5 x 1 6 //m. Six slim R2 cephalic setae; 1 2-5 //m long in cf
and 10-10'5 //m long in 99. Four slim R3 cephalic setae situated just posterior to R2 setae;
13'5 jum long in d, 1 1 //m long in 99. In the d, two medial holes in the cephalic capsule were
observed just posterior to the level of the R3 sensilla (Fig. 5b): possibly positions of sub-
cuticular sensilla. Amphids an elongated loop, 17-1 9 /zm long. Buccal cavity absent.
Oesophagus narrow but widens slightly at the posterior end. Nerve ring at 52-54% of
oesophagus length. Excretory ampullae situated anterior and posterior to oesophageal-
intestinal junction. Tail elongated, 7 a.b.d. in d, 1 1-12 a.b.d. in 99. In the d, the terminal
cone is 14- 5 //m long and bears two lateral 8 //m setae.
Spicules paired, slightly curved and 24 //m long (chord). Gubernaculum almost straight,
1 5 um long. No supplements. Testes not clear.
Ovaries paired, opposed and reflexed.
DIFFERENTIAL DIAGNOSIS. Ceramonema yunfengi sp. nov. can be distinguished from all
other known species of the genus by the presence of two lateral sub-terminal caudal setae in
the d and a relatively longer cephalic capsule. The only other species described which have
an elongated cloacal annule are C. carinatum Wieser, 1959 and possibly, according to the
figure, C. pisanum Gerlach, 1952. However, both these species have shorter cephalic setae
and are not described as having subcuticular vacuolisation. In addition, the tail of C.
pisanum is longer than that of C. yunfengi (1 1 a.b.d. vs. 7 a.b.d.).
DISCUSSION. Haspeslagh (1973) recently revised the superfamily Ceramonematoidea and, on
the basis of the presence or absence of vacuolisation in the cuticle, divided Ceramonema into
three genera. However, Lorenzen (1981) did not accept this argument and synonomized
Haspeslagh's new genera, Ceramonemoides and Cyttaronema, with Ceramonema.
Currently, there are eleven species contained in the genus including C. yunfengi, which can
be distinguished using the data contained in Table 1 and Fig. 6.
Apart from C. pisanum Gerlach, 1952, which is described from a male and a juvenile
female, C. yunfengi is the only species where both male and adult female characters are
known. Four species are known from females only and one from a juvenile only. Many of the
important characters used to distinguish marine nematodes in general are vested in the male,
so current practice is to consider species described from females or juveniles only as dubious.
However, it is still possible to separate the eleven species contained in Table 1 on the
combination of the relative length of the cephalic capsule, presence or absence of
vacuolisation and its pattern of distribution, relative position of R2 and R3 sensilla and
cuticle structure. However, as more species become known, it may become impossible to
separate the taxa on these characters alone so that C. attenuatum, C. chitwoodi, C. rectum, C.
sculpturatum and C. undulatum may have to be designated species dubiae, particularly as
certain characters display sexual dimorphism including length of R2 and R3 sensilla and
number of body annules, at least in C. yunfengi. The apparent sexual dimorphism in the
amphid reported for C. pisanum (Fig. 6e, 0 may be simply due to the female not being adult:
the amphids are similar in C. yunfengi.
Acantholaimus ewensis sp. nov.
Figs 7-8
MATERIAL STUDIED. Holotype: d, BM(NH) 1 98 1 .4. 1 5.
Allotype: 9, BM(NH) 1981.4.16. Paratypes: d2 BM(NH) 1981.4.17; d3 BM(NH) 1981.4.18;
d4, d5 in collection of Z.N.Z.
238
H. M. PLATT & Z. N. ZHANG
c c c c
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00
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^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
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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 <J (here designated) Schouten
Island (FS, Frankfurt am Main) [examined]. Strand, 1915:262. Roewer, 1954:936. Bonnet,
1956 : 1 172. Proszynski, 1971 : 390. Syn. n.
Cocalodes plebejus Szombathy, 1915:468, cf, 9, immatures. Syntypes (presumably in TM,
Budapest) [not examined]. Roewer, 1954 : 936. Bonnet, 1956 : 1 173. Syn. n.
REMARKS. Szombathy (1915) supplied good figures of the palpal organs in his original
description of C. plebejus and there is little doubt that plebejus and papuanus are
conspecific.
268 F. R. WANLESS
DIAGNOSIS. Separated from other species of Cocalodes by the cluster of cheliceral
setae and truncate median apophysis (Fig. 2C, F) in males, and by the large rounded
epigynal openings in females (Fig. 3D).
MALE from Madang, Papua New Guinea. In good condition. Carapace (Fig. 2A, B):
pale yellow-brown grading to orange-brown in eye region with dark brown bands
around margins and from PL's to posterior margin of thorax; pale areas clothed in
creamy white hairs with orange-brown or black hairs elsewhere. Eyes: with black
surrounds except AM; fringed in creamy white and pale amber hairs. Clypeus:
clothed in very fine whitish hairs. Chelicerae: elongate, porrect and diverging; orange-brown
with dorsal cluster of stout spines; shiny under some angles of illumination; promargin with
four teeth, retromargin with two; cheliceral horn small, not protruding beyond clypeal
margin. Maxillae and labium: pale yellow grading to whitish yellow along inner margins of
maxillae. Sternum: more or less as in female; pale yellow, shiny. Coxae: pale yellow to
whitish yellow with black lateral bands on coxae I. Abdomen: pale yellow with sooty
markings and blackish lateral bands above, greyish black below; clothed in fine iridescent
setae; spinnerets pale yellow heavily tinged grey-black except for whitish medians. Legs: legs
I pale yellow to amber with blackish lateral markings on femora and metatarsi; legs II
similar, but lighter with less conspicuous dark markings; legs III pale yellow grading to light
amber distally; legs IV as III, but with black lateral markings on patellae, tibiae and
metatarsi; spines numerous, strongest on legs I. Spination of legs I: metatarsi v 2-2-2,
p 1-0-1, r 1-0-1; tibiae v 2-2-2, p 1-1-1, r 1-1-1, d 1-0-1; patellae 1-0-0, r 1-0-0;
femora p 1-1-1, d 1-1-3, r 1-2-1. Palp (Figs 2C, E, F, H; 18C, E): the truncate dorsal
prong of the median apophysis and pronounced functional conductor are characteristic of
this species.
Dimensions (mm): total length 7*88; carapace length 3*02, breadth 2-22, height
1-44; abdomen length 4-84; eyes, anterior row 1-96, middle row 1*68, posterior row
1-68; quadrangle length 1-6 (52% of carapace length). Ratios: AM : AL :
PM: PL:: 16:9:6:9; AL-PM-PL :: 10-13; AM : CL (clypeus) :: 16 : 3 (18% of AM
diameter).
FEMALE (formerly undescribed) from Madang, Papua New Guinea. In good condition.
Carapace (Fig. 3A): pale yellow-brown, shiny; clothed in fine whitish hairs with
bands composed of amber hairs from posterior lateral eyes to posterior thoracic
margin. Eyes: as in cf. Clypeus: below anterior median eyes and inner sides of anterior
laterals fringed in pale amber hairs with outer sides of anterior laterals densely white
haired. Chelicerae: robust, porrect and diverging; yellow-brown, shiny; thinly clothed in
yellow-brown hairs; both margins with three teeth. Maxillae and labium: pale yellow, shiny.
Sternum (Fig. 3B): pale yellow, shiny. Coxae: pale yellow, shiny. Abdomen: whitish yellow;
irregularly clothed (?partly rubbed) in very fine iridescent setae with longitudinal bands,
composed of dark amber hairs, flanked externally by white haired bands on the sides;
spinnerets pale yellow tinged grey. Legs: pale yellow-brown with blackish lateral streaks on
metatarsi IV. Spination of legs I: metatarsi v 4-0-2, p 1-0-1 , r 1-0-1 ; tibiae v 2-3-1 , p 1-1-1 ,
r 1-1-1, d 1-0-1; patellae p 1-0-0, r 1-0-0; femora p 0-2-1, d 1-1-2, r \-2-\.EpigyneQr\&
3C-E; 19 A, B): clothed in testaceous hairs.
Dimensions (mm): total length 10*8; carapace length 3-76, breadth 2-8, height 1-64;
abdomen length 6*8; eyes, anterior row 2'26, middle row 1'96, posterior row 1'96;
quadrangle length 1-8 (47% of carapace length). Ratios: AM : AL :
PM : PL :: 19 : 10 : 5-5 : 10; AL-PM-PL :: 1 1-16; AM : CL :: 19 : 5 (26% of AM diameter).
VARIATION, rf total length varies from 4-88 to 10*4 mm, carapace length 2-6-3-96 mm
(seven specimens). 9 total length 8-48-10-8 mm, carapace length 2-92-3-76 mm (three
specimens).
In males the number of peg-like spines on the cymbium varies from two to four. In
one specimen (lectotype of C. papuanus) the left palp has two spines on the cymbium while
SPIDER GENUS COCALODES
269
the right has three. In females the epigyne varies slightly, the dorsal rims of the rounded
openings sometimes lie across the posterior margins of the spermathecae.
DISTRIBUTION. Papua New Guinea; Irian Jaya.
MATERIAL EXAMINED. Papua New Guinea: Madang Province, Madang, 22.iii.1979 (H.
W. Levi, Y. Lubin, B. Robinson) (MCZ, Harvard): 19, garden, night collection; Icf,
40 Km S. of Madang, swamp forest. Astrolabe Bay: (R. Rohde} (MNHU, Berlin.
17795). Irian Jaya: Dorey, Lectotype rf, (A. Raffray) (MNHN, Paris. 5479); Schouten Island,
H
Fig. 2 Cocalodes papuanus Simon, cf; A, dorsal; B, lateral; C, palp, retrolateral; D,
cleared fang; E, palp, prolateral; F, median apophysis and tip of functional conductor;
G, cheliceral teeth inner view; H, palp, ventral.
270
F. R. WANLESS
Fig. 3 Cocalodes papuanus Simon. 9: A, dorsal; B, sternum; C, vulva ventral; D, epigyne; E,
vulva, dorsal.
Woges, [lectotype rf of C armatissimus], 1909, (E. Wolf) (FS, Frankfurt am Main. 2431).
Sukarnapura (Hollandia) AMNH, New York): Irf, v. 1945 (Borys Malkin); 19, rainforest,
250 ft. xii. 1944 (H. Hoogstraal); Irf, on foliage, rainforest, 300 ft. xii.1944 (L. W. Saylof).
Sukarnapura: 2rf, 1936 (L. E. Cheeseman) (BMNH. 1937.12.13.117); 19, 1936 (L. E.
Cheeseman)(]BMNH. 1937.12.13.161).
Cocalodes leptopus Pocock
(Figs4A-H;19C,D)
Cocalodes leptopus Pocock, 1897:628, 9. Holotype 9, Indonesia (BMNH) [examined]. Simon,
1901 : 403^05. Petrunkevitch, 1928: 181. Roewer, 1954:936. Bonnet, 1956: 1172. Proszynski,
1971 :390.
Cocalodes melanognathus Pocock, 1897:629, rf. Holotype rf, Indonesia (BMNH) [examined].
Simon, 1901 : 403^04. Roewer, 1954 : 936. Bonnet, 1956 : 1 172. Proszynski, 1971 : 390. Syn. n.
REMARKS. As C. leptopus and C. melanognathus both originate from Halmahera and
are only known from separate sexes, they are regarded here as being conspecific.
DIAGNOSIS. From females of C. cignatus and C. expers by the apparent absence of a
notch or notches on the posterior margin of the epigynal plate (Fig. 4E). From male
cygnatus by the lack of a white moustache below the anterior median eyes and by the
curved tip of the functional conductor (Fig. 4H). Males of expers are unknown.
SPIDER GENUS COCALODES 27 1
FEMALE HOLOTYPE. In poor condition. Carapace: brown-black with orange-brown eye
region; irregularly clothed in whitish hairs, especially dense below lateral eyes. Eyes:
with black surrounds; irregularly fringed in whitish hairs. Clypeus: densely white
haired. Chelicerae: robust, inclined anteriorly and slightly diverging; brown with
lighter brown markings; pro- and retromargins with three teeth. Maxillae (Fig. 4D):
brownish orange grading to yellow-brown along inner margins. Labium: brownish
orange tipped yellow-brown. Sternum: greenish yellow with thin brownish margins;
shiny. Coxae: dark grey tinged greenish yellow. Abdomen: greenish yellow with
brownish lateral markings and two pairs of impressed spots dorsally; clothed in
whitish hairs (mostly rubbed on dorsum) with irregular longitudinal red haired bands
on each side; spinnerets greenish yellow tinged black. Legs: brownish orange tinged
greyish green, femora IV and patellae IV with black spots; spines strong and numerous.
Spination of legs I: metatarsi v 2-0-0, p 1-1-1, d 0-0-2, r 1-1-1; tibiae v 4-4-4; patellae p
1-0-0, r 1-0-0; femora p 1-0-2, d 0-2-2, r 0-1-1 . Epigyne (Figs 4E-G; 19C, D): a low dark
mound with indistinct lateral openings (arrowed in Fig. 4F).
Dimensions (mm): total length lO'O; carapace length 3'56, breadth 3'0, height 2*24;
abdomen length 6'56; eyes, anterior row 2'52, middle row 2*12, posterior row 2*16;
quadrangle length 1'92 (53% of carapace length). Ratios: AM : AL :
PM:PL:: 19: 11 :7: 10; AL-PM-PL :: 13-15; AM : CL :: 19 : 9 (47% of AM diameter).
MALE (holotype of C. melanognathus). In poor condition. Carapace (Fig. 4A, B):
dark chocolate brown with orange-brown eye region; badly rubbed. Eyes: with black
surrounds except AL and AM. Clypeus: irregularly and scantily clothed in fine
whitish hairs. Chelicerae: elongate, porrect and diverging; dark brown with bluish
sheen under some angles of illumination; pro- and retromargins with three teeth;
horn small, dislocated by a pin pushed through the specimen, a method frequently
used by Pocock for orientating larger spiders. Maxillae, labium, sternum and coxae:
more or less as in 9. Abdomen: similar to 9. Legs: broken and detached, otherwise
similar to 9. Palp (Fig. 4C, H): similar to that of C. cygnatus, but readily distinguished
by the curved tip of the functional conductor.
Dimensions (mm): total length 8'8; carapace length 3*4, breadth 2'72, height 2'08;
abdomen length 5'36; eyes, anterior row 2*48, middle row 2*08, posterior row 2*16;
quadrangle length l-94 (57% of carapace length). Ratios: AM : AL :
PM:PL:: 19-5: 11-5 ::7: 11; AL-PM-PL:: 12-15; AM : CL ::19'5 : 5 (28% of AM
diameter).
DISTRIBUTION. Indonesia: Halmahera.
MATERIAL EXAMINED. Halmahera: Patani, holotype 9, ii.1894 (Kukenthal); Soah Konorah,
holotype rf [of C. melanognathus] 1 894 (Kukenthal) (BMNH. 1981.1.22.1-2.)
Cocalodes expers sp. n.
(Fig. 5A-C)
DIAGNOSIS. From C. leptopus and C. cygnatus by the presence of a deep median notch
on the posterior margin of the epigynal plate (Fig. 5C).
MALE. Unknown.
FEMALE HOLOTYPE. In good condition. Carapace (Fig. 5A, B): orange-brown with
vague darker markings; irregularly clothed in short, fine shinning hairs which appear
whitish or pale amber under varying angles of illumination. Eyes: with black surrounds
except AM; fringed in whitish and pale amber hairs. Clypeus: densely clothed in creamy
white hairs. Chelicerae: robust, divergent and inclined anteriorly; orange-brown, thinly
clothed in clear amber hairs; pro- and retromargins with three teeth. Maxillae and labium:
amber, shiny. Sternum: light amber with darker margins; thinly covered in light brownish
272
F. R. WANLESS
H
Fig. 4 Cocalodes leptopus Pocock. 3 (holotype of C. melanognathus): A, dorsal; B,
lateral; C, palp, retrolateral; H, palp, ventral. Holotype 9: D, sternum, coxae and mouth
parts; E, epigyne; F, vulva, ventral; G, vulva, dorsal.
hairs. Coxae: light amber. Abdomen: pale orange-brown with fine shinning hairs, scattered
spots composed of amber hairs, and dark reddish brown lateral bands covered in dark amber
hairs; spinnerets pale orange-brown. Legs: orange-brown with blackish apices on femora IV;
spines numerous. Spination of legs I: metatarsi v 2-2-2, p 1-0-1, r 1-0-1; tibiae v 2-2-2,
p 1-1-1, d 0-1-0, r 1-1-1; patellae p 0-1-0, r 0-1-0; femora p 2-2-1, d 1-1-3, r 2-1-1.
Epigyne (Fig. 5C): similar to that of C. leptopus, but readily separated by the median notch
on the posterior margin of the epigynal plate.
SPIDER GENUS COCALODES
273
..
A /7BT C
Fig. 5 Cocalodes expers sp. n. Holotype 9: A, dorsal; B, carapace lateral; C, epigyne.
Dimensions (mm): total length 10'9; carapace length 4-5, breadth 3*8, height 2*8;
abdomen length 6*24; Eyes, anterior row 3*04, middle row 2-68, posterior row 2*68;
quadrangle length 2*28 (50% of carapace length). Ratios: AM : AL :
PM : PL :: 22 : 14 : 8 :14; AL-PM-PL :: 14 : 19; AM : CL :: 22 : 8 (36% of AM diameter).
DISTRIBUTION. Papua New Guinea.
MATERIAL EXAMINED. Papua New Guinea, D'Entrecasteaux Is., Fergusson Island, lamelele
about 1*5 miles from Seymour Bay, 15m., camp 3. holotype 9, 1956 (Fifth Archbold Exp. to
New Guinea, L. J. Brass) (AMNH, New York).
REMARKS. This species has been recorded from the same locality (Fergusson Island)
as C. longicornis sp. n.
Cocalodes cy gnat us sp. n.
(Figs6A-G;19E,F)
DIAGNOSIS. From females of C. leptopus and C. expers by the presence of two small
notches on the posterior margin of the epigynal plate (Fig. 6D). From male leptopus
by the white moustache below the anterior median eyes and by the cygniform tip of
the functional conductor (arrowed, Fig. 6G). Males of expers are unknown.
MALE HOLOTYPE. In fair condition. Carapace (Fig. 6A, B): orange-brown with dark
274
F. R. WANLESS
Fig. 6 Cocalodes cygnatus sp. n. Holotype rf: A, dorsal; B, carapace, lateral; C, palp,
retrolateral; G, palp, ventral. Paratype 9: D, epigyne; E, vulva, ventral; F, vulva, dorsal.
brown margins and vague bands behind PL's; thinly clothed in whitish and pale
amber hairs with scanty white haired patches on posterior declivity. Eyes: with black
surrounds except AM; fringed by white hairs. Clypeus: densely white haired below
AM. Chelicerae: elongate, porrect and diverging; dark reddish brown; shiny; thinly
covered in pale amber hairs; pro- and retromargins with three teeth; horn small, not
protruding beyond clypeal margin. Maxillae: brownish with inner margins orange-
brown. Labium: dark brown tipped orange-brown. Sternum: yellow-brown with darker
margins. Coxae: yellow-brown with black lateral stripes on coxae I. Abdomen: mottled
grey-black with creamy white dorsal markings; spinnerets yellow-brown tinged black. Legs:
legs I femora brown-black with yellow-brown markings, patellae and tarsi yellow-brown,
tibiae dark brown with yellow-brown annuli, metatarsi basally yellow-brown grading to dark
brown distally; other legs similar, but markings becoming paler except for brown-black
blotches on femora IV; spines strong and numerous. Spination of legs I: metatarsi v 2-0-0,
p 1-1-1, d 0-1-2, r 1-1-1; tibiae v 2-2-2, p 1-1-1, d 1-1-0, r 1-1-1; patellae p 1-0-0,
SPIDER GENUS COCALODES 275
r 1-0-0; femora p 1-1-2, d 0-2-2, r 0-1-1. Palp (Fig. 6C, G): the tip of the functional
conductor is sometimes obscured by the tip of the median apophysis. Also, in lateral view
the ventral prong of the median apophysis is hardly evident, c.f. (C. leptopus).
Dimensions (mm): total length c. 8-2; carapace length 3' 16, breadth 2-6, height 2-0;
abdomen length 5'0; eyes, anterior row 2-36, middle row 1'98, posterior row 2-04;
quadrangle length 1'84 (58% of carapace length). Ratios: AM:AL:
PM:PL:: 19: 11 : 5-5 : 10; AL-PM-PL :: 1 1-15; AM : CL :: 19 : 8 (42% of AM diameter).
FEMALE PARATYPE. In poor condition. Carapace: generally yellow-brown with blackish
bands from PL's to thoracic margin; sparsely and irregularly clothed in creamy white hairs.
Eyes: with black surrounds except AM; fringed in silky white hairs. Chelicerae: robust,
diverging and inclined anteriorly; amber; shiny under some angles of illumination; with
scattered pale amber hairs along inner margins; pro- and retromargins with three teeth.
Maxillae, labium, sternum and coxae: yellow-brown. Abdomen: pale yellow-brown; clothed
in fine whitish hairs with irregular lateral bands composed of amber hairs; spinnerets pale
yellow-brown. Legs: generally yellow-brown, but with black spots on patellae IV and femora
IV; spines strong and numerous on anterior legs becoming weaker and fewer on posteriors.
Spination of legs I: metatarsi v 2-0-0, p 1-1-1, d 0-1-0, r 1-1-1; tibiae v 2-2-2, p 1-1-1,
d 1-0-1, r 1-1-1; patellae r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-1-1. Epigyne (Figs
6D-F;19E, F): a low mound as is leptopus, but pale and with more apparent detail.
Dimensions (mm): total length c. 8'0; carapace length 3' 16, breadth 2-68, height
2*0; abdomen length 4-8; eyes, anterior row 2*44, middle row 2'07, posterior row
2-10; quadrangle length 1-88 (59% of AM diameter). Ratios: AM:AL:
PM: PL:: 19: 11-5:6: 11; AL-PM-PL :: 12 : 15; AM:CL::19:c. 8 (c. 42% of AM
diameter).
VARIATION. Paratype cf measures c. 7-2 mm total length, 2-8 mm carapace length.
DISTRIBUTION. Indonesia: Halmahera Island.
MATERIAL EXAMINED. Halmahera Island: Edkor, holotype rf, paratype $ (MNHN,
Paris. 7682). Paratype cf, same data as holotype (BMNH, 198 1 .5. 14.1).
REMARKS. Simon misidentified the above specimens as C. leptopus, but as far as I am
aware there has been no reference to them in the literature.
Cocalodes macellus (Thorell)
(Figs 7A-I; 20A, B)
Cocalus macellus Thorell, 1878:287, 311, 9 and juvenile. Holotype $, juvenile, Amboina
(MCSN, Geneva) [examined]. Thorell, 1881 : 493, 706. Simon, 1901 : 407. Roewer, 1954:934.
Bonnet, 1956: 1 173. Proszyriski, 1971 : 391. Wanless, 1981 : 256.
DIAGNOSIS. From other species of Cocalodes by the long slender prongs of the median
apophysis (Fig. 7H, I) in males, and the appearance of the epigyne (Figs 7C-F;
20A, B) in females.
HOLOTYPE FEMALE. In fair condition. Carapace: yellow-brown with pale yellowish
eye region. Eyes: with black surrounds except AM; fringed in whitish hairs. Clypeus:
densely white haired. Chelicerae: yellow-brown thinly clothed in testaceous hairs;
pro- and retromargins with three teeth. Maxillae: yellow-brown with pale yellow
blades. Labium: yellow-brown tipped yellow. Sternum: yellow-brown. Abdomen: long and
tapered; pale yellow. Legs: generally yellow-brown; spines strong and numerous on legs I
becoming weaker and fewer on posterior legs. Spination of legs I: metatarsi v 2-2-2, p 1-0-0,
d 0-2-2, r 1-0-0; tibiae v 2-4-0, p 1-1-2, d 1-1-0, r 1-1-2; patellae p 1-0-0, r 1-0-0; femora
p 1-0-1 , d 1-1-3, r 0-1-0. Epigyne (Fig. 7D): somewhat translucent; the ducts will probably
be less evident in freshly preserved specimens.
276
F. R. WANLESS
Dimensions (mm): total length c. 7'8; carapace length 2*88, breadth 2'34, height
1'72; abdomen length 4'88; eyes, anterior row 2'2, middle row l-92, posterior row
2-0; quadrangle length 1*64 (56% of carapace length). Ratios: AM : AL :
PM : PL :: 17 : 1 1 : 6 : 10; AL-PM-PL :: 9-12; AM : CL :: 17 : 3-5 (20% of AM diameter).
MALE (formerly undescribed) from Draeger Harbor, Papua New Guinea. In fair
condition. Carapace (Fig. 7A, B): dark mahogany grading to orange-brown in eye
region; irregularly clothed in short amber hairs (?rubbed) with white haired marginal
band from level of coxae II to IV. Eyes: with black surrounds except AM; fringed by
shining pale amber, and white hairs. Clypeus: fringed in light brownish hairs. Chelicerae:
moderately elongate and porrect, more or less parallel; dark orange-brown, lighter distally;
thinly clothed in fine light brownish hairs; teeth not examined; horn not evident. Maxillae:
dark orange-brown with yellow-brown blades. Labium: dark orange-brown. Sternum:
yellow-brown with amber margins, shiny; sparsely clothed in fine clear hairs. Coxae: coxae I
yellow-brown below, dark mahogany above; other coxae yellow-brown. Abdomen: badly
H
j^r
Fig. 7 Cocalodes macellus (Thorell). d: A, dorsal; B, carapace, lateral; G, palp, ventral;
H, median apophysis; I, palp, retrolateral. Holotype 9: D, epigyne. C, epigyne of
another specimen; E, vulva, ventral; F, vulva, dorsal.
SPIDER GENUS COCALODES 277
rubbed; yellow-brown with a dark patch anteriorly and ventral grey longitudinal band. Legs:
legs I with tarsi and patellae yellow-brown, metatarsi yellow-brown suffused with black
distally, tibiae dark mahogany with apices yellow-brown, femora dark mahogany; legs II-III
yellow-brown; legs IV missing; spines strong and numerous. Spination of legs I:
metatarsi v 2-2-2, p 1-0-0, d 1-2-2, r 1-0-0; tibiae v 3-2-4, p 0-1-1, d 1-1-0,
r 1-1-1; patellae p 1-0-0, r 1^0-0; femora p 0-1-1, d 0-2-2. Palp (Fig. 7G-I): femora
brown-black with distal third light yellow, other segments light yellow; both prongs of
median apophysis relatively long and slender.
Dimensions (mm): total length 5'68; carapace length 2*32, breadth 1'92, height
1-48; abdomen length 3'2; eyes, anterior row 1-85, middle row 1'56, posterior row
1-72; quadrangle length 1-4 (60% of carapace length). Ratios: AM : AL :
PM : PL :: 15 : 9 : 4 : 8-5; AL-PM-PL :: 9*5-10; AM : CL :: 15 : 4-5 (30% of AM diameter).
VARIATION. 9 from Papua New Guinea measures c. 6*9 mm total length, 2-76 mm
carapace length.
The epigyne (Fig. 7C) is slightly less translucent and the looped ducts are less
conspicuous.
DISTRIBUTION. Indonesia: Amboina; Papua New Guinea.
MATERIAL EXAMINED. Amboina: holotype 9, juvenile, (O. Beccari) (MCSN, Geneva). Papua
New Guinea: Draeger Harbour, 5 miles E. of Finschhafen, 19, vi.1944 (G. H. Penn) (AMNH,
New York); Nr. River Song, 6 miles W. of Finschhafen, Icf, viii-ix. 1944, (T. F. Delaney)
(AMNH, New York).
Coca I odes longicornis sp. n.
(Figs8A-H;21A)
DIAGNOSIS. From males of C. longipes and C. innotabilis by the absence of peg-like
cymbial spines, and from male C. platnicki by the thickened dorsal prong of the
median apophysis (Fig. 8B, C). Females, presumed to be of this species, most closely
resemble females of C. macellus, but may be separated by the lack of a dense covering
of white hairs on the clypeus and by the laterally extended introductory ducts (Figs
8F, G, H; 2 1 A) of the epigynum.
MALE HOLOTYPE. In fair condition. Carapace (Fig. 8A, D): dark rufose with eye
region dark amber; sparsely clothed in short fine amber hairs with whitish hairs in
foveal region and a narrow marginal band of white hairs extending from level of coxae
II to IV. Eyes: with black surrounds except AM; fringed by shining amber hairs with
white hairs outside AL's. Clypeus: densely white haired below AM, sparsely amber
haired below AL. Chelicerae: elongate, porrect and diverging; rufose; shiny; pro- and
retro-margins evidently with three teeth; horn well developed and protruding well
beyond clypeal margin. Maxillae: orange-brown grading to yellow-brown along inner
margins. Labium: orange-brown. Sternum: orange-brown with darker margins; sparsely
clothed in coarse black hairs marginally with fine pale brown hairs centrally. Coxae:
orange-brown. Abdomen: dull orange-brown with blackish lateral bands above and central
blackish band below; spinnerets orange-brown. Legs: legs I-II dark rufose with orange-
brown tarsi, other legs rufose grading to orange-brown distally; spines strong and numerous.
Spination of legs I: metatarsi v 2-2-2, p 1-0-1, d 0-1-0, r 1-0-1; tibiae v 2-2-2, p 1-1-1,
d 1-1-0, r 1-1-1; patellae p 1-0-0, r 1-0-0; femora r 1-1-1, d 0-2-3, p 1-1-1. Palp (Fig.
8B, C, E).
Dimensions (mm): total length 10' 16; carapace length 4*28, breadth 3'5, height
2-64; abdomen length 5'84; eyes, anterior row 2*84, middle row 2*48, posterior row
2-5; quadrangle length 2-12 (49% of carapace length). Ratios: AM : AL :
PM :PL:: 21 : 13-5:8: 13; AL-PM-PL :: 12-18; AM : CL :: 21 : 7-5 (35% of AM
diameter).
278
F. R. WANLESS
hEMALE PARATYPE. In fair condition. Carapace: orange-brown with reddish brown
eye region and dark red-brown markings from PL's to thoracic margin; clothed in fine
shining hairs (mostly rubbed). Eyes: with black surrounds except AM; fringed by
whitish and pale amber hairs. Clypeus: fringed in whitish hairs. Chelicerae: robust,
inclined anteriorly and slightly diverging; reddish with darker markings; shiny; sparsely
fringed in pale amber hairs; pro- and retromargins with three teeth. Maxillae and labium:
amber. Sternum: amber with darker margins; shiny; thinly clothed in fine light amber hairs.
Abdomen: long and tapered; dull yellow-brown with blackish lateral markings; spinnerets
yellow-brown. Legs: legs I amber with darker metatarsi; other legs amber to pale amber with
black spots on femora IV; spines strong and numerous. Spination of legs I: metatarsi v
2-2-2, p 1-0-1 , d 0-1-1 , r 1-0-1 ; tibiae v 3-2-2, p 0-1-1 , d 0-1-0, r 1-1-1 ; patellae p 1-0-0,
Fig. 8 Cocalodes longicornis sp. n. Holotype d: A, dorsal; B, palp, retrolateral; C,
median apophysis; D, carapace and chelicera, lateral; E, palp, ventral. Paratype 9: F, epigyne;
G, vulva, ventral; H, vulva dorsal.
SPIDER GENUS COCALODES
279
r i-O-O; femora p 1-1-1, d 0-2-3, r 0-1-0. Epigyne (Fig. 8F-H; 21 A): relatively simple, the
openings lead into narrow introductory ducts which extend and loop laterally before joining
the spermathecae.
Dimensions (mm): total length c. 6-4; carapace length 3'32, breadth 2'88, height
2-08; abdomen length c. 3'0; eyes, anterior row 2'6, middle row 2-32, posterior row
2-44; quadrangle length 2*0 (60% of carapace length). Ratios: AM : AL :
PM : PL :: 19 : 12 : 7 : 12; AL-PM-PL :: 13 : 15-5; AM : CL :: 19 : 5 (26% of AM diameter).
VARIATION, <S total length varies from lO'O to 10-16 mm, carapace length 3-76-4'16mm
(three specimens). Another 9 measures c. 8'0 mm total length, 3*0 mm carapace length.
DISTRIBUTION. Papua New Guinea.
MATERIAL EXAMINED. Papua New Guinea: D'Entrecasteaux Islands, Normanby Island, Mt.
Pabinama, about 3 miles ENE of Cape Prevost, 820m, camp 2, 1956 (J. L. Brass, 5th
Archbold Exp., to New Guinea) holotype d1, (AMNH, New York), paratype d (BMNH.
1981.2.2.1); Fergusson Island, Agamoia about 3 miles S. of Lake Ruaba, 200m, camp 5,
18-24. vi.1956 (J. L. Brass, 5th Archbold Exp., to New Guinea) paratype d1, paratype 9,
(AMNH, New York); New Britain, Ralum, (nr. Kokopo) Lowan, on tree trunk, paratype 9,
27.xii. 1 896 (F. Dahl) (MNHU, Berlin. 1 7799).
Cocalodes protervus (Thorell)
(Fig.9A-B)
Cocalus protervus Thorell, 1881 :493, 9. Holotype 9, West Irian (MCSN, Geneva) [examined].
Simon, 1901:407. Roewer, 1954:935. Bonnet, 1956:1174. Proszynski, 1971:391. Wanless
1981 :256.
B
Fig. 9 Cocalodes protervus (Thorell). Holotype 9: A, dorsal; B, epigyne.
280 F. R. WANLESS
DIAGNOSIS. Similar to C. macellus and C. longicornis from which it differs by the
large darkened spermathecae (Fig. 9B).
MALE. Unknown.
FEMALE HOLOTYPE. In poor condition. Carapace (Fig. 9A): brownish with scattered
white hairs, rubbed. Eyes: with black surrounds except AM; fringed by whitish hairs.
Clypeus: densely white haired. Chelicerae: robust, inclined anteriorly and slightly
diverging; brownish with sooty markings; sparsely clothed in light brown hairs; pro-
and retromargins with three teeth. Maxillae and labium: pale brown. Sternum: pale
brown with darker margins. Coxae: generally brownish. Abdomen: greyish yellow
with darker somewhat mottled longitudinal lateral markings; clothed in short recumbent
amber hairs. Legs: dark brown; spines long, slender and numerous. Spination of legs I:
metatarsi v 2-2-2, p 1-2-1, d 1-0-0, r 1-0-1; tibiae v 4-3-3, p 0-2-1, d 1-1-0, r 0-0-1;
patellae p 1-0-0, r 1-0-0; femora p 0-1-1 , d 0-2-3, r 0-2-0. Epigyne (Fig. 9B): rather dark,
openings separated by distinct septum with black disc-like spermathecae posteriorly.
Dimensions (mm): total length 6*8; carapace length 3*04, breadth 2-46, height 1-84;
abdomen length 3'84; eyes, anterior row 2-32, middle row 1*98, posterior row 2*08;
quadrangle length 1-72 (56% of carapace length). Ratios: AM:AL:
PM: PL ::18: 11:6: 10; AL-PM-PL:: 11-13; AM : CL :: 18 : c. 7 (c. 38% of AM
diameter).
DISTRIBUTION. West Irian.
MATERIAL EXAMINED. West Irian: Pulo Faor, holotype d, 1872, (L. M. D'Albertis)
(MCSN, Geneva).
Cocalodes thoracicus Szombathy
(Figs 10A-E; 1 1 A-E; 18D; 20C, D)
Cocalodes thoracicus Szombathy, 1915:470, cf, 9. Syntypes, Papua New Guinea (?in
Termeszettudomanyi Muzeum, Budapest) [not examined]. Roewer, 1954 : 936. Bonnet, 1956 : 1 173.
REMARKS. Although it was not possible to examine the type specimens of C. thoracicus it
may be recognized with a fair degree of confidence from Szombathy 's original figures, which
show the distal cheliceral lobes (arrowed, Fig. 10A) to be more pronounced in this species
than in others of the genus.
DIAGNOSIS. From other species of Cocalodes by the more or less equal development of
the prongs of the median apophysis (Figs IOC, D; 18D) in males, and by the relatively
broad epigynal septum (Fig. 1 1 B, C) in females.
MALE from Morobe Province. In good condition, but legs detached. Carapace brown-
black with dark orange-brown eye area and central yellow-brown stripe from foveal
region to posterior margin; clothed dorsally in fine shining pale amber and whitish
hairs with short coarse black hairs around thoracic margins. Eyes: with black surrounds
except AM; fringed by pale amber and whitish hairs. Clypeus: sparsely covered in pale
amber and blackish hairs. Chelicerae: elongate and porrect with distal lobes; black with
bluish metallic sheen under some angles of illumination; proximally sparsely clothed in stiff
black hairs; promargin with four teeth, retromargin with two; horn protruding just beyond
clypeal margin. Maxillae: brownish black grading to yellow-brown distally. Labium:
brown-black tipped yellow-brown. Sternum: yellow-brown faintly tinged black with
brownish orange margins. Coxae: pale yellow. Abdomen: with four impressed spots; mottled
grey-black with a dorsal greyish yellow band having vague sooty markings; ventrally a
longitudinal black stripe; irregularly clothed in minute setae with scattered long blackish
hairs; spinnerets mottled yellow-brown and black. Legs: legs I tarsi pale yellow, metatarsi
black with pale yellow basal region, tibiae black distally grading to dark yellow-
brown to black proximally, patellae black distally grading to yellow-brown proximally,
SPIDER GENUS COCALODES
281
Fig. 10 Cocalodes thoracicus Szombathy. d: A, dorsal; B, carapace, lateral; C, palp,
retrolateral; D, median apophysis; E, palp, ventral.
femora yellow to yellow-brown; legs II as I, but tibiae and patellae yellow-brown; legs III
yellow-brown to pale yellow-brown; legs IV similar, but with vague darker markings; spines
strong and numerous. Spination of legs I: matatarsi v 2-2-2, p 1-0-0, r 1-0-1 ; tibiae v 2-2-2,
p 1-1-1, d 1-1-0, r 1-1-1; patellae p 1-0-0, r 1-0-0; femora p 1-1-2, d 0-2-1, r 0-1-2. Palp
(FigslOC-E;18D).
Dimensions (mm): total length 9*28; carapace length 3-48, breadth 3*16, height
2'28; abdomen length 5'36; eyes, anterior row 2*56, middle row 2*28, posterior row
2-37; quadrangle length 2*08 (59% of carapace length). Ratios: AM : AL :
PM : PL :: 20 : 12 ; 8 : 12; AL-PM-PL :: 13-15-5; AM : CL :: 20 : 5 (25% of AM diameter).
FEMALE from Madang Province. In good condition. Carapace (Fig. 11 A): yellow-
brown with orange-brown eye region, a thin dark brown border line, lateral brownish
stripes and dark brown bands from PL's to thoracic margin; clothed in very fine
shining amber and whitish hairs. Eyes: with black surrounds except AM; fringed in
shining pale amber and whitish hairs. Clypeus: sparsely covered in white hairs.
Chelicerae: robust, inclined anteriorly and diverging; yellow-brown; shiny; thinly clothed in
testaceous hairs; promargin with four teeth, retromargin with three. Maxillae and labium:
pale yellow-brown. Sternum: pale yellow-brown with darker margins; sparsely clothed in
282 F. R. WANLESS
testaceous hairs. Coxae: pale yellow-brown. Abdomen: similar to d" except for pale yellow
venter and covering of dark amber hairs over lateral bands. Legs: legs I yellow-brown with
darker markings and a black streak on inside of femora; legs II yellow-brown with brownish
markings; legs III yellow-brown; legs IV yellow-brown with brownish markings and black
spots on femora and patellae; spines strong and numerous. Spination of legs I: metatarsi v
2-2-2, p 1-0-1, r 1-0-1; tibiae v 2-2-2, p 1-1-2, r 1-1-1; patellae p 1-0-0, r 1-0-0; femora p
0-2-2, d 0-2-1, r 0-2-0. Epigyne (Figs 1 1B-E; 20C, D): relatively simple, but variable; the
obscure openings (arrowed, Fig. 1 1 D) lead into wide ducts which narrow and loop
posteriorly before entering the spermathecae.
Dimensions (mm): total length 9'28; carapace length 3*32, breadth 2'84, height
1-96; abdomen length 5-6; eyes, anterior row 2-44, middle row 2'2, posterior row
2-28; quadrangle length 1*92 (57% of carapace length). Ratios:
AM : AL: PM : PL :: 18: 12:7: 1 1; AL-PM-PL :: 12-5-15; AM : CL :: 18 : 5 (27 per cent of
AM diameter).
VARIATION, d1 total length varies from 6*9 to 11 -5 mm, carapace length 2-72-4-4 mm
(14 specimens). 9 total length 7-28-10-8 mm, carapace length 2-84^-08 mm (nine
specimens).
The cheliceral horn, possibly an allometric growth character, only protrudes well
beyond the clypeus in large specimens. The epigynal septum varies from rectangular
to triangular in outline (Fig. 1 1 B, C) and the rod-like extensions of the spermathecae
(in reality part of the introductory ducts), are not always evident in uncleared
epigynes.
DISTRIBUTION. Papua New Guinea; Irian Jaya.
W. :-^c- 'i' ' JtfaMttb •..-.•.' - - ~~Zi&r /• •
Fig. 11 Cocalodes thoracicus Szombathy. 9: A, dorsal; B, epigyne: C, epigyne of another
specimen; D, vulva, ventral; E, vulva, dorsal.
SPIDER GENUS COCALODES
283
MATERIAL EXAMINED. Papua New Guinea: Morobe Province, Wau: Ip, iv.1979 (H. W.
Levi); 19, iv.1979 (M. Robinson); 19, iv.1979, beating vegetation in forest, (H. W.
Levi); 3dtf, 19, Me Adam Park, 4.iv.l979 (H. W. Levi, Y. Lubin, M. Robinson);
Madang Province, 25 Km, N. of Madang, teak forest, night collection, Id1, 21.iii.1979
(H. W. Levi, Y. Lubin. B. Robinson) (MCZ. Harvard); Draeger Harbour: 5 miles E. of
Finschhafen, Id1, vi.1944 (G. H. Penn) (AMNH, New York); Maffm Bay: Id1, xii.1944
(Lt. Geo. B. Sirotiak) (AMNH. New York); Astrolabe Bay, 19, (R. Rohde) (MNHN,
Berlin. 17794); Ramu, Id1, (F. Dahl, Ramu Expedition) (MNHN, Berlin. 17801); Jagei
River, Id, 2.viii.l896 (Lauterbach) (MNHN, Berlin, 17796). Irian Jaya: Sukarnapura
(Hollandia): 2dtf, 5. i. 1945 (L. W. Saylor); Icf, iv.1945 (Borys Malkin); Id, 14.V.1945
Borys Malkin); rain forest, 250ft. 2dtf, xii.1944 (H. Hoogstraat) (AMNH, New York);
Humboldts Bay, 200ft. 19, iv.1936 (L. E. Cheesman) (BMNH. 1937.12.13.425); Cyclops
Mts.Sabron,2,200ft. I$,v.l936 (L. E. Cheesman) (BMNH. 1937.12.13.541).
Coca lodes turgidus sp. n.
(Figsl2A-D;20E,F)
DIAGNOSIS. Like C. thoracicus, but differs by the median swelling of the epigynal
septum (Fig. 12B).
MALE. Unknown.
FEMALE HOLOTYPE. In fair condition. Carapace (Fig. 12 A): yellow-brown with orange-brown
eye region, dark brown lateral margins and dark brown stripes below the lateral eyes to
posterior margin; clothed in minute creamy white setae with light brownish hairs around
thoracic margins. Eyes: with black surrounds except AM; irregularly fringed by creamy
ai±ii*i£rj&^
Fig. 12 Cocalodes turgidus sp. n. Holotype 9: A, dorsal; B, epigyne. Paratype 9: C, vulva,
ventral; D, vulva, dorsal.
284 F. R. WANLESS
white hairs. Chelicerae: robust, inclined anteriorly and diverging; amber with brownish
markings; shiny; thinly clothed in clear pale amber hairs; pro- and retromargins with three
teeth. Maxillae: yellow-brown with inner distal margins whitish yellow. Labium:
yellow-brown. Sternum: light yellow-brown with darker margins; shiny; thinly clothed in
testaceous hairs. Coxae: pale yellow-brown. Abdomen: with four impressed spots; pale
yellow with black lateral stripes and a black ventral band. Legs: legs I with tarsi
yellow-brown, metatarsi blackish grading to light orange-brown proximally, tibiae
yellow-brown with blackish annuli, patellae yellow-brown, femora yellow-brown with
blackish streaks; other legs similar, but darker markings less extensive especially on legs
III-IV; spines strong and numerous. Spination of legs I: metatarsi v 2-2-2, p 1-0-0, d 0-1-0,
r 1-0-1 ; tibiae v 2-2-2, p 0-1-1 , d 0-1-0, r 1-1-1 ; patellae p 1-0-0, r 1-0-0; femora p 1-1-0,
d 0-2-3, r 0-1-1. Epigyne (Figs 12B-D;20E, F).
Dimensions (mm): total length 8*72; carapace length 3'32, breadth 2'88, height
1'96; abdomen length 5'28; eyes, anterior row 2*44, middle row 2- 18, posterior row
2-32; quadrangle length 2*02 (60% of carapace length). Ratios: AM : AL :
PM : PL :: 19 : 12 : 8 : 12; AL-PM-PL :: 1 1-16; AM : CL :: 19 : 5 (26% of AM diameter).
VARIATION. A paratype 9 measures 8*64 mm total length, 3'28 mm carapace length.
DISTRIBUTION. Irian Jaya.
MATERIAL EXAMINED. Irian Jaya: Sansapor, holotype 9, paratype 9, viii-ix.1944 (R. B.
Burrows, A.P.O. 1 59) (AMNH, New York).
Cocalodes longipes (Thorell)
(Fig. 13A-G)
Cocalus longipes Thorell, 1881:494, 707, 9, d. LECTOTYPE 9, Yule Isl. PARALECTO-
TYPE cf, Ceram Isl. (here designated) (MCSN, Geneva) [examined]. Simon, 1901:407.
Roewer, 1954 : 935. Bonnet, 1956: 1 173. Proszyriski, 1971 : 391. Wanless 1981 : 256
DIAGNOSIS. From other species of Cocalodes by the scimitar-shaped dorsal prong of the
median apophysis (Fig. 13E) in males, and by the comma-like introductory ducts (Fig. 13B)
in females.
FEMALE LECTOTYPE. In fair condition. Carapace (Fig. 13D, F): orange-brown with
sooty markings; irregularly clothed in short recumbent white hairs. Eyes: with black
surrounds except AM; fringed by whitish hairs. Clypeus: thinly clothed in fine
whitish hairs. Chelicerae: robust, inclined anteriorly and slightly diverging; dark amber;
thinly clothed in white and light amber hairs; pro- and retromargins with three teeth.
Maxillae: orange-brown with lighter inner margins. Labium: orange-brown. Sternum: amber
with darker margins. Coxae: generally yellowish orange to amber. Abdomen: pale
yellow-orange with faint sooty lateral markings outlining a central pale band; spinnerets pale
yellow-brown. Legs: brownish amber grading to light amber distally; spines numerous and
robust. Spination of legs I: metatarsi v 2-1-1, p 1-1-1, d 0-0-2, r 1-0-0; tibiae v 4-4-2,
p 0-1-1, d 0-1-0; patellae r 1-0-0, p 1-0-0; femora p 1-0-1, d 0-2-3, r 0-1-1. Epigyne (Fig.
1 3B): similar toplatnicki and signatus, but readily separated by the comma-like ducts.
Dimensions (mm): total length c. 10*0; carapace length 3*76, breadth 3*32, height
2-36; abdomen length 5*92; eyes, anterior row 2*6, middle row 2*36, posterior row
2-41; quadrangle length 2'04 (54% of carapace length). Ratios: AM : AL :
PM : PL ::20 : 12 : 7'5 : 12; AL-PM-PL :: 11-5-1 7; AM : CL :: 20 : 5 (25% of AM diameter).
MALE PARALECTOTYPE. In fair condition. Carapace (Fig. 13 A): amber with vague
brownish markings; rubbed. Eyes: with black surrounds except AM; irregularly fringed by
creamy white hairs. Clypeus: fringed in greyish white hairs. Chelicerae: elongate, porrect and
diverging distally; amber, shiny; very sparsely clothed in light amber hairs; pro- and
retromargins with three teeth. Maxillae and labium: yellow-brown. Sternum: pale
yellow-brown with darker margins. Coxae: yellow-brown. Abdomen: yellow-brown with
SPIDER GENUS COCALODES
285
Fig. 13 Cocalodes longipes (Thorell). Paralectotype d1: A, dorsal; C, palp, retrolateral; E,
median apophysis; G, palp, ventral. Lectotype 9: B, epigyne; D, carapace, lateral; F, dorsal.
faint sooty lateral markings; spinnerets yellow-brown. Legs: generally orange-brown; spines
strong and numerous. Spination of legs I: metatarsi v 2-2-2, p 1-1-1 , r 1-0-1 ; tibiae v 4^-0,
p 0-1 -l,d 0-1-0, 1-1-1; patellae p 1-0-0, r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-2-0. Palp
(Fig. 1 3C, E, G): the triangular tip of the functional conductor is distinctive.
Dimensions (mm): total length c. 9*9; carapace length c. 4*28, breadth 3*64, height
2*52; abdomen length 6-24; eyes, anterior row 2*88, middle row 2'55, posterior row
2*68; quadrangle length 2-32 (54% of carapace length). Ratios: AM : AL :
PM: PL:: 22-5: 14:8.5: 14:5; AL-PM-PL :: 13-17; AM : CL :: 22-5 : 8 (35% of AM
diameter).
286 F. R. WANLESS
DISTRIBUTION. Indonesia: Ceram Island; Papua New Guinea: Yule Island.
MATERIAL EXAMINED. Indonesia, Wahai, Ceram Island, paralectotype d1, 1872, (L. M.
D'Albertis). Papua New Guinea, Yule Island (Roro), lectotype 9, 1872 (L. M. D'Albertis)
(MCSN, Geneva).
REMARKS. To judge from the geographical distribution, the male and female described above
may not be conspecific.
Coca lodes platnicki sp . n .
(Figsl4A-J;21C-F)
DIAGNOSIS. From other species of Cocalodes by the combined absence of peg-like
cymbial spines and curved tip of the functional conductor (Fig. 14F) in males.
Females are readily separated by the absence of an epigynal septum and broad
posterior ledge (Fig. 14H).
MALE HOLOTYPE. In fair condition. Carapace (Fig. 14B, C): yellow-brown with dark
brownish sides and thin yellow-brown border line; pale areas including border line
clothed in short whitish hairs, elsewhere covered in short blackish hairs. Eyes: with
black surrounds except AM; fringed by whitish hairs. Clypeus: fringed in long white
hairs. Chelicerae: moderately elongate, robust, inclined anteriorly and slightly divergent;
pale orange-brown; sparsely fringed by long white hairs along outer basal margin; pro- and
retromargins with three teeth. Maxillae and labium: yellow-brown. Sternum (Fig. 14G):
yellow-brown with darker margins; shiny. Coxae: yellow-brown; shiny. Abdomen: with two
pairs of impressed spots: pale yellow with blackish lateral bands; spinnerets pale yellow-
brown tinged black. Legs: legs I amber grading to pale yellow-brown distally with blackish
markings on femora, distal half of tibiae and metatarsi; legs II similar to I, but markings less
distinct; legs III-IV yellow-brown with vague sooty annuli on metatarsi and darkening
towards apices of tibiae; spines strong and numerous. Spination of legs I: metatarsi v 2-2-2,
p 1-1-1 , r 1-0-1 ; tibiae v 2-2-2, p 1-1-2, d 0-0-1 , r 1-1-2; patellae p 1-0-0, r 1-0-0; femora
p 2-1-2, d 0-3-2, r 1-2-2. Palp (Fig. 14D-F).
Dimensions (mm): total length 7*2; carapace length 2*98, breadth 2-48, height 1'76;
abdomen length 4*24; eyes, anterior row 2*28, middle row 2'02, posterior row 2' 12;
quadrangle length 1-8 (60% of carapace length). Ratios: AM : AL :
PM : PL :: 18 : 1 1 : 7 : 1 1-5; AL-PM-PL :: 10-14; AM : CL :: 18 : 4 (22% of AM diameter).
FEMALE PARATYPE. In good condition. Carapace (Fig. 14 A): dark orange-brown with
yellow-brown markings; dark areas clothed in short blackish hairs with whitish hairs
elsewhere. Eyes: more or less as in d. Clypeus: fringed by white hairs. Chelicerae:
robust and moderately porrect; amber; sparsely clothed in whitish and pale amber
hairs; pro- and retro-margins with three teeth. Maxillae and labium: pale yellow-
brown. Sternum: pale yellow-brown; thinly covered in testaceous hairs. Coxae: pale
yellow-brown. Abdomen: similar to d, but ventrally with a pale greyish band from
epigyne to spinnerets. Legs: legs I-II yellow-brown with vague darker markings and
blackish femoral stripes; other legs yellow-brown with sooty spots on femora, darkened tibial
apices and sooty annuli of metatarsi; spines strong and numerous. Spination of legs I:
metatarsi v 2-2-2, p 1-0-0, d 0-1-2, r 1-0-0; tibiae v 2-2-2, p 1-1-1, d 0-1-0, r 1-1-1;
patellae p 1-0-0, r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-2-0. Epigyne (Figs 14H-J; 21C-F):
only species in genus known to have secondary spermathecae.
Dimensions (mm): total length 8*96; carapace length 3-44, breadth 2-84, height 2'0;
abdomen length 5*28; eyes, anterior row 2'52, middle row 2' 18, posterior row 2-28;
quadrangle length 2*0 (58% of carapace length). Ratios: AM : AL :
PM:PL:: 19 : 12 : 8 : 12; AL-PM-PL :: 13-16-5; AM : CL :: 19 : 4 (21% of AM diameter).
SPIDER GENUS COCALODES
287
Fig. 14 Cocalodes platnicki sp. n. Holotype cf: B, carapace, lateral; C, dorsal; D, palp,
retrolateral; E, median apophysis; F, palp, ventral; G, sternum. Paratype 9: A, dorsal;
H, epigyne; I, vulva, dorsal; J, vulva, ventral.
VARIATION. Paratype cf measures 9*6 mm total length, 3-72 mm carapace length. Paratype 9
c. 9-1 mm total length, 3'24 mm carapace length.
DISTRIBUTION. Irian Jaya.
MATERIAL EXAMINED. Irian Jaya, Sukarnapura (Hollandia): tropical rain forest, holotype d1,
paratype 9, 31.xii.1944 (L. W. Saylor) (AMNH, New York); paratype 9, 5. i. 1945 (L. W.
Saylor) (AMNH, New York); paratype rf, rain forest, xii.1944, (H. Hoogstraat) (AMNH,
New York).
ETYMOLOGY. This species is named after Dr N. I. Platnick, American Museum of
Natural History, New York.
288
F. R. WANLESS
Cocalodes in not a hi I is sp. n.
(Figsl5A-F;18A, B)
DIAGNOSIS. Similar to males of C. longipes and C. longicornis, but separated by the
combined presence of peg-like cymbial spines and the finger-like dorsal prong of the
median apophysis (Fig. 1 5D, E).
FEMALE. Unknown. C. expers from Fergusson Island may belong here or possibly the
female of C. longipes, from Yule Island (Papua New Guinea), which seems doubtfully
matched with the male known only from Seram (Indonesia).
MALE HOLOTYPE. In fair condition. Carapace (Fig. 15 A, B): orange-brown with dark
reddish brown sides and light orange margins; weakly iridescent under some angles of
illumination; rubbed. Eyes: with black surrounds except AM; fringed in whitish and
pale amber hairs. Clypeus: sparsely fringed by fine pale brownish hairs. Chelicerae:
moderately long, inclined anteriorly; dark reddish brown, weakly iridescent blue under some
angles of illumination; thinly covered in stiff brownish hairs; pro- and retromargins with
three teeth; horn moderately long, but not protruding beyond clypeus. Maxillae:
orange-brown to light yellowish along inner and distal margins. Labium: orange-brown
tipped light yellow. Sternum: pale amber with darker margins; shiny. Coxae: pale amber.
Abdomen: with two pairs of impressed spots; yellow-brown with brownish lateral bands
above and central blackish band below; rubbed; spinnerets light orange-brown tinged black.
Legs: tarsi light amber, metatarsi dark brown with light brownish hairs grading to amber with
Fig. 15 Cocalodes innotabilis sp. n. Holotype rf: dorsal; B, carapace, lateral; D, palp,
retrolateral; E, median apophysis; F, palp, ventral. Paratype cf: C, carapace, dorsal,
showing cheliceral horn.
SPIDER GENUS COCALODES 289
white hairs proximally, tibiae dark brown, patellae brownish grading to amber proximally,
femora amber; other legs amber; spines strong and numerous. Spination of legs I: metatarsi
v 2-2-2, p 1-0-0, d 0-1-2, r 1-0-1; tibiae v 2-2-2, p 1-1-1, d 1-1-0, r 1-1-1; patellae
p 1-0-0, r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-1-1. Palp (Figs 15D-F; 18A, B); the lobe
associated with the tip of the functional conductor (arrowed, Fig. 15F) is also
characteristic of this species.
Dimensions (mm): total length 6'96; carapace length 2*88, breadth 2*4, height 1'84;
abdomen length 4-0; eyes, anterior row 2*26, middle row 2*04, posterior row 2- 18;
quadrangle length l-72 (59% of carapace length). Ratios: AM:AL:
PM : PL :: 18 : 1 1 : 6 : 1 1; AL-PM-PL :: 10-13; AM : CL :: 18 : 5 (27% of AM diameter).
VARIATION, rf total length varies from 5'9 to 8-2 mm, carapace length 2'28-4-4 mm
(eight specimens).
In one male, having the same carapace length as the holotype, the cheliceral horn
protrudes beyond the clypeus and may be clearly seen in dorsal view (Fig. 15C). Also,
as in the case of C. papuanus the cymbium bears either two or three peg-like spines.
DISTRIBUTION. Papua New Guinea.
MATERIAL EXAMINED. Papua New Guinea: Louisiade Archipelago, Sudest Island, (L.
J. Brass, Fifth Archbold Expedition to New Guinea) (AMNH, New York): Rambuso,
on north coast, 0-100 m, holotype d1, camp 11, xiii.1956; Mt. Riu, paratype rf,
250-350 m, 1956. Kokoda, paratype d1, viii.1933 (L. E. Cheesman) (BMNH.
1934.12.14.269); Morobi Province, Buro river, paratype d, ix.1979 (/. H. Martin}
(BMNH. 1981.7.31.1); Milne Bay, paratype cf, autumn 1944 (R. B. Burrows, A.P.O.
928) (AMNH, New York); Oro Bay; paratypes 2c?d", (Lt. B. Struck, A.P.O. 503)
(AMNH, New York), paratype d1, vii-viii. 1 943 (Sidney Sandier) (AMNH, New York).
Genus ALLOCOCALODES gen. n.
DEFINITION. Based on two species known only from males.
Spiders of medium size (i.e. 4*0 to 8'0 mm in length). Male with elongate and
porrect chelicerae bearing a median horn which is sometimes visible in dorsal view;
not hirsute. Carapace: profile as in Figs 16 A, B; 17A, B); high, longer than broad,
with steep thoracic slope, widest at level of coxae II-III; fovea long, sulciform, centre
situated just behind posterior margin of posterior lateral eyes. Eyes: large with black
surrounds except anterior medians; posterior median and posterior laterals set on
pronounced tubercles; set in three transverse rows comprised of anterior medians
(AM) and anterior laterals (AL), posterior medians (PM) and posterior laterals (PL);
anteriors 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 large, positioned nearer to anterior laterals than to posterior
laterals and more or less on optical axis of anterior laterals; posterior laterals about as large as
anterior laterals and set inside lateral margins of carapace when viewed from above;
quadrangle formed by posterior medians and posterior laterals broader than long and widest
posteriorly; entire quadrangle, measured from between bases of anterior medians to posterior
margins of posterior laterals, occupying between 60-65% of carapace length. Clypeus:
between 25 and 37% of diameter of anterior median eyes. Chelicerae: elongate and porrect,
sometimes diverging, with a median horn arising basally; promargin with four teeth,
retromargin with two. Maxillae (Fig. 1 6C): long and diverging with rounded apices. Labium:
oblong about half maxillae length. Sternum (Fig. 16C): scutiform. Coxae: I and II
largest. Pedicel: short. Abdomen: elongate ovoid; spinnerets moderately long, posteriors
longest with moderately 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 scanty tuft of fine hairs; anal tubercle a broad
cone. Legs: long and slender; spines moderately strong and numerous; claws pectinate; tufts
290
F. R. WANLESS
present; scopulae absent. Male palps: moderately simple and similar in appearance; femora
slightly bowed; patellae moderately long; tibiae with small dorsal tubercle, bifid retrolateral
apophysis, the ventral element broad and rounded, the dorsal slender with translucent tip;
cymbium with distal finger-like extension and shallow dorsally ridged excavation on retro-
lateral basal margin; embolus (e) short, moderately stout and curved, arising subapically;
functional conductor (c) a finger-like extension of the tegulum the tip of which lies either
above or below that of the embolus; also arising from tegulum a lobe-like median apophysis
(m); tegulum (t) ovoid with peripheral seminal ducts and what appears to be an inner folded
duct. Expanded palps not examined.
TYPE SPECIES. Allococalodes alticeps sp. n.
AFFINITIES. Closely related to Cocalodes, see remarks on p. 264.
DIAGNOSIS. From Cocalodes by the bifid retrolateral tibial apophysis, lobe-like functional
conductor and median apophysis (Figs 16E; 17C).
Fig. 16 Allococalodes alticeps sp. n. Holotype rf: A, dorsal; B, carapace lateral; C,
sternum, coxae and mouth parts; D, palpal tibia, retrolateral; E, palp, ventral; F, palp,
retrolateral.
SPIDER GENUS COCALODES 29 1
List of species in the genus Allococalodes gen. n.
Allococalodes alticeps sp. n.
A. cornutus sp. n.
Allococalodes alticeps sp. n.
(Fig. 16A-F)
DIAGNOSIS. From A. cornutus by the pointed tip of the median apophysis (Fig. 16E).
FEMALE. Unknown.
MALE HOLOTYPE. In fair condition. Carapace (Fig. 16 A, B): dark chestnut brown with
central yellow-brown thoracic band and vague yellow-brown markings in eye region;
sides clothed in short recumbent amber hairs, iridescent under some angles of illumination,
with whitish hairs on thoracic band and eye region. Eyes: with black surrounds except AM;
fringed by whitish hairs. Clypeus: fringed in white hairs. Chelicerae: elongate and porrect;
dark brownish orange with blackish mottling; thinly clothed in stiff light amber hairs;
promargin with four teeth, retromargin with two; cheliceral horn protruding just beyond
clypeal margin, but not visible in dorsal view. Maxillae and labium: brownish orange
suffused black with inner margin of maxillae and labial tip dull yellow-brown. Sternum:
brownish orange suffused black. Coxae: yellow-brown. Abdomen: mottled black with dorsal
light yellow band somewhat broken and having blackish markings; clothed in minute dull
whitish hairs; spinnerets tinged black, the posteriors with lighter terminal articles. Legs:
generally yellow-brown with blackish annuli; spines strong and numerous. Spination
of legs I: metatarsi v 2-0-1, p 1-1-1, r 1-1-1; tibiae v 2-2-2, p 1-1-1, d 1-1-0,
r 1-1-1; patellae p 0-1-0, r 0-1-0; femora p 1-1-1, d 0-2-2, r 0-1-0. Palp (Fig.
16D, E): when viewed from below (i.e. Fig. 16E), the tip of the embolus lies above the
tip of the functional conductor (c.f. A. cornutus).
Dimensions (mm): total length 4-84; carapace length 2*2, breadth 2*04, height 1'64;
abdomen length 2-56; eyes, anterior row 1'96, middle row 1*68, posterior row 1-8;
quadrangle length 1'44 (65% of carapace length). Ratios: AM:AL:
PM : PL :: 16 : 10 : 6'5 : 10; AL-PM-PL :: 8-10; AM : CL :: 16 : 4 (25% of AM diameter).
DISTRIBUTION. Irian Jaya.
MATERIAL EXAMINED. Irian Jaya: Sukarnapura (Hollandia), rain forest, holotype d1,
3 1 .xii. 1 944 (L. W. Saylor) (AMNH, New York).
Allococalodes cornutus sp. n.
(Fig. 17A-E)
DIAGNOSIS. From A. alticeps by the obtuse tip of the median apophysis (Fig. 17C).
FEMALE. Unknown.
MALE HOLOTYPE. In poor condition. Carapace (Fig. 17A, B): dark orange-brown with
pale eye region and median light yellowish brown thoracic band from foveal area to
posterior margin; sides clothed in short recumbent amber hairs, shining under some
angles of illumination, with dull whitish hairs on thoracic band and eye region. Eyes:
with black surrounds except AM; fringed by dull whitish hairs. Clypeus: White
haired. Chelicerae: elongate, porrect and diverging; dark orange-brown with blackish
mottling; sparsely clothed in fine clear whitish hairs; promargin with four teeth,
retromargin with two; cheliceral horn spiralled and protruding beyond clypeal margin.
Maxillae: brown-black to dirty yellow-brown. Labium: dark brownish. Sternum: amber with
irregular blackish mottling. Coxae: yellow-brown. Abdomen: damaged; elongate ovoid;
yellow-brown with irregular blackish markings. Legs: generally light orange-brown grading
292
F. R. WANLESS
Fig. 17 Allococalodes cornutus sp. n. Holotype d: A, carapace lateral; B, carapace,
dorsal; C, palp, ventral; D, palpal tibia, retrolateral; E, palp, retrolateral. Abbreviations: c,
functional conductor; e, embolus; m, median apophysis; t, tegulum.
to yellow-brown distally with blackish annuli; spines moderately strong and numerous.
Spination of legs I: metatarsi v 2-1-1, p 1-1-1, d 0-0-1, r 1-0-1; tibiae v 2-4-1, p 1-1-1,
d 1-1-0, r 1-1-0; patellae p 1-0-0, r 1-0-0; femora p 1-1-0, d 0-2-2. Palp (Fig. 17C-E):
when viewed from below the functional conductor lies above the embolic tip (Fig. 1 7C).
Dimensions (mm): total length c. 4*8; carapace length 2-32, breadth 2*02, height
l-66; abdomen length c. 2*6; eyes, anterior row 1*94, middle row l-62, posterior row
1-76; quadrangle length 1-4 (60% of carapace length). Ratios: AM : AL :
PM : PL :: 15: 9 : 6-5 : 9'5; AL-PM-PL :: 9-9; AM : CL :: 15 : 5'5 (37% of AM diameter).
DISTRIBUTION. Irian Jaya.
MATERIAL EXAMINED. Irian Jaya, holotype d1, (Dr. Moszkowski, no 54) (MNHU, Berlin.
17800).
SPIDER GENUS COCALODES 293
Taxonomic summary
1 . Allococalodes gen. n., is proposed.
2. Eight new species are described:
Allococalodes alticeps sp. n.
Allococalodes cornutus sp. n.
Cocalodes cygnatus sp. n.
Cocalodes expers sp. n.
Cocalodes innotabilis sp. n.
Cocalodes longicornis sp. n.
Cocalodes platnicki sp. n.
Cocalodes turgidus sp. n.
3. Three species are newly synonymized:
Cocalodes armatissimus Strand, 1913, and Cocalodes plebejus Szombathy, 1915, are
junior synonyms of Cocalodes papuanus Simon, 1900.
Cocalodes melanognathus Pocock, 1897, is a junior synonym of Cocalodes leptopus
Pocock, 1897.
Acknowledgements
I wish to thank the following colleagues for providing specimens for study. Dr Gianna
Arbocco, Museo Civico di Storia Naturale, Geneva, Italy (MCSN, Genova); Dr M.
GrasshofT, Forschungsinstitut Senckenberg, Natur-Museum, Senckenberg, Frankfurt am
Main, West Germany (FS, Frankfurt am Main); Dr M Hubert, Museum National
d'Histoire Naturelle, Paris, France (MNHN, Paris); Dr H. W. Levi, Museum of
Comparative Zoology, Harvard, U.S.A. (MCZ, Harvard); Dr M. Moritz, Museum fur
Naturkunde der Humboldt-Universitat, Berlin, East Germany (MNHU, Berlin); Dr
N. I. Platnick, American Museum of Natural History, New York, U.S.A. (AMNH,
New York).
I also wish to thank Mr D. Macfarlane (CIE, London) for reading the manuscript
and Mr P. York (BMNH. Photo unit) for the cleared epigyne photographs.
References
Bonnet, P. 1956. Bibliographia Araneorum. 2 (2) : 919-1925. Imprimerie Douladoure, Toulouse.
Forster, R. 1980. Unidentified objects. /. Arachnol. 8 (1) : 91-92.
Neave, S. A. 1939. Nomencl. Zool. Vol I A-C: 957 pp. Zoological Society, London.
Petrunkevitch, A. 1928. Systema Aranearum. Trans. Conn. Acad. Arts Sci. 29 : 270 pp.
Platnick, N. I. & Shadab, M. U. 1975. A revision of the spider genus Gnaphosa (Araneae:
Gnaphosidae) in America. Bull. Am. Mus. nat. Hist. 155 : 3-66.
Pocock, R. I. 1897. Spinnen (Araneae). In Kiikenthal, W., Ergebnisse einer zoologische
Forschungsreise in dem Molukken und Borneo. Abh. senckenb. naturforsch. Ges. 23 : 591-629.
Prdszyriski, J. 1971. Catalogue of Salticidae (Aranei) specimens kept in major collections of
the world. Annls zool. Warsz. 28 : 367-5 19.
Roewer, C. F. 1954. Katalog der Araneae. 2 Abt. B: 924-1290. Institut Royal des Sciences
Naturelle de Belgique, Bruxelles.
Simon, E. 1900. Etudes arachnologiques 30e Memoire (1) XLVII Descriptions d'especes nouvelles de
la famille des Attidae. Annls Soc. ent. Fr. 69 : 27-6 1 .
1901. Histoire Naturelle des Araignees. 2 (3) : 381-668. Roret: Libraire Encyclopedique,
Paris.
Strand, E. 1913. Neue indoaustralische und polynesische Spinnen des senckenbergischen Museums.
Arch, naturgesch. 79A (6) : 1 13-123.
1915. Indoaustralische, papuanische und polynesische Spinnen des senckenbergischen Museums,
gesammelt von Dr E. Wolf, Dr J. Elbert u. a. In wissenschaftliche Ergebnisse de Hanseatischen
Siidsee Expedition 1909. Abh. senckenb. naturforsch. Ges. 36 (2) : 181-274.
294 F. R. WANLESS
Szombathy, K. 1915. Attides nouveaux appartenant aux collections du Musee National Hongrois.
Annls hist.-nat. Mus. natn. hung. 13 : 468-490.
Thorell, T. 1878. Studi sui ragni Malesi e Papuani. Part II. Ragni di Amboina raccolti da Prof.
O. Beccari. Mus. civ. Stor. nat. Giacomo Doria 13:317 pp.
1881. Studi sui ragni Malesi e Papuani. Part III. Ragni dell Austro-Malesia e del Capo
York, conservati nel Museo Civico di Storia Naturale di Geneva. Mus. civ. Stor. nat.
Giacomo Doria 17 : 720 pp.
Wanless, F. R. 1978. A revision of the spider genera Belippo and Myrmarachne (Araneae:
Salticidae) in the Ethiopian region. Bull. Br. Mus. nat. Hist. (Zool.)33 (1) : 139 pp.
1981. A revision of the spider genus Cocalus (Araneae: Salticidae). Bull. Br. Mus. nat.
Hist.(Zool) 41 (5): 253-261.
Waterhouse, C. O. 1 902. Index zoologicus 42 1 pp. London.
Manuscript accepted for publication 5 October 198 1
SPIDER GENUS COCALODES
295
Fig. IS (A, B) Cocalodes innotabilis sp. n. cT palp: A, peg-like cymbial spines, x500; B,
median apophysis and functional conductor, xlOO. (C, E) C. papuanus Simon, rf palp: C,
median apophysis, xlOO; E, ventrolateral view, x60. (D) C. thoracicus Szombathy. d
palp, median apophysis, xl 50.
296
F. R. WANLESS
B
Fig. 19 (A, B) Cocalodes papuanus Simon. Vulva: A, ventral; B, dorsal, xlOO. (C, D) C.
leptopus Pocock. Vulva: C, ventral; D, dorsal, x!40. (E, F) C. cygnatus sp. n. Vulva: E,
ventral; F, dorsal; x 140.
SPIDER GENUS COCALODES
297
Fig. 20 (A, B) Cocalodes macellus (Thorell). Vulva: A, ventral; B, dorsal; xlOO. (C, D) C.
thoracicus Szombathy. Vulva: C, ventral; D, dorsal; xlOO. (E, F) C. turgidus sp. n.
Vulva: E, ventral, xlOO; F, dorsal, x!35.
298
F. R. WANLESS
B
Fig. 21 A, Cocalodes longicornis sp. n., vulva dorsal view, x!40. B, C. papuanus Simon,
cheliceral horn, x85. (C, D) C. platnicki sp. n. Vulva: C, ventral; D, dorsal; xlOO. (E, F)
spherical bodies in vulva of C. platnicki; E, x280; F, x!200.
Anatomy and evolution of the jaws in the
semiplotine carps with a review of the Genus
Cyprinion Heckel, 1843 (Teleostei: Cyprinidae)
Gordon Howes
Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7
5BD
Introduction
Scant attention has been paid to the jaw anatomy of cyprinid fishes. Takahasi (1925)
described the jaw musculature of various cyprinoids, but only Matthes (1963) and Alexander
(1966) have described the osteo-myological anatomy of the cyprinid jaw. These authors, and
Ballantijn (1969), have also attempted a functional interpretation of particular jaw types.
Howes (1978, 1979, 1980, 1981) described and commented on jaw types of (mostly) non-
barbelled carps ('Leuciscinae'). But, in these studies the potential value of the jaws as
indicators of phyletic relationship was not recognised. The more diverse jaw morphology of
barbelled carps ('Cyprininae') has, however, focussed attention on this aspect of jaw
anatomy.
From among the 'Cyprininae' the genus Semiplotus is chosen as a model for detailed
description because member species possess what is an obviously highly derived osteo-
myological jaw structure, manifest externally in the size and shape of the mouth (see below).
Semiplotus species are large (according to Day (1889) S. semiplotus attains at least 2 feet in
length), distinctive carps whose geographical range extends through northern India and
Burma. The distinctiveness of these fishes lies in the size and shape of the head and extreme
body depth (Figs 1A-C). Likewise, the mouth is of great width (probably the widest mouth
in relation to head length of any cyprinid fish), the lower jaw is edged with a cornified
epithelium and is overhung by the upper jaw (Figs 1 B & C).
Many cyprinid taxa possess broad, ventral mouths with cornified cutting edges, the
so-called 'sector mouth' (see p. 313 for definition). Only in Onychostoma, Varicorhinus,
Capoeta and Cyprinion does the morphology of the mouth approach that of Semiplotus,
both in width and complex development of the lower lip.
Members of these genera are compared here in order to test the hypothesis that a sector
mouth is synapomorphic for the group and also to elucidate the possible evolutionary
pattern of semiplotine jaw modification.
Nomenclatural note
In the anatomical descriptions, Scaphiodonichthys burmanicus is referred to as Semiplotus
burmanicus. The existence in both 'genera' of synapomorphic osteological and myological
characters indicates their generic unity (see p. 312 for a list of characters, and p. 331 for
complete synonymy). No specimens of Semiplotus modestus have been examined and
reference to 'Semiplotus' in the anatomical text refers only to the species semiplotus and
burmanicus.
List of specimens used in anatomical description
Capoeta capoeta BMNH 1968.12.13 : 405^16
Capoeta capoeta BMNH 1879.1 1.14 : 19 (skeleton)
Bull. Br. Mm. not. Hist. (Zool.)42 (4) : 299-335 Issued 24 June 1982
300
G. J. HOWES
Fig. 1 Semiplotus semiplotus. A, in lateral view. B & C, ventral and anterior aspects of the head.
Scale = 10 mm. Drawn from specimen 1889.2.1 :365-6.
Capoetafusca
Cyprinion acinaces
Cyprinion kais
Cyprinion macrostoma
Cyprinion microphthalmum
Cyprinion microphthalmum
Cyprinion milesi
Cyprinion watsoni
Cyprinion watsoni
Onychostoma laticeps
Onychostoma varus
Semiplotus burmanicus
Semiplotus burmanicus
Semiplotus burmanicus
Semiplotus burmanicus
Semiplotus semiplotus
Semiplotus semiplotus
Semiplotus semiplotus
Squaliobarbus curriculus
BMNH 1899.7. 15 26
BMNH 1977.8.25 1-5
BMNH 1974.2.22 1115-8
BMNH 1974.2.22 118-95
BMNH 1883.8.2:4-9
BMNH 1977. 12. 13: 532-545; 546-556
BMNH 1889.2.1 : 263^
BMNH 1889.2.1 : 387-396
BMNH 1913.4.15:90-99
BMNH 1969.4.15: 105-110
BMNH 1935.4.18:22
BMNH 1893.2.16 : 32 (skeleton)
BMNH 1889. 10.4: 2
BMNH 1920.9.8: 1-3
Uncat. Smithsonian Colin.
BMNH 1889.2.1 : 365-6
BMNH 1889.2.1 : 367 (skeleton)
BMNH 1972.7.26: 10
BMNH 1 888.5. 1 5 : 29 (skeleton)
JAW ANATOMY OF SEMIPLOTINE CARPS 30 1
Varicorhinus beso BMNH 1968.7.24 : 2
Varicorhinus beso BMNH 1968.7.24 : 17-19 (alizarins)
Varicorhinus beso BMNH 1902.12.13 : 377 (skeleton)
Varicorhinus tanganicae BMNH 1906.9.6 : 1 1 (skeleton)
Abbreviations used in text figures
A(a, A,D, A2, A3,
Aw Divisions of the adductor mandibulae muscle
Aa Anguloarticular
Aamp Aponeurosis of adductor mandibulae and pro-
tractor hyoideus muscles
Aap Premaxillary anterior ascending process
Afl, 2, 3 Articulatory facets of anguloarticular
Ahy Anterohyal
Ama Aponeurosis of adductor mandibulae A 1 muscle
Ao Adductor operculi muscle
Apl, 2 Articulatory processes of anguloarticular
Br Branchiostegal ray (numbered)
Cm Coronomeckelian bone
Cp Coronoid process of dentary
Ctls Labial shelf connective tissue
Cts-fef Connective tissue strand linking maxilla to frontal-
ethmoid foramen
De Dentary
Dhy Dorsohyal
Do Dilatator operculi muscle
Ect Ectopterygoid
Elt Epithelial layer of lower lip
Ent Entopterygoid
Epo Epioccipital
Etb Epithelial tendinous layer
Exo Exoccipital
Fr Frontal
Fsef Frontal-supraethmoid foramen
Hy Hyomandibula
Ic Intercalar
Ihy Interhyal
Im Intermandibularis muscle
lop Interoperculum
lopf Interopercular facet
Ke Kinethmoid
Ks Keratinized lip surface
Lac Ligament linking retroarticular with anterohyal
Lai Ligament linking retroarticular with
interoperculum
Lap Levator arcus palatini muscle
Lapo Ligament linking retroarticular with preoperculum
Laq Ligament linking anguloarticular with quadrate
Le Lateral ethmoid
Lkee Ligament linking kinethmoid to ethmoid
Lkmx Ligament linking kinethmoid to maxilla
302 G. J. HOWES
Lkp Ligament linking kinethmoid to palatine
Lkpm Ligament connecting kinethmoid to premaxilla
LI Lower lip
Llm Lateral lamellae
Lo Levator operculi muscle
Lpe Ligament linking palatine to ethmoid
Lpm Ligament linking palatine to maxilla
Lqu Ligament linking quadrate to interoperculum
Ls Labial shelf
Mdp Maxillary distal process
Me Mesethmoid
Met Metapterygoid
Mfp Medial fatty partition
Mkl Medial keratinized layer of lower lip
Mlp Maxillary mid-lateral ascending process
Mlt Medial tissue layer of lower lip
Mmp Medial maxillary process
Mn Medial notch
Mv Maxillary valve
MX Maxillary
Mxf Maxillary foramen
Mxn Maxillary notch
Nc Neural complex
Ns Neural spines
Op Operculum
Pa Parietal
Pal Palatine
Pe Preethmoid
Ph Protractor hyoideus muscle
Pha Medial aponeurosis of protractor hyoideus
Phat Anterior tendinous segment of protractor hyoideus
Phav Antero- ventral segment of protractor hyoideus
Phbt Basal tendinous layer of protractor hyoideus
Phd Dorsal segment of protractor hyoideus
Phi Lateral segment of protractor hyoideus
Phlt Lateral tendon of protractor hyoideus
Phpv Postero- ventral segment of protractor hyoideus
Phy Posterohyal
Pop Preoperculum
Pro Prootic
Ps Parasphenoid
Pte Pterotic
Pty Pterygiophore (dorsal)
Q Quadrate
Qaf Quadrate anterior facet
Qlf Quadrate lateral facet
Qmb Quadrate medial buttress
Ra Retroarticular
Se Supraethmoid
Sh Sternohyoideus muscle
Sj Synarthritic mandibular joint
Sn Supraneurals (numbered)
Soc Supraoccipital
Socr Supraoccipital crest
JAW ANATOMY OF SEMIPLOTINE CARPS
303
Subtemporal fossa
Symplectic
Symplectic facet
Tendon connecting A2 with A,b
Insertion tendon of adductor mandibulae A3
Urohyal
Vomer
Ventral epithelial lip tissue
Ventrohyal
Ventral medial tissue layer of lower lip
Mmp
Mmp
IP ,MnMdp
Fig. 2 Semiplotus semiplotus, upper jaw bones. A, premaxilla, lateral view. B, maxilla, lateral
view. C, maxilla, medial view of symphysial portion; dashed line indicates meniscus. D, maxilla,
dorsal view. Scale = 10 mm.
Jaw anatomy of Semiplotus semiplotus (McClelland, 1839)
Upper jaw bones
The premaxilla (Fig. 2A) is exceedingly thin and shallow, with only a slight anterior
ascending process. The posterior portion of the bone is directed ventrally at a right angle to
its anterior part, with its tip compressed and shallowly bifurcated.
The maxilla (Fig. 2B) is deep, its dorsal border convex. The anterior ascending process
(Aap) is shallow and thick, its dorsal border notched (the palatine ligament lies across this
notch). The upper symphysial border of the ascending process is turned so that its face forms
a 45° angle to the midline. The lower portion is strongly concave and extends antero-
medially to form the symphysial part to the maxilla, syndesmotically contacting its partner.
The medial (rostral) maxillary process (Mmp, Fig. 2C) is almost square and it too makes
304 G. J. HOWES
syndesmotic contact with its partner. A mid-lateral ascending process is represented by a
slight convexity of the maxillary dorsal border (Mlp). This part of the maxilla is separated
from the posterior part by a medial notch (Mn). The posterior rim of the notch acts also as
the leading border of a broad medial triangular ledge.
Lkee
I
Cts-fef
Fsef
Lkpm
Lkmx
Lpm
Fig. 3 Semiplotus semiplotus, upper jaw articulation in oblique dorso-lateral aspect.
Scale = 5 mm.
Articulation of the upper jaw (Fig. 3)
The upper jaw bones articulate with the cranium via the kinethmoid, the palatine and the
premaxillary meniscus.
The kinethmoid (Ke) is attached via a thick ligament from its head to the ascending
process of the premaxilla (Lkpm). A broad, bifurcated ligament (Lkee) extends from the
posterior face of the kinethmoid to attach to the lateral edges of the supraethmoid and,
dorsally, to the connective tissue overlying the ethmoid region. A strand of this connective
tissue (Cts-fef) inserts onto the rim of the foramen between the frontals and the
supraethmoid. A mid-lateral ligament (Lkp) attaches the kinethmoid to the palatine, and a
ventro-lateral ligament (Lkmx) links it to the maxilla.
A paired ligament (Lpm) attaches the lateral face of the maxilla with the lateral shelf of the
palatine (Pal). The palatine, in turn, is connected with the mesethmoid by a thick medially
directed ligament (Lpe).
Interposed between the medial face of the maxilla, the preethmoid and part of the vomer is
a cartilaginous meniscus (sensu Alexander, 1966). This is thick and ellipsoidal and lies
between a ventral prominence of the premaxillary ascending process and the medial (rostral)
process (Fig. 3).
Muscles attaching to the upper jaw (Fig. 4)
The adductor mandibulae A, muscle occupies only the lower half of the cheek, the area
below the orbit and between it being filled with fatty connective tissue. The muscle is
divisible into ventral and dorsal sections. The lower section, A,b, originates from the lower
border of the preoperculum, is thin and triangular, but with its ventral border thickly
tendinous. Fibre alignment varies from 45° (ventrally) to almost horizontal (dorsally). At the
posterior tip of the maxilla, the muscle section is constricted into a bifurcated tendinous
JAW ANATOMY OF SEMIPLOTINE CARPS
Ama
305
Lap
Lo
A-|3
Fig. 4 Semiplotus semiplotus. A, lateral view of superficial jaw and cranial musculature. B, detail
of jaw muscle insertions. Scale = 5 mm.
area, the outer tendon inserting on a small lateral maxillary process, the inner on the medial
maxillary notch (see above, p. 304). The lateral border of the inner tendon (Fmd, Fig. 4B)
extends mesially to attach to the lateral face of the dentary coronoid process. The medial
fascia of adductor A,b is connected with the lateral body of adductor mandibulae muscle A2
by a thick tendon (see below).
The dorsal section of the adductor mandibulae, A,a, originates from the central area of the
preoperculum. This portion of the muscle is even thinner than A,b, being only a few fibres in
thickness at its origin. The segment is oblong, with an almost parallel arrangement of fibres.
The muscle terminates via a narrow tendon which forms the lateral segment of a broad, thick
aponeurosis (see below). The tendon of insertion reaches the first of the palatine-maxillary
ligaments.
The aponeurosis into which muscle A,a inserts is complex (Ama, Fig. 4). Laterally it is
joined to the medial face of the 1st infraorbital, and medially to the lateral ethmoid and to
the palatines. Ventro-medially the sheet has a posterior extension in the form of a tendinous
band which joins onto the anterior edge of the entopterygoid and then expands to become
continuous with the medial fascia of muscle A3 (Fig. 8A).
Upper lip and buccal cavity (Fig. 9)
Epithelial tissue covering the upper jaw is thin and lacks a cornified edge. The inside of the
mouth is bordered by a single row of leaf-like lamellae; above this row is a thin, narrow
maxillary valve (Mv). The tissue covering the antero-lateral buccal roof is formed into a near
longitudinal series of broad leaf-like lamellae (Llm). The surfaces of these lamellae are highly
papillose. The vomerine tissue is transversly rugose; there is no vomero-palatine organ
(sensu Matthes, 1963).
306
Af2
Cp
Ap2
Fig. 5 Semiplotus semiplotus, lower jaw bones. A, lateral; C, dorsal; and D, posterior views. B,
medial aspect of dentary joint. Dotted lines indicate the course of a ligamentous connection.
Scale = 10 mm.
Lower jaw bones (Fig. 5)
Each dentary (Figs 5 A & B) articulates with its partner via a complex symphysial joint (Sj).
This is basically a peg and socket joint, the central peg being on the dentary of the right side
and inserting into a deep notch in the left dentary. Above the joint there is a strong ascending
process on each bone, joined syndesmotically to that of its partner. Ventrally, the
articulation is more complex (Fig. 5B). Each dentary is extended as a ventral process which is
joined to its counterpart via a bifurcated, curved articulation. Strong, vertical ligaments
connect each facet of this joint.
The wall of the dentary is thick, its dorsal part tapering to a thin edge and curving mesially
so as to form a sloped labial surface. On the medial underside of this lip is a deep cavity into
which the intermandibularis muscle inserts. The lateral face of the dentary bears a deep ridge
which tapers posteriorly to a V-shaped process. The coronoid process is high and almost
square, its anterior edge curved outward so that when viewed dorsally (Fig. 5C) the plane of
the dorsal margin of the coronoid is coincident with that of the body axis. The posterior
border of the dentary is blunt with a rounded ventral process corresponding in position to the
coronoid process (Dvp, Fig. 5 A).
The anguloarticular (Aa, Figs 5C & D) is a large, triangular bone, its posterior dorsal edge
extending someway up the medial face of the dentary coronoid process, following that
processes' outward curvature. The articulatory surface is complex. Postero-laterally there is
a triangular facet (Afl, Fig. 5D), its posterior border extending only slightly beyond that of
the dentary. The medially directed portion of the anguloarticular is bifurcated. The dorsal
process (Apl) is thickly rounded and directed somewhat dorsally, its posterior face forming a
facet (Af2). The ventral process (Ap2) is triangular and is directed slightly ventrally. The
medial dorsal process (Apl) is separated from the outer wall of the bone by a deep, concave
channel. The posterior aspect of the anguloarticular is V-shaped and there is a large, figure of
8-shaped facet situated between the dorsal and ventral medial processes (Af3). In lateral
JAW ANATOMY OF SEMIPLOTINE CARPS
307
view, only a narrow portion of the anguloarticular is visible beyond the posterior dentary
margin.
The retroarticular (Ra, Fig. 5D) is a thick wedge lying below the ventral anguloarticular
process (Ap2). Laterally, the retroarticular is almost entirely covered by the ventral process
of the dentary and only a narrow portion is visible (Fig. 5 A).
The coronomeckelian (Cm, Fig. 5C) is an almost oblong bone with a rounded posterior
border. Its anterior portion projects medially to form a deep, sloped shelf. There is a slight
depression on the shelf into which insert the tendons of the adductor mandibulae
musculature.
Articulation of the lower jaw (Fig,. 6)
The complex articulatory surface of the anguloarticular is matched by an equally complex
quadrate surface.
The quadrate facet occupies the lateral anterior and medial portions of the bone (Figs 6B &
C). The lateral facet (Lqf) is aligned at 45° to the horizontal and apposes the medial face of
the lateral anguloarticular process (Afl). The anterior facet is also aligned at 45° and apposes
Ect i
Met
Cp
Lapo
Qaf
Fig. 6 Semiplotus semiplotus, lower jaw articulation. A, medial view of posterior jaw elements
and their connections with suspensorial and opercular bones. B, anterior and C, lateral aspects of
quadrate and inferior opercular elements. Scale = 5 mm.
308
G. J. HOWES
the central anguloarticular facet (Af3). The medial part of the quadrate is formed into a
buttress with an elliptical anterior face which apposes the dorso-medial anguloarticular facet
(Af2). The ventro-medial process of the anguloarticular (Ap2) lies below the medial quadrate
buttress and is attached to it by a thick ligament (Laq).
Ventrally a complexly divided ligament attaches the jaw to certain opercular and hyal
elements. Immediately from its point of attachment to the posterior face of the retroarticular
the ligament trifurcates and attaches, (1) to the anterior face of the interoperculum (Laio)
which is medially curved and formed into a facet; (2) to the medial face of the preoperculum
(Lapo) which is formed into a condyle; and (3) to the posterohyal via a fascial attachment to
the ceratohyal (Lac).
The lateral articulations between the anguloarticular, preoperculum and interopercular
bones is covered by a broad ligament which stems from the ventro-lateral face of the
quadrate (Lqi, Fig. 8A).
Muscles attaching to the lower jaw (Fig. 7)
The muscles having their attachment to the lower jaw are the adductor mandibulae A2 and
A3 (Fig. 7B). The upper portion of muscle A2 originates from the hyomandibula and the
Aamp
Aa
Ph B
Fig. 7 Semiplotus semiplotus, lower jaw musculature. A, lateral view of the deeper muscle
elements. B, medial view of jaw muscle insertions and their connection with hyoid musculature.
Scale = 5 mm.
JAW ANATOMY OF SEMIPLOTINE CARPS
309
lower lateral portion of the preoperculum. All its fibres insert into a medial aponeurosis.
From the lateral part of this aponeurosis a narrow tendon runs antero-ventrally into the
medial fascia of the adductor muscle (A,b (Fig. 7A). A bundle of fibres continues forward
from the aponeurosis and almost immediately inserts into a thick tendon which terminates
on the coronomeckelian bone (sesamoid articular). Insertion of A2 is via a broad tendon onto
the dorso-posterior rim of the dentary coronoid process. A thin, inner branch of the tendon
continues ventrally down the medial face of the coronoid process to insert on the
coronomeckelian bone. Attaching to this tendon is a triangular sheet of muscle fibres
crossing from A3; this most probably represents section Aw of the adductor mandibulae
series (see Fig. 7B & p. 3 1 3).
LI
Phav
Pha
Fig. 8. Semiplotus semiplotus, hyoid musculature. A, ventral aspect of head with right side
dissected to display major hyoid muscle elements. Dotted lines indicate path of the inter-
mandibularis muscle. B, detailed view of protractor hyoideus insertions on hyoid elements of the
right side. Scale = 5 mm.
310 G.J.HOWES
Adductor mandibulae muscle A3 originates partly from the hyomandibula (where it is
separated from A2 by the levator arcus palatini muscle) and partly from the deeply concave
lateral face of the metapterygoid. Its fibres run almost at right angles to the midline to join
the aponeurosis of muscle A2. Antero-ventrally, A3 inserts via a strong tendon which runs
through the deep cleft between the medial and lateral portions of the anguloarticular, to join
an aponeurosis on the coronomeckelian bone (Fig. 7B). Extending anteriorly from this
aponeurosis is a set of thin, tendinous fibres which spread along the inner aspect of the
dentary upper margin. These tendinous bands are connected by fibrous tissue to the
overlying loose, fatty connective tissue of the lower lip (see below). From the lateral part of
the aponeurosis a tendon extends ventro-posteriorly, across the medial face of the angulo-
articular, to join the lateral portion of the protractor hyoideus muscle (Tcph, Fig. 7B).
The protractor hyoideus muscle is a short and thick element divisible into dorsal and
ventral sections (Fig. 8A). The ventral portion is X-shaped, the two anterior arms separated
by a thick nodule of fatty tissue and each attaching to the medial face of the dentary. Their
posterior junction is a thick pad (Pha) from which radiate wide, tendinous bands to form the
ventral face of the muscle. These bands insert into the posterior margin of the tissue forming
the lower lip (see below). The posterior arms of the muscle are almost entirely tendinous and
attach to the branchiostegal rays, and the inner faces of the posterohyal and the interhyal.
Each dorsal section of the protractor hyoideus is broad and extends from the ceratohyal and
the 1st branchiostegal ray to join the ventrohyal. The more medial fibres run into the body of
the muscle and insert into the tendinous area just anterior to the ventrohyal (Fig. 8B).
The intermandibularis is a thin, cylindrical muscle crossing between the dentaries where it
is contained in a small cavity. The muscle separates, transversly, the anterior portions of the
protractor hyoideus (Figs 8B & 9B).
The lower lip (Fig. 9)
The lower lip of Semiplotus semiplotus is a complex structure composed of several tissue
types which extend well beyond the outer face of the dentary. Its core is a deep shelf of dense
collagenous tissue that surrounds the dentary (Ls). This labial shelf is enveloped in loose,
stratified epithelium. The tissue that covers the dorsal surface of the labial shelf is comprised
of a thick, fatty epidermal layer (Elt), and an underlying, more fibrous tissue (Mlt). The
ventral surface of this tissue adheres to tendinous bands originating from the aponeurosis
that incorporates the insertions of the adductor mandibulae and protractor hyoideus muscles
(see above). Thus, toward the lateral part of each dentary the labial epidermis becomes
united with jaw and hyoid tendinous elements and only at the symphysis is it firmly attached
to each dentary. Anteriorly, where it passes around the leading edge of the labial shelf, the
epidermis is thin and cornified, forming a chisel-edge to the jaw (Ks). Ventrally, the
epidermis is attached to the labial shelf by thin strands of connective tissue. As it passes
beneath the dentary the tissue becomes tendinous and contiguous with the ventral face of the
protractor hyoideus (see above).
Apomorphic jaw characters in Semiplotus
Before considering whether, or to what degree, the jaws of Semiplotus semiplotus may be a
derived feature, it is necessary to present a model of the plesiomorph cyprinid jaw and its
associated articular elements for comparison. The following model is modified from Howes
(1980, 1981) and is reckoned to be the plesiomorph type on grounds of its wide-spread
occurrence amongst various cyprinid taxa.
The plesiomorph maxilla has a well-developed mid-lateral ascending process with convex
anterior and concave posterior borders; the premaxilla L-shaped with a moderately
developed anterior ascending process; dentary with a vertical, high, long coronoid process
situated posteriorly; the anguloarticular with its dorsal border confluent with that of the
dentary coronoid process but not extending to cover the medial face of that process, its
JAW ANATOMY OF SEMIPLOTINE CARPS
311
Ks
Ls
Phbt
Fig. 9 Semiplotus semiplotus. A, ventral view of upper lip and buccal cavity. B, sagittal section
through the lower lip and associated hyoid musculature. Scale = 5 mm.
posterior border reaching to well beyond the posterior margin of the dentary, its articulatory
facet transversly concave. All those features are found in the jaws ofSqualiobarbus, Fig. 10.
The plesiomorph cyprinid quadrate is regarded as one essentially broadly triangular in
outline and with an anteriorly positioned articulatory facet.
The plesiomorphic condition of the adductor mandibulae A, muscle in cyprinids would
appear an undivided element having its insertion on the outer, posterior portion of the
maxilla (as in aspinine cyprinids; see Howes, 1979). Commonly in cyprinids there are two
medial sections of the adductor mandibulae. The outer (A2) inserting on the rim of the
coronoid process of the dentary, and the inner (A3) inserting on the coronomeckelian bone
(see Takahasi, 1925 : 20). Primitively, the mentalis section (Aw) of the adductor complex is
present as a well-developed element (as in the bariliines, chelines, aspinines and cultrines)
but in the majority of cyprinid taxa it is reduced or lacking entirely (see Takahasi, 1925 : 20).
According to Takahasi (1925) and Matthes (1963) the protractor hyoideus ( = genio-
hyoideus) muscle is usually attached to the 1st and/or 2nd branchiostegal rays, rarely to the
312
G. J. HOWES
Aa
Fig. 10 Squaliobarbus curriculus, jaw bones. A, maxilla. B & C, dentary in lateral and dorsal
views. Scale = 5 mm.
3rd and never to the interhyal. A tendinous lateral border of the muscle is not uncommon in
cyprinids and in many taxa the lateral part of the muscle appears as a separate element.
Semiplotus semiplotus and S. burmanicus show markedly derived states of the above
hypothesized plesiomorph jaw and muscle morphology. The following features uniquely
characterize the jaws of Semiplotus:
Upper jaw
1. Premaxilla: shallow, thin, with acute posterior curvature and reduced ascending
process.
2. Maxilla: deep, with broad anterior ascending process, reduced mid-lateral process and
depressed, notched posterior portion.
3. Bifurcated tendon of muscle A, attaching to lateral and medial faces of the maxilla.
Some amplification of this character is required. It was noted above that the plesio-
morph condition is for muscle A, to insert on the outer face of the maxilla. Most often
this insertion is tendinous but in bariliine, cheline and aspinine cyprinids it may be
musculose across a broad fascia that cover both the dorsal rim of the dentary as well as
its lateral aspect (see Howes, 1976 Fig. 23). This, often tendinous, fascia may represent
the 'ligamentum primordium' as might the connective tissue in other cyprinids which
links the maxilla to the face of the anguloarticular and enwraps the insertion area of the
lower jaw musculature. A divided tendinous insertion of the superficial adductor
muscle is present in Cyprinus, Carassius and Gobio. However, unlike the semiplotine
JAW ANATOMY OF SEMIPLOTINE CARPS 313
condition, the tendons cross over so that the insertion of the lower element (A,b) is
above that of the upper (A,a). Takahasi (1925) regarded Cyprinus and Carassius as
representing Transitional stages . . .' between the simple bariliine (Opsariichthys) and
complex gobioine (Gobio) types of insertion. Crossing of A, tendons also occurs in
Aspidoparia, a genus with bariliine affinities (see Howes, 1980), Cosmocheilus,
Cyclocheilichthys, Puntioplites and Amblyrhynchichthys. Whilst it is possible that these
four latter genera are closely related, such a relationship appears unlikely between them
and Aspidoparia. All five taxa have a short, deep ethmoid region and the mouth is
inferior in position. It is likely that the crossed tendon arrangement is functionally more
efficient, in activating jaws of this kind and has been derived independently in the
various taxa.
4. Kinethmoid attached to connective tissue overlying the ethmoid bloc.
Lower jaw
5. Dentary: with a sloped labial surface, lateral ridge, outwardly curved coronoid process,
ventral posterior process and a synarthritic joint.
6. Anguloarticular rising dorsally to cover the medial face of the dentary coronoid
process, only a small portion of the bone extending beyond the posterior border of the
dentary; the medial portion extended and formed into stout and complex articular
processes.
7. A medial tendon of muscle A2 to which is attached muscle Aw. Some amplification of
this character is required: As noted above, the common, and presumed plesiomorph
condition in cyprinids is for muscle A2 to insert on the rim of the coronoid process of
the dentary. In Semiplotus and some Cyprinion species (see p. 309) the muscle is
bifurcated (see above) with both an outer tendinous insertion on the coronoid process
and an inner one on the coronomeckelian bone. A segment of muscle stretches from the
lower part of the tendon of muscle A3 to the anguloarticular and should almost
certainly be identified as an Aw. Although a similarly developed muscle stems from the
tendon of A3 in Varicorhinus beso (p. 317) and some Barbus species (Banister, pers.
comm.) it does not insert on a medial tendon of A2 but on the medial face of the
anguloarticular.
8. Protractor hyoideus muscle connected anteriorly, via an aponeurosis, with the adductor
mandibulae insertions; its posterior part tendinous, attaching to all branchiostegal rays,
posterohyal and interhyal.
9. Lateral quadrate facet apposing lateral anguloarticulatory process. Quadrate short and
tall.
Comparisons of the jaws in Semiplotus with those in Onychostoma, Varicorhinus,
Capoeta and Cyprinion
Taxa with 'sector mouths', i.e. an inferior mouth which is wide and has an exposed cornified
mandibular cutting edge; the dentary invariably has a broad, sloping or deflected labial
surface, occur frequently amongst the Cyprinidae. Examples are Onychostoma,
Varicorhinus, Capoeta, Cyprinion, Barbus and some species of Schizothorax. Although a
cornified lower jaw occurs in Labeo and Garra it is invariably covered by a thick
labial fold.
The jaw morphology of Onychostoma, Varicorhinus, Capoeta and Cyprinion species is
remarkably similar to that of Semiplotus. The similarities may be independently derived, i.e.
they are convergent, or inherited through a shared common ancester, i.e. they are synapo-
morphic. These hypotheses are tested by comparative analysis.
Onychostoma laticeps Gunther, 1896 (Figs 11A-C)
The premaxilla is deeper than that of Semiplotus and has no distal ventral curvature, its
ventral border is also thick and lacks the sharp, thin edge of the semiplotine bone.
314
G. J. HOWES
Fig. 11 Lower jaw bones of A, B & C, Onychostoma laticeps in lateral, dorsal and posterior
views; D, Varicorhinus beso, dorsal view; E & F, Capoeta capoeta in medial and posterior views.
Scale = 5 mm.
The maxilla is deep with a broad anterior ascending process. As in Semiplotus the mid-
lateral process is curved mesially, and although there is a posterior depression of the
premaxilla there is no medial notch.
The lower jaw resembles that of Semiplotus in a number of features, cf. Figs 5 & 1 1 A. The
dentary has a bevelled labial surface, a prominent lateral ridge and a postero-ventral process.
The coronoid process differs from the semiplotine one and that in other compared taxa in
having a markedly concave anterior edge with a long dorso-anterior process. There is no
synarthritic joint. The anguloarticular is broad, its posterior face shaped into a figure-8 facet.
Although the medial portion of the bone forms an articular surface it is not posteriorly
extended, nor does it appose a separate facet.
The superficial jaw musculature is more complex than in Semiplotus. Adductor
mandibulae A,b is extensive, its lower, preopercular part almost entirely tendinous;
insertion is via a thick tendon to the lateral maxillary face. Muscle A,a is a parallel fibred
element that inserts musculosly on the maxilla above A,b. The anterior part of the adductor
JAW ANATOMY OF SEMIPLOTINE CARPS
315
muscle complex is crossed by a band of ligamentous tissue attached dorsally to the inner face
of the 1st infraorbital and ventrally to the skin covering the preoperculum (Fig. 12A). Muscle
A2 is a narrow element, its posterior part overlying A,a (Fig. 12B). The muscle's ventral
border is tendinous and insertion is via a thick tendon on to the posterior rim of the coronoid
process. The area of insertion is invested by a thick connective tissue sheath which extends
around the coronoid process and laterally attaches to the maxilla. Muscle A3 can be
distinguished from A2 by its posterior separation by the levator arcus palatini muscle, and its
separate insertion onto the coronomeckelian bone.
MX De
A2
Fig. 12 Jaw musculature of A & B, Onychostoma laticeps, A, superfcial and B, deeper muscle
layers; C, Cyprinion microphthalmum superfcial musculature. Scale = 5 mm.
The protractor hyoideus (Fig. 13 A) is complex and composed of several sections. The
muscle is organised around a tendon that originates from the interoperculum and meets its
partner at a median raphe. The anterior part of the muscle runs from the transverse tendon
to the inner rim of the dentary. The symphysial area of the muscle grades into a thick, fatty
connective tissue which becomes confluent with the lower lip tissue. Laterally, there is a
separate segment of the protractor running to the hyoid arch. A dorsal portion of the muscle
extends from the anterohyal and becomes confluent with the ventral section of the muscle.
The posterior body of the protractor extends from the branchiostegal rays to the central,
transverse tendon; it is dense, with its fibres, or strands of fibres widely separated by a fatty
and tendinous tissue. The basal layer of the muscle strongly adheres to the dermis of the
hyoid area.
316
G. J. HOWES
De /Phl Pop
Phav
LI
Phat
Pha
Phpv
Fig. 13 Hyoid musculature of A, Onychostoma laticeps in oblique ventral view. Scale = 10 mm;
B, Varicorhinus beso in oblique ventral view, and C, Capoeta capoeta direct ventral view of left
side. B & C are semi-diagrammatic.
The lower lip (Fig. 1 4)
As in Semiplotus, the core of the lip is a deep, broad labial shelf of dense tissue, its dorsal
layer (Dls) being of a more fibrous nature. Overlying, and completely free from its dorsal and
anterior surfaces is a thick tissue envelope composed of a thin basal layer (Mlt) and a thicker,
denser upper layer (Elt) whose lingual portion rises above the dentary rim. Anteriorly the
epidermal layer is transformed into a keratinized sheath. Sandwiched between the labial
shelf and the basal layer is a thin keratinized band (Mkl) that extends along the central
portion of the labial shelf. Ventral to the shelf are layers of tissue which are the counterparts
of those lying dorsally. The ventral dermal and epidermal layers (Vmlt, Velt) are, however,
firmly attached to the shelf. Unlike Semiplotus, there are no tendinous bands connecting the
dorsal lip tissue directly with the protractor hyoideus muscle and neither, apart from the
epidermis, is the ventral tissue contiguous with that muscle.
JAW ANATOMY OF SEMIPLOTINE CARPS
Mkl Elt
317
Ls
Elt
Fig. 14 Sagittal sections through the lower lips of A, Onychostoma laticeps; B, Varicorhinus
beso; C, Capoeta capoeta; D, Cyprinion macrostomum. Scale = 10 mm.
Varicorhinus beso Ruppell, 1836
The premaxilla is shallow and lacks a prominent anterior ascending process; the maxilla has
broad and thick anterior, and long, mesially curved mid-lateral processes. Unlike
Semiplotus, the posterior portion of the maxilla is not expanded, compressed or medially
notched.
The dentary (Fig. 1 ID) has an abrupt mesial curvature with a broad labial surface and a
laterally protruding process at the rictus; its coronoid process is tall and slopes outward. The
anguloarticular is short and broad, its anterior tip extending beyond the base of the coronoid
process. Unlike Semiplotus the articular facet is not developed into medial processes, and it
is transversly aligned.
The quadrate, like that in Semiplotus, is short and deep with an expanded articulatory
facet. The facet is a double ellipsoid inclined at 45° to the transverse horizontal. In contrast
to Semiplotus the facet is confined to the anterior face of the bone, no part extending to the
lateral surface.
The jaw musculature differs from Semiplotus in that adductor mandibulae muscle A! has
a single tendinous insertion on the maxillary outer face. A small Aw portion fans out from
the tendon of A3 across the anguloarticular and part of the coronoid process. The protractor
hyoideus is strongly developed, its main portion attaching tendinously to the 1st and 2nd
318
G. J. HOWES
branchiostegal rays and to the posterohyal (Fig. 13B). A lateral part attaches via a strong
tendon (almost ossiFed in large specimens) to the anterior tip of the interoperculum, a
feature not found in Semiplotus. Its anterior part becomes tendinous and fans out to attach
along the posterior rim of the dentary (Fig. 1 3B).
The lower lip, although having the same basic structure as in Semiplotus has a less
complex histological structure (Fig. 14B). The differences are a shallower labial shelf, lack of
a basal dermal tissue layer, the dermis united at its central portion with the labial shelf, and
more extensive keratinization of the anterior epidermis. There are no tendinous connections
between the lip tissue and the protractor hyoideus muscle.
Mxn -T-
Fig. 15
Premaxillae in dorsal view of A, Cyprinion watsoni; B, Capoeta capoeta posterior portion
only. Scale = 5 mm.
Capoeta capoeta Guldenstadt, 1773
The premaxilla lacks an anterior ascending process; the maxilla has a marked mesial
curvature, a thick and wide anterior and a shallow mid-lateral ascending process. The
posterior arm of the maxilla, as in Semiplotus, is compressed and medially notched (Fig 1 5).
The dentary has the same abrupt medial curvature and antero-ventrally sloped labial
surface as in Varicorhinus beso (Fig. HE). Unlike Semiplotus, Onychostoma and
Varicorhinus, the coronoid process is aligned at 45° to the midline, so that it is set at the
curve of the dentary. The ventro-posterior surface of the dentary is broad, as is the depressed
anguloarticular which overlies it. The articulatory facet is ovate and transversly aligned (Fig.
11F).
The jaw musculature of Capoeta is similar to that of Semiplotus and Varicorhinus.
Adductor mandibulae A, is a single muscle with a central anterior aponeurosis; insertion is
via a thick tendon onto a lateral maxillary process. A thin tendinous strip of tissue runs from
the dorso-anterior part of the muscle to insert on the connective tissue covering the medial
face of the maxilla. Muscle A2 inserts entirely on the posterior border of the dentary coronoid
process; a separate group of fibres leaves the main tendon to run ventrally across the medial
coronoid face. Tendon A3 inserts on the coronomeckelian bone.
The protractor hyoideus is extensively tendinous in its posterior part and it is attached to
all the hyoid elements except the interhyal. There is no lateral portion stemming from the
interoperculum as in Varicorhinus. Anteriorly the left and right arms of the muscle are
widely separated so that the intermandibularis is exposed (Fig. 1 3C).
The lower lip differs from that in the other taxa compared here in having a narrow labial
shelf, no layered organisation of the tissue overlying the shelf, and this tissue continuous with
JAW ANATOMY OF SEMIPLOTINE CARPS
319
that lining the mouth. The outer, keratinized sheath lies along the upper edge of the labial
shelf rather than around its anterior face.
Cyprinion Heckel, 1843
In Cyprinion species, the premaxilla usually lacks an ascending process, and when one is
present it is but slightly developed. Maxillary shape is interspecifically variable and in only
one species, C. watsoni, does it closely resemble Semiplotus in having a reduced, medially
directed mid-lateral ascending process and a depressed, medially notched posterior arm (Fig.
15).
The lower jaw of C. macrostomum and C. microphthalmum resembles that of Semiplotus
more closely than does that of any other Cyprinion species. This resemblance is in the
B
Fig. 16 Lower jaw bones (right side) of A & B, Cyprinion acinaces in dorsal and medial views; C,
Cyprinion watsoni in dorsal view; D & E, Cyprinion microphthalmum in dorsal and posterior
views; F, quadrate in lateral view; G & H, Cyprinion macrostomum in dorsal and posterior
views; I, quadrate in anterior view; J & K Cyprinion kais in dorsal (right jaw bone) and lateral
(left jaw bone) views. Scale = 5 mm.
320 G. J. HOWES
curvature of the dentary, a complex synarthritic joint and the broad articular surface of the
anguloarticular. There is a slight medial process, equivalent to process Apl in Semiplotus,
separated from the lateral part of the bone by a deep cleft. Both Cyprinion species also
possess a quadrate bearing part of its articulatory facet laterally (Fig. 16F). Cyprinion watsoni
has a more elongate jaw than C. microphthalmum and C. macrostomum, but the angulo-
articular has a similar broadening of its facet.
With the exception of C. microphthalmum the muscles attaching to the upper jaw are
undivided or only partially divided. In that species the adductor mandibulae A,a is separated
from A,b for almost its entire length but shares a common insertion with it on both the outer
and inner aspects of the maxilla (Fig. 12C). The lower jaw muscles are also interspecifically
variable with respect to their proportions and fibre orientation, but their insertions are
essentially similar. There is some variability in the size of muscle Aw. In C. microphthalmum
and C. macrostoma, as in Semiplotus it inserts on a medial tendon of muscle A2.
The lower jaws of C. acinaces, C. milesi and C. watsoni are less derived with respect to
their labial and articulatory surfaces but all have a synarthritic joint of varying complexity,
the weakest being in C. acinaces (Fig. 16A). The jaw of C. acinaces is regarded as the most
plesiomorph of all Cyprinion species with respect to its moderately expanded labial surface,
mid-laterally placed coronoid process and small, transverse articulatory facet. The lower jaw
of C. kais is deeper posteriorly than in any other Cyprinion species, and its labial surface is
much narrower (Fig. 16J); as in Semiplotus, the anguloarticular covers a substantial area of
the medial face of the dentary coronoid process. In all Cyprinion species, the anguloarticular
extends posteriorly only marginally beyond the dentary.
The lower lip of Cyprinion is of the same layered type found in Semiplotus (Fig. 14D),
with that of C. macrostoma having the greatest resemblance in terms of dimensions of the
labial shelf, tissue layering, and tendinous connections with the protractor hyoideus muscles.
The degree of keratinization is interspecifically variable but reaches its greatest development
in C. macrostomum.
Onychostoma, Varicorhinus and Capoeta all lack the Semiplotus features of a synarthritic
mandibular joint, medial anguloarticular processes, lateral quadrate facet, aponeurotic
connection of the adductor mandibulae and protractor hyoideus muscles, interhyal
connection of the protractor hyoideus, and complexity of lower lip structure. Onychostoma
shares with Semiplotus a lateral dentary ridge and posteroventral process, but the nature of
the coronoid process and the lack of the apomorphies cited above favour the hypothesis that
the derived jaw characters held in common have been independently acquired. Likewise, the
sector type of mouth morphology of Varicorhinus and Capoeta is believed to be a convergent
feature.
The phyletic relationships of Onychostoma, Varicorhinus and Capoeta have yet to be
determined, but other apomorphies (absent in Semiplotus and Cyprinion) suggest the
following affinities: Onychostoma is related to Ageniogarra and possibly to a group of
Chinese 'Schizothorax*; Varicorhinus forms a close relationship with certain African Barbus
(Banister, in preparation); Capoeta is the likely sister-group to Cyprinus and Carassius (see
Howes, 1981).
As noted above all Cyprinion species possess a synarthritic mandibular joint, aponeurotic
connection of the adductor mandibulae and protractor hyoideus muscles and similarly
developed lower lip structure. Synapomorphies other than those of the jaws also suggest a
close relationship between Semiplotus and Cyprinion.
Other apomorph characters of Semiplotus
In addition to the jaws, Semiplotus species are characterized by other and presumed
apomorphic features.
1 . Condylar articulation of the interhyal with the symplectic. The usual condition in
cyprinids is for the symplectic to be an elongate bone. It is rarely short and deep, and
JAW ANATOMY OF SEMIPLOTINE CARPS
321
only in a minority of taxa is the posterior face concave and in articulation with the
interhyal. The common condition of the interhyal is a lamellate bone, sometimes
incompletely ossified.
In Semiplotus the symplectic is short and buttress-like (Figs 6 & 1 7A), its posterior
surface is concave and articulates directly with the interhyal, thereby forming part of
the ball and socket joint between the two elements. The interhyal is short and
cylindrical, and articulates with the dorsal surface of the posterohyal via a cartilaginous
meniscus.
Br1
Fig. 17 Hyoid and suspensorial elements of A & B, Semiplotus semiplotus, medial and
dorsal views of posterior hyoid bar; C, Cyprinion macrostomum, medial view of posterior hyoid
bar; D & E, Varicorhinus beso, medial and dorsal views of posterior hyoid bar; F-I, symplectic
bones of F, Cyprinion microphthlamum; G, C. kais; H, C. acinaces; I, C. watsoni; J, Cyprinion
kais, anterior view of 1st branchiostegal ray and hyoid bar. Scale = 3 mm.
322 G. J. HOWES
Amongst the taxa considered here, only in Onychostoma and some species of
Cyprinion does the symplectic approach that of Semiplotus in size and shape. In
Onychostoma, the condylar surface of the bone faces ventrally whereas in Semiplotus
and Cyprinion it is angled postero- ventrally. The interhyal in all these genera, and in
Varicorhinus, is hypertrophied and articulates with the dorsal aspect of the postero-
hyal, but since there are no other synapomorphies of the hyoid elements shared by
Onychostoma, Varicorhinus and Semiplotus, these are considered to be independently
derived features (see p. 320).
2. Ventral and lateral articulation of the 3rd branchiostegal ray with the posterohyal (Figs
17A & B). Plesiomorphically the 3rd branchiostegal is attached ligamentously to the
lateral face of the posterohyal. In Semiplotus, the posterohyal is short and thick, and
the branchiostegal ray expanded medially so as to form a shelf underlying the ventral
border of the posterohyal with whose posterior portion it articulates. The anterior part
of the branchiostegal shelf is attached ligamentously to the medial face of the postero-
hyal whilst the proximal lamellate area of the branchiostegal ray is attached to the
lateral face of the hyal.
A slight medial shelf is present on the 3rd branchiostegal rays of Varicorhinus and
Capoeta. It is, however, barely developed in these taxa and the posterohyal is not
distally expanded as in Semiplotus (Fig. 17). Such an expansion of the branchiostegal is
lacking in Onychostoma. Only in Cyprinion is there a substantial 3rd branchiostegal
shelf, which in C. macrostomum is developed much as in Semiplotus (Fig. 1 7).
3. Ventrally directed supraethmoid and hypertrophied vomer (Figs 1 8A-C). Although the
rostral curvature of the ethmoid bloc is variable in the Cyprinidae (see Howes, 1980)
nowhere is it more extreme than in Semiplotus where the supraethmoid is almost
vertically aligned. An apparent correlate of this condition is the hypertrophied vomer,
whose enlargement also results in a steep angle forming between it and the para-
sphenoid (Fig. ISA).
In both Onychostoma and Varicorhinus the ethmoid is strongly curved ventrally and
there is a corresponding hypertrophy of the vomer. In these genera the ventrally
directed part of the supraethmoid is confined to the lower part of the bone. In
Cyprinion macrostomum the curvature of the supraethmoid more closely resembles the
condition in Semiplotus where the entire bone slopes ventrally.
4. Frontal- supraethmoid foramen (Figs 18B-C). Synapomorphic for Semiplotus semi-
plotus and S. burmanicus is a foramen between the frontals and the supraethmoid (see
p. 304). The posterior border of the foramen indents each frontal to form a notch in
much the same way as the antero-medial supraethmoid notch occurs in other cyprinids.
Likewise, the frontal notch of Semiplotus performs a similar function, namely in
providing a hold-fast for the kinethmoid ligaments. Attachment of kinethmoid
ligaments to the frontal instead of the supraethmoid is unknown in any other cyprinid.
5. Elliptical subtemporal fossa with major contribution from the pterotic (Fig. 18D). The
plesiomorph subtemporal fossa is shallow with a round or oval outline (see Howes,
1981). In Semiplotus the fossa is deep and markedly ellipsoidal, its long axis transverse.
Only in Onychostoma does the subtemporal fossa approach the shape of that in S.
semiplotus and S. burmanicus, but it lacks the extreme depth of those species.
Typically the posterior wall of the subtemporal fossa in cyprinids is formed princi-
pally by the exoccipital, with a minor contribution from the pterotic (see, for example,
Howes, 1978, Figs 6 & 33; 1980, Fig. 13). In Semiplotus the greater part of the posterior
wall is formed by the pterotic (Fig. 18D); only in Cyprinion is there a similarly
substantial contribution of the pterotic.
6. Proximal part ofsupraoccipital crest formed by the parietals (Fig. 1 9 A).
Semiplotus species have a high supraoccipital crest; the parietals are short and slope
upwards, their medial portions extending posteriorly and embracing the base of the
supraocipital crest. With the exception of Cyprinion, in no other cyprinid taxon, even
JAW ANATOMY OF SEMIPLOTINE CARPS
So
323
Se
Fig. 18 Semiplotus semiplotus, cranium. A, lateral view of ethmo-vomerine and orbital areas; B,
dorsal & C, anterior views of ethmoid region; D, ventral view of posterior cranial area; E & F,
basioccipital process in lateral view of E, S. semiplotus and F, S. burmanicus. Scale = 10 mm.
those with a relatively high supraoccipital crest, does the parietal extend postero-
medially. In Cyprinion there is some variability in the length of the parietal, but in all
species it extends postero-medially to form part of the supraoccipital crest (Figs
19B-C).
Reduced basioccipital process and masticatory plate (Figs 1 8E & F).
Although a reduction of the basioccipital process and masticatory plate occurs in
several groups of cyprinids (see Howes, 1981), in none is the reduction so extreme as in
Semiplotus. In S. burmanicus, the basioccipital (masticatory) plate is entirely absent
and the ventral (aortic) ossification is transversly convex. Only in some Cyprinion
species (C. kais and C. microphthalmum) is a truncated basioccipital process associated
with marked reduction of the ventral plate.
Interoperculum-preoperculum articulation (Figs. 6B-C).
In Semiplotus the anterior tip of the preoperculum is formed into a condyle and is
curved medially so as to overlie the anterodorsal part of the interoperculum; the
324
G. J. HOWES
Socr
B
Fig. 19 Dorsal posterior cranium of A, Semiplotus semiplotus; B, Cyprinion macrostomum; C,
Cyprinion kais. Scale = 5 mm.
anterior tip of the interoperculum is formed into a facet. The two elements are firmly
articulated and the medial curvature of the preoperculum beneath the quadrate appears
to reinforce the rigidity of the quadrate-lower jaw articulation. The interoperculum
also bears a medial facet with which the rounded tip of the posterohyal articulates.
In Varicorhinus there is a similar development of the terminal point of the intero-
perculum, but only in Cyprinion do both elements articulate as in Semiplotus (Fig.
161). A medial interopercular facet is variously developed in the Cyprinidae, but in
Cyprinion alone is it of similar depth and position.
Development of the supraneurals and articulation with 1st dorsal pterygiophore (Fig. 20).
The supraneurals of Semiplotus are hypertrophied and each is in contact with its
respective neural spine. The 'neural complex' of the Weberian apparatus is upright and
has an almost straight anterior edge; dorsally it contacts the posteriorly directed supra-
occipital process. This is contrary to the usual cyprinid condition where the anterior
border of the neural complex is concave, giving it an axe-shaped appearance. In
Semiplotus semiplotus there are 5 or 6 supraneurals, each articulating with its neigh-
bour; the posterior element is bifurcated and overlaps the anterior margin of the 1 st
dorsal pterygiophore. Semiplotus burmanicus has 7 supraneurals, all somewhat thinner
than in S. semiplotus but similarly and sequentially joined, each bearing a vertical ridge
laterally. The 1st supraneural is narrowly separated from the neural complex.
Although enlarged supraneurals occur in other cyprinid taxa they are never so tall as
those in Semiplotus semiplotus. Apart from S. burmanicus, interlocking of the last
supraneural with the 1st pterygiophore occurs only in Cyprinion macrostomum, and a
straight-edged neural complex is shared only with Cyprinion microphthalmum.
JAW ANATOMY OF SEMIPLOTINE CARPS
Sn6
325
Nc
Sn1
Fig. 20 Anterior vertebral column of A, Semiplotus semiplotus; B, S. burmanicus. Scale = 5 mm.
The relationship between Semiplotus and Cyprinion
In addition to the jaw synapomorphies (p. 320), Semiplotus shares with Cyprinion species a
series of derived hyoid, cranial and vertebral features (enumerated above as 1-9), some of
which are present in all Cyprinion (I, 2, 5, 6, 8), others in only some species of that genus
(3,7,9).
At present, the genus Cyprinion cannot be defined on the basis of even a single autapo-
morphy. Derived characters such as the expansion of the proximal portion of the pelvic fin
rays, interpelvic papillate flaps (see Banister & Clarke, 1 977) and a naked pre-dorsal ridge
(see Mirza, 1969) although synapomorphic for some species do not occur in all members of
the genus. If, however, the concept of Cyprinion is expanded so as to include Semiplotus,
then the taxon so formed is recognisable as a monophyletic unit on the basis of several
synapomorphies (the principal one being the presence of a synarthritic dentary joint), and
the problem posed by those apomorphies shared by Semiplotus and some Cyprinion species
is also overcome. Thus (following Heckel, 1843; see p. 331), Semiplotus is regarded as a
junior synonym of Cyprinion and any references to Cyprinion from here onwards must be
taken to include Semiplotus semiplotus and S. burmanicus. (A synopsis of included taxa is
given on p. 331.)
326 G. J. HOWES
Discussion
Apomorph character distribution in Cyprinion species
If the jaws of Cyprinion semiplotum are seen as the terminal stage of increasing complexity,
then there is a possibility that an evolutionary or 'transitional sequence' of jaw types might
reflect the pattern of interspecific relationships within the genus Cyprinion. However, such
an arrangement of increasingly derived jaw morphology (in terms of hypertrophy or
reduction of various elements) is not congruent with polarized morphoclines in other
characters amongst Cyprinion species.
Jaw synapomorphies possessed by C. semiplotum and C. burmanica unite them with other
Cyprinion species in the following patterns:
(i) A greatly increased complexity of the mandibular synarthritic joint, a broad angulo-
articular with a medial process and a cleft between the lateral and medial faces of the
bone. Shared with C. macrostomum, C. microphthalmum and C. watsoni.
(ii) A lateral quadrate facet and a medial A2 tendon. Shared with C. macrostomum and C.
microphthalmum.
(iii) A medial maxillary notch. Shared with C. watsoni.
(iv) A divided insertion of adductor mandibulae A, muscle. Shared with C. micro-
phthalmum.
Any 'transitional series' present amongst this character complex is interrupted by the
distribution of characters (iii) and (iv). The following synapomorphies uniting C. semi-
plotum and C. burmanica among other Cyprinion species display incongruent patterns
when set against those of the jaw:
(v) A deep symplectic and a truncated basioccipital process with a reduced masticatory
plate. Shared with C. kais and C. microphthalmum.
(vi) A straight-edged neural complex. Shared with C. microphthalmum.
(vii) Articulation of the last supraneural with the 1 st pterygiophore.
Shared with C. macrostomum.
Synapomorphies which are not present in C. semiplotum and C. burmanica unite
the following species:
(viii) 1st branchiostegal ray articulating ventrally with the ceratohyal through a
long medial process (Fig. 17). Shared only by C. macrostomum and C. kais.
(Cyprinion kais, it should be remembered has neither of the derived jaw
characters (i) nor (ii) which C. macrostomum shares with other species).
(ix) Pelvic flaps and papillate interpelvic region. Shared by C. watsoni, C. acinaces and
C. milesi. The two latter species are conservative in their cranial and jaw
morphologies (see p. 320) and are thought to comprise the plesiomorph lineage
within the genus. Thus, unlike C. watsoni, C. acinaces and C. milesi share none of
those characters that distinguish the more derived lineage (see (i) and (ii) above).
In virtually all monophyletic assemblages of cyprinid fishes so far identified there are sets
of incongruent apomorphies, a state of affairs which makes it impossible to apply a rigidly
dichotomous system of relationships. Such incongruencies usually can be ascribed to some
form of homoplasy, particularly when the characters in question occur in outgroups, as for
example the medial maxillary notch present in Cyprinion semiplotum, C. burmanica, C.
watsoni and also in Capoeta. Here the most likely explanation for its presence in Capoeta
and Cyprinion is one of parallelism as no other synapomorphies unite Capoeta with
Cyprinion; see p. 320. However, its distribution within the three Cyprinion species can only
be accounted for most parsimoniously by assuming its loss in two other species (C. micro-
phthalmum and C. macrostomum).
Other character sets also reveal incongruency when treated dichotomously. For example,
in Cyprinion kais one must postulate the loss of all jaw synapomorphies plus the parallel
evolution in this species and in C. semiplotum, C. burmanica and C. microphthlamum of
similarly derived symplectic and basioccipital characters. Furthermore, the loss of a
JAW ANATOMY OF SEMIPLOTINE CARPS
327
supraneural-pterygiophore articulation must be assumed in C. kais and C. micro-
phthalmum, and interpelvic papillate flaps must be treated either as a plesiomorph character
or assumed to be secondarily derived in C. watsoni. Such solutions result in a confusing
mixture of assumed parallelism and reversal, with the acceptance of one alternative being no
more parsimonious than the acceptance of another; see cladogram, Fig. 2 1 .
Bremer & Wanntorp (1979) have discussed the difficulties posed by conflicting synapo-
morphies and show that the situation may best be displayed as reticulate cladograms. Such a
device is used here to show character distribution in Cyprinion (Fig. 22).
Although sister species cannot be identified with this synapomorphy scheme, both
apomorph character weighting and the simplistic, essentially numerical approach of using
Fig. 21 Most parsimonious cladogram of Cyprinion characters. Synapomorphies; nodes 1,
Cyprinion synapomorphies (including expanded pelvic fin rays); 2, jaw synapomorphies (see
text, p. 320); 3, lateral quadrate facet, medial A2 tendon, truncated basioccipital, deep
symplectic; 4, upright neural complex; 5, jaw and cranial synapomorphies (see text); 6, inter-
pelvic papillate flaps; 7, method of articulation of 1st branchiostegal with ceratohyal (see text,
p. 326). Parallelisms and reversals; lineage A, scaleless interpelvic region (parallel to 6); B, loss
of deep symplectic and truncated basioccipital in C. macrostomum, parallelism of
pterygiophore-supraneural articulation to C. semiplotus and C. burmanicus; C, loss of jaw
synapomorphies in C. kais; D, loss of supraneural-pterygiophore articulation in C.
microphthalmum; E, loss of pelvic fin ray expansion in C. semiplotus and C. burmanicus.
328
G. J. HOWES
Fig. 22 Reticulate synapomorphy scheme for Cyprinion characters Level 1, Cyprinion
synapomorphies (see text, p. 325); 2, expanded pelvic fin rays; 3, increased complexity of
synarthritic joint, broadened anguloarticular with medial process Ap3; 4, lateral quadrate facet,
medial A2 tendon, interopercular-preopercular articulation; 5, jaw and cranial synapomorphies
(see text, p. 312). Conflicting synapomorphies: (a) interpelvic papillate flaps; (b) articulation of
1st branchiostegal ray; (c) upright neural complex; (d) divided adductor mandibulae A,; (e)
medial maxillary notch; (h) supraneural-pterygiophore articulation. Black boxes indicate the
presence of that character on a particular lineage, a break in the horizontal indicates its absence.
JAW ANATOMY OF SEMIPLOTINE CARPS 329
the greatest number of synapomorphies as an assessment of relationship is avoided
(Tarsitano & Hecht, 1980: 178). Likewise, the alternative approach, using unresolved
polychotomies, serves only to indicate that there is an absence of characters rather than the
presence of conflicting ones.
It may be argued that the choice of character sets employed here is itself a weighting
procedure, and that the use of other sets would reveal dichotomous sister-group relation-
ships. The counterclaim to these arguments is that the characters employed are those whose
intraspecific variability is known, and indeed are the only definitive characters recognizable
at this level of investigation. Cyprinids are 'notorious' for their conservative characters (see
for example Regan, 1911). Banister (1980) held that this conservatism '. . . coupled with the
occasional small saltatory morphological changes . . . has the effect of making it difficult, if
not impossible to distinguish between parallelism and the sharing of characters derived
through common ancestry'. In each of the cyprinid groups so far identified as monphyletic
(the aspinines, chelines, bariliines and abramines) one lineage appears as the bearer of a large
number of autapomorphies and thus leaves a considerable 'morphological gap' between it
and the more conservative corpus of lineages comprising that particular group. Examples are
Macrochirichthys + other chelines and Hypophthalmichthys + other abramines (see Howes,
1979; 1981). Such also appears to be the case within Cyprinion where C. semiplotum and C.
burmanica represent a highly derived lineage characterized by many jaw synapomorphies.
Presumably the accumulation of synapomorphies in one such lineage is the result of a
cumulative series of speciation events, lacking in the other, more generalized lineages (i.e.
'the occasional small saltatory changes' of Banister, 1980).
Whether the reticulate pattern of relationships is an artifact resulting from incomplete
character analysis or a reflection of polymorphic descent through populations of the intra-
group lineages is a problem as yet unresolved.
Evolution of the Cyprinion jaw and the 'sector mouth'
Even though no 'transitional series' of jaw types emerges from the reticulate pattern
presented (Fig. 22), particular synapomorphies indicate a possible evolutionary course
leading to the specialized morphology of the Cyprinion semiplotum-C. burmanica lineage.
Comparison of these apomorphies with similar but independently derived characters in
other cyprinid lineages indicates those features which are prerequisites for the development
of a sector mouth.
In the upper jaw a decisive step towards a sector mouth appears to have been taken with
the reduction of the premaxillary ascending process. Matthes (1963) remarks that
reduction of this process probably occurred independently in several genera and is not there-
fore indicative of close relationship. Matthes was referring particularly to similarities
between Varicorhinus, Labeo and Garra. Whilst this study endorses Matthes' view that this
particular feature is not necessarily a synapomorphy for all three genera, it does not refute its
possible synapomorphic status in Labeo and Garra, which share other derived characters
(Reid, unpublished thesis).
At first sight it would seem that reduction of the premaxillary processes should lead to
restricted protrusibility of the upper jaw. However, this may be compensated for by a
pronounced downward curvature of the ethmoid bloc as seen in Cyprinion and
Onychostoma and, to a lesser extent, in Varicorhinus and Capoeta. The depressed ethmoid
region in Labeo and Garra possibly fulfills the same function.
The reduced and medially curved mid-lateral ascending maxillary process of Cyprinion
semiplotum appears otherwise only in Onychostoma. In both taxa this feature is apparently
correlated with an extensive area of connective and tendinous tissue attaching to the inner
face of the 1st infraorbital (see p. 305). The medial notch on the posterior arm of the maxilla
in some Cyprinion species and in Capoeta performs a similar function as a hold-fast for the
thick tissue connecting the maxilla with the coronoid process of the dentary (see p. 305).
Concerning the Cyprinion lower jaw, the most important evolutionary innovation appears
330 G. J. HOWES
to have been the development of a synarthritic mandibular joint. Such a joint usually
suggests some degree of lateral movement, as in the 'knuckle joint' of cynodontine
characoids (Howes, 1976), but in the case of Cyprinion it may act as a compensatory device
to the torque induced by the contraction of the thickly ligamentous protractor hyoideus
muscle. The absence of any lateral jaw movement is also suggested by the strongly produced
lateral and medial anguloarticular processes which provide a rigid coupling with the
quadrate.
Changes associated with increased medial curvature of the dentary appear to be manifest
in the outward slope of the coronoid process so that the axis and insertion angle of the
adductor mandibulae A2 and A3 muscles remain unchanged. This is contrary to the
condition in Capoeta where the coronoid process and muscle fibres are set at an angle to the
body axis. In Cyprinion there is a noticeable change in the relative position of the coronoid
process from its mid-lateral point in the plesiomorph C. acinaces to a posterior position in C.
semiplotum. This apparent positional shift is due to morphological changes in the
surrounding elements, i.e. broadening and posterior shortening of the anguloarticular and
the medial curvature of the dentary (Fig. 1 6 A-E, G-K).
The rigid articulation of the lower jaw in Cyprinion semiplotum and C. burmanica
appears to restrict its vertical movement and it seems merely to act as a firm platform for the
highly mobile lip.
The lower lip of Cyprinion differs from that in other taxa in being a mobile structure; the
epidermis is so loosely attached to the underlying labial shelf that it is free to move antero-
posteriorly, and its mobility appears to be controlled by the action of the protractor hyoideus
complex via dorsal and ventral tendinous linkages. This additional function of the protractor
hyoideus is also indicated by a multi-directional joint between the hyoid and the
suspensorium.
Thus, in the more derived Cyprinion species, increased complexity of the jaw-quadrate
articulatory surfaces and of the lower lip is correlated with increased mobility of the hyoid
bar and hypertrophy of the protractor hyoideus.
Where it occurs, a sector mouth appears to have the same basic function i.e. for scraping or
ploughing epilithic material (see Matthes, 1963). Certainly, there are some 'necessary design
components' common to all lineages of taxa with a sector mouth, viz. marked medial
curvature of the anterior portion of the dentary; a broadened anguloarticular and barely
extending beyond the posterior border of the dentary; greater mobility of the hyoid-
suspensorial joint; 3rd branchiostegal ray shifting ventrally to the posterohyal, and an hyper-
trophy of the protractor hyoideus musculature.
Apart from Cyprinion such specializations can be found among Barbus-Varicorhinus
species. Doubtless, cranial and vertebral apomorphies (deepening of the posterior cranial
region and increased rigidity of the anterior vertebral column) form a functional unity
contributing to epilithic feeding activity. Just how these components interact functionally
with those of the jaws, the hyoid and branchial arches, are problems to be solved.
An earlier remark (Howes, 1981) that the '. . . morphology of the lower jaw is possibly the
most variable unit in Cyprinidae . . .' is borne out by this study. That the cyprinid jaw
appears to be so adaptable suggests that, as in cichlids, the pharyngeal bones perform a
dominant functional role of food manipulation and mastication, so freeing the jaws for
specialized food gathering. No comparative experimental functional data are available for
cyprinid taxa and it is not known whether the pharyngeal apparatus is as versatile as that of
cichlids and whether it exerts a primary dominant influence on jaw modification. Liem
(1980) has pointed out that in cichlids epilithic feeding patterns are of a kind requiring
specializations that exceed those of simple inertial suction feeding. Certainly this appears to
be true for sector-mouthed cyprinids as well, where (in the more derived forms) upper jaw
protrusion is minimal and the manipulative function of the jaw in food gathering has been
transferred, in part, to the hyoid arch.
JAW ANATOMY OF SEMIPLOTINE CARPS 33 1
Synopsis of Cyprinion species
The synonymies include only the original citation of the species.
CYPRINION deckel, 1843
Semiplotus Bleeker, 1863
Scaphiodonichthys Vinciguerra, 1889
Scaphiodontopsis Fowler, 1934
The reasons for synonymising Semiplotus with Cyprinion have been explained in this paper
p. 299).
Scaphiodonichthys was separated from 'Semiplotus' on the differences in the number of
branched dorsal fin rays (9-12 in Scaphiodonichthys, 20-25 in 'Semiplotus'). Smith
(1945 : 206) quotes from a letter he received from Dr S. L. Hora who suggested that inter-
mediate forms were '. . . yet to be discovered with a number of branched rays intermediate
between 12 and 20. When such forms are found Semiplotus and Scaphiodonichthys will have
to be regarded as congeneric . . .' To my knowledge no such 'intermediate' forms have been
found. The existence of synapomorphic osteological and myological characters in species of
both 'Scaphiodonichthys'' and Cyprinion indicates their generic unity (see p. 312 for a list of
characters). When viewed as species of Cyprinion, then 'intermediate forms' linking
''Semiplotus'' and 'Scaphiodonichthys' are in fact present, and indeed the number of branched
dorsal fin rays does range from 10-25.
Species occurring east of the Himalayas
Cyprinion semiplotum McClelland, 1839
Cyprinus semiplotus McClelland, 1839 Asiat. Reschs. 19 : 374
Cyprinion semiplotus Heckel, 1843 in Russegger's Reisen 1 : 1015
Semiplotus semiplotus Bleeker, 1 863 Atlas Ichth. 3 : 25
Semiplotus mcclellandi Bleeker, 1863 Atlas Ichth. 3 : 25
Semiplotus cirrhosusChaudhuri, 1919 Rec. Ind. Mus. 16 : 280
DISTRIBUTION. Nepal, north Bengal (see Hora, 1937; Hora & Gupta, 1940).
Cyprinion modestum Day, 1870
Semiplotus modestus Day, 1870 Proc. Zool. Soc. Lond. : 101
Day (1870) considered this species as '. . . intermediate between the genera Semiplotus and
Cyprinion'. Unlike C. semiplotum but in common with other Cyprinion species, C.
modestum has a serrated dorsal fin spine. No specimens have been examined in the course of
this study (the presumed types are in Calcutta) but according to Hora (1937 : 46) Day's
species is 'quite distinct'.
DISTRIBUTION. Akyab Hills, Upper Burma.
Cyprinion burmanica Vinciguerra, 1889
Scaphiodonichthys burmanicus Vinciguerra, 1 889 Ann. Mus. Civ. Storia Nat. Geneva (2) 9 : 285
Scaphiodontopsis acanthopertus Fowler, 1934 Proc. Acad. Nat. Sci. Philad. 86 : 119
Onychostoma macroacanthus Pellegrin & Chevey, 1936 Bull. Soc. Zool. France 6\ (1) : 18-27
Taki (1975) demonstrated an overlap of lateral line scale numbers between C. burmanicus
and C. acanthopterum and stated that the two species were distinguishable only by the
number of dorsal fin rays. I find that in a sample of 16 specimens identified as C. burmanica
(28-65 mm SL uncat. Smithsonian Coll. Upper Thailand), there are nine specimens with 1 1
and seven with 10 branched dorsal rays. There are thus no meristic differences separating the
two 'species'. Since I can find no morphological differences between the syntypes of C.
burmanica and the series of C. acanthopterum studied by Smith (1945), I consider the
species to be synonymous.
332 G. J. HOWES
DISTRIBUTION. Burma, Thailand, Laos, Vietnam.
Species west of the Himalayas
Despite the revision of Berg (1949) and the work of Mirza (1969), many taxonomic problems
remain concerning the species in this area. As yet, no author has examined all the nominal
types. Also, too few specimens have been examined to judge the degree of intraspecific
variability in certain characters.
Cyprinion acinaces Banister & Clarke, 1 977
Cyprinion acinaces Banister & Clarke, 1977 J. Oman Studies : 123-126
DISTRIBUTION. Saudi Arabia
Cyprinion macrostomum Heckel, 1843; type species of the genus
Cyprinion macrostomum Heckel, 1843 Ichthyologiein Russegger's Reisen 1 : 1065
?Cyprinion neglectum Heckel, 1 846 Die Fische Persiens in Russegger's Reisen 2 (3) : 223
?Cyprinion tenuiradius Heckel, 1 846 Die Fische Persiens in Russegger's Reisen 2 (3) : 26 1
Berg (1949) included C. neglectum in synonymy without having seen the type and regarded
C. tenuiradius as a valid species. However, from Heckel 's figure C. tenuiradius would seem
to be simply a 'variant' of C. macrostomum.
DISTRIBUTION. Tigris-Euphrates (see Banister, 1980).
Cyprinion kais Heckel, 1 843
Cyprinion kais Heckel, 1843 Ichthyologiein Russegger's Reisen 1 : 1066
?Cyprinion cypris Heckel, 1843 Ichthyologie in Russegger's Reisen 1 : 1067
Berg (1949) included C. kais in the synonymy of C. macrostomum on the grounds of minor
variation in position of the dorsal fin with respect to the pelvic fin insertion (the character
Heckel had used to separate the species). Although Berg (1949) recognised differences in
mouth shape between the taxa, he obviously considered them to be of little taxonomic worth.
I have seen the types of neither C. macrostomum nor C. kais, and the recognition of these
species is based on a comparison of Heckel's (1843) figures with two recognisably distinct
taxa represented in the BMNH collections. In C. kais the mouth is small, with the
keratinized lip of the lower jaw prominent and more upwardly directed than in C. macro-
stomum. Other differences involving the morphology of the jaws and hyoid elements are
given elsewhere in this paper.
From Heckel's (1 843) figure (Fig. 3, pi. 7) it seems likely that C. cypris is synonymous with
C. kais rather than with C. macrostomum as indicated by Berg (1949), and that it represents
a juvenile specimen in which the keratinization of the jaw is still incomplete.
DISTRIBUTION. Tigris-Euphrates.
Cyprinion microphthalmum Day, 1880
Scaphiodon microphthalmus Day, 1880 Proc. Zool. Soc. Lond. : 227
?Scaphiodon muscatensis Boulenger, 1887 Proc. Zool. Soc. Lond. : 665
Cirrhina afghana Gunther, 1889 Trans. Linn. Soc. Zool. 5 : 106
Cirrhina afghana nikolskii Berg, 1905 Ann. Mus. Zool. Acad. Sci. 10 : 106
Scaphiodon baluchiorum Jenkins, 1910 Rec. Ind. Mus. 5 : 124
This synonymy is that of Berg (1949) but appears suspect. The type specimens of
Scaphiodon microphthalmus are possibly lost (see Banister & Clarke, 1977) and until such
time as they reappear certain identification of this species is not possible. However, Mirza's
(1969) description based on specimens from Quetta, the type locality, appears to represent
the species Day (1880) described. Specimens I have examined from Baluchistan (BMNH
JAW ANATOMY OF SEMIPLOTINE CARPS 333
1883.8.2 : 20-26) and the syntypes of Cirrhina afghana (BMNH 1886.9.12 : 155-159; 21)
conform to Mirza's description and that given by Jenkins (1910) for Scaphiodon
baluchiorum.
Banister & Clarke (1977) followed Berg (1949) in regarding Cyprinion muscatensis as
synonymous with C. microphthalmum.
DISTRIBUTION. Baluchistan, Afghanistan, the Indus Plain and Saudi Arabia.
Cyprinion milesi Day, 1 880
Barbus milesi Day, 1 880 Proc. Zool. Soc. Lond : 228
Barbus bampurensis Nikolsky (1899) 1900 Ann. Mus. St. Petersb. 4:410
Scaphiodon daukesi Zugmayer, \9\2Ann. Mag. nat. Hist. 8 : 596
Barbus baschakirdi Holly, 1929 Anz. Akad. Wiss Wienl : 1
Cyprinion milesi Berg, 1949 Trudy zool. Inst. Leningr. 8 (4) : 821
DISTRIBUTION. West Pakistan and Iran (see Mirza, 1969): BMNH specimens from
Afghanistan (1889.2.1 : 263-4) and Dizak, Baluchistan (1883.8.2 : 2-3).
Cyprinion watsoni Day, 1872
Scaphiodon watsoni Day, 1872 J. Asiatic Soc. Bengal 41 : 324
Scaphiodon irregularis Day, 1872 J. Asiatic Soc. Bengal 4\ : 321
ICyprinion kirmanse Nikolski, 1899 Ann. Mus. St. Petersb. 4:412
Scaphiodon macmahoni Regan, 1906 J. Asiatic Soc. Bengali : 8
Scaphiodon watsoni var. belensis Zugmayer, \9\2Ann. Mag. nat. Hist. (8) 10 : 596
Scaphiodon readingi Hora, 1923 Rec. Ind. Mus. 25 : 379-382
Cyprinion watsoni Berg, 1949 Trudy zool. Inst. Leningr. 8 (4) : 814
The above synonymy mainly follows that of Mirza (1969) who found a complex overlap of
morphometric and meristic characters between C. watsoni and Scaphiodon irregularis.
Certainly, there appear to be no osteological or myological differences between these
'species'.
Scaphiodon macmahoni was included by both Berg (1949) and Mirza (1969) in the
synonymy of Cyprinion microphthalmum. However, I find that S. macmahoni possesses the
same mouth morphology and degree of dorsal fin spine serration as does Cyprinion watsoni
and thus it is included in the synonymy of that species.
DISTRIBUTION. West Pakistan, Iran and Afghanistan, and the Helmand basin of Baluchistan.
The taxonomic status of Semiplotus dayi Fowler, 1958
Fowler (1958) considered that Scaphiodon aculeatus of Day (1880) was a misidentification
and that the specimens concerned represented a new species of Semiplotus for which Fowler
(1958) created the species S. dayi.
Although Day's specimens can no longer be traced, his description (1880) gives no cause
to think that he was not describing Scaphiodon aculeatus. Since Scaphiodon aculeatus (Val.
1844) is a synonym oiCapoeta capoeta (see Karaman, 1969) it follows that Semiplotus dayi
must also become a synonym of that taxon.
Relationship of the genus Cyprinion
In this study the presence of a sector mouth in various cyprinid taxa is seen both as a
character indicating relationship (synapomorphy) and as one independently evolved as a
parallelism. Taki (1975) used mouth morphology, degree of dorsal fin spine serration and
ossification, and the presence or absence of barbels as the criteria for determining relation-
ships amongst 'semiplotine' genera. Taki considered Onychostoma closely related to
'Semiplotus', "Scaphiodonichthys", Scaphiodon and Scaphiognathops. On the basis of these
supposed relationships he recognised an Onychostoma group and constructed an elaborate
dispersal hypothesis to explain the distribution of the included genera.
334 G. J. HOWES
Taki (1975) supposed that the onychostomine genera were '. . . derived from Barbus-
Varicorhinus stock'. As he indicated the group's 'origin' to have been in northern Asia, I take
it that he was referring to Euroasiatic Barbus and that his ' Varicorhinus' was in fact Capoeta
(see Karaman, 1969). Taki further confuses the issue by referring both the middle Asian and
western Indian species to Scaphiodon thereby implying their monophyly. In fact, Taki's
middle Asian 'Scaphiodon' are all Cyprinion species and those along the western Ghats of
India are Osteocheilus (see Hora, 1942, for synonymies).
The remaining genus included in Taki's Onychostoma group is Scaphiognathops.
Osteological data on this taxon are completely lacking; examination of the type specimen of
S. stejnegeri (USNM 90303) reveals a quite different jaw morphology to that of any
Cyprinion species (see figures in Smith, 1931 and Taki, 1974), or for that matter, from any
Onychostoma species.
Taki's (1975) hypothesis of relationships and dispersal are falsified by the conclusions of
this study, namely: 1. That Semiplotus, Scaphiodonichthys and Scaphiodon (part) are all
synonymous with Cyprinion; 2. That there are no synapomorphies linking Cyprinion with
Onychostoma or with Capoeta.
The relationships of Cyprinion are still in doubt and the sister-group remains unidentified.
This is due to lack of comparative anatomical data for the corpus of species currently
referred to the genus Barbus. Initial researches cast doubt on the assumed monophyly of
Barbus and also indicate one group within the complex which could be the sister lineage to
Cyprinion. The taxa involved are the north African species B. luteus, B. paytoni, B. callensis,
B. waldoi, B. rothschildi and B. sharpyei. In particular 'Barbus' paytoni possesses cranial and
jaw features similar to those of Cyprinion viz. deep and ventrally curved -ethmoid bloc,
transverse dilatator fossa, an almost identical posterior neurocranial morphology, broad and
deflected labial surface on the dentary, and a broad anguloarticular facet. The shape of the
quadrate, the hypertrophy of the interhyal, and the inward curvature of the interoperculum
appear to be further indications of relationship. More research is however, necessary to test
the hypothesis that the north African ' Barbus' species group is itself monophyletic.
Acknowledgements
I wish to express deep gratitude to Drs Humphry Greenwood and Keith Banister. To the
former for his critical reading of the manuscript and the many helpful suggestions that have
made for its improvement, and to the latter for spending so many hours in discussing the
vagaries of the carps.
My thanks are due to my former colleague Margaret Clarke for her assistance in compiling
the species synopsis, to Robert Travers for preparing alizarin specimens and Gina Sandford
for typing the manuscript.
It is a pleasure also to thank Dr George Lauder for our many stimulating discussions on
cyprinoid myology and functional anatomy.
Special thanks go to Dr Christine Karrer for translating, from the Russian, passages of L. S.
Berg's 'Freshwater Fishes of Iran'.
Finally, my thanks are due to Dr Richard Vari for his assistance in locating H. M. Smith's
specimens in the Smithsonian Institution and for his hospitality during my visit there.
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Zool. J. Linn. Soc. 69 : 149-182.
Manuscript accepted for publication 23 September 198 1
East African Cichlid Fishes
The haplochromine fishes of the east African lakes
P. H. Greenwood, British Museum (Natural History)
May 1981, 840 pp, 3 plates, 350 figures Hardback Price £25.00
This volume brings together, for the first time, Dr Greenwood's various papers (some
now out of print) on the taxonomy and biology of the species of haplochromine fishes
from Lake Victoria, east Africa. Also reprinted are his papers on the haplochromines
from Lakes Nabugabo, George and Turkana (Rudolph), two papers dealing with
the classification of the genus Haplochromis, and a recent essay on the explosive
evolution of cichlid fishes in Africa.
An index to the 200 species dealt with in this book (and their current generic place-
ment), and a general introduction to the evolutionary and taxonomic problems posed
by these biologically complex and fascinating fishes, are included.
Published jointly by British Museum (Natural History)
(exclusive U.K. rights) and Kraus-Thomson Organization Ltd,
(all other rights). ISBN 3 601 00438 6
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British Museum (Natural History)
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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. Keirans
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
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