Skip to main content

Full text of "Bulletin of the British Museum (Natural History)"

See other formats


1 4  MAR  /983 

LIBRARY 


Bulletin  of  the 

xcgdlti 

British  Museum  (Natural  History 


Zoology  series    Vol  42    1982 


British  Museum  (Natural  History) 
London  1983 


Dates  of  publication  of  the  parts 

No  1 25  March  1982 

No  2 29  April  1982 

No  3 27  May  1982 

No  4  24  June  1982 


ISSN  0007-1498 


Printed  in  Great  Britain  by  Henry  Ling  Ltd,  at  the  Dorset  Press,  Dorchester,  Dorset 


Contents 
Zoology  Volume  42 

Page 

No  1      The   tick  collection   (Acarina:    Ixodoidea)   of  the   Hon.    Nathaniel 
Charles  Rothschild  deposited  in  the  Nuttall  and  general  collections  of 
the  British  Museum  (Natural  History) 
By  James  E.  Keirans 1 

No  2     Hydroids  and  medusae  of  the  family  Campanulariidae  recorded  from 
the  eastern  North  Atlantic,  with  a  world  synopsis  of  genera 
By  P.  F.  S.  Cornelius 37 

No  3     Miscellanea 

A  new  species  of  free-living  nematode  from  the  Firth  of  Clyde,  Scotland 

By  P.  J.  D.  Lambshead 149 

Notes  on  Atlantic  Asteroidea  2.  Luidiidae 

By  A.  M.  Clark 157 

New  and  little  known  species  of  Oncaeidae  (Cyclopoida)  from  the 

Northeastern  Atlantic 

By  S.  J.  Malt 185 

Larval  and  post-larval  development  of  the  Slender-legged  Spider  Crab, 
Macropodia  rostrata  (Linnaeus),  (Oxyrhyncha:  Majidae:  Inachinae), 
reared  in  the  laboratory 
By  R.  W.  Ingle 207 

No  4     Miscellanea 

New  species  of  marine  nematodes  from  Loch  Ewe,  Scotland 

By  H.  M.  Platt  and  Z.  N.  Zhang 227 

The  larval  development  of  Crangon  crangon  (Fabr.  1 795)  (Crustacea  : 

Decapoda) 

By  A.  R.  Gurney 247 

A  revision  of  the  spider  genus  Cocalodes  with  a  description  of  a  new 

related  genus  (Araneae:  Salticidae) 

By  F.  R.  Wanless 263 

Anatomy  and  evolution  of  the  jaws  in  the  semiplotine  carps  with  a 
review  of  the  Genus  Cyprinion  Heckel,  1843  (Teleostei:  Cyprinidae) 
By  G.  J.  Howes .299 


Bulletin  of  the 

British  Museum  (Natural  History) 

The  tick  collection  (Acarina:  Ixodoidea) 
of  the  Hon.  Nathaniel  Charles  Rothschild 
deposited  in  the  Nuttall  and  general 
collections  of  the  British  Museum 
(Natural  History) 


James  E.  Keirans 

With  a  Foreword  by  the  Hon.  Miriam  Rothschild 


Zoology  series    Vol  42  No  1    25  March  1982 


The  Bulletin  of  the  British  Museum  (Natural  History),  instituted  in  1949,  is  issued  in  four 
scientific  series,  Botany,  Entomology,  Geology  (incorporating  Mineralogy)  and  Zoology,  and 
an  Historical  series. 

Papers  in  the  Bulletin  are  primarily  the  results  of  research  carried  out  on  the  unique  and 
ever-growing  collections  of  the  Museum,  both  by  the  scientific  staff  of  the  Museum  and  by 
specialists  from  elsewhere  who  make  use  of  the  Museum's  resources.  Many  of  the  papers  are 
works  of  reference  that  will  remain  indispensable  for  years  to  come. 

Parts  are  published  at  irregular  intervals  as  they  become  ready,  each  is  complete  in  itself, 
available  separately,  and  individually  priced.  Volumes  contain  about  300  pages  and  several 
volumes  may  appear  within  a  calendar  year.  Subscriptions  may  be  placed  for  one  or  more  of 
the  series  on  either  an  Annual  or  Per  Volume  basis.  Prices  vary  according  to  the  contents  of 
the  individual  parts.  Orders  and  enquiries  should  be  sent  to: 

Publications  Sales, 

British  Museum  (Natural  History), 
Cromwell  Road, 

London  SW7  5BD, 
England. 


World  List  abbreviation:  Bull.  Br.  Mus,  not.  Hist.  (Zool.) 


©  Trustees  of  the  British  Museum  (Natural  History),  1982 


The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 
Keeper  of  Zoology     :    Dr  J.  G.  Sheals 
Editor  of  Bulletin       :    Dr  C.  R.  Curds 
Assistant  Editor         :    Mr  C.  G.  Ogden 


ISSN  0007-1498  Zoology  series 

Vol  42  No  1  pp  1-36 

British  Museum  (Natural  History) 

Cromwell  Road 

London  SW7  5BD  Issued  25  March  1982 


GENERAL 


The  tick  collection  ( Acarina:  Ixodoidea)  of  thV  ^ 1982 

\  -4     LIBRARY     "^» 

Hon.  Nathaniel  Charles  Rothschild  deposited  in  ^zy 
the  Nuttall  and  general  collections  of  the  BritisBT 
Museum  (Natural  History) 

James  E.  Keirans 

Department  of  Health  and  Human  Services,  Public  Health  Service,  National  Institutes  of 
Health,  National  Institute  of  Allergy  and  Infectious  Diseases,  Rocky  Mountain  Laboratory, 
Hamilton,  Montana  59840,  U.S.A. 

With  a  Foreword  by  the  Hon.  Miriam  Rothschild 


Contents 

Synopsis 1 

Foreword 3 

Introduction 4 

PART  I  Rothschild  specimens  donated  to  Professor  G.  H.  F.  Nuttall ...  5 
Species  of  Ixodoidea  represented  in  the  Nuttall  collection,  British  Museum 

(Natural  History),  donated  by  the  Hon.  N.  C.  Rothschild     ....  5 

Rothschild  specimens  in  the  Nuttall  collection  with  type  status      ...  6 

Rothschild  specimens  in  the  Nuttall  collection 6 

A    single    tick    collection    given    by    the    Hon.    Walter    Rothschild    to 

G.  H.  F.  Nuttal  via  Guy  A.  K.  Marshall 17 

Rothschild  specimens  missing  from  Nuttall  collection 17 

PART  II  Rothschild  specimens  donated  to  British  Museum  (Natural  History).  20 
Species  of  Ixodoidea  represented  in  the  collection  of  the  British  Museum  (Natural 

History),  donated  by  the  Hon.  N.  C.  Rothschild 20 

Rothschild  specimens  in  the  British  Museum  (Natural  History)  collection  with 

type  status 21 

Rothschild  specimens  in  British  Museum  (Natural  History)  collection  .  .  22 

Rothschild  specimens  missing  from  British  Museum  (Natural  History)  collection  32 

Locality  list  for  ticks  in  the  Rothschild  collection 32 

Host  list  for  ticks  in  the  Rothschild  collection 34 

Acknowledgements 35 

References 36 

Synopsis 

The  Hon.  N.  C.  Rothschild  achieved  worldwide  fame  for  his  study  of  the  Siphonaptera. 

During  the  early  part  of  this  century  he  also  made  a  significant  collection  of  ticks  which  he 
gave  to  Professor  George  H.  F.  Nuttall  at  Cambridge  University.  Upon  Nuttall's  death,  the 
collection  was  given  to  the  British  Museum  (Natural  History)  and  is  kept  as  a  separate  entity. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  42(  1 ):  1  -36  Issued  25  March  1 982 


J.  E.  KEIRANS 


Male  Ixodes  uriae  White,  1852.  (Ixodoidea:  Ixodidae).  A  tick  with  a  circumpolar  distribution  in 
both  the  northern  and  southern  hemispheres.  It  is  ectoparasitic  on  a  variety  of  seabirds  and  to 
date,  twenty  different  arboviruses  have  been  recovered  from  this  species. 


ROTHSCHILD  TICKS  IN  BM(NH)  3 

In  addition,  Rothschild  on  numerous  occasions  between  1911  and  1923  deposited  ticks  in 
the  British  Museum  (Natural  History).  Herewith  are  recorded  the  197  tick  collections  of  the 
Hon.  N.  C.  Rothschild  plus  one  of  the  Hon.  Walter  Rothschild  deposited  in  the  Nuttall 
collection  and  the  156  collections  in  the  general  collection  of  the  British  Museum  (Natural 
History). 


Foreword 

In  1913  N.  C.  Rothschild  presented  his  collection  of 'Siphonaptera  and  other  parasitic 
insects'  to  the  British  Museum  (Natural  History).  One  of  the  conditions  attached  to  the  gift 
read  as  follows:  'The  Trustees  shall  so  soon  as  the  said  collection  comes  into  their  hands  or  is 
placed  under  their  control  and  as  soon  as  the  funds  placed  at  their  disposal  by  Parliament 
will  allow  cause  to  be  made  and  published  a  catalogue  giving  the  names  and  full  number  of 
specimens  of  each  species  of  parasitic  insect  contained  in  the  said  collection.' 

Over  60  years  has  now  elapsed  since  the  collection  was  accepted,  but  the  catalogue  has  not 
yet  been  completed.  Although  ticks  are  arthropods,  they  are  not  insects,  but  this  group  of 
animals,  along  with  a  number  of  mites,  was  included  in  the  collection. 

During  my  period  of  Trusteeship  of  the  British  Museum  (Natural  History)  I  had  the 
opportunity  of  discussing  this  problem  with  Harry  Hoogstraal,  and  he  suggested  that  James 
Keirans  might  undertake  to  catalogue  the  Rothschild  ticks.  To  my  great  delight  he  agreed  to 
do  this,  although  the  task  proved  more  time-consuming  than  we  had  at  first  anticipated.  This 
was  because  the  Rothschild  Collection  had  not  been  preserved  as  a  single  unit,  and  the  ticks, 
along  with  the  Nycteribiidae,  Cimicidae,  Hippoboscidae  and  other  smaller  groups,  had  been 
distributed  in  the  general  collection  and,  in  addition,  had  not  all  received  Museum  accession 
numbers. 

Although  these  circumstances  added  enormously  to  James  Keirans'  labours,  they  made 
the  compilation  of  this  Catalogue  even  more  necessary  and  valuable.  Until  those  who  are 
engaged  in  ecological  and  medical  research  come  to  use  museum  material  such  as  this,  no 
one  can  really  appreciate  the  immense  boon  of  well-catalogued  individual  collections. 

Since,  between  1900  and  1910,  N.  C.  Rothschild  had  given  Professor  Nuttall  the  bulk  of 
his  tick  material*,  it  was  considered  more  appropriate  to  merge  the  two  catalogues  into  one, 
and  the  manuscript  was  therefore  divided  into  two  parts:  I,  the  specimens  collected  by 
Rothschild,  now  in  the  Nuttall  collection;  and  II,  the  specimens  donated  by  Rothschild  to 
the  British  Museum  (Natural  History).  The  ticks  in  the  general  collection  have  been 
added — a  course  we  followed  when  cataloguing  the  fleas  (G.  H.  E.  Hopkins  &  M.  Rothschild, 
Vols.  I-V,  1953-1 971)  and  the  Nycteribiidae  (Oscar  Theodor,  1967). 

It  will  be  noted  that  Walter  Rothschild  contributed  one  specimen  to  this  collection.  It  has 
not  usually  been  appreciated  that  N.  C.  Rothschild's  brother  also  added  considerably  to  the 
flea  material,  although  he  did  not  describe  any  species  himself.  The  N.  C.  Rothschild 
donations  of  Nycteribiidae  to  the  British  Museum  contained  56  undescribed  species,  and  the 
Ixodoidea  collections  10.  The  two  brothers  and  their  collaborators  described  over  5000  new 
species,  but  they  probably  collected  twice  that  number  of  animals  new  to  science. 

James  Keirans  has  added  greatly  to  the  value  of  this  compilation  by  providing  information 
concerning  the  viruses  and  in  certain  cases  rickettsiae  of  which  the  Rothschild  ticks  are 
known  vectors. 

The  illustration  depicts  a  male  specimen  of  Ixodes  uriae  White,  one  of  the  most 
spectacular  species,  adorned  with  a  handsome  fringe  along  the  posterior  margin,  which  has 
the  dubious  distinction  of  carrying  at  least  20  known  viruses  in  five  different  serogroups. 


*After  the  sudden  death  of  Professor  Nuttall  in  December  1937  his  collection  was  presented  19  the  British  Museum 
(Natural  History)  by  the  Molteno  Institute.  In  accordance  with  the  terms  of  the  gift  this  collection  is  maintained  as  a 
discrete  entity. 


4  J.  E.  KEIRANS 

Altogether  the  meticulous  care  and  accuracy  with  which  James  Keirans  has  completed 
this  catalogue  is  beyond  praise. 

Miriam  Rothschild 


Introduction 

Early  in  the  twentieth  century,  Nathanial  Charles  Rothschild  acquired  a  large  tick 
collection,  primarily  from  collectors  who,  knowing  of  his  great  interest  in  fleas,  would  send 
him  not  only  that  group  but  also  other  ectoparasites  including  flies,  bugs,  and  ticks. 

He  deposited  his  tick  collection  in  two  institutions;  the  Molteno  Institute  for  Research  in 
Parasitology,  Cambridge  University,  where  Professor  G.  H.  F.  Nuttall  made  the  original 
determinations  of  all  tick  material,  and  the  British  Museum  (Natural  History),  where  Mr  A. 
S.  Hirst  made  the  tick  determinations.  As  mentioned  earlier,  the  Nuttall  collection  is  now  at 
the  BM(NH). 

Because  the  Nuttall  collection  is  kept  as  a  discrete  entity,  separate  from  but  housed  with 
the  BM(NH)  collection,  parts  of  Rothschild's  tick  collection  are  in  each  of  the  above 
collections.  This  catalogue  is  also  divided  into  two  sections:  Part  I,  Rothschild  tick  donations 
to  the  Nuttall  collection;  Part  II,  Rothschild  tick  donations  to  the  BM(NH). 

The  systems  for  numbering  tick  collections  are  different  for  the  two  sections.  In  Part  I,  the 
number  used  is  that  given  by  Professor  Nuttall  to  a  collection.  The  first  entry  is  N  (Nuttall) 
72  followed  by  N73,  N74,  etc.,  in  ascending  sequence,  first  for  existing  specimens,  then  for 
specimens  which  are  now  missing.  Each  Nuttall  number  is  followed  by  a  Rothschild 
collection  number  or  the  word  'None'  if  the  collection  received  no  Rothschild  number.  A 
Nuttall  number  refers  to  a  handwritten  entry  in  Professor  Nuttall's  tick  catalogue,  which  is 
now  kept  in  the  Arachnida  and  Myriapoda  Section  of  the  BM(NH).  The  locality  and  host 
data  are  cited  in  their  modern  forms,  but  where  these  could  not  be  established  Nuttall's  own 
entries  are  quoted. 

One  additional  point  should  be  made  relating  to  the  Nuttall  collection  forming  Part  I. 
Nathaniel  C.  Rothschild  donated  all  specimens  recorded  in  Part  I  with  the  single  exception 
of  N888  which  was  donated  by  his  brother,  the  Hon.  Walter  Rothschild,  to  the  Entomo- 
logical Research  Committee  and  subsequently  to  Professor  Nuttall. 

Collections  in  Part  II  are  entered  alphabetically  by  genus,  first  Argasidae,  then  Ixodidae. 
All  information  is  presented  as  in  Part  I  with  the  exception  that  replacing  a  Nuttall  and 
Rothschild  number  will  be  a  British  Museum  (Natural  History)  registration  number.  Where 
no  registration  number  was  given  to  a  collection,  the  word  'None'  will  appear.  I  have 
given  each  collection  a  set  of  Keirans  numbers,  K1-K156.  These  numbers,  along 
with  the  Nuttall  numbers  from  Part  I,  will  allow  the  reader  to  use  the  tick  species, 
locality,  and  host  lists  and  refer  back  to  individual  collections  within  Parts  I  and  II. 

Throughout  the  text,  citations  are  given  to  publications  which  mention  a  particular  taxon. 
My  comments  are  presented  within  square  brackets  [  ];  those  comments  by  Nuttall  or  some 
other  investigator  are  cited  within  square  brackets  enclosed  by  quotation  marks '[  ]'. 


ROTHSCHILD  TICKS  IN  BM  (NH) 

PARTI 


Rothschild  specimens  donated  to  the  Nuttall  collection 


Species  of  Ixodoidea  represented  in  the  Nuttall  collection,  British  Museum 
(Natural  History),  donated  by  the  Hon.  N.  C.  Rothschild 

Nuttall  numbers  follow  each  species.  Numbers  in  parentheses  refer  to  missing  collections. 
Thus,  validity  of  original  determinations  is  unverified. 


ARGASIDAE 

Argas  boueti  Robaud  and  Colas-Belcour  1 164 
Argas persicus  (Oken)  589,  590,  (3536)) 
Argas  reflexus  (Fabricius)  (3 1 59) 
Argas  vespertilionis  (Latreille)  591,   592,   593, 

1151,1152,1153, 1163 
Argas  sp.  11 54,  (1388) 

Ornithodoros  capensis  Neumann  group  594 
Ornithodoros  moubata  (Murray)  595 

Otobius  megnini  (Duges)  (596) 

IXODIDAE 

Amblyomma  albolimbatum  Neumann  (78),  567, 

569 

Amblyomma  australiense  Neumann  547 
Amblyomma  cajennense (Fabricius)  557, 648% 
Amblyomma  clypeolatum  Neumann  (75) 
Amblyomma  compressum  Macalister  564 
Amblyomma  cyprium  Neumann  73,  565 
Amblyomma  decoratumC.  L.  Koch  (93) 
Amblyomma  dissimile  C.  L.  Koch  568 
Amblyomma  gemma  Donitz  558b 
Amblyomma  geoemydae  (Cantor)  545 
Amblyomma  hirtum  Neumann  (397) 
Amblyomma  longirostre(C.  L.  Koch)  560 
Amblyomma  marmoreum  C.  L.  Koch  (77),  570 
Amblyomma  ovaleC.  L.  Koch  562,  566 
Amblyomma pecarium  Dunn  557 
Amblyomma  tholloni  Neumann  559 
Amblyomma  triguttatum  C.  L.  Koch  546,  (551), 

552,553 

Amblyomma  varium  C.  L.  Koch  (3351) 
Amblyomma  sp.  74,  (76),  (544),  633,  1 166,  1386, 

1391, 1392, 1393, 1394, 1396,  1397 

Aponomma  decorosum  (L.  Koch)  550 
Aponomma  gervaisi  (Lucas)  (92),  (548) 
Aponomma  gervaisi  var.  lucasi  (Warburton)  (90), 

(91) 

Aponomma  hydrosauri  (Denny)  556 
Aponomma  varanensis  (Supino)  554 
Aponomma  sp.  (57 1 ),  3 1 38 

Boophilus  decoloratus  (C.  L.  Koch)  (3157) 
Boophilus  microplus  (G.  Canestrini)  588 


Dermacentor   albipictus   (Packard)   666,    1105, 

1497, 1498, 1499, 1500, 1501, 1502, 1503 
Dermacentor  andersoni  Stiles  1387,  1390,3502 
Dermacentor  reticulatus  (Fabricius)  3 1 60 
Dermacentor  rhinocerinus  (Denny)  555,  563 
Dermacentor  sp.  3330 

Haemaphysalis  bispinosa  Neumann  574,  582, 

583,892 

Haemaphysalis  campanulata  Warburton  579 
Haemaphysalis  celebensis  Hoogstraal,  Trapido 

and  Kohls  72 

Haemaphysalis  elongata  Neumann  670 
Haemaphysalis  erinacei  Pavesi  3537,  3538,  3539 
Haemaphysalis  humerosa  Warburton  and  Nuttall 

669 
Haemaphysalis  hystricis  Supino  572,  573,  575, 

576 

Haemaphysalis  indica  Warburton  575x 
Haemaphysalis  leachi  (Audouin)  (539),  577,  578 
Haemaphysalis    punctata    G.    Canestrini    and 

Fanzago  580 

Haemaphysalis  tiptoni  Hoogstraal  670 
Haemaphysalis  sp.  58 1 

Hyalomma  aegyptium  (Linnaeus)  549,  56 1 
Hvalomma  rufipes  C.  L.  Koch  558a,  888  (W. 

Rothschild) 
Hyalomma  syriacum  C.  L.  Koch  (3139),  (3140), 

(3141) 

Hyalomma  truncatum  C.  L.  Koch  1 167 
Hyalomma  sp.  1398,  (3 136) 

Ixodes  australiensis  Neumann  645, 646 

Ixodes  boliviensis  Neumann  637 

Ixodes  brunneus  C.  L.  Koch  (396) 

Ixodes  cookei  Packard  1 504 

Ixodes  fecialis  Warburton  and  Nuttall  650,  1211, 

1213 

Ixodesfrontalis  (Panzer)  523 
Ixodes   hexagonus   Leach   (216),   (217),   (298), 

(322),  (324),  597,  598,  599,  600,  601 ,  602,  603, 

604,  606,  607,  608,  609,  610,  611,  612,  613, 

(674),  (746),  1067,  1159,  1160 
Ixodes  holocyclus  Neumann  643,  (644),  1212 
Ixodes  lividus  C.  L.  Koch  605,  1068 


J.  E.  KEIRANS 


Ixodes  loricatus  Neumann  638, 639,  640, 64 1 

Ixodes  luciae  Senevet  647 

Ixodes  nitens  Neumann  360 

Ixodes    putus    (O.    Pickard-Cambridge)    (516), 

(5 17),  (5 18) 
Ixodes   ricinus  (Linnaeus)  (509),   (628),   (629), 

(630),  (631),  (632),  (656) 
Ixodes  rothschildi  Nuttall  and  Warburton  634 
Ixodes  rubidus  Neumann  826 
Ixodes  scapularis  Say  626 
Ixodes  texanus  Banks  1 399 
Ixodes  trianguliceps  Birula  619,  (620),  621,  622, 

623,624,625,1069,  1161 


Ixodes  unicavatus  Neumann  (395),  652, 667 
Ixodes  uriae  White  617,618 
Ixodes  vespertilionis  C.  L.  Koch  649 
Ixodes  vestitus  Neumann  642 
Ixodes  sp.  1066,  1 162,  1 167,  1389 


Rhipicephalus  evertsi  Neumann  (3 1 57) 
Rhipicephalus   haemaphysaloides   Supino   584, 

(3158) 

Rhipicephalus  muehlensi  Zumpt  585 
Rhipicephalus  sanguineus  (Latreille)  (586),  (587) 
Rhipicephalus  sp.  1 395 


Rothschild  specimens  in  the  Nuttall  collection  with  type  status 


NUTTALL  ROTHSCHILD 


No. 
72 

360 
565 

634 
647 
650 
669 


No. 
916 

None 
243 

159 
248 
203 
178 


Haemaphysalis  celebensis  Hoogstraal,  Trapido,  and  Kohls,  1965.  J. 

Paras  it.  51:  1001,  figs.  1-9. 

Ixodes  nitens  Neumann,  1904.  Archs.  Parasit.  8:  459. 

Amblyomma  quasicyprium  Robinson,  1926.  Ticks.  A  Monograph  of  the 

Ixodoidea  Pt.  4:  237,  fig.  1 17. 

Ixodes  percavatus  rothschildi  Nuttall  and  Warburton,  1911.  Ticks.  A 

Monograph  of  the  Ixodoidea  Pt.  2:  22 1 . 

Ixodes  loricatus  spinosus  Nuttall,  1910.  Parasitology,  Cambridge  3: 

411,  fig.  5. 

Ixodes  fecialis  Warburton  and  Nuttall,  1909.  Parasitology,  Cambridge 

2:  58,  figs.  1-2. 

Haemaphysalis  humerosa  Warburton  and  Nuttall,  1909.  Parasitology, 

Cambridge  2:  60,  figs.  4-5. 


Rothschild  specimens  in  the  Nuttall  collection 


N72        R916 

1    9   Haemaphysalis   hystricis   (Haemaphysalis 

celebensis  HOLOTYPE) 
ex.  Sus  celebensis  (Sus  verrucosus  celebensis) 
Celebes.  No  date  (Sulawesi  02WS,   121WE, 

Indonesia) 

Presented  as  mounted  specimen  No.  916  which 
we  unmounted  4.1. 1 9 1 5,  C.  W.  [Cecil  Warburton] 
det.  8.1.1915. 

Publications:  Nuttall  &  Warburton  (1915:  425, 
426);  Hoogstraal  el  al.  (19656:  1001);  Hoogstraal 
etal.  (1973:  556). 

N73        R915 

1  d  Amblyomma  caelaturum  [This  species  epithet 

crossed       out       and       cyprium       inserted] 

(Amblyomma  cyprium) 


ex.  Sus  celebensis  (Sus  verrucosus  subsp.) 
Celebes.  No  date  (Sulawesi  02WS,   121°00'E, 

Indonesia) 

Presented  as  slide  No.  9 1 5  which  was  unmounted, 
being  defective  5.1.1915. 
3  d1  Amblyomma  cyprium  mounted  as  slides,  2 

transparent,  1  opaque — see  slides  No.  73  (i)  (ii) 

(iii)  (Nos.  917,  919,  920)  (3   slides)  (L.   E. 

Robinson  det.  26.1. 1915)  redet.  26.111. 1919. 
Note:  The  3  slide-mounted  males  were  missing 
from  the  Nuttall  collection  in  1977. 
Publication:  Robinson  (1926:  236). 

N74        R918 

1    N    Amblyomma    (?  caelaturum    Cooper    & 
Robinson)    (Amblyomma    sp.).     Unmounted 


ROTHSCHILD  TICKS  IN  BM(NH) 


from  slide  labelled  as  off  Sus  celebensis  (Sus 

verrucosus  subsp.) 
Celebes.  No  date  (Sulawesi  02WS,   121WE, 

Indonesia 
(L.  E.  Robinson  det.  26.1. 1915). 

N360        None 

1    9    Ixodes    nitens    COTYPE    (Ixodes    nitens 

SYNTYPE) 

ex.  Mus  macleari  (Rattus  macleari) 
Christmas     Island,     Pacific     Ocean     (10°39'S, 

105°40'E) 

(ex.  Shipley  coll.  No.  2) 

Publications:  Neumann  (1904:  460);  Nuttall  & 
Warburton(1911:  185). 

N523        None 

1  9  Ixodes  brunneus  (minus  hypostome,  digits) 

(Ixodes  fron  tails) 
ex.  Passer  montanus 
Saint-Genies-de   Malgoires   (43°57'N,   04°13'E), 

Card,  France 
16.XI.  1908  Albert  Hugues 
Publication:  Nuttall  &  Warburton  (1911:1 92). 

N545        R258 

Amblyomma  sp.  [sex/stage  not  given]  (1  9 
Amblyomma  geoemydae)  (tentative  deter- 
mination) 

Found  on  ground  at  Paquil,  Luzon,  Philippine 
Islands  (locality  not  verified) 

15.1.1895  A.  Everett 

Publication:  Robinson  (1926:  48)  recorded  thfs  as 

A.  americanum. 

N546        R174 

Amblyomma  triguttatum  [sex/stage  not  given]  (4  9 

Amblyomma  triguttatum) 
ex.  kangaroo 
Barrow    Island    (20°48'S,     115'23'E),    N.     W. 

Australia 

XI.  1900  C.  J.  T.  Tanney.  det.  L.  E.  Robinson 
Publication:  Robinson  (1926:  57)  indicated  d's 
present  but  not  in  Nuttall  collection  in  1977. 

N547        R274 

6  d,  1  9  Amblyomma  australiense 

ex.  Echidna  aculeata(Tachvglossus  aculeatus] 

Mt  Anderson  ( 1 7°58'S,  1 24;04'E),  W.  Australia 

[No  date]  C.  J.  T.  Tanney 

Publications:      Robinson      (1926:      135,      136) 

illustrated  d  and  9  from  this  lot.  Taylor  (1946: 

100)     repeated     Robinson's    descriptions    and 

illustrations. 

N549        R254 

Hyalomma  syriacum  [sex/stage  not  given]  (4  cf 

Hyalomma  aegyptium) 
ex.  tortoise 
No  data 
Note:  An  additional  1  d,  1  9  of  this  collection  sent 


to  New  York  National  Museum,  New  York 
City,  18.IV.1913. 

N550        R196 

Aponomma  decorosum  [sex/stage  not  given]  (2  d 

Aponomma  decorosum) 
ex.  Australian  monitor 

Note:  A.  decorosum  is  found  only  in  Australia  and 
ranges  from  Queensland  south  to  Victoria. 

N552        R179 

Amblyomma  triguttatum  [sex/stage  not  given]  (2 

d,  3  9  Amblyomma  triguttatum) 
ex.  kangaroo 
Barrow    Island    (20°48'S,     115'23'E),    N.    W. 

Australia 

20.XI.1900C.J.  T.  Tanney 
Publication:  Robinson  (1926:  57). 

N553        R211 

Amblyomma  triguttatum  [sex/stage  not  given]  (4 

9,  13  N  Amblyomma  triguttatum) 
ex. kangaroo 
Barrow    Island    (20°48'S,     115'23'E),    N.    W. 

Australia 

18.XI.1 900  C.J.T.  Tanney 
Publication:  Robinson  (1926:  57). 

N554     R236 

Aponomma  exornatum  [sex/stage  not  given]  (2  d 

Aponomma  varanensis) 
ex.  Monitor  strix  [name  not  verified] 
?  Australia 

Note:  Collection  probably  not  from  Australia. 
The  only  collection  of  A.  varanensis  from 
Australia  is  the  type  d  of  A.  quadratus  (an 
inornate  form  of  A.  varenensis).  This  single 
record  may  have  been  an  accidental  importation 
of  tick  and  host. 

N555        R259 

Dermacentor  rhinocerotis  [sex/stage  not  given]  (7 

d  Dermacentor  rhinocerinus) 
No  other  data 

Note:  There  is  an  additional  d  of  this  species 
mounted  on  a  slide. 

N556        R287 

Aponomma  hydrosauri  [sex/stage  not  given]  (2  d, 

1  9  Aponomma  hydrosauri) 
?  Adelaide  (34°55'S,  1 38°35'E),  W.  Australia 

N557        R198 

Amblyomma  cajennense  [sex/stage  not  given]  (1 

d,     2     9     Amblyomma     cajennense;     3     d 

Amblyomma  pecarium) 
ex.  peccary  Tayassu  sp. 
Santa  Andrea,  Tabasco 
30. V.? '[No  particulars]' 

Note:  The  above  locality  information  was  taken 
from  the  vial  label.  It  should  be  San  Andres, 
Estado  de  Tabasco  ( 1 8WN,  92°40'W),  Mexico. 


8 


J.  E.  KEIRANS 


Publication:  Robinson  (1926:  53)  incorrectly  gave 
the  Nuttall  number  as  577. 
Amblyomma   cajennense   is   a   vector   of  Wad 
Medani    virus    and    the    rickettsia    of    Rocky 
Mountain  spotted  fever. 

N558a        R263 

1  d,    1    9   Hyalomma   aegyptium   (Hyalomma 
rufipes) 

ex.  giraffe,  Giraffa  camelopardalis 

No  other  data 

Hyalomma  rufipes  is  the  vector  of  the  viruses 

Dugbe,    Tete,    Matruh,    and    Crimean    Congo 

hemorrhagic  fever. 

N558b        R263 

2  d  Amblyomma  gemma 

ex.  giraffe,  Giraffa  camelopardalis 

No  other  data. 

Amblyomma   gemma   is   a   vector   of  Nairobi 

sheep  disease  virus. 

N559        R285 

1  9  Amblyomma  tholloni 

No  host 

Mt    Ruwenzori    (00°23'N,    29°54'E),    on    the 

Uganda-Zaire  border,   Africa    1906   A.    F.    R. 

Wollaston 

N560        R260 

1  d  Amblyomma  longirostre 

ex.  Coendou  simonsi  (Coendou  bicolor  simonsi) 

Charapaya(17°22'S,  66°45'W),  Bolivia 

22.VI.1901  P.O.Simons 

Publication:  Robinson  (1926:  140). 

N561        None 

Hyalomma  syriacum  [sex/stage  not  given]  (4  d 

Hyalomma  aegyptium) 

ex.  Algerian  tortoise 

N.  C.  Rothschild 

N562        R261 

Amblyomma  ovale  [sex/stage  not  given]   (2  d 

Amblyomma  ovale) 
ex.  Speothas  venaticus 
Charapaya(17°22'S,66'45'W),  Bolivia 
27.VI.1901  P.O.Simons 

N563        R266 

10  d  '[small  and  large]'  Dermacentor  rhinocerotis 
(7  cf  Dermacentor  rhinocerinus) 
ex.  rhinoceros 
No  other  data 

Note:  Nuttall,  in  his  notebook  containing  infor- 
mation on  ticks  sent  to  various  individuals  and 
institutions,  indicated  that  this  collection  was 
from  Nyasaland,  British  Central  Africa  [Malawi] 
in  1907.  One  additional  d  of  this  collection  was 
sent  to  Dr  J.  G.  Parham,  Marromeu,  Zembezia, 
Portuguese  East  Africa  [People's  Republic  of 
Mozambique],  18.IV.1913  and  one  additional  d 


was  sent  to  New  York  National  Museum,  New 
York  City,  18.IV.1913. 

N564        R218 

Amblyomma  badium  [The  species  epithet  crossed 

out  and  sublaeve  inserted]  (6  d  Amblyomma 

compressum) 

ex.  white-bellied  pangolin,  Manis  tricuspis 
No  other  data 
L.  E.  Robinson  det.  1926 

N565        R243 

1  9  Amblyomma  quasicyprium  Robinson,  1926 

TYPE  (Amblyomma  cyprium) 
ex.  Spider  monkey  Ateles  melanochoerus  [This 

species  epithet  is  crossed  out  and  melanochir 

inserted]  (Ateles  g.  geoffroyi) 
Frontera,  ( 1 8°32'N,  92°38'W),  Tabasco,  Mexico 
Notes:  Both  host  and  locality  are  incorrect  for 
Amblyomma  cyprium  although  this  9  appears  to 
be  true  A.  cyprium. 

Nuttall  575  and  3330  also  bear  Rothschild 
number  R243. 

Publications:  Robinson  (1926:  237,  238) 
incorrectly  gave  the  Nuttall  number  564  for  this 
collection.  Taylor  (1946:  116)  repeated 
Robinson's  error  for  the  Nuttall  number. 

N566        R270 

1  9  Amblyomma  ovale 

ex.  Speothas  venaticus 

Charapaya  (17'22'S,  66'45'W),  Bolivia 

Publication:  Robinson  (1926:  29)  gave  the  date  as 

1900  and  indicated  that  the  collection  contained 

only  males. 

N567        R282 

Amblyomma  albolimbatum  [sex/stage  not  given] 

(2  d1,  1  9  Amblyomma  albolimbatum) 
ex.  Morelia  variegata  (Morelia  spilotes  variegatd) 
Perth  (3 1  °56'S,  1 1 5°50'E),  W.  Australia. 
Woodward 
L.  E.  Robinson  det.  1926 

N568        R231 

Amblyomma  [species  epithet  integrum  crossed 

out  and  dissimile  inserted]:  [sex/stage  not  given] 

(7  d,  2  9, 7  N  Amblyomma  dissimile) 
ex.  large  snake 
Probably  Frontera  (18'32'N,  92°38'W),  Tabasco, 

Mexico 
May  1 .  1  .V.  1 90 1  [The  full  date  is  on  vial  label  but 

not  in  catalogue] 

Publication:  Robinson  (1926:  167)  indicated  the 
date  for  this  collection  was  V.I 900  and  that  only 
one  female  was  present. 

N569        R267 

Amblyomma  moreliae  [sex/stage  not  given]  [This 
species  epithet  crossed  out  and  albolimbatum 
inserted]  (1  9  Amblyomma  albolimbatum) 


ROTHSCHILD  TICKS  IN  BM(NH) 


ex.  Diemenia  superciliosa  (Diemenia  textilis) 
Herdman's  Lake,  near  Perth  (31'56'S,  1 15'15'E), 

W.  Australia 

Det.  Neumann  &  Warburton 
Publication:  Robinson  (1926:  226)  figured  this  9; 
on   p.  227    he   incorrectly   gave   the   collection 
number  as  N 1569. 

N570        R182 
2  cf  Amblyomma  marmoreum 
leopard  tortoise 
[Africa] 
No  other  data 
L.  E.  Robinson  det. 

Note  from  Mr.  Robinson  17. XI.  1922:  'They 
originally  showed  the  characteristic  ornamen- 
tation, sufficient  of  which  is  still  visible,  especially 
if  examined  under  alcohol,  to  leave  no  doubt.  The 
coarse  punctations  are  larger  than  those  seen  in 
specimens  off  Rhinoceros  in  East  Africa,  and  the 
dark  ornamentation  is  more  salient,  but  I  do  not 
consider  these  differences  to  be  of  specific  impor- 
tance. The  other  morphological  characters  are 
typical.' 

N572        R288 

Haemaphysalis  hystricis  [sex/stage  not  given]  (3  9 

Haemaphysalis  hystricis) 
ex.  Canis  familiaris 
Lhassia,  Assam,  India  (coordinates  for  Assam — 

26°00'N,93°00'E) 
1907.R.  A.  Lorrain 

Note:  Nuttall  sent  1  cf,  1  9  of  this  collection  to  Or 
L.  O.  Howard  of  the  U.S.  Department  of 
Agriculture.  These  specimens  are  now  deposited 
in  the  Rocky  Mountain  Laboratory  collection 
(RML  109001). 

Publications:  Nuttall  &  Warburton  (1915:  425); 
Hoogstraal  et  al.  (19650:  476)  gave  combined 
collection  data  from  Nuttall  572  and  573. 

N573        R293 

Haemaphysalis  hystricis  [sex/stage  not  given]  (2  9 

Haemaphysalis  hystricis) 
ex.  Canis  familiaris 
Lhassia,  Assam,  India  (coordinates  for  Assam — 

26WN,  93WE) 
1907.R.  A.  Lorrain 

Publication:  Hoogstraal  et  al.  (19650:  476)  gave 
combined  data  from  Nuttall  572  and  573. 

N574        R294C 

Haemaphysalis  bispinosa  [sex/stage  not  given] 

(10  cf,  6  9,  1  N  Haemaphysalis  bispinosa) 
No  other  data 

Haemaphysalis  bispinosa  is  a  vector  of  Kyasanur 
forest  disease  virus. 

N575        R243 

9  Haemaphysalis  hystricis  (2  9  Haemaphysalis 
hystricis)) 


Note:  Nuttall  565  and  3330  also  bear  Rothschild 
number  R243. 

N575x        R163 

Haemaphysalis  leachft  [sex/stage  not  given]  (1  cf 

Haemaphysalis  indica)' 
ex.  Hyaena  crocuta  (Hyaena  hyaena) 
Darie  (locality  not  verified) 
(Exp.  Erlanger  Hilgert) 

Note:  The  Nuttall  number  for  this  collection  is 
575x  in  his  catalogue  but  575a  on  the  vial  label. 
Nuttall  1395  also  bears  Rothschild  number  R 163. 

N576        R294a 

Haemaphysalis  hystricis  [sex/stage  not  given]  (1  9, 

1  N  Haemaphysalis  hystricis) 
ex.  Canis  familiaris 

Lushai  Hills  (23'10'N,  92°50'E),  Assam,  India 
1907.R.  A.  Lorrain 

Publications:  Nuttall  &  Warburton  (1915:  425) 
indicated  that  N576  was  composed  of  1  cf,  3  9. 
Hoogstraal  et  al.  (1965a:  476)  saw  the  9  labeled 
N576ii.  The  nymph,  in  a  separate  vial  bears  the 
label  N576i. 

N577        R294 

Haemaphysalis  leachi  [sex/stage  not  given]  (1  cf,  1 

9  Haemaphysalis  leachi) 
ex.  Ictonyx  capensis  (Ictonyx  striatus) 
Bulawayo  (20°09'S,  28°35'E),  Zimbabwe 
3.IV.1907E.C.Chubb 

Notes:  The  genus  Ictonyx  is  also  known  in  the 
literature  as  Zorilla.  I.  Geoffrey,  1826. 
Nuttall  595  also  bears  Rothschild  number  R294. 
Publication:  Nuttall  &  Warburton  (1915:471). 

N578        R154 

Haemaphysalis    leachi    (?)    (not    long    enough) 

[sex/stage  not  given]  (2  cf,  1  9  Haemaphysalis 

leachi) 

Deelfontein  (20'17'S,  32°38'E),  Zimbabwe 
1. IV.  1 902  C.J.B.  Grant 

Publication:  Nuttall  &  Warburton  (1915:  472) 
indicated  the  genus  as  Suricata.  However,  the 
collection  locality  appears  to  be  out  of  the  known 
range  of  the  meerkat. 

N579        R212 

Haemaphysalis     campanulata     [sex/stage     not 

given]  (2  cf,  1  9  Haemaphysalis  campanulata) 
ex.  Canis  familiaris 
Yokohama  (35°26'N,  139837'E),  Japan 
24. IV.  1902  Allan  Owston 

Note:  Nuttall  609  also  bears  the  Rothschild 
number  R212  but  the  data  do  not  correspond  to 
N579. 

Publication:  Nuttall  &  Warburton  (1915:  472) 
indicated  the  collection  contained  2  cf,  2  9  from  a 
dog. 


10 


J.  E.  KEIRANS 


N580        R160 

1  cf  1  9  Haemaphysalis  punctata 

[no  host  data] 

Cambridge  (52°  1 3'N,  00'08'E),  England 

A.  E.  Shipley  (Nuttall  det.) 

Haemaphysalis   punctata    is    a    vector   of  the 

viruses  Bhanja,  tickborne  encephalitis,  Crimean 

Congo  hemorrhagic  fever,  and  perhaps  Tribec. 

N581        R183 

1  N  Haemaphysalis  spinulosal  [The  species 
epithet  crossed  out  and  sp.  inserted]  (Probably 
H.  leachi  var.  indica)  (1  N  Haemaphysalis  sp.) 
(Bad  condition,  very  queer.  C.  Warburton) 

ex.  flying  squirrel 

Bunguran  (possibly  Bungura  26°20'N,  84'05'E), 
India 

N582        R284 

Haemaphysalis  bispinosa  [sex/stage  not  given]  (2 

9  Haemaphysalis  bispinosa) 
ex.  Equus  caballus 

Lushai  Hills  (23°10'N,  92°50'E),  Assam,  India 
26. VI.  1 907  R.  A.  Lorrain 
Publication:  Nuttall  &  Warburton  (1915:431). 

N583        R291 

Haemaphysalis  bispinosa  [sex/stage  not  given]  ( 1 

cf,  1  9  Haemaphysalis  bispinosa) 
ex.  Talpa  sp. 

Lushai  Hills  (23°10'N,  92°50'E),  Assam,  India 
1907R.  A.  Lorrain 
Publication:  Nuttall  &  Warburton  ( 1 9 1 5:  43 1 ). 

N584        R294b 

Rhipicephalus  haemaphysaloides  [sex/stage  not 

given]       (3       cf,        1        9       Rhipicephalus 

haemaphysaloides) 
No  data 

N585        R242 

Rhipicephalus    appendiculatus    [sex/stage    not 

given]  (2  cf  Rhipicephalus  muehlensi) 
No  data 

N588        R255 

Boophilus  sp.  incert.  [sex/stage  not  given]  (10  9 
Boophilus  microplus) 

No  host  data 

Entre-Rios,  Argentina.  Probably — Provincia  de 
Entra  Rios  (32WS,  59WW),  Argentina. 
(There  are  also  two  small  populated  areas  with 
the  name  Entra  Rios  at  23°18'S,  64°  1 1'W  and 
28°50'S,  64°11'W,  which  could  be  the  collec- 
tion locality.) 

IV.  1 897  Dr  C.  Berg  [C.  Warburton  det.  1918] 

Note:  This  collection  also  contained  1  cf  of  the 

human  sucking  louse  Pediculus  humanus. 

Boophilus  microplus  is  a  vector  of  the  viruses 

Wad    Medani,    Seletar,    and    Crimean    Congo 

hemorrhagic  fever. 


N589        R290 

Argas  persicus  [sex/stage  not  given]  (9  cf,  6  9,  5  N 

Argas  persicus)) 
No  history 

N590        R292 

Argas  persicus  [sex/stage  not  given]  (11  cf ,  4  9,  11 

N  Argas  persicus) 
No  history 

Note:  An  additional  1  cf ,  1  9  of  this  collection  were 
presented  to  J.  de  Meza  of  Zomba,  Malawi, 
19.11.1916. 

N591         R214 

1 6L  Argas  vespertilionis 

ex.  bat  (also  parasites  off  mouse) 

No  particulars 

N592        R195 

3L  Argas  vespertilionis  (&  Nycteribia) 

ex.  Scot 'us  wroughtoni  (Scotophilus  temmincki 

wroughtoni) 
Helwak(17°22'N,  73'44'E),  Maharashtra,  India. 

N593        R224 

Argas  vespertilionis  [sex/stage  not  given]  (3  L 

Argas  vespertilionis) 
ex.         Vesperugo       pipistrellus        (Pipistrellus 

pipistrellus) 

Puttenham  (5 1  °49'N,  00°44'W),  England 
28.VII.1899 

N594        R240 

Ornithodoros  talaje  [sex/stage  not  given]  (14  N 

Ornithodoros  capensis  group) 
ex.  nestlings 
Arrows  Sand?  Puhekiox,   Hawaii  (locality  not 

verified) 
30.VI.1900H.W.  Henshaw 

N597        R153 

Ixodes  hexagonus  [sex/stage  not  given]  (8  N  1  L, 

Ixodes  hexagonus) 
ex.  Erinaceus  europaeus 
Belgium.  E.  A.  Boulenger 
Publication:  Nuttall  &  Warburton  (1911:183). 
Ixodes  hexagonus  is  a  vector  of  tickborne 
encephalitis  virus. 

N598        R206 

Ixodes  hexagonus  [sex/stage  not  given]  (1  9,  6  N 

Ixodes  hexagonus) 
ex.  otter  (Lutra  vulgaris)  (Lutra  lutra) 
No  particulars 

Publication:  Nuttall  &  Warburton  (1911:  182) 
indicated  that  the  locality  was  England. 

N599        R209 

Ixodes  hexagonus  [sex/stage  not  given]  (3  9,  4  N 

Ixodes  hexagonus) 
ex.  Erinaceus  europaeus 
Cambridge  (52°  1 3'N,  00°08'E),  England 
Publication:  Nuttall  &  Warburton  (1911:182). 


ROTHSCHILD  TICKS  IN  BM(NH) 


11 


N600        R221 

Ixodes  hexagonus  [sex/stage  not  given]  (3  9,  1  L 

Ixodes  hexagonus) 
ex.  polecat  Mustela  putorius 
Boxworth  (52'16'N,  00°02'W),  England 
30.IV.1901 
Publication:  Nuttall  &  Warburton  (1911:1 82). 

N601        R208 

Ixodes  hexagonus  [sex/stage  not  given]  (1  9,  41  L 

Ixodes  hexagonus) 
ex.  otter  (Lutra  vulgaris)  (Lutra  lutrd) 
No  other  data 

Note:  One  larva  (RML  16476)  of  this  collection 
was  donated  to  the  Rocky  Mountain  Laboratory 
by  P.  A.  Buxton,  London  School  of  Hygiene  and 
Tropical  Medicine,  in  1940.  (See  also  Nuttall 
1159). 

Publication:  Nuttall  &  Warburton  (1911:  182) 
indicated  that  the  locality  was  England. 

N602        R222 

1  9 Ixodes  hexagonus 

ex.  Mustela  erminea 

Tring  (5 1  '48'N,  00°40'W),  England 

19.V.1903 

Publication:  Nuttall  &  Warburton  (1911:1 82). 

N603        R179a 

Ixodes  hexagonus  [sex/stage  not  given]  (2  9,  1  N 

Ixodes  hexagonus} 
ex.  Mustela  erminea 
Wigginton  (5 1  °47'N,  00'38'W),  England 
4. V.I 903  Barrett  coll. 
Publication:  Nuttall  &  Warburton  (1911:1 82). 

N604        R143 

Ixodes   hexagonus   [sex/stage  not  given]   (4   N 

Ixodes  hexagonus) 
ex.  Mustela  vulgaris  (Mustela  nivalis) 
Tring  (5 1  °48'N,  00°40'W),  England 
20.111.1902. 
Publication:  Nuttall  &  Warburton  (1911:1 82). 

N605        R164 

Ixodes   hexagonus   [sex/stage   not   given]   (3    9 

Ixodes  lividus) 

ex.  Cotile  riparia  (Riparia  riparid) 
Worsborough      Bridge      (53°32'N,      01'28'W), 

England 

6.VII.1906A.  Whitakercoll. 
Publication:  Arthur  (1963:  92) 
Ixodes     lividus     is     a     vector     of    tickborne 
encephalitis  virus. 

N606        R225 

d"s,  9's,  &  larvae  Ixodes  hexagonus  (3  9,  6  N,  8  L 

Ixodes  hexagonus) 
ex.  polecat  (Mustela  putorius) 
Aberystwyth  (52°25'N,  04°05'W),  Wales 
26.XI.1902. 


Publication:  Nuttall  &  Warburton  (1911:  183) 
gave  the  date  as  1902. 

N607        R201 

Ixodes  hexagonus  [sex/stage  not  given]  (1  N,  7  L 

Ixodes  hexagonus) 
ex.  Mustela  erminea 
Tring  (5 1  °48'N,  00°40'W),  England 
3.IX.1902 

Publication:  Nuttall  &  Warburton  (1911:  182) 
indicated  that  the  collection  was  composed  only 
ofnymphs. 

N608        R215 

Ixodes  hexagonus   [sex/stage  not  given]   (4  N 

Ixodes  hexagonus) 
ex.  Erinaceus  europaeus 
Durnberg(48°13'N,  1 1'55'E),  Federal  Republic  of 

Germany 
3.IX.1898 

Publication:  Nuttall  &  Warburton  (1911:  183) 
gave  the  spelling  as  Dernberg  rather  than  the 
catalogue  spelling  of  Dermberg. 

N609        R212 

1  N  Ixodes  hexagonus 

ex.  polecat  (Mustela  putorius) 

Aberystwyth  (52°25'N,  04°05'W),  Wales 

23.XII.1902 

Note:   Nuttall   579   also  bears  the   Rothschild 

number  R2 1 2  but  the  data  do  not  correspond  to 

N609. 

Publication:  Nuttall  &  Warburton  (1911:   183) 

indicated  that  more  than  one  nymph  was  in  this 

collection. 

N610        R219 

N's,  larvae  Ixodes  hexagonus  (1  N,  15  L  Ixodes 

hexagonus) 
ex.  Mustela  erminea 
Tring  (5 1  °48'N,  00°40'W),  England 
7.V.1902 
Publication:  Nuttall  &  Warburton  (1911:1 82). 

N611        R145 

1  N,  1  L  Ixodes  hexagonus 

ex.  Mustela  erminea 

Lyndhurst  (50'52'N,  0 1  °34'W),  England 

20.III.1901G.  Tate 

Publication:  Nuttall  &  Warburton  (1911:   182) 

indicated  that  multiple  nymphs  and  larvae  were 

in  this  collection. 

N612        R180 

9's,   N's  Ixodes  hexagonus  (1    9,   7   N  Ixodes 

hexagonus) 
ex.  Mustela  erminea 
Inismore  (54°  1 8'N,  07'34'W),  Ireland 
23.111.1902 
Publication:  Nuttall  &  Warburton  (1911:1 82). 


12 


J.  E.  KEIRANS 


N613        R202 

9's,  N's  Ixodes  hexagonus  (5  N  Ixodes  hexagonus) 

ex.  stoat  (Mustela  ermined) 

No  other  data 

N617        R177 

1  9  Ceratixodes  putus  (Ixodes  uriae) 

ex.  wild  duck 

No  other  data 

Publication:  Nuttall  &  Warburton  (1911:261). 

Ixodes  uriae  is  a  vector  of  the  viruses  tickborne 

encephalitis,       Tuleniy,       Oceanside,       Zaliv 

Terpenya,  Sakhalin,   Tillamook,   Avalon,  Clo- 

Mor,   Taggert,   Bauline,  Cape   Wrath,   Nugget, 

Tindholmur,  Mykines,  Great  Island,  Okhotskiy, 

Yaquina  Head,  Poovoot,  Paramushir,  and  Runde. 

N618        R220 

1  cf,  1  9, 2  N  Ceratixodes  putus  (Ixodes  uriae) 

British.  E.  G.  Wheler 

No  particulars 

N619        R147 

1  d1,  2  9  Ixodes  tenuirostris  (2  rf,  2  9  Ixodes 
trianguliceps) 

ex.      Hypudaeus      glareolus      (Clethrionomys 

glareolus) 

Campfer  (46°28'N,  09°48'E),  Switzerland 
7.VII.1904  [full  date  on  vial  label].  K.  Jordan. 
Publication:  Nuttall  &  Warburton  ( 1 9 1 1 :  248). 
Ixodes  trianguliceps  is  a   vector  of  tickborne 
encephalitis  virus. 

N621        R204 

N's  Ixodes  tenuirostris  (1  N  Ixodes  trianguliceps) 

ex.  Sorex  vulgaris  (Sorex  araneus) 

Bevendean,  (50°5 1  'N,  00°05'E),  England 

VI.1899. 

Publication:  Nuttall  &  Warburton  (1911:  248), 

indicated  that  more  than  one  nymph  was  in  the 

collection. 

N622        R166 

9  Ixodes  tenuirostris  (1  9  Ixodes  trianguliceps) 

ex.  Arvicola  amphibius  (Arvicola  terrestris) 

Reservoir 

6.IV.1892 

Publication:  Nuttall  &  Warburton  (1911:  248) 

indicated  the  collection  was  from  England. 

N623        R197 

2  9  Ixodes  tenuirostris  (Ixodes  trianguliceps) 
ex.  Arvicola  agrestis  (Microtus  agrestis) 
Tring  (5 1  '48'N,  00°40'W),  England 
Publication:  Nuttall  &  Warburton  (1911:  248). 

N624        R149 

1  9  Ixodes  tenuirostris  (Ixodes  trianguliceps) 

ex.  Arvicola  arvalis  (Microtus  arvalis) 

Campfer  (46°28'N,  09°48'E),  Switzerland 

20.  VII.  1904  K.Jordan 

Publication:  Nuttall  &  Warburton  (1911:  248). 


N625        R148 

3  9  Ixodes  tenuirostris  (Ixodes  trianguliceps) 

ex.  Arvicola  arvalis  (Microtus  arvalis) 

Campfer  (46°28'N,  09°48'E),  Switzerland 

5. VII.  1904  K.Jordan 

Publication:  Nuttall  &  Warburton  ( 1 9 1 1 :  248). 

N626        R226 

d"s,   9's   Ixodes   scapularis   (4   rf,   4   9   Ixodes 

scapular  is) 

ex.  Cariacus  virginianus  (Odocoileus  virginianus) 
Craven  Co.  (35'07'N,  77°03'W),  North  Carolina, 

U.S.A. 
13.XI.1907  k[det.  C.  Warburton  5. 11.1909.  Found 

with  N666]' 

Publication:  Nuttall  &  Warburton  (1911:  158) 
gave  the  date  of  collection  as  XI.  1897.  The  date 
1 3.XI.  1 897  is  also  on  the  vial  label. 

N633        R169 

N's  Amblyomma  sp.  (8  N  Amblyomma  sp.) 

ex.  Macropus  sp. 

No  other  data 

Note:    Nuttall   644   also  bears  the   Rothschild 

number  R169  and  gives  the  additional  data:  W. 

Australia  28.VII.  1900. 

N634        R159 

Ixodes  percavatus  (2  9,  2  N  Ixodes  rothschildi 

TYPES) 

ex.  Puffin  Fratercula  arctica.  Wrongly  labelled  Ix. 
putus  '(Pocock)'  On  back  of  label;  no  parti- 
culars.   (Sent    to    Neumann    who   confirmed 
the  determination  II.  1909.) 
No  other  data. 

Publications:  Nuttall  &  Warburton  (1911:  221) 
described  the  9  and  N  of  Ixodes  percavatus 
rothschildi.  Zumpt  (1952:  17)  raised  this  taxon  to 
specific  rank;  Arthur  (1953:  222-226)  indepen- 
dently raised  /.  percavatus  rothschildi  to  specific 
rank;  Arthur  ( 1 963 :  1 34)  referred  to  2  9, 3  N  in  the 
type  lot. 

There  was  (IX.  1977)  a  note  in  the  vial  with  the 
words '1  N  Missing'. 

N637        R213 

Ixodes  bicornis  [sex/stage  not  given]  ( 1 1  9  Ixodes 

boliviensis) 
ex.  Felis  concolor 
'[American]'  No  other  data 
Publication:  Nuttall  &  Warburton  (1911:   187) 
indicated  only  one  9  in  the  collection. 

N638        R276 

1  cf,  1  9  Ixodes  loricatus 

ex.  Didelphia  (Didelphis  sp.) 

Sapucay.  Foster  coll. 

Note:    Locality    is    probably    Sapucai    (25°  1 9'S, 

56°55'W),  Paraguay,  which  is  in  the  range  of  /. 

loricatus.  [See  also  Nuttall  1 39 1]. 

Publications:  Nuttall  &  Warburton  (1911:  269); 

Cooley&  Kohls  (1945:  190). 


ROTHSCHILD  TICKS  IN  BNi(NH) 


13 


N639        R256 

Ixodes  loricatus  [sex/stage  not  given]  (1  d  Ixodes 

loricatus) 
ex.  'Piquoti' 
No  other  data 

N640        R241 

2  rf,  2  9,  2  N  Ixodes  loricatus  (2  cf,  2  9  Ixodes 

loricatus) 

ex.  A  teles  melanochir  (A  teles  g.  geoffroyi) 
Frontera  (18'32'N,  92°38'W),  Tabasco,  Mexico. 

Note    following    locality    '(Compared    with 

Neumann's  types  at  Toulouse,   1910;  agrees 

fully  but  Nn's  type  larger).' 
Publications:  Nuttall  &  Warburton  (1911:  269); 
Cooley&  Kohls  (1945:  190). 

N641        R161 

Ixodes  loricatus  [sex/stage  not  given]  (2  rf,  1  L 

Ixodes  loricatus) 

ex.  Didelphys  aurita  (Didelphis  marsupialis) 
Brazil  24.XI.1901 

Publications:  Nuttall  &  Warburton  (1911:  269); 
Cooley&  Kohls  (1945:  190). 

N642        R165 

1  9,  1  N,  6  L  Ixodes  vestitus  (1  9,  5  N  Ixodes 
vestitus) 

ex.  Diemenia  superciliosa  (Diemenia  textilis) 
Herdman's  Lake,  near  Perth  (31'56'S,  1 15'15'E), 

W.  Australia 
B.  H.  Woodward 

Note:  Nuttall  628  also  bears  Rothschild  numfjer 
R165. 

Publications:  Nuttall  &  Warburton  (1911:  255) 
indicated  larvae  as  did  the  catalogue  entry.  These 
were  missing  7.X.1977.  Taylor  (1946:  59,  60) 
repeated  Nuttall  &  Warburton's  description  and 
figures.  Roberts  ( 1 960:  45 1 , 452). 

N643        R265 

2  N  Ixodes  holocyclus 

Perth  (3 1  °56'S,  1 1 5°  1 5'E),  W.  Australia 

B.  H.  Woodward  coll. 

Publication:  Nuttall  &  Warburton  (1911:238). 

N645        R275 

3  9  Ixodes  australiensis 

ex.  tame  dog  (Canis  familiar  is) 

Cranbrook  (34°  1 8'S,  1 1 7°32'E),  West  Australia 

1. IV.  1 900  J.T.  Young 

Publications:  Nuttall  &  Warburton  (1911:  252). 

Taylor    (1946:    57,    58)    repeated    Nuttall    & 

Warburton's  descriptions  and  figures. 

Note:  This  collection  was  seen  by  R.  A.  Cooley  in 

1946    who    made    3    slides    of   dissections    (9 

spiracular  plate;  9  legs  I  &  IV;  9  hypostome)  all  in 

the  Nuttall  collection  at  the  British   Museum 

(N.H.).    He   gave   them   the    Rocky   Mountain 

Laboratory  (RML)  number  22284. 


N646        R210 

Ixodes  australiensis  [sex/stage  not  given]  (8  9 

Ixodes  australiensis) 
ex.  Bettongia  lescuri  (Bettongia  lesueuri) 
Kojonup  (33°50'S,  1 1 7°09'E),  W.  Australia 
4.XI.1900 

Publications:  Nuttall  &  Warburton  (1911:  252). 
Taylor  (1946:  57,  58)  repeated  Nuttall  & 
Warburton's  description  and  figures.  Roberts 
(1960:  440);  Roberts  (1970:  40)  gave  the  correct 
spelling  of  the  locality. 

N647        R248 

3  9  Ixodes  loricatus  var.  spinosus  n.  var.  Nuttall, 

1 9 1 0  (2  9  Ixodes  luciae) 
ex.  large  opossum  (Didelphis  sp.) 
Frontera  ( 1 8°32'N,  92'38'W),  Tabasco,  Mexico 
May.  Found  with  648.  Note  long  spine  on  coxa  I. 
Note:  These  are  the  TYPE  SPECIMENS  of  /. 
loricatus  spinosus. 

Publications:  Nuttall  (1910:  411),  original  de- 
scription. Nuttall  &  Warburton  (1911:  269), 
repeated  original  description  and  figures. 

N648        R248 

N's  &  larvae  Amblyomma  Icajennense  (16  N 

Amblyomma  cajennense) 
ex.  large  opossum  (Didelphis  sp.) 
Frontera  ( 1 8°32'N,  92°38'W),  Tabasco,  Mexico 
May 
Note:  Found  with  Nuttall  647 

N649        R185 

1  N  Ixodes  vespertilionis. 

ex.  Vespertilio  tricolor  (Myotis  tricolor) 

Kei  Road,  W.  Kingwilliamstown,  S.  Australia  [S. 

Australia  crossed  out  and  S.  Africa  inserted]. 

Kei   Road,    King  William's   Town   (32'51'S, 

27°22'E),  South  Africa. 
Purchased  of  Capt.  H.  Trevelyan. 
Publications:  Nuttall  &  Warburton  (1911:  277). 
Taylor  (1946:   41)  considered  /.    vespertilionis 
'very     doubtfully     Australian'     and     'original 
specimen  is  unfortunately  lost'.  Hoogstraal  (1956: 
569,  570)  considered  this  collection  to  be  of  South 
African  origin. 

Ixodes  vespertilionis  is  a  vector  of  the  viruses 
Issyk  Kul  and  tickborne  encephalitis. 

N650        R203 

1  9  Ixodes  fecialis  TYPE  (HOLOTYPE) 

ex.  Dasyurus  geoffroyi 

Cranbrook  (34°  1 8'S,  1 17'32'E),  W.  Australia 

5.III.1900 

Publications:  Warburton  &  Nuttall  (1909:  58) 

original  description.  Nuttall  &  Warburton  (1911: 

248)  repeated  the  original  description  and  figures 

as  did  Taylor  (1946:  56).  Roberts  (1960:  445) 

incorrectly  gave  the  collection  date  as  March 

1910. 


14  J.  E.  KEIRANS 

N652        R168 

2  9  Ixodes  unicavatm(\  9,  1  N  Ixodes  unicavatus) 

ex.  shag  (Phalacrocorax  aristotelis) 

Plymouth  (50'23'N,  04°10'W),  England 

IX.1897 

Publications:  Nuttall  &  Warburton  (1911:  266) 

indicated  that  the  collection  consisted  of  only  one 

nymph.  Arthur  ( 1 963:  1 26)  repeated  this  error. 

N666        R226 

1  9  Dermacentor  nigrolineatus  (1  9  Dermacentor 

albipictus) 

ex.  Cariacus  virginianus  (Odocoileus  virginianus) 
Craven  Co.  (35807'N,  77°03'W),  North  Carolina, 

U.S.A.  Found  with  N626  q.v.  for  details. 
C.  Warburton  det.  1917 

Dermacentor  albipictus  is  a  vector  of  Colorado 
tick  fever  virus. 

N667        R250 

9's,  N's  Ixodes  unicavatus  (39,  10  N,  3  L  Ixodes 

unicavatus) 
ex.     Phalacrocorax     graculus     (Phalacrocorax 

aristotelis) 

Cromarty  (57°40'N,  04°02'W),  Scotland. 
10.111. 1898  W.Roy 

Publications:  Nuttall  &  Warburton  (1911:  266): 
Arthur  (1963:  126). 

N669        R178 

Haemaphy sails  humerosa  [sex/stage  not  given] 

TYPES  (2  d1,  1  9,  1  N  Haemaphysalis  humerosa 

TYPES) 

ex.  Parameles  macrura  (Isoodon  macrourus) 
Barrow    Island    (20°48'S,     115°23'E),    N.    W. 

Australia 

Note:  An  additional  1  cf  of  this  collection  was  sent 
to  Dr  L.  O.  Howard,  U.S.  Dept.  of  Agriculture, 
Washington,  D.C.,  U.S.A.  2 1  .V.  1 909. 
Publications:  Warburton  &  Nuttall  (1909:  60) 
original  description.  Nuttall  &  Warburton  (1915: 
496,  497)  and  Taylor  (1946:  82)  repeated  original 
description  and  figures.  Roberts  (1963:  39)  dis- 
cussed the  type  series  and  the  note  inserted  in  the 
tube  by  Nuttall  that  one  specimen  was  missing 
20.X.1934. 

N670        R172 

Haemaphysalis  elongata  [sex/stage  not  given]  (1  d1 
Haemaphysalis  elongata,  4  d1,  2  9  Haema- 
physalis tiptoni) 

ex.  Centetes  ecaudatus  (Tenerec ecaudatus) 

Madagascar 

Note:  The  male  of  H.  elongata  has  been  given  a 

collection    number    by    the    Rocky    Mountain 

Laboratory     (RML     71004)     and     by     Harry 

Hoogstraal(HH44251). 

Publications:  Nuttall  &  Warburton  (1915:  500). 

Hoogstraal   (1953:    59,   64)   discussed   the   two 

species  represented  in  Nuttall  &  Warburton's 

(1915)  figures  and  descriptions. 


N826        R189 

2  9  Ixodes  hexagonus  (Ixodes  rubidus) 

ex.    Coypu,    Myopotamus    coypus    (Myocaster 

coypus) 

S.  America.  No  further  particulars 
Publication:  Nuttall  &  Warburton  (1911:1 83). ' 

N892        R295 

Haemaphysalis  bispinosa  [sex/stage  not  given]  (7 

d,  7  9  Haemaphysalis  bispinosa) 
ex.  Canis  familiaris 
Luthaia  Mts,  Assam 

Note:  The  locality  is  questionable.  There  are  no 
Luthaia  Mts  as  written  in  the  catalog  or  Southaia 
Mts  as  in  Nuttall  &  Warburton  (1915:  431)  to  be 
found  in  gazetteers  of  India.  I  believe  the  locality 
to  be  Lushai  Hills  (23°10'N,  92°50'E),  Assam, 
India  as  in  N576,  (R294a)  and  N582  (R284). 
Publication:  Nuttall  &  Warburton  ( 1 9 1 5:  43 1 ). 

N1066        R245 

5  9   Ixodes   texanus   var.    (Ixodes   sp.    not   /. 
texanus  or  /.  hearlei) 

No  other  data 

N1067        R200 

1  9 Ixodes  hexagonus 

ex.  fox  ( Vulpes  vulpes) 

Boxworth  (52°  1 6'N,  00°02'W),  England 

22.1.1898 

Publication:  Nuttall  &  Warburton  (1911:1 82). 

N1068        R190 

6  d,  6  9  Ixodes  canisuga  (4  d1, 4  9  Ixodes  lividus) 
Lyndhurst       (50°52'N,       01°34'W),       England 

11. VII.  1900 

Publications:  Nuttall  &  Warburton  (1911:  214). 
Arthur  (1 963:  92)  indicated  6  d,  6  9  present.  There 
were  4  d,  4  9  in  the  collection  as  of  IX.  1977. 

N1069        R146 

1    d,    2    9,    3    N    Ixodes    tenuirostris   (Ixodes 

trianguliceps) 

ex.  Arvicola  arvalis  (Microtus  arvalis) 
Campfer  (46°28'N,  09°48'E),  Switzerland 
1 1.  VII.  1 904  Dr  K.Jordan 
Publication:  Nuttall  &  Warburton  (1911:  248). 

N1105        None 

3  d1,  3  9, 6  N  Dermacentor  albipictus 

Blucher     Hall     (51'06'N,     120°01'W),     British 

Columbia,  Canada 
III. 1910.    Rec'd   2.IV.1910    from    Hon.    N.    C. 

Rothschild  (London). 

N1151         R296 

10  L  Argas  vespertilionis 

ex.  P ipistrellus  pipistrellus 

Oristano  (39'54'N,  08°35'E),  Sardinia 

May  1907  DrA.  Krausse 


ROTHSCHILD  TICKS  IN  BM(NH) 


15 


N1152        R175 

6  LArgas  vespertilionis 

ex.         Vesperugo       pipistrellus        (Pipistrellus 

pipistrellus) 

Tring  (5 1  °48'N,  00°40'W),  England 
1. IX.  1898 

Nil  53        R244 

2  L  Argas  vespertilionis 

No  other  data 

Ml  154        R271 

2  L  Argas  vespertilionis  (Argas  (Carios)  sp.) 
Perth  (3 1  °56'S,  1 1 5°50'E),  W.  Australia 
Note:   These  specimens  may  represent  a  new 
species.  It  is  not  Argas  australiensis,  Argas  daviesi 
or  Argas  dewae,  the  currently  known  representa- 
tives of  the  subgenus  Carios  in  Australia. 

N1159        R142 

N's  Ixodes  hexagonus  (3  N  Ixodes  hexagonus) 

ex.  Mustela  erminea 

Tring  (5 1  °48'N,  00°40'W),  England 

10.IV.1902. 

Note:  One  nymph  (RML  16476)  of  this  collection 

was  donated  to  the  Rocky  Mountain  Laboratory 

collection  by  P.  A.  Buxton,  London  School  of 

Hygiene  and  Tropical  Medicine,  in  1940.  (See 

also  Nuttall  601). 

Publication:  Nuttall  &  Warburton  (1911:1 82). 

N1160        R194 

L's  Ixodes  hexagonus  (17  L  Ixodes  hexagonus) 
ex.  fox  Vulpes  vulpes  (cub) 
Boxworth  (52°  16'N,00°02'W),  England 
30.111.1900 

Note:  2  additional  larvae  from  this  collection 
were  sent  to  Prof.  R.  Matheson,  Cornell  Univer- 
sity, Ithaca,  N.Y.,  U.S.A. ,  27. V.  1929. 
Publication:  Nuttall  &  Warburton  (1911:182). 

N1161         R278 

9  Ixodes  tenuirostris  (Ixodes  trianguliceps) 

Tarasp(46°38'N,  10'25'E),  Switzerland 

summer  1901 

Publication:  Nuttall  &  Warburton  (1911:  248) 

N1162        R207 

2  L  Ixodes  sp. 

ex.  Dromicia  concinna  (Cercartetus  concinnus) 

Note:   The  nocturnal   marsupial   C.   concinnus 

inhabits  the  southern  part  of  western  and  central 

Australia. 

N1163        R247 

L's  Argas  vespertilionis  (1 3  L  Argas  vespertilionis) 

ex.         Vesperugo       pipistrellus        (Pipistrellus 

pipistrellus} 

Yalding  (5 1  °  1 4'N,  00°26'E),  England. 
8. VIII.  1 897  [However,  1 997  was  written] 
W.  R.  Ogilvie-Grant 


N1164        R250,251 

L's  Argas  sp.  (5  L  Argas  boueti) 

ex.  either  Phalacrocorax  graculus,  Cromarty,  or 

Megaderma  cor,  Salamana,  Havish,  4,500  ft 

(Number  on  cork  defaced) 
ex.  Megaderma  cor. 

Salamana,  Havish  1,372  m  (locality  not  verified) 
Notes:  Megaderma  cor  ranges  in  East  Africa  from 
Ethiopia  to  Tanzania. 

The  correct  Rothschild  number  for  this  collection 
is  R25 1 .  For  R250  see  Nuttall  667. 

N1166        R230 

10     nymphs     [genus     &     species     not    given] 

(Amblyommasp.) 

ex.  Mus  escularis  (name  not  verified) 
St   Aignan.    Possibly    Misima    Island   (10'41'S, 

152°42'E),  Louisade  Archipelago.  This  island 

was  formerly  known  as  St  Aignan. 
VIII-XI.  1897  (Meek) 

N1167        R186 

N1167a 

2  N  Ixodes  sp.  [1 167],  1  d  Hyalomma  aegyptium 

[1 167a]  (Hyalomma  truncatum) 
ex.  a  'Sassaby'  a  bastard  hartebeest 
(Damaliscus  lunatus)  Republic  of  South  Africa. 

Date? 

Hyalomma  truncatum  is  a  vector  of  the  viruses 
Dugbe,  Bhanja,  Jos,  and  Crimean  Congo 
hemorrhagic  fever. 

N1211         R237 

2  N  Ixodes  sp.  (2  N,  1  L  Ixodes  fecialis) 

ex.   Tarsipes  rostratus  or  Smiothopsis  murinus 

[vial    label    reads    S.    murinus]    (Sminthopsis 

murina) 

Albany  (35'02'S,  1 1 7°53'E),  W.  Australia 
2 1. XII.  1 900  B.  H.  Woodward 

N1212        R272 

5  N  Ixodes  sp.  (Ixodes  holocyclus) 

Perth  (3 1  °56'S,  1 1 5°50'E),  W.  Australia 

B.  H.  Woodward 

No  further  details 

N1213     R216 

1  9  Ixodes  fecialis  var.  aegrifossus  (Ixodes  fecialis) 

ex.  Perameles  obesula  (Isoodon  obesulus) 

Bannister  (32°40'S,  1 16'33'E),  W.  Australia 

21.VIII.1900 

Publication:  Nuttall  &  Warburton  (191 1:  250). 

N1386        R249 

N's,  L's  Amblvomma  sp.  (14  N,  2  L  Amblyomma 

sp.) 
ex.  Cariacus  virginianus  mexicanus  (Odocoileus 

virginianus) 

San  Rafael  (20'12'N,  96°5 1  'W),  Veracruz,  Mexico 
27. III. 1903 


16 


J.  E.  KEIRANS 


N1387        R264 

1  N  ?  genus  (Dermacentor  andersoni) 

ex.  Lagomys  princeps  (Ochotona  princeps) 
Canadian  National  Park,  ?  Ottawa,  Canada 
5. VIII.  1 899  J.  F.  Dippie 

Note:  Ottawa,  Ontario  is  far  to  the  east  of  the 
ranges  of  D.  andersoni  and  the  pika.  The 
Canadian  National  Park  and  the  date  1899 
suggests  a  relationship  to  N1390  (R280),  Banff 
National  Park,  which  is  within  the  range  of  both 
O.  princeps  and  D.  andersoni. 
Dermacentor  andersoni  is  a  vector  of  the  viruses 
Powassan  and  Colorado  tick  fever,  the  bacterium 
of  tularemia,  and  the  rickettsia  of  Rocky 
Mountain  spotted  fever. 

N1389        R155 

larvae  Ixodes  sp.  (3  N  Ixodes  sp.) 

ex.       Crysomys       longicaudatus       (Oryzomys 

longicaudatus) 

Valparaiso  (33°02'S,  7 1  °38'W),  Chile 
Date?J.  A.Wolffshon 

N1390        R280 

2  N  ?  genus  '(has  eyes!)'  (2   N  Dermacentor 
andersoni) 

Banff  (5 1  °38'N,  1 1 6°22'W),  Alberta,  Canada 
19.VII.1899 

N1391        R269 

4  N  ?  genus  (3  N  Amblyomma  sp.) 

ex.  Didelphys  sp.  (Didelphis  sp.) 

Sapucai  (25'19'S,  56°55'W),  Paraguay 

?Date 

N1392        R187 

N's  Amblyomma  sp.  (10  N  Amblyomma  sp.) 
ex.  Hare  wallaby  (Lagorchestes  conspicillatus) 
Barrow    Island    (30'48'S,     115'23'E),    N.    W. 
Australia 

N1393        R176 

larvae  Amblyomma  sp.  (64  L  Amblyomma  sp.) 

ex.  Bettongia  penicillata 

Bokerup,  W.  Australia.  Possibly  Bokarup  Swamp 

(34'24'S,  116°50'E). 
20. IV.  1900 

N1394        R227 

2  N  Amblyomma  sp. 

ex.  Ontalis  vetula  macalii  (Neotoma  alleni  vetula) 

Frontera       (18'32'N,       92'38'W),       Tabasco, 

Switzerland  [sic.]  Mexico 
18.V.- 

N1395        R163 

N's  Rhipicephalus  sp.  (2  N  Rhipicephalus  sp.) 

ex.  Hyaena  crocuta  (Hyaena  hyaena) 

Darie  (locality  not  verified) 

?  Date.  Von  Erlanger 

N1396        R246 
1  N  Amblyomma  sp. 


ex.  Black  squirrel  (Sciurus  sp.) 

Frontera  ( 1 8°32'N,  92°38'W),  Tabasco,  Mexico 

19.V.- 

N1397        R171 

larvae  Amblyomma  sp.  (26  L  Amblyomma  sp.) 

ex.  Macropus  eugenii  (Macropusfuligenosus) 

Cranbrook  (34°  1 8'S,  1 1 7°32'E),  W.  Australia 

6.III.1910 

N1398        R150 

1  larvae?  genus  (1  LHyalommasp.) 

ex.  Gerbillus  gerbillus 

Zaghig* 

1  .III.  1903  Hon.  N.  C.  Rothschild 

""Locality  not  verified.  However,  N.  C.  Rothschild 
was  in  the  Nile  delta  early  in  1903.  Quite 
possibly  the  locality  is  Zagazig  (30°35'N, 
31*3  l'E),  Egypt. 

N1399        R162b 

N's  Ixodes  sp.  (4  N,  4  L  Ixodes  texanus) 

ex.  Putorius  energermanis  (Mustela  vison) 

Sumas,  B.C.,  Canada* 

III.  1903 

*Either  Sumas  Lake  (49'07'N,  122°02'W),  British 

Columbia,     Canada     or     Sumas     (49°00'N, 

122813'W),  Washington,  U.S.A. 

N1497        None 

1  N  Dermacentor  sp.  (Dermacentor  albipictus) 

ex.  Moose  (Alces  alces) 

New  Brunswick,  (47WN,  66WW),  Canada 

30.XI.1911  J.  Birrellcoll. 

N1498        None 

1  cf  Dermacentor  albipictus 

ex.  man  (Homo  sapiens)  (Dr  Walker) 

New  Brunswick  (47WN,  66WW),  Canada 

29.XI.1911  J.  Birrellcoll. 

N1499        None 

2  c?  Dermacentor  albipictus 
ex.  Moose  (Alces  alces) 

New  Brunswick  (47WN,  66WW),  Canada 
30.XI.1911  J.  Birrellcoll. 

N1500        None 

2  cf,  1  9,  1  N  Dermacentor  albipictus 

ex.  carabou  [sic.]  (Rangifer  tarandus) 

New  Brunswick  (47WN,  66WW),  Canada 

25.XI.1911  J.  Birrellcoll. 

N1501        None 

1  N  Dermacentor  sp.  (Dermacentor  albipictus) 

ex.  carabou  [sic.]  (Rangifer  tarandus) 

New  Brunswick  (47WN,  66'00'W),  Canada 

25. XI. 1911  J.  Birrell  coll. 

N1502        None 

1  N  Dermacentor  sp.  (Dermacentor  albipictus) 

ex.  carabou  [sic.]  (Rangifer  tarandus) 

New  Brunswick  (47'00'N,  66WW),  Canada 

25. XI.  19 11  J.  Birrellcoll. 


ROTHSCHILD  TICKS  IN  BM(NH) 


17 


N1503        None 

1  d  Dermacentor  sp.  (5  N  Dermacentor  albipictus) 

ex.  carabou  [sic.]  (Rangifer  tarandus) 

New  Brunswick  (47'00'N,  66WW),  Canada 

25.XI.1911  J.  Birrellcoll. 

N1504        None 

1    9   Ixodes   hexagonus   (Ixodes   cookei)   '[Not 

typical — bad  condition;  possibly  /.  rubidus]' 
ex.  black  cat 

New  Brunswick  (47WN,  66WW),  Canada 
19.XII.1911  J.  Birrellcoll. 
Ixodes  cookei  is  a  vector  of  Powassan  virus. 

N3138        R926 

1  d  Aponomma  laeve  var.  capensis  (Aponomma 
sp.) 

ex.  Echidna  sp. 
No  further  data 
Gift  of  Hon.  N.  C.  Rothschild  "(unmounted  by 

me  from  a  slide  R926)".  L.  E.  Robinson  del. 
Note:   This  is  not  Aponomma  concolor  or  A. 
oudemansi.  Coxal  spurring  is  similar  to  A.  pattoni 
but  the  specimen  lacks  cervical  pits.  It  is  a  species 
near  A.  pattoni. 

N3160        R238 

2  d1,  1  9,  1  N  Dermacentor  reticulatus  (  1  d1,  1  9,  1 
N  Dermacentor  reticulatus) 

Revelstoke,  and  Revelstoke  Point,  now  known  as 

Stoke  Point  (50°  1 8'N,  04°0 1  'W),  England 
III.1902. 

Dermacentor  reticulatus  is  a  vector  of  the  viruses 
tickborne  encephalitis,  Russian  spring  summer 
encephalitis,  and  Omsk  hemorrhagic  fever. 

N3330        R243 

1  9  Dermacentor  auratus  (Dermacentor  sp.) 

No  other  data 

Note:  Nuttall  565  and  575  also  bear  Rothschild 

number  R243. 

N3502        None 

d's,    9's    Dermacentor    andersoni    (7    d,    9    9 

Dermacentor  andersoni) 
Okanagan    Falls   (49'21'N,    119'31'W),    British 

Columbia,  Canada 
V.  19 13  Rothschild  coll. 
Gift  of  Seymour  Hadwen,  1922 


N3537        None 

N's  Haemaphysalis  leachi  (ION  Haemaphysalis 

erinacei) 

ex.  Dipodillus  campestris  (Gerbillus  campestris) 
Timgad  (35°30'N,  06°35'E),  Algeria 
1 5. IV.  1920  N.  C.  Rothschild  &  K.  Jordan,  coll. 
Publication:  Hoogstraal  (1955:222, 223)  (see  note 
under  3539). 

N3538        None 

N's  Haemaphysalis  leachi  (UN  Haemaphysalis 

erinacei) 

ex.  Dipodillus  campestris  (Gerbillus  campestris) 
Timgad  (35'30'N,  06835'E),  Algeria 
15.IV.  1920  N.  C.  Rothschild  &  K.  Jordan  coll. 

N3539        None 

N's  Haemaphysalis  leachi  (4  N  Haemaphysalis 

erinacei) 

ex.  Dipodillus  campestris  (Gerbillus  campestris) 
Timgad  (35°30'N,  06°35'E),  Algeria 
1 7.IV.  1 920  N.  C.  Rothschild  &  K.  Jordan  coll. 
Publication:    Hoogstraal   (1955:   223)  gave  the 
Nuttall   number  3739   for  this  collection.   He 
indicated  the  total  number  of  nymphs  as  1 7  in 
combined  collections  3537  and  3539.  In  1977  the 
total  number  of  nymphs  in  these  two  collections 
was  14. 

A  single  tick  collection  given  by  the  Hon. 
Walter  Rothschild  to  G.  H.  F.  Nuttall  via  Guy 
A.  K.  Marshall 

N888        None 

2   d,    1    9   Hyalomma   aegyptium   (Hyalomma 

rufipes) 

ex.  giraffe  (Giraffa  camelopardalis  subsp.) 
Ogo,  Senegal 
ex.    Riggenbach    coll.    given    by    Hon.    Walter 

Rothschild       to       African       Entomological 

Committee  (rec'd  19.X.1909  from  Guy  A.  K. 

Marshall). 

Notes:  There  are  two  populated  places  and  one 
campsite  with  the  name  Ogo  in  Senegal:  PPL. 
14°35'N,  15°50'W;  PPL.  15°33'N,  13°17'W; 
CMP.  14°45'N,  15°03'W. 

18  specimens  of  this  collection  were  received 
11. X.I  909  by  the  Entomological  Research 
Committee  (Anon.  1910). 


Rothschild  specimens  missing  from  Nuttall  collection 


N75        R1015,  1017,  1019 

3  d  Amblvomma  clvpeolatum  ( =  atrogenatum  N. 

&W.) 
ex.  tortoise 
Zoological        Society's        Gardens,        London, 


11. X. 1895 

Presented  by  N.  C.  Rothschild 
Mounted  (i)  opaque  &  (ii)  (iii)  transparent  on  3 

slides  nos.  1015,1017,1019 
L.  E.Robinson  det.  26.111. 19 19 


18 


J.  E.  KEIRANS 


N76        R964 
1  $  Amblyomma  sp. 
ex  Monitor 

Kalao  Island.  No  further  data 
Mounted  opaque  on  slide  No.  964  from  N.  C. 
Rothschild 

N77        R1033,  1034,  1035 

3  cf  Amblyomma  marmoreum 

ex.  Testudo  pardalis  (leopard  tortoise) 

No    further    data.     Mounted    as    transparent 

specimens  on  slides  Nos.  1033,  1034,  1035 
Presented  by  N.  C.  Rothschild 
L.  E.  Robinson  det.  26. III.  1919 

N78        R996,997,999 
1  cf,  2  9  Amblyomma  albolimbatum 
ex.  black  &  white  snake 

30.1.1896.  No  further  data.  Mounted  as  trans- 
parent specimens  Nos.  997, 996, 999 
Presented  by  N.  C.  Rothschild 
L.  E.  Robinson  det.  26.111.1919. 

N90        R938-941 

4  d1  Aponomma  gervaisi  var.  lucasi 

ex.  Ophiophagus  bungarus  (Hamadryad) 

Presented  by  N.  C.  Rothschild 

No  particulars.  Mounted  on  slides  938-941;  2  in 

glycerine,  2  in  balsam 
C.  Warburton  det.  1.1915 

N91        R900 

1  cf  Aponomma  gervaisi  var.  lucasi 
ex.  Varanus  civitatus  (Monitor) 
Presented  by  N.  C.  Rothschild 

No  particulars.   Mounted  as  slide  900  in  dil. 
glycerin.  C.  Warburton  det.  1. 1 5 

N92        R962,965 

2  c?  Aponomma  gervaisi 

ex.  Varanus  civitatus  (Monitor) 
Presented  by  N.  C.  Rothschild 
No  particulars.  Mounted  as  slides  962,  965  in 
glycerin.  C.  Warburton  det.  1.1915 

N93        R959,963,964,966 

4  d1, 1  9  Amblyomma  decoratum 

ex.  Monitor 

Kalas  Island 

Presented  by  N.  C.  Rothschild 

No  particulars.  5  slides,  3  in  glycerin  2  cf  in 

balsam:    R959,    961,    963,    964,    966.    C. 

Warburton  det.  1.1915 

N216        R96 1-973 

9's,  N's,  L's  Ixodes  hexagonus 

ex.  Lulra  vulgaris 

No  doubt  British 

N.  C.  Rothschild  7  slides 

N217        R930,931 
2  9 Ixodes  hexagonus 
ex.  fox  (Canis  vulpes) 


No  doubt  British 

Gift  of  N.  C.  Rothschild  (2  slides  R930-93 1 ) 

N298        R1000-1011 

6  9,  5  N,  1  L  Ixodes  hexagonus 

ex.  stoat  (Putorius  ermined) 

No  doubt  British 

Gift  of  N.  C.  Rothschild  12  slides 

N322        R929 
1  9 Ixodes  hexagonus 
ex.  Mustela  putorius  (ferret) 
Probably  British 

Gift  of  N.  C.   Rothschild.   R929   mounted  in 
glycerin  in  glass  cell.  G.  H.  F.  Nuttall  det. 

N324        R944-946 
3  N  Ixodes  hexagonus 
ex.  Erinaceus  europaeus 
Probably  British 

Gift  of  N.  C.  Rothschild  (3  slides  R944-946).  G. 
H.  F.  Nuttall  det. 

N395        R988-990 
3  L  Ixodes  unicavatus  slightly  fed 
ex.  Phalacrocorax  graculus  (shag) 
No  locality  etc.  Probably  British 
Gift  of  N.  C.  Rothschild.  Mounted  on  3  slides 
R988-90.  G.  H.  F.  Nuttall  det.  13.11.1915 

N396        R1032 

1  N Ixodes  brunneusl 

ex.  Erithacus  rubecula  (robin) 

No  data.  Probably  British 

Gift  of  N.  C.  Rothschild 

Mounted  on  a  slide  R 1032 

N397        R911 
1  cf  Amblyomma  hirtum 
ex.  bird  (in  spirit) 
Galapagos  Islds,  Pacific 
Ocean  (W.  of  Ecuador) 
Gift  of  N.C.Rothschild 
Mounted  on  a  slide  R9 1 1 
6.III.1915 


L.  E.  Robinson  det. 


N509        R982, 984-987 

4  9, 2  N  Ixodes  ricinus 

ex.  Cervus  capreolus  (roe  deer) 

?  British 

Gift  of  N.C.Rothschild 

Mounted  on  5  slides  R982, 984-987 

N516        R901-3,905 

5  N  Ixodes  putus 

ex.  Diomedea  salvinii  (albatross) 

No  data 

Gift  of  N.  C.  Rothschild  5  slides 

R901-3,905 

N517  R934 
1  9  Ixodes  putus 
ex.  Uria  lachrymans  (ringed  guillemot) 


ROTHSCHILD  TICKS  IN  BM(NH) 


19 


No  data 

Gift  of  N.  C.  Rothschild  1  slide 

N518        R975-979 

5L  Ixodes  putus 

ex.  Puffin 

No  data.  Probably  British 

Gift  of  N.  C.  Rothschild  5  slides 

G.  H.  F.  Nuttall  del.  12.11.1915 

N539        R1036 

d  Haemaphysalis  leachi  1  slide 

ex.  Erinaceus  albiventris 

Gebel  Auli,  White  Nile,  Egypt 

9.V.1900S.  &F.  Witherby 

Received  from  N.  C.  Rothschild  1908 

Det.G.H.F.  Nuttall  27.1. 1909 

N544        R253 

Amblyomma  sp.  not  varium;  bad  condition 

ex.  3-toed  sloth  (Bradypus  tridactylus) 

Brazil 

N548        R257 

Aponomma  gervaisi 

ex.  Boa 

Feb. 1887 

Given  to  D.  S.  MacLagan  14/10/38 

N551        R181 
Amblyomma  triguttatum 
ex.  Kangeroo(sic.) 
Barrow  Island,  N.  W.  Australia 
20.XI.1900C.J.  T.  Tanney 

N571         R279 

Aponomma  sp. 

ex.  Diemenia  superciliosa 

Herdman's  Lake,  Perth,  West  Australia 

N586        R193 

d  Rhipicephalus  sanguineus 

ex.  Camel 

Locality  unknown 

N587        R151 

2  d  Rhipicephalus  sanguineus 

Zaghig,  25.11. 1908 

N.C.Rothschild  coll. 

N595        R294 
Ornithodoros  moubata 
No  history 

Note:  Nuttall  577  also  bears  Rothschild  number 
R294 

N596        R157 
N's  Ornithodoros  megnini 
Okanagan,  British  Columbia 
1 2. XII.  1902  Allan  Brooks 

Note:  Otobius  is  the  correct  genus  for  this  tick 

N620        R942 

1  9  Ixodes  tenuirostris 


ex.  Mus  minutus  (Harvest  mouse) 

No  further  particulars  '[Mounted  on  a  slide]' 

N628        R165 

9  &  N  Ixodes  ricinus 

No  particulars 

Note:  Nuttall  642  also  bears  Rothschild  number 

R165 

N629        R233 
4  d  Ixodes  ricinus 
ex.  stag 
No  particulars 

N630        R217 

3  N  Ixodes  ricinus 
ex.  Sciurus  vulgaris 

17. IV.  1901  Timmer,  Ireland 

N631         R223 

4  9  Ixodes  ricinus 

ex.  Erinaceus  europaeus 
Belgium  E.  A.  Boulengercoll. 

N632        R173 
2  9  Ixodes  ricinus 
ex.  fox 
No  particulars 

N644        R169 

2  9  Ixodes  holocyclus 

ex.  Macropus  sp. 

W.Australia  28.  VII.  1900 

Note:  Nuttall  633  also  bears  Rothschild  number 

R169 

N656        R205 

2  L  Ixodes  ricinus 
ex.  Mustela  erminea 
Lyndhurst,  Hants.,  England 
25. VII.  1 900  G.  Tatecoll. 

N674        R922-924 

3  9 Ixodes  hexagonus 
ex.  Myopotamus  coypu 

?  locality,  host  a  rodent  from  S.  America 
Presented  by  N.  C.  Rothschild  (R922-924)  '[see 
record  of  2  other  9'$  in  Ticks  Part  II,  p.  183]' 
G.H.F.  Nuttall.det.il.  19 15 

N746        R232 
larvae  Ixodes  hexagonus 
ex.  polecat 
Aberystwyth,  Wales 
VIII.  1900 

N1388        R162 

1  LArgassp. 

ex.  Putorius  energermanis 

Sumas,  B.C.,  Canada 

III.1903 

N3136        R958 
1  d  Hyalomma  sp. 


20  -J-  E.  KEIRANS 

ex.  Capra  falconer i  (the  Markhoor) 

No  further  data 

Gift  of  N.  C.  Rothschild  (slide  R958) 

Note:  The  host  occurs  in  Kashmir,  Afghanistan, 

Gilgit,  etc. 

N3139        R1012-1020 

Hyalomma  syriacum 

ex.  tortoise 

No  further  data 

Gift    of    N.    C.     Rothschild       (5     slides    i-v 

R1012-1020) 


N3140        R1022, 1023 

2  9  Hyalomma  syriacum 

ex.  Testudo  pardalis 

No  further  data 

Gift     from     N.     C.     Rothschild 

R1022— 1023 


(2     slides, 


N3141         R906,  1031 

Hyalomma  syriacum 

ex.  Araconda  [sic] 

No  further  data  (evidently  bought  slides  from  M. 
Pillischer,  optician,  88  New  Bond  St.,  London 
W.)  Gift  of  N.  C.  Rothschild  (2  slides  R906, 
1031) 

N3157a        R1026-1029 

N(3 1 57)  3  d  Rhipicephalus  evertsi 

(a)  1  cf  Boophilus  decoloratus 

ex.  Zebra  No  locality 

Gift  of  N.C.Rothschild  1908 

Mounted  on  slides  in  balsam  (R 1026-1 029) 


R.  evertsi  det.  C.  Warburton  18.111.1915 
B.  decoloratus  det.  C.  Warburton  &  G. 
Nuttall  1915 


H.  F. 


N3158        R907 

1  9  Rhipichephalus  haemaphysaloides 

India 

Mounted  on  slide  in  glycerin 

Gift  of  N.C.Rothschild  1908 

(Prep,  bought  from  Fredk.  Enock*) 

*Spelled  Enock  in  this  collection,  but  see  N3 1 59 

N3159        R908-910 

3  N  Argas  reflexus 

India 

3  slides,  glycerine  in  cells  G.  H.  F.  Nuttall  det. 

111.1915 

Gift  of  N.C.Rothschild  1908 
(3  slides  Nos.  908-9 10)  Fredk.  Enoch  prep. 
Cost  2  shillings  each! 

N3351         None 

9  Amblyomma  varium 

ex.  Bradypus  tridactylus  (sloth) 

Gift  of  N.  C.  Rothschild  years  ago. 

Scutum,  legs,  spiracle  on  3  slides 

L.  E.  Robinson  det.  26. III.  1919 

N3536        None 

1  9  Argas  persicus 

Found  on  wall 

Djama,  Algeria 

25.11.1920 

N.      C.      Rothschild      &      K.      Jordan      coll. 


PART  II 
Rothschild  specimens  donated  to  British  Museum  (Natural  History) 


Species  of  Ixodoidea  represented  in  the  collection  of  the  British  Museum  (Natural 
History),  donated  by  the  Hon.  N.  C.  Rothschild 

K  numbers  follow  each  species.  Numbers  in  parentheses  refer  to  missing  collections.  Thus, 
validity  of  original  determinations  is  unverified. 


ARGASIDAE 

Argas  persicus  (Oten)Kl,  K2,(K148) 
Argas  reflexus  (Fabricius)  K3 
Argas  sanchez i  Duges  K.4 

Ornilhodoros  talaje  Guerin-Meneville  K5 

IXODIDAE 

Amblyomma  albolimbatum  Neumann  K6 
Amblyomma  americanum  (Linnaeus)  K7 
Amblyomma  brasiliense  Aragao  K.8 


Amblyomma  cajennense  (Fabricius)  K9 
Amblyomma  calcaratum  Neumann  K10 
Amblyomma  compressum  Macalister  Kl  1 
Amblyomma  gemma  Donitz  K12,  K13,(K149) 
Amblyomma  humeraleC.  L.  Koch  K14,  K15 
Amblyomma  incisum  Neumann  K16 
Amblyomma  longirostre(C.  L.  Koch)  K17,  K18 
Amblvomma  maculatum  C.  L.  Koch  K19,  K20, 

K2'l,K22 

Amblyomma  naponense  (Packard)  K23 
Amblyomma  nodosum  Neumann  K8,  K24,  K25 


ROTHSCHILD  TICKS  IN  BM(NH) 


21 


Amblyomma  ovale  C.  L.  Koch  K8,  K23,  K26, 

K27,  K28,  K29,  K30,  K3 1 ,  K32 
Amblyomma postoculatum  Neumann  K33 
Amblyomma pseudoconcolor  Aragao  K34 
Amblyomma  testudinarium  C.  L.  Koch  K35 
Amblyomma  variegatum  (Fabricius)  K36,  K37 
Amblyomma  sp.  K38,  K39 

Aponomma  concolor  Neumann  K40,  K41,  K42, 

K43 
Aponomma  decorosum   (L.    Koch)   K44,   K45, 

K46,  K47 

Aponomma  exornatum  (C.  L.  Koch)  K48 
Aponomma  fimbriatum  (C.  L.  Koch)  K49 
Aponomma  gervaisi  var.  trimaculatus  (Lucas) 

(K150) 

Aponomma  latum  (C.  L.  Koch)  K50 
Aponomma  trimaculatum  (Lucas)  K5 1 

Boophilus  decoloratus  (C.  L.  Koch)  K52 
Boophilus  microplus  (G.  Canestrini)  K53 
Dermacentor  atrosignatus  Neumann  group  K54 
Dermacentor      reticulatus      (Fabricius)      K55, 

K56 
Haemaphysalis  bancrofti  Nuttall  and  Warburton 

K57,  K58,  K59,  K60 
Haemaphysalis  calcarata  Neumann  K6 1 
Haemaphysalis  juxtakochi  Cooley  K53 
Haemaphysalis  lagostrophi  Roberts  K33 
Haemaphysalis  leachi  (Audouin)  K62,  K63,  K64, 

K66,  K67,  K68,  K69,  K70,  K71,  K72,  K73, 

K74,  K75,(K151) 

Haemaphysalis  leachi  indica  Warburton  (Kl  52} 
Haemaphysalis  leachi  (Audouin)  group  K76 
Haemaphysalis  novaeguineae  Hirst  K77,  K78, 

K79 

Haemaphysalis  parmata  Neumann  K80 
Haemaphysalis  spinigera  Neumann  K8 1 
Haemaphysalis  spinulosa  Neumann  K82,  K83 
Haemaphysalis  sulcata  G.  Canestrini,  and 

Fanzago  K84,  K85 


Haemaphysalis  tibetensis  Hoogstraal  K86 

Hyalomma  aegyptium  (Linnaeus)  K87 
Hyalomma  anatolicum  excavatum  C.  L.  Koch 

K88 

Hyalomma  marginatum  C.  L.  Koch  K89 
Hyalomma  truncatum  C.  L.  Koch  K90,  (Kl  53) 
Hyalomma  sp.  K9 1 ,  K92,  K93 

Ixodes  alluaudi  Neumann  K94,  K95,  K96,  K97, 

K98 

Ixodes  cordifer  Neumann   K99,    K100,   K101, 

K102 
Ixodes  fecialis  Warburton  and  Nuttall  K103 

Ixodes  hexagonus  Leach   K104,   K105,   K106, 

K107,  K108,K109,K110 
Ixodes  loricatus  Neumann  Kl  1 1 ,  Kl  12 
Ixodes  luciae  Senevet  Kl  1 3 
Ixodes  ornithorhynchi  Lucas  K 1 1 4 
Ixodes  cumulatimpunctatus  Schulze  Kl  15 
Ixodes  ricinus (Linnaeus)  Kl  16,  (Kl 54) 
Ixodes  tasmani  Neumann  Kl  1 7 
Ixodes  thomasae  Arthur  and  Burrow  K69 
Ixodes  trianguliceps  Birula  Kl  18 
Ixodes  vespertilionis  C.  L.  Koch  K119,  K120, 

K121,  K122,  K123,  K124,(K155) 

Rhipicephalus  haemaphysaloides  Supino  K125 
Rhipicephalus  jeanneli  Neumann  K126 
Rhipicephalus  kochi  Donitz  K127 
Rhipicephalus  kochi  Donitz  group  K128 
Rhipicephalus  longicoxatus  Neumann  K127 
Rhipicephalus  pravus  Donitz  K127,  K129,  K130, 

K131 

Rhipicephalus pulchellus  (Gerstacker)  K93,  K132 
Rhipicephalus     sanguineus     (Latreille)     K133, 

K134,  K135,  K136,  K137,  K138,  K139,  K140, 

K141,(K156) 

Rhipicephalus simusC.  L.  Koch  K142,  K143 
Rhipicephalus  sp.  K144,  K145,  K146,  K147 


Rothschild  specimens  in  British  Museum  (Natural  History)  collection  with  type 

status 


B.M.(N.H.)  Accession  No. 
None 

19 14. 1 1.1 7. 1-46. 

None 


Dermacentor  reticulatus  aulicus  Hirst,  1916.  Ann.  Mag.  nat.  Hist. 

(8),  17:308. 

Haemaphvsalis  spinigera  novaeguineae  Hirst,  1914.  Trans,  zool. 

Soc.  Lorn/ 20:  325,  fig.  16. 

Haemaphvsalis  tibetensis  Hoogstraal,  1965.  J.  Parasit.  51:  452, 

figs.  1-26." 


22 


J.  E.  KEIRANS 


Rothschild  specimens  in  British  Museum  (Natural  History)  collection 


ARGASIDAE 

Kl  1912.6.21.21 

1  cf  Argas  persicus 

In  railway  carriage 

El  Kantara  (35°  13'N,05'43'E),  Algeria 

17.111.1911 

Hon.  Walter  Rothschild  &  Ernst  Hartert 

Argas  persicus  is  a  vector  of  Crimean  Congo 

hemorrhagic  fever  virus. 

K2  1919.8. 14.1-70  (pt) 

1  $  Argas  persicus 

No  host 

Ouargla  house  (locality  not  verified) 

E.  Hartert  and  C.  Hilgert 

Note:  The  locality  is  probably  a  house  in  Ouargla 

(3  r57'N,05°20'E),  Algeria. 

K3  1912.6.21,1-2 

1  9,  1  N  Argas  reflexus 

ex.  Columba  livia 

Leipzig  (51°18'N,  12°20'E),  German  Democratic 

Republic 
V.I  9 10 
O.  Fritsche 

Argas  reflexus  is  a  vector  of  the  viruses  Ponteves 
and  Grand  Arbaud. 

K4  1912.12.4.1-8 

Argas  persicus  (1  cf ,  5  9,  3  N  Argas  sanchezi) 

ex.  chicken 

Price,  Pinal  Co.  (33°05'N,  1 1 1'30'W),  Arizona, 

U.S.A. 
27.VII.1911 
Dr  R.  E.  Kunzr 

K5  None 

1  cf ,  3  9,  3  N  Ornithodoros  talaje 

ex.  native  house 

Colombia,  S.  America 

IXODIDAE 

K6  1912.6.21.304-318 

Amblyomma  albolimbatum  (19  cf,  6  9 
Amblyomma  albolimbatum;  1  9  Haema- 
physalis  sp.;  1  9  Haemaphysalis  sp.) 

ex  Trachysaurus  rugosus 

Bernier  Is.  (24°52'S,  1 13°08'E),  W.  Australia 

1910 

B.  H.  Woodward 

Note:  1  cf,  1  9  included  in  the  above  total  were 

seen  by  R.  A.  Cooley  in  1946  who  made  six  slides 

of  dissections  (RML  22306). 

The  Haemaphysalis  specimens  represent  either 

two  species  or  a  single  exceptionally  variable 

species  (Hoogstraal,  personal  communication). 

K7  None 

Amblyomma  sp.  (1  N  Amblyomma  americanum) 

ex  Scuirus  niger 


Mount    Pleasant    (32°47'N,    79°52'W),    South 

Carolina,  U.S.A. 
1909 

A.  T.  Wayne 

Amblyomma  americanum  is  a  vector  of  Lone  Star 
virus. 

K8  None 

Amblyomma  nodosum  (6  cf,   1   9  Amblyomma 

brasiliense;  25  cf,  5  9  Amblyomma  nodosum;  1  9 

Amblyomma  ovale) 
Host  unknown 
Joinvile  (26°18'S,  48'50'W),  Humboldt,  Estadode 

Santa  Catharina,  Brazil 
25.VI.1912 
W.  Ehrhardt 

K9  1912.6.21.326-337 

6  cf,  1 1  9,  3  N  Amblyomma  cajennense 

ex.  cattle,  Canis  familiaris  &  Homo  sapiens 

St.  Ann's  ( 1 8°2 1  'N,  77°  1 6'W),  Jamaica 

22-29.  III.1911 

F.  E.  Sherlock 

KlONone 

Amblyomma  concolor!  '9  etc.'  (79,  1   N,  4  L 

Amblyomma  calcaratum) 
ex.  Tapir  americanus  ( Tapirus  terrestris) 
Joinvile  (26°  1 8'S,  48°50'W),  Humboldt,  Estado  de 

Santa  Catarina,  Brazil 
W.  Ehrhardt  15. IX.  19 12 

Kll  1912.6.21.299-302 

Amblyomma  cuneatum  (Species  epithet  crossed 

out    and    compressum    inserted)    (1    cf,    4    9 

Amblyomma  compressum) 
ex.  Pangolin  (or  Scaly  Ant-Eater)  Manis  sp. 
Bibianha,    Gold    Coast.     Bebianiha    (05°43'N, 

00°27'W)  or  other  populated  places  of  this 

name  nearby.  Ghana 
9. XII. 1911 
H.  G.  F.  Spurrell 

Note:    Vial    label    gives   accession    number   as 
1912.6.21.299-303 

K12  1911. 12.9.1-320  pt 

2  9  Amblyomma  gemma 

ex.  Oxen 

Voi  (03°23'S,  28°34'E),  Kenya 

25. IV. 1910 

Robin  Kemp 

K13  191 1.12.9. 1-320  pt 

5cf ,  4  9  Amblyomma  gemma 

ex.  Oxen 

Voi  (03°23'S,  38°34'E),  Kenya 

30.IV. 1910 

Robin  Kemp 


ROTHSCHILD  TICKS  IN  BM(NH) 


23 


K14  1912.10.31.18-20 
1  cf,  3  9  Amblyomma  humerale 
ex.  donkey  (Equus  asinus) 
Locality  unknown 

K15None 

4  cf  Amblyomma  humerale 

ex.  turtle 

Guyana 

K16None 

1  cf  Amblyomma  incisum 

ex.  tapir  ( Tapirus  sp.) 

Guyana 

Note:  There  is  1  9  in  this  collection  which  is 

tentatively  being  called  A.  incisum  but  which  has 

a  3/3  hypostome. 

K17  1912.6.21.338 

1  cf  Amblyomma  longirostre 

ex.  Galera  barbara  (Eira  barbara) 

Mar  de  Espanha  (21'52'S,  43'00'W),  Minas 

Gerais,  Brazil 
Z.  F.  Zikan 

K18  1912.6.21.339-343 

2  cf,  3  9  Amblyomma  longirostre 

ex.  Synetheres  prehensilis  (Coendou  prehensilis) 
Mar   de    Espanha    (21'52'S,   43WW),    Minas 

Gerais,  Brazil 
6.III.1911 
Z.  F.  Zikan 

K19  1912.10.31.13 

1  cf  Amblyomma  maculatum 

ex.  Canis  sp. 

Buenos  Aires  (34°36'S,  58°27'W),  Argentina 

30.VI.1912 

Miss  Idina  G.  Runnael 

K20  1912. 12.4.12-14 

1  cf,  1  $  Amblyomma  maculatum 

ex.  Small  Chubut  Fox 

BahiaBlanca(38°43'S,  62°  17'W),  Argentina 

15.11.1911 

E.  Weiske 

K21  1919.8. 14.1-70  pt 

4  cf,  5  9  Amblyomma  maculatum 

ex.  Blastocerus  dichotomus  (B.  bezoarcticus) 

Parana    Delta   (vicinity   of  33°43'S,    59'15'W), 

Argentina 
25.11.1917 
Robin  Kemp 

K22  None 

3  cf  Amblyomma  maculatum 

ex.      Pseudalopex     gymnocerus      (Dusicyon 

gymnocerus) 
Laguna  Alsina  (36°49'S,  62'13'W),  to  Buenos 

Aires  (34°36'S,  58°27'W),  Argentina 
V.1916 
Robin  Kemp 


K23  1919.8. 14. 1-70  pt 

Amblyomma  mantiquirense  (1   cf  Amblyomma 

naponense) 

ex.  Hydrochaerus  hydrochaeris 
Eastern  Ecuador 

This  collection  also  contains  1  9  of  Amblyomma 
ovale  or  a  closely  related  species. 

K24  None 

1  cf,  1  $  Amblyomma  nodosum 

ex.  Anteater 

Colombia,  S.  America 

1912 

K25  1912.6.21.325 

1  cf,  3  N,  3  L  Amblyomma  nodosum 

ex.  Myrmecophaga  tetradactla  (Myrmecophaga 

tridactyld) 

Misantla  (19°56'N,  96°50'W),  Mexico 
1.1911 

W.  Engelmann 
Note:  Immature  specimens  tentatively  this  sp. 

K26  1912.6.21.344-352 

8  cf,  4  9,  3  N  Amblyomma  ovale 

ex.  Nasua  socialis  (Nasua  nasud) 

Mar  de  Espanha  (21°52'S,  43'00'W),  Estado  de 

Minas  Gerais,  Brazil 
15.X.1910 
Z.  F.  Zikan 

Note:    Also    present    were    1    cf,    2    9    of  the 
mallophagan  Neotrichodectes  pallidus 

K27  1912.6.21.353-358 

7  cf ,  3  9  Amblyomma  ovale 

ex.  Galera  barbara  (Eira  barbara) 

Mar  de   Espanha   (20'52'W,   43WW),    Minas 

Gerais,  Brazil 
Z.  F.  Zikan 

K28  1921.12.9.10-11 

1  cf,  1  9  Amblyomma  ovale 

ex.  Galictes  vittata  (Grison  vittatd) 

9.VIII.1916 

W.  Ehrhardt 

K29  1921.12.9.12-20 

8  cf ,  2  9  Amblyomma  ovale 

ex.  Nasua  familiar  is  (Nasua  nasua) 
Joinvile(26°18'S,48°50'W),  Humboldt,  Estado  de 

Santa  Catharina,  Brazil 
5.VIII.1918 
W.  Ehrhardt 

K30  None 

3  cf ,  2  9  Amblyomma  ovale 

ex.  Procyon  cancrivorus 

Joinvile  (26'18'S,  48'50'W),  Humboldt,  Estado  de 

Santa  Catharina,  Brazil 
24.VII.1912 
W.  Ehrhardt 


24 


J.  E.  KEIRANS 


K3 1  None 

1  rf  A  mblyomma  ovale 

ex.  Nasua  socialis  (Nasua  nasua) 

Mar  de  Espanha  (21'52'S,  43WW),  Estado  de 

Minas  Gerais,  Brazil 
18.VIII.1911 
Z.  F.  Zikan 

K32  None 

1 2  rf,  5  9  Amblyomma  ovale 

ex.  Nasua  socialis  (Nasua  nasua) 

Mar  de  Espanha  (21'52'S,  43WW),  Estado  de 

Minas  Gerais,  Brazil 
18.VIII.1911 
Z.  F.  Zikan 

K33  1912.6.21.319-324 

Amblyomma  albolimbatum   (2   9  Amblyomma 

postoculatum;  5  9  Haemaphysalis  lagostrophi) 
ex.  Lagostrophus  fasciatus 
Bernier  Island  (24°52'S,  1 1 3°08'E),  W.  Australia 
1910 
B.  H.  Woodward 

K34  None 

Amblyomma  concolor  (2  rf,  7  N,  3  L  Amblyomma 

pseudoconcolor) 
ex.  Armadillo 

La  Rioja  (29°26'S,  66°  1 5'W),  Argentina 
E.  Giacomelli 

K35  1912.12.4.15-16 

2  d  Amblyomma  testudinarium 

ex.  Sheep 

Sri  Lanka 

K36  None 

1  N  Amblyomma  variegatum 

ex.  Raven 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 

17.11.1912 

G.  Kristensen 

Amblyomma  variegatum  is  a  vector  of  the  viruses 

Dugbe,      Thogoto,      Jos,      Crimean      Congo 

hemorrhagic  fever,  Nairobi  sheep  disease,  Bhanja 

and  Somone. 

K37  None 

1  d  Amblyomma  variegatum 

Host  unknown 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 

9.II.1912 

G.  Kristensen 

K38  None 

1  N  Amblyomma  sp. 
ex.  Canis  familiar  is 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 

4.XII.1911 

G.  Kristensen 

K39  None 

2  N  Amblyomma  sp. 


ex.  Homo  sapiens 

Gola  National  Forest  (06°56'N,  10°45'W),  Liberia 

26.V.1910 

R.  H.  Bunting 

K40  1912.6.21.370-380 

Amblyomma  concolor  (3  9,  23   N  Aponomma 

concolor) 

ex.  Echidna  aculeata  (Tachyglossus  aculeatus) 
Gippsland  District  (38°00'S,  147WE),  Australia 
12.IX.1910 
A.  Coles 

K41  None 

Amblyomma     concolor     (11      N     Aponomma 

concolor) 

ex.  Echidna  sp.  (Tachyglossus  aculeatus) 
Australia 
A.  Coles 

K42  1912.6.21.381-383 

Amblyomma  concolor  (3   9,   3   N  Aponomma 

concolor) 

ex.  Echidna  aculeata  (Tachyglossus  aculeatus) 
Traralgon  (38°12'S,  146°32'E),  Australia 
29.X.1910 
A.  Coles 

K43  1912.6.21.384-386 

Amblyomma  concolor  (3  9  Aponomma  concolor) 

ex.  Echidna  sp.  (Tachyglossus  aculeatus) 

Australia 

27.11.1911 

A.  Coles  coll. 

K44  None 

Aponomma  decorosum  (Species  epithet  crossed 

out  and  undatum  inserted)  (22  cf,  3  9,  3  N 

Aponomma  decorosum) 
Ex.  Varanus  sp. 
Kelsey  Creek  (20°26'S,   148°27'E),  Queensland, 

Australia 
1911 
A.  Fontaine 

K45  1912.6.21.391^01 

Aponomma  decorosum  (Species  epithet  crossed 

out  and  undatum  inserted)  (22  cf ,  6  9,  18  N,  2  L 

Aponomma  decorosum) 
ex.  Brown  Iguana  ( Varanus  sp.) 
Kelsey  Creek  (20°26'S,   148°27'E),  Queensland, 

Australia 
A.  Fontaine 
Publication:  Kaufman  (1972:  369) 

K46  19 12.6.2 1.402^10 

Aponomma  decorosum  (Species  epithet  crossed 

out  and  undatum  inserted)  ( 1 5  cT,  1 3  9, 2 1  N,  2  L 

Aponomma  decorosum) 
ex.  Black  Iguana  ( Varanus  sp.) 
Kelsey  Creek  (20°26'S,  148°27'E),  Queensland, 

Australia 


ROTHSCHILD  TICKS  IN  BM(NH) 


25 


A.  Fontaine 

Publication:  Kaufman  (1972:  369) 

K47  None 

Aponomma  decorosum  (Species  epithet  crossed 

out  and  undatum  inserted)  (1   d1,    1   9,  4  N 

Aponomma  decorosum) 
ex.  Iguana  ( Varanus  sp.) 
Australia 
11.1.1911 
Publication:  Kaufman  (1972:  370) 

K48  1912.10.31.21-25 

2  cf  Aponomma  exornatum 

ex.  Egyptian  monitor  (Varanus  sp.) 

Bibianha    [Bebaianiha    (05'43'N,    00°27'W)    or 

other  populated  place  of  this  name  nearby], 

Ghanaa 
17.XII.1911 
H.  G.  F.  Spurrell 

K49  1912.6.21.387-390 

Aponomma    ecinctum    (2    d1,    2    9   Aponomma 

fimbriatum) 

ex.  Tiger  snake  (Notechis  scutatus) 
Melbourne  (37°49'S,  144°58'E),  Australia 
12.VII.1911 
A.  Coles 

Note:  Very  poor  specimens;  hypostomes  lacking 
in  all  specimens.  Capitulum  present  in  only  one  9 
but  from  porose  areas  and  scutal  configuration  of 
9  and  scutum  and  tarsi  of  cf  these  appear  to  be 
good  A.  fimbriatum. 

K50  1912.6.21.416 

1  d  Aponomma  latum 
ex.  Hystrix  sp. 

Mduna  River,  Hlabisa  District,  (28°10'S, 
32°10'E),  Zululand  Region,  Republic  of  South 
Africa 

9.III.1910 

F.  Toppin 

Received  from  Natal  Museum,  Pietermaritzberg 

H.  Hoogstraal  det.  1952  'unusual  host' 

K51  1912.6.21.359-369 

Aponomma  gervaisi  var.  trimaculatus  (11  d,  2  9, 

1  N  Aponomma  trimaculatum) 
ex.  Python  sp. 
Satte'lberg,     Huon    Gulf    (07°10'S,     147'25'E), 

Papua  New  Guinea 
1911 
Prof.  F.  Forster 

K52  None 

2  9  Boophilus  decoloratus 
Host  unknown 
Tanzania 

Dr  E.  J.  Baxter 

Boophilus  decoloratus  is  a  vector  of  the  viruses 
Dugbe,  Thogoto,  Bhanja,  Jos,  Somone  and 
Crimean  Congo  hemorrhagic  fever. 


K53  1912.10.31.26-29 

Haemaphysalis  kochi(\  9  Boophilus  microplus;  2 

d,  1  9,  1  N  Haemaphysalis  juxtakochi) 
ex.  Deer 
Caparo   (10'27'N,    61°20'W),    Trinidad    Island, 

Trinidad  and  Tobago 
22.V.1912 
S.  M.  Klages 

K54  1912.12.4.17 

1  cf  Dermacentor  atrosignatus  group 
ex.  Sheep 

Sri  Lanka 

K55  None 

Dermacentor  reticulatus  var.  aulicus  SYNTYPE 

( 1  9  Dermacentor  reticulatus) 
Host  unknown 
Valescure,  nr.  St.  Raphael  (43°26'N,  06°46'E), 

France 
2.IV.1910 

Publication:      Hirst      (1916:      308)      original 
description. 

K56  1912.6.21.77-97 

2  d,  8  9  Dermacentor  reticulatus 
ex.  Canis  familiaris 

Kuybyshev  (55°40'N,  78°12'E),  Tomsk,  U.S.S.R. 

27.VIII.1911 

W.  M.  Thomas 

K57  1912.6.21.69-71 

1  9  Haemaphysalis  bancrofti 

ex.  Horse  Equus  caballus 

Kelsey  Creek  (20°26'S,   148°27'E),  Queensland, 

Australia 
1911 
A.  Fontaine 

K58  1912.6.21.76 

1  d  Haemaphysalis  bancrofti 

ex.  Black  Wallaby  [Macropus  dorsalis,  the  black- 
striped  wallaby  or  Wallabia  bicolor,  the  black, 
or  black-tailed  wallaby,  both  of  which  occur  in 
Queensland]. 

Kelsey  Creek  (20°26'S,  148°27'E),  Queensland, 
Australia 

1911 

A.  Fontaine 

K59  None 

2  d  Haemaphysalis  bancrofti 

ex.  Wallaby  ( Wallabia  sp.  or  Macropus  sp.) 
Kelsey  Creek  (20°26'S,   148°27'E),  Queensland, 

Australia 
1911 
A.  Fontaine 

K60  None 

2  d  Haemaphysalis  bancrofti 
ex.  Canis  familiaris 


26 


J.  E.  KEIRANS 


Kelsey  Creek  (20'26'S,   148°27'E),  Queensland, 

Australia 
1911 
A.  Fontaine 

K61  None 

2  rf  Haemaphysalis  calcarata 

Host  unknown 

Voi  (03°23'S,  38°34'E),  Kenya 

14.IV.1910 

Robin  Kemp 

K62  None 

Haemaphysalis  leachi  leachi  (2  d1,  2  9  Haema- 
physalis leachi) 
Host  unknown 
Tanzania 
Dr  E.  J.  Baxter 

K63  1912.10.31.30-35 

Haemaphysalis  leachi  leachi  (9  d1,  2  9  Haema- 
physalis leachi) 
ex.  Felis  sp. 

Lake  Suai,  Ethiopia  (locality  not  verified) 
[Hirst  original  det.  Haemaphysalis  sp.] 

K64  1912.6.21.33-35 

2  rf,  2  9  Haemaphysalis  leachi 

ex.  Hyaena  (Hyaena  hyaena) 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 

31.XII.1911 

G.  Kristensen 

K65  1912.6.21.36^4 

1 0  d1,  8  9  Haemaphysalis  leachi 

ex.  Felis  catus 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 

22.1.1912 

G.  Kristensen 

K66  19 12.6.2 1.4  5-49 

5  cf  Haemaphysalis  leachi 

ex.  Marten 

Harar  (09°  1 9'N,  42'09'E),  Ethiopia 

18.1.1912 

G.  Kristensen 

K67  1912.6.21.50-55 

5  cf,  3  9  Haemaphysalis  leachi 

ex.  'Badlington'  dog 

Gola  National  Forest  (06'56'N,  10°45'W),  Liberia 

17.VI.1910 

R.  H.  Bunting 

K68  1912.6.21.56-58 

1  d1,  2  9  Haemaphysalis  leachi 

ex.  Felis  serval 

Mpwapwa  (06°2 1  'S,  36°29'E),  Tanzania 

15.X. 1911 

Marschner.  Rec'd.  from  W.  Fasius 


K69  None 

Haemaphysalis  leachi  (4  d  Haemaphysalis  leachi; 

2  9  Ixodes  thomasae) 

Host  unknown 

Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
7.VI.1910 
Robin  Kemp 

K70  None 

7  rf,  109  Haemaphysalis  leachi 

Host  unknown 

Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
6.VI.1910 
Robin  Kemp 

K71  None 

4  rf,  4  9  Haemaphysalis  leachi 

Host  unknown 

Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
8.VI.1910 
Robin  Kemp 

K72  None 

2  cf  Haemaphysalis  leachi 

ex.  Domestic  Rabbit  (Oryctolagus  sp.  probably.) 

The  genus  Oryctolagus  does  not  exist  south  of 

the  Sahara  unless  bred  in  captivity. 
Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
20.  VI. 1910 
Robin  Kemp 

K73  None 

1 1  d1, 8  9,  1  N  Haemaphysalis  leachi 

Host  unknown 

Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
11.VI.1910 
Robin  Kemp 

K74  None 

25  d1,  1 1  9  Haemaphysalis  leachi 

Host  unknown 

Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
9.VI.1910 
Robin  Kemp 

K75  None 

1  cf  Haemaphysalis  leachi 

ex.  Genetta  stuhlmanni 

Fennek,  Mutoragwa 

Aberdare  Mts,  Kenya  (locality  not  verified*) 

23. HI.  19 10 

Robin  Kemp 

*There  is  a  Mutarakwa  Hill  (00°47'S,  35°14'E)  in 

the  Aberdares  that  is  a  possible  locality  for  this 

collection. 


ROTHSCHILD  TICKS  IN  BM(NH) 


27 


K76  1912.6.21.59-65 

Haemaphysalis  leachi!  (7  cf,  4  9  Haemaphysalis 

leachi  group) 
ex.         Paradoxurus        niger        (Paradoxurus 

hermaphroditus) 

Hoshangabad  District  (22°35'N,  77°40'E),  India 
1912 

C.  A.  Crump 
Received  from  Bombay  Nat.  Hist.  Soc. 

K77  1914.11.17.16-35 

Haemaphysalis    spinigera    var.    novae-guineae 
TYPE  (19  cf,  4  9,  1  N  Haemaphysalis  novae- 
guineae  paralectotypes) 
ex.  Kangaroo 
Sattelberg,  Huon  Gulf  (07°10'S,  147'25'E),  Papua 

New  Guinea 
1911 

Prof.  F.  Forster 

Publications:  Hirst  (1914:  328),  original  descrip- 
tion; Nuttall  &  Warburton  (1915:  452);  Roberts 
(1963:57) 

K78  1914.11.17.1-15 

Haemaphysalis    spinigera    var.    novae-guineae 

TYPE  (1  cf  Haemaphysalis  novaeguineae  lecto- 

type;  9  cf,  1 1  9  Haemaphysalis  novaeguineae 

paralectotypes) 
ex.  Perameles  sp.  ? 
Sattelberg,  Huon  Gulf  (07'10'S,  147°25'E),  Papua 

New  Guinea 
Prof.  F.  Forster 

Publications:  Hirst  (1914:  328)  original  descrip- 
tion; Nuttall  &  Warburton  (1915:  452);  Roberts 
(1963:  57)  selected  lectotype  and  paralectotypes. 

K79  1914.11.17.36-46 

Haemaphysalis    spinigera    var.    novae-guineae 
TYPE  (6  cf,  10  9,  1  N  Haemaphysalis  novae- 
guineae paralectotypes) 
ex.  Hydromys  sp. 
Sattelberg,  Huon  Gulf  (07°10'S,  147°25'E),  Papua 

New  Guinea 
1911 

Prof.  F.  Forster 

Publications:  Hirst  (1914:  328),  original  descrip- 
tion; Nuttall  &  Warburton  (1915:  452);  Roberts 
(1963:  57)  designated  paralectotypes. 

K80  1912.6.21.66 

1  cf  Haemaphysalis  parmata 

ex.  Neotragus  pygmaeus 

Bebaianiha     (05°43'N,     00°27'W)     or     other 

populated  places  of  this  name  nearby,  Ghana 
10.XI.1911 
H.  G.  F.  Spurrell 

K81  None 

13  N  Haemaphysalis  spinigera 

ex.  Lepussp. 


Hoshangabad  District  (22°35'S,  77°40'E),  India 

1912 

C.  A.  Crump 

Received  from  Bombay  Nat.  Hist.  Soc. 

Haemaphysalis  spinigera  is  a  vector  of  the  viruses 

Kaisodi  and  Kyasanur  forest  disease. 

K82  1912.10.31.36-46 

Haemaphysalis  sp.  (28  cf,  7  9  Haemaphysalis 

spinulosa) 
ex.  Wild  cat 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 
6. 11.1912 
G.  Kristensen 

K83  None 

Haemaphysalis    leachi]    (1     9    Haemaphysalis 

spinulosa) 
ex.  Canis  familiar  is 

Himo  River  (03°33'S,  37°30'E),  Tanzania 
30.V.1910 
Robin  Kemp 

K84  19 12.6.21. 22-32  pt 

Haemaphysalis  cinnabarina  var.  punctata  (2  cf,  3 

9  Haemaphysalis  sulcata) 
ex.  Sheep 

Larnaca  (34°55'N,  33°38'E),  Cyprus 
XI.1911 
C.  B.  Horsbrugh 

K85  19 12.6.21. 22-32  pt 

2  cf ,  3  9  Haemaphysalis  sulcata 

ex.  Sheep 

Larnaca  (34°55'N,  33°38'E),  Cyprus 

XI.1911 

C.  B.  Horsbrugh 

Note:  A  portion  of  this  collection  was  originally 
entered  under  Haemaphysalis  cinnabarina  var. 
punctata. 

K86  None 

Haemaphysalis    warburtonil   (1    cf,    2    9,   2    N 

Haemaphysalis  tibetensis  TYPES 
1  cf  (holotype)  2  9,  2  N  (paratypes) 
ex.  Canis  familiar  is 
Yatung  (27°25'N,  89  WE)  (ca.  3896  m)  mouth  of 

Chumbi  Valley,  Tibet 

D.  Macdonald 

Publication:  Hoogstraal  (1965:  452),  original 
description. 

K87  1912.6.21.247-250 

Hyalomma    syriacum    (3    cf,    1    9    Hyalomma 

aegyptium) 

ex.  Testudo  graeca  ibera 
Hammam  Rirha  (36°26'N,  02°28'E),  Algeria 
24. V. 1911 
Hon.  Walter  Rothschild  &  Ernst  Hartert 


28 


J.  E.  KEIRANS 


K88  1912.6.21.251-253 

Hyalomma  syriacum  (Species  epithet  crossed  out 

and    aegyptium    inserted)    (4    9    Hyalomma 

anatolicum  excavatum) 
Host  unknown 
North  West  Morocco 
Received  from  Hon.  W.  Rothschild 

K89  1912.6.21.240-6 

Hyalomma   aegyptium   (2   cf,    1    9   Hyalomma 

marginatum) 
Host  unknown 

Hammam  Rirha  (36°26'N,  02°28'E),  Algeria 
1911 

C.  Hilgert 

Hyalomma  marginatum  is  a  vector  of  the  viruses 
Dhori,  West  Nile,  tickborne  encephalitis,  Bhanja, 
Crimean  Congo  hemorrhagic  fever,  Tamdy  and 
Batken. 

K90  19 11. 12.9. 1-320  (pt) 

Hyalomma      aegyptium      (2      9      Hyalomma 

truncatum) 

ex.  Aged  donkey  (Equus  asinus) 
Voi  (03°23'S,  38'34'E),  Kenya 
29.IV.1910 
Robin  Kemp 

K91  None 

8  N  Hyalomma  sp. 

Host  unknown 

Biskra  (34°50'N,  05°40'E),  Algeria 

25.111.1909 

Hon.  L.  W.  Rothschild 

K92  19 19.8. 14. 1-70  (part) 

3  N  Hyalomma  sp. 

ex.?  Dipodillus  (gerbil)  (Gerbillus  (Dipodillus)  sp.) 

('host  similar  to  108') 

Guelt-es-Stel  (35°09'N,  03°02'E),  Algeria 

IV.1912 

Hon.  Walter  Rothschild  and  Karl  Jordan 

K93  1912.6.21.202 

Rhipicephalus  sp.   (4   N   Hyalomma   sp.;    1    9 

Rhipicephalus  pulchellus;  10  N  Rhipicephalus 

sp. 

ex.  Lepus  sp. 

Harar  (09°  1 9'N,  42°09'E),  Ethiopia 
9.1.1912 
G.  Kristensen 

Rhipicephalus  pulchellus  is  a  vector  of  the 
viruses  Dugbe,  Crimean  Congo  hemorrhagic 
fever,  and  Nairobi  sheep  disease. 

K94  None 

Ixodes  var.  offecialis  (2  9  Ixodes  alluaudi) 

ex.  Lophuromysjlavopunctatus  zena 

Aberdare  Mts  (00°25'S,  36°38'E),  Kenya 

14.11.1910 

Robin  Kemp 


K95  None 

Ixodes  var.  offecialis  ( 1  9  Ixodes  alluaudi) 

ex.  Lophuromys  fJavopunctatus  zena 

Aberdare  Mts  (00°25'S,  36°38'E),  Kenya  1 1 ,000  ft 

(3,353  m) 
21.11.1910 
Robin  Kemp 

K96  None 

Ixodes  tenuirostris  (2  9  Ixodes  alluaudi) 

ex.  Otomys  tropicalis  elgonis 

Mt  Ilkinangop  (00°38'S,  36°42'E),  Aberdare  Mts, 

Kenya 
24.11.1910 
Robin  Kemp 

K97  None 

Ixodes  tenuirostris  (1  9  Ixodes  alluaudi) 

Host  unknown 

Rombo,       Kilimanjaro      (03°05'S,       37°20'E), 

Tanzania 
15.IV.1910 
Robin  Kemp 
Note:  This  9  has  been  previously  pinned. 

K98  None 

[undetermined  genus  &  species]   (1    N   Ixodes 

alluaudi) 
ex.      Mus     jacksoni     (Possibly     Anomalurus 

derb  ianus  Jackson  i) 
Mutaragwa,  Aberdare  Mts  (locality  not  verified*), 

Kenya 
13.111.1910 
R.  Kemp 

*There  is  a  Mutarakua  Hill  (00°47'S,  35°14'E)  in 
the  Aberdares  that  is  a  possible  locality  for  this 
collection. 

K99  1912.6.21.280-2 

4  9  Ixodes  cordifer 

ex.  Pseudoircornia  (probably  Pseudocheirus  sp.) 

Sattelberg,  Huon  Gulf  (07°10'S,  147°25'E),  Papua 

New  Guinea 
1911 
Prof.  F.  Forster 

K100  1912.6.21.283 

1  9  Ixodes  cordifer 

ex.  Dactylopsida  trivirgata 

Sattelberg,  Huon  Gulf  (07°10'S,  147°25'E),  Papua 

New  Guinea 
Prof.  F.  Forster 

Note:  label  in  vial — seen  by  Prof.  R.  A.  Cooley 
1946.  3  slides  of  dissections  (RML  22285). 

K101  1912.6.21.284-8 

4  9  Ixodes  cordifer 

ex.  Phalanger  maculatus 

Sattelberg,  Huon  Gulf  (07°10'S,  147'25'E),  Papua 

New  Guinea 
1911 
Prof.  F.  Forster 


ROTHSCHILD  TICKS  IN  BM(NH) 


29 


K102  1912.6.21.279 

1  d1  Ixodes  cordifer 

ex.  Perameles  sp. 

Sattelberg,  Huon  Gulf  (07°10'S,  147°25'E),  Papua 

New  Guinea 
1911 
Prof.  F.  Forster 

K103  1912.12.4.11 

1  9, 4  N  Ixodes  fecialis 

cx  Fclis  so 

Studley   Park,   Melbourne  (37°49'S,    144°58'E), 

Australia 
19.V.1911 
A.  Coles 

Note:    Reverse    of   the    vial    label    bears    the 
inscription — Native  cat. 

K104  1912.6.21.254— 6 

5  9, 2  N  Ixodes  hexagonus 

ex.  Mustela  vulgaris  (Mustela  nivalis) 

Tring  (5 1  °47'N,  00°39'W),  England 

12.111.1912 

J.  F.  Cox 

K105  1912.6.21.259-61 

1  9,  2  N,  7  L  Ixodes  hexagonus  (1  9,  2  N  Ixodes 

hexagonus) 
ex.  Mustela  erminea 
Tring  (5 1  °47'N,  00°39'W),  England 
III.1912 
J.  F.  Cox 

K106  1912.6.21.262-3 

4  N  Ixodes  hexagonus  (4  N,  8  L  Ixodes 

hexagonus) 

ex.  Meles  taxus  (Meles  meles) 
Senj  (45WN,  14°54'E),  Yugoslavia 
5.XII.1911 
F.  Dobiasch 

K107  None 

22  N,  6  L  Ixodes  hexagonus 

ex.  Mustela  erminea 

Loch  Tay  (56°28'N,  04°  1 8'W),  Lawes,  Scotland 

2.III.1911 

L.  G.  Esson 

KlOSNone 

1  9 Ixodes  hexagonus 
Host  unknown 
Faugeres*,  S.  France 
8.X.1913 

W.  Davy 

N.  C.  Rothschild 

*Three  possible  localities  for  Faugeres,  France: 

43°34'N,  03'11'E;  44°28'N,  03°58'E;  44°58'N, 

04°35'E. 

K 109  None 

2  N  Ixodes  hexagonus 
ex.  Mustela  putorius 


Carmarthen  (5 1°52'N,  04°  1 8'W),  Wales 

1914 

F.  W.  Frohawk 

Kl  10  1919.8. 14.1-70  (part) 

1  9  Ixodes  hexagonus 

ex.  dog  Canis  familiaris 

Bridgnorth  (52°32'N,  02°25'W),  England 

28.X.1916 

Miss  Frances  Pitt 

Kill  1912.12.4.9-10 

1  rf,  1  9  Ixodes  loricatus 

ex.  Didelphis  marsupialis  aurita 

Mar  de  Espanha  (20°52'S,  43'00'W),  Estado  de 

Minas  Gerais,  Brazil 
6.X.1910 
Z.  F.  Zikan 

K112  1919.8. 14.1-70  pt 

2  <?,  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 
scientific  series,  Botany,  Entomology,  Geology  (incorporating  Mineralogy)  and  Zoology,  and 
an  Historical  series. 

Papers  in  the  Bulletin  are  primarily  the  results  of  research  carried  out  on  the  unique  and 
ever-growing  collections  of  the  Museum,  both  by  the  scientific  staff  of  the  Museum  and  by 
specialists  from  elsewhere  who  make  use  of  the  Museum's  resources.  Many  of  the  papers  are 
works  of  reference  that  will  remain  indispensable  for  years  to  come. 

Parts  are  published  at  irregular  intervals  as  they  become  ready,  each  is  complete  in  itself, 
available  separately,  and  individually  priced.  Volumes  contain  about  300  pages  and  several 
volumes  may  appear  within  a  calendar  year.  Subscriptions  may  be  placed  for  one  or  more  of 
the  series  on  either  an  Annual  or  Per  Volume  basis.  Prices  vary  according  to  the  contents  of 
the  individual  parts.  Orders  and  enquiries  should  be  sent  to : 


Publications  Sales, 

British  Museum  (Natural  History), 
Cromwell  Road, 

London  SW7  5BD, 
England. 


World  List  abbreviation:  Bull.  Br.  Mus.  not.  Hist.  (Zool.) 


<g)  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  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 


o 

JS  .S 

o 

i—  — 
c^  "O 

>>               _£- 

6  c 

ta 

o  2 

c/) 

o 

•a  S3             3 

"O                           ^   to 

£* 

"O           O 

o-S           8 

8  S      w  t          S          6 

i-    c    _^<2T3          0                0 

1     I  II 

C                  "^     M-^ 

O 
O 

2i 

c 

0         0 

^    tt2 

«-        c 

JS         •- 

included. 

M 

O 

'S  1  45  b  8  "1    1         c  «« 
o5>,o^uiS      £  C      ~  * 

"5        ^     o  ."g 

1    ll!| 

CO 

J? 
u 

03 

to 
a> 

o        o  .  

i     |B 

LL          O  *-• 
*            f»    <u 
to           5«    « 

«     >r>.a 

V 
CO 

(75 

O 

U 

UJ   CX  i/j  «/•>  UJ   o        -J  tr        c/)   E 

1        nllS 

~ 

CO 

U 

•  «•           CO  ^^ 

-    18 

W        <   CO 

"2 

o 
o 

SO 

o 

ro 

UN 

Os    53          13 

•j2 

10               0  §      -a          -a 

•o-o      •^otrno^-o 

rsT^ 

C^l          <N  ^o 

00 

^O  "^  ^Bii^   4)                  ^ 

O     O            sJ  »_  i  <^>        ,     O 

(N  <^ 

O         —  OS 

c 

fli 

^^            ftj            M        rf^                                                  ,.f^ 

f^    «^           C          Os           <""] 

Os         Os  — 

w 

C 

•o"              ""  5  S  •-          •- 

t/3    t/n         V3  *-T  -  ~  *-T  t/5 

—  — 

1 

0 

2 

Sill        1 

£               IS  '-J  a  ex           ex 

•§•§       |  o  §"  0-3 
a  ex       c  ^  o  ^  ex 

"2  §• 
^3  o 

"H     "H-- 

c 

•<-> 

CO                       O    &o  "^    C                C 

C    C          3    <u  *u    O    C 

""  "o 

?5      S  'S 

^ 

3 

X               at  3-  c>cD          D 

D  D      X  >  J  >  D 

03  -J 

03        03  H 

< 

• 

TT  (N                                    ^, 

CO 
<L> 

SO                       so                2                "" 

—  —                                    <N 

J= 

-               °i          d          «N 

c"  c"                       ^" 

•^ 

-^                                        ^^      l         ^^ 

O    O 

O 

rf^                       ON                iX      •         iX 

so 

Z 

i/^                  \^             r^.  ^        r*** 

S  C                          t~- 

-1    J                                                    OJN 

c 

_--  i 

^^                                                                       ^^                                                  ^^       ^-^                             W-* 

u 

.2 

PC               X          SCO      35 

x"x"               x 

-C 

o 

X-<                                                    *      '                                     '      -      O^                      '      ' 

zz                z 

O 

08 

2 

OQ                    GQ              CO  6C        CO 

0^  0^                    e^.  ^..  OQ 

{>• 

«>•        o- 

T3 

C 

CO 

r^4 

O 

"o 

c^ 

C 

c 

.                    3 

•**, 

PJ 

**->         1                  IS                              rr, 

o 

CO 

o  ,0      13            {2 

>,  3  X)    C 

6  >, 

U 

w- 
O 

C                     .t?C/3    3    C               ^ 

a  |  ^ 

01 

•a      -a  .Ji 

— 

o 

aj    O         ^               »^ 

Q      M          Jj 

3 

CO              CO     (/5 

•  22 

>  T3    2  C/5 

"™          ^ 

_O 

—           -^  'o 

"ob 
U 

"o 
U 

=      'sj|i     2 

(-.<-•        e-        W  CU  U 

'C 

CO 

03        03  H 
<        <O 

<u 

_c 

oo       oo                 so 

6 
o 

so 

SO                ^"i,        ^^ 

<N          <N                        rt 
§1          >          S^E 

so" 

O 

>>         C 

<i: 

O                                              '"^  ^^  f^\ 

PH( 

X3            ° 

—     ^           O          ^^  ^^    Z3 

1 

CO          — 

t3 

u 

~*                                                                                    f*^     *^*            A  ^^ 

^<          *7          ^^  ^^  <f* 

(N    Tt 

g 

"^                '"'             I  i  —  2P  ex 

2^                             C    i 

O  — 

"o  o  o^ 

C 

D 

—                        —                 <N  —  <  C/2 

<^i      2      5  a^ 

Os  OS 

£5  5= 

-5 

•a 

c 

•2 

c 

CO 

J 

-c 
O 

.M 

1                6     =         »s 

"§"&"§  «""!* 

aT 

3 

<*- 
o 

C/5 
I 

Is     ^  1    | 

J2^^  gwf  ^^ 

ttj  »^J     o    UN  >        ^    r*    r^ 

^^3?.^^Z|| 

<uT 

1    * 

•r      "^ 

•7         Os 

O 
o 

(U 

a: 

•55      C/5                         HH      .  ^               C                                      O 

3    *             S      2          <u  c 

»o!r;5N     -J2NN 

•S  -^  •-       'Z  S    -  - 
2  cx-^  "41       o  "  " 

B 

i-s 

c  c 
C  <u 

3?S 

1  ofc" 

i/) 

-i 
Z 

es 

*CO 

8 

fill  si  ll  II 

O    ^  .  i«  MM    O           C    C    C 
g    O    3    .       UN  i.  i    O    o    O 

s  -^  ts 
I  go 

^S£ 

CO  -C    O 
It  *^    R    9 

g    >      (U      (/5 

c  ^  -c  9 

I  c/5  uoi 

H 

u 

z 

00 

u» 

N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  1 1 7 

HABITAT.  Substrates  not  recorded  in  my  \915a  paper  include  an  isopod,  Anilocra  physodes 
(Linnaeus),  from  Naples  (Stechow,  19216,  1925,  as  Obelia  bifurca)  and  a  commercial 
shrimp,  Pennaeus  aztecus  Ives,  from  Florida  (Kruse,  1959;  via  A.  A.  Fincham,  pers. 
comm.);  ships  hulls,  hermit  crabs  gastropod  shells  and  algae  (Millard,  1975).  O.  bidentata  is 
highly  tolerant  of  brackish  water.  It  has  been  recorded  at  salinities  of  18'6%o  (Netherlands; 
Vervoort,  19460),  10-20%o  (Italy;  Morri,  19796)  and  even  so  low  as  l%o  (S  Carolina,  U.S.A.; 
Calder,  1976).  The  species  has  often  been  recorded  in  shallow  water  and  Millard  (1978) 
listed  it  as  intertidal. 

REMARKS.  A  detailed,  world-wide  treatment  has  appeared  recently  (Cornelius,  \915a)  and 
the  notes  given  here  are  mainly  intended  to  supplement  that  account.  O.  bidentata  is  a  nearly 
cosmopolitan  species,  and  a  possible  explanation  is  given  above  (pp.  44-45). 

Differences  from  the  superficially  similar  Hartlaubella  gelatinosa  are  given  under  that 
species,  and  from  the  hydroid  stages  of  other  Obelia  species  by  Cornelius  (\915a  :  table  1). 

Mayer  (1910),  Stechow  (1925)  and  recently  Stepanyants  (1979)  regarded  Obelia  austro- 
georgiae  Jaderholm,  1904#,  as  conspecific.  I  do  not  disagree  (cf.  Cornelius,  1975a)  but  have 
no  fresh  evidence.  However,  I  cannot  yet  agree  with  Stepanyants  that  Clytia  paulensis 
Vanhoffen,  1910,  is  identical  with  O.  bidentata.  The  young  medusae  of  both  are  sufficiently 
well  described  to  indicate  that  two  species  are  involved  (see  notes  on  Dispersive  stage  under 
the  two  species,  pp.  89,  113).  It  could  be  argued  that  since  Vanhoffen's  material  was 
infertile  it  was  merely  young  O.  bidentata:  but  this  was  not  Stepanyants'  argument.  I  believe 
neither  Stepanyants  nor  I  have  seen  the  type  material  of  C.  paulensis.  If  it  did  prove  to  be 
young  O.  bidentata  then  a  new  name  would  have  to  be  found  for  C.  paulensis  auct.  (see 
synonymy  in  this  paper).  In  that  event  it  would  be  useful  to  apeal  to  the  International 
Commission  on  Zoological  Nomenclature  to  validate  the  widely  used  name  paulensis  sens, 
auct. 

Clytia  arborescens  Pictet,  1 893  :  34-35,  pi.  2,  figs  30-3 1 ,  originally  based  on  material  from 
Indonesia,  was  reported  south  of  Madeira  at  100  m  by  Billard  (1907);  but  Stechow  (1925) 
included  no  further  records  in  his  synoptic  list  of  warm  water  W  Atlantic  hydroids.  Billard's 
material  lacked  gonothecae  but,  like  the  type  material,  had  a  compound  stem  and  long 
hydrotheca  (560-700  /zm)  with  sharply  pointed  cusps.  It  is  likely  that  his  material  was  O. 
bidentata;  and  it  is  relevant  that  he  had  wrongly  identified  O.  bidentata  from  the  English 
Channel  (see  Distribution,  above).  O.  bidentata  was  not  well  known  at  the  time  he  published 
the  arborescens  record.  The  type  material  of  arborescens  was  probably  not  O.  bidentata, 
however,  since  Pictet  reported  hemispherical  medusa  buds  in  the  gonotheca,  indicating 
Clytia  sp. 


Obelia  dichotoma  (Linnaeus,  1758) 
(For  illustrations  see  Cornelius,  1975a) 

Sertularia  dichotoma  Linnaeus,  1758  :  812. 

ICymodocea  simplex  Lamouroux,  1816  :  2 1 6,  pi.  7,  fig.  2  (see  p.  121). 

ITubularia  clytioides  Lamouroux,  in  Freycinet,  1824  :  620,  pi.  95,  figs  6-8. 

Lomedea  gracilis  C.  Pickering,  in  Dana,  1846  :  689, -pi.  61,  figs  7,  7a-b  (lapsus  pro  Laomedea;  see  p. 

78). 

Obelia  dichotoma:  Hincks,  1868:  156-157,  pi.  28;  Cornelius,  1975a  :  265-272,  figs  3-4. 
Obelia plicata  Hincks,  1868  :  159,  pi.  30,  figs  1,  la. 
Campanularia  cheloniae  Allman,  1 888  :  22,  pi.  11,  figs  2, 2a. 
Obelia  dischotoma  Billard,  1927  :  332  (lapsus  pro  dichotoma). 

lObelia  commensuralis:  Gudger,  1937:  1-6  (?lapsus  pro  0.  commissuralisMcCrady,  1857). 
Obelia  hyaliana  Vannucci,  1955  :  56. 
Laomedea  plicata:  Rasmussen,  1973  :  30,  fig.  9. 

Eulaomedajlexuosa:  Rees  &  Thursfield,  1965  :  102-103  (lapsus  pro  Eulaomedea\  redet.;  see  p.  105). 
For  further  synonymy  see  Remarks,  Table  4  and  Cornelius  ( 1 975a). 


118  P.  F.  S.  CORNELIUS 

DISPERSIVE  STAGE.  Medusae  of  O.  dichotoma  and  O.  geniculata  reared  from  the  hydroids  in 
the  laboratory  still  cannot  be  distinguished  and  some  taxonomic  confusion  continues 
(Russell,  1953;  Cornelius,  1975a;  Arai  &  Brinckmann-Voss,  1980;  see  also  Remarks). 
Russell's  book  includes  the  best  account  of  the  biology  of  Obelia  spp.  medusae. 

The  alternation  of  generations  characteristic  of  medusoid  coelenterates  was  actually  first 
appreciated  in  scyphozoans,  in  1829.  It  was  reported  in  Obelia  as  early  as  1836,  first  among 
the  hydrozoans;  but  several  still  earlier  workers  had  come  close  to  its  discovery  in  this  genus. 
This  and  other  historical  aspects  have  been  reviewed  recently  (Winsor,  1976;  Cornelius, 
\911a,  b).  A  paper  by  Desor  (1849)  was  missed  by  these  reviewers.  Desor  provided  an  early 
confirmation  of  medusa  release  in  Obelia,  probably  O.  dichotoma.  However,  like  several 
contemporary  workers  (see  p.  97)  he  assigned  his  material  to  Hartlaubella  gelatinosa 
which  has  no  medusa.  He  probably  identified  his  material  using  the  well  illustrated  work  of 
van  Beneden  (1844),  who  made  the  same  error.  Several  other  workers  have  assigned  an 
Obelia-\ike  medusa  to  H.  gelatinosa  and,  despite  the  early  clarification  by  Couch  (1844), 
there  was  confusion  for  many  years  between  O.  dichotoma,  O.  geniculata,  H.  gelatinosa  and 
Gonothyraea  loveni  (see  notes  under  H.  gelatinosa  (p.  97),  and  Cornelius,  1977a). 

Couch  (1844)  was  in  addition  the  first  to  note  medusa  release  in  Obelia  geniculata  (p.  120). 
Other  early  records  in  that  species  were  contributed  by  F.  W.  L.  Thomas  (in  Johnston, 
1847: 467),  Hincks  (1852  :  85)andGosse(1853  :  84-90,  pi.  4). 

Illustrations  of  the  medusae  of  0.  dichotoma  were  first  provided  by  Baster  (1762,  pi.  5,  fig. 
7),  but  the  earliest  of  0.  geniculata  appeared  nearly  a  century  later,  in  Gosse's  (1853)  book. 

Naumov  (1969,  as  O.  longissima,  here  referred  to  O.  dichotoma)  reported  that  'the 
medusae  were  apparently  not  liberated  from'  Black  Sea  populations.  But  the  species  of 
Obelia  are  exceptionally  well  documented  and  since  there  is  no  other  report  of  medusa 
retention  in  the  genus  a  repeat  of  the  observations  would  be  desirable. 

REPRODUCTIVE  SEASON.  Medusae  of  Obelia  spp.  have  been  reported  around  the  British  Isles 
in  all  months  but  are  commonest  from  'spring  to  late  autumn'  (Russell,  1953).  Fertile 
colonies  were  recorded  on  the  coast  of  Egypt  between  6  Sep  and  4  Oct,  1933  (Billard,  1936). 

DISTRIBUTION.  Found  throughout  European  and  African  coastal  waters  with  the  exception 
of  the  Black  Sea,  from  which  there  are  no  confirmed  records  (but  see  last  paragraph  of 
Dispersive  stage).  The  species  is  remarkable  in  being  nearly  cosmopolitan  in  coastal  waters 
and  is  one  of  the  most  widely  distributed  of  hydroids.  Aspects  of  this  are  discussed  on  page 
45;  and  a  detailed  summary  of  locality  records  has  appeared  elsewhere  (Cornelius,  1975a). 

HABITAT.  Tolerance  of  12%o  salinity  recorded  by  Calder  (1976)  in  S  Carolina.  Unusual 
substrates  noted  in  the  literature  included  a  sponge  and  the  fins  of  a  shark  (Couch, 
1 844  :  XV,  as  'the  sea  thread  of  Ellis'),  a  turtle  (Allman,  1 888,  as  Campanularia  cheloniae),  a 
crab  (Rasmussen,  1973:  the  colony  was  thought  to  be  feeding  on  debris  in  the  exhalent 
current);  on  the  parasitic  copepod  Lernaeocera  on  the  gills  of  a  cod  (Gadus);  and  on  the 
back  and  claws  of  an  anomuran  'crab',  Lithodes  maia  (in  Evans,  1978).  Probably  the  record 
by  Gudger  (1937),  of 'Obelia  commensuralis'  on  the  blueback  herring  (Alosa  aestivalis), 
referred  to  this  species;  but  his  account  suggested  that  the  hydrorhizae  entered  the  muscles  of 
the  fish  and  the  identification  is  not  certain. 

REMARKS.  A  revision  of  this  species  has  appeared  recently  (Cornelius,  1975a)  and  only 
additional  notes  are  given  here.  Identification  of  the  hydroid  stage  is  discussed  above  (p. 
1 13).  Nematocysts  were  described  by  Ostman  (1979)  and  by  others  whom  she  cited. 

Tubularia  clytioides  Lamouroux,  in  Freycinet,  1824,  was  based  on  live  material  collected 
from  algae  in  the  Azores.  Probably  it  was  Obelia  dichotoma.  Rees  &  White  (1966)  evidently 
thought  Silicularia  gracilis  Meyen,  1834,  also  described  from  Azores  material,  to  be 
conspecific  but  while  the  clytioides  type  specimen  was  an  erect  colony  that  of  gracilis  was 
reptant.  Rees  &  White  presented  a  muddled  synonymy,  referring  to  the  'species' 
(clytioides  +  gracilis)  as  Orthopyxis  clytioides  (Lamouroux);  and  including  in  its  synonymy 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  1 19 

the  entry  "Silicularia  clytioides  Meyen,  1834',  evidently  a  lapsus  for  "S.  gracilis  Meyen'. 
Nutting  (1915)  had  made  the  same  error  fifty  years  earlier  (p.  50). 

The  once  widely  recognized  nominal  species  O.  commissuralis  McCrady,  1857,  based  on 
the  hydroid  stage,  was  referred  to  O.  dichotoma  first  by  Bohm  (1878),  and  again  by  me 
(19750).  Bohm's  paper  has  been  overlooked;  and  I  know  of  no  other  works  giving  this 
synonymy. 

Obelia  plicata  Hincks,  1868,  described  from  infertile  hydroid  material,  type  locality 
'Shetland',  was  recently  referred  to  the  present  species  without  comment  (Cornelius,  1975a). 
I  have  not  been  able  to  locate  type  material.  The  species  deserves  close  evaluation  as  it  has 
been  recognized  by  several  authors  (Marktanner-Turneretscher,  1890;  Nutting,  1915,  as 
?O.  plicata;  Billard,  1927;  Kramp,  1935,  as  Laomedea  plicata;  Fraser,  1944;  Teissier,  1965; 
Rasmussen,  1973)  although  not  by  Broch  (19 126).  O.  plicata  was  said  by  Hincks  to  resemble 
H.  gelatinosa  in  having  compound  stems  but  the  hydrothecal  rims  were  even,  not  castellated. 
The  hydrotheca  was  said  by  Hincks  'very  much  [to]  resemble  in  form  that  of  O.  dichotoma". 
Gonothecae  were  first  assigned  to  O.  plicata  by  Marktanner-Turneretscher  (1890)  who  said 
they  were  the  same  as  those  of  O.  dichotoma.  Indeed,  the  two  species  seem  conspecific.  Some 
older  colonies  of  O.  dichotoma  (e.g.  BMNH  1948.9.8.14,  Plymouth,  21  Apr  1899,  coll.  E.  T. 
Browne)  exhibit  compound  stems  resulting  from  overgrowth  of  up  to  10  young,  pale  stems 
along  the  older,  thicker  and  darker  ones.  The  specimen  mentioned  resembles  closely  the 
colony  illustrated  by  Hincks  as  O.  plicta,  and  synonymy  seems  justified.  Godeaux  (1941,  as 
H.  gelatinosa)  described  similar  O.  dichotoma  material,  from  the  North  Sea;  while  Billard 
(1927)  maintained  O.  plicata  distinct  on  the  basis  of  the  original  characters.  Broch  (19126) 
alone  raised  doubts,  but  was  non-committal. 

The  factors  causing  overgrowth  in  occasional  O.  dichotoma  specimens  are  not  known, 
most  colonies  being  solitary.  However,  such  overgrowth  has  been  described  in  other  hydroids 
and  called  'auto-epizoism'  (Millard,  1973).  If  an  overgrown  specimen  results  from  second 
and  subsequent  planulae  settling  on  the  original  colony  then  the  whole  should  be  regarded  as 
an  aggregation  of  colonies,  and  not  as  a  single  colony.  The  phenomenon  is  discussed  above 
(p.  40). 

Campanularia  cheloniae  Allman,  1888,  was  based  on  infertile  material  collected  during 
the  'Challenged  expedition  from  the  back  of  a  turtle;  but  the  locality  of  the  turtle  was  not 
recorded.  The  original  description  and  what  remains  of  the  type  series  (BMNH 
1 888. 1 1.13.15,  small  colony  in  spirit  +  microslide  preparation)  are  probably  O.  dichotoma. 

Laomedea  sargassi  Broch,  1913,  was  recorded  from  Ghana  by  Buchanan  (1957  :  360). 
The  name  sargassi  was  a  nom.  nov.  for  O.  hyalina  Clarke,  1879,  introduced  just  to  prevent 
confusion  with  Gonothyraea  hyalina  auct.,  and  is  therefore  inadmissible  under  the  current 
conventions  of  nomenclature.  Congdon  (1907  :  468)  described  material  from  Bermuda  as  O. 
hyalina,  but  Hargitt  (1909)  gave  that  material  specific  status,  as  Laomedea  congdoni.  I  have 
previously  (Cornelius,  1975a)  referred  O.  hyalina  Clarke  and  O.  congdoni  (Hargitt)  to 
O.  dichotoma,  but  overlooked  the  nom.  nov.  sargassi.  Recent  examination  of  the  type 
material  of  O.  congdoni  (U.S.  nat.  mus.  cat.  no.  42647,  microslide  preparation  of  infertile 
fragments)  confirms  that  it  too  is  simply  O.  dichotoma.  Obelia  hyaliana  Vannucci,  ?  in  ms, 
was  a  name  given  to  some  "O.  hyalina'  material  which  Vannucci  (1955)  later  considered 
conspecific. 

Obelia  geniculata  (Linnaeus,  1758) 
(For  illustrations  see  Cornelius,  19750) 

Sertularia  geniculata  Linnaeus,  1758  :  812. 

Obelia  geniculata:  Hincks,  1868:  149-151,  pi.  25,  fig.  1,  la;  Cornelius,  1 9750:272-278, figs  1,5. 

For  further  synonymy  see  Cornelius  ( 1 91 5a),  Remarks  and  Table  4. 

DISPERSIVE  STAGE.  A  medusa.  See  Remarks  and  notes  under  Obelia  dichotoma. 

REPRODUCTIVE  SEASON.  Russell  (1953)  recorded  the  following  in  British  waters:  Obelia  sp. 
medusae  found  nearly  throughout  year,  especially  'spring  to  late  autumn'  (Russell);  medusae 


120  P.  F.  S.  CORNELIUS 

released  at  Plymouth,  March-November  (Orton,  1920);  some  evidence  of  lunar  periodicity 
in  release  times  (Elmhirst,  1925). 

In  the  Mediterranean,  Lo  Bianco  (1909)  recorded  medusa  release  at  Naples  March-June 
and  October-January. 

DISTRIBUTION.  An  almost  cosmopolitan  shallow  water  species  found  nearly  throughout  the 
eastern  North  Altantic. 

The  species  was  not  recorded  from  the  Black  Sea  by  Naumov  (1969).  He  listed  the  species 
as  absent  from  the  Baltic  as  well  but  Stechow  (1927)  reported  it  from  Stockholm.  It  was 
recorded  from  Iceland  and  the  Faeroes  but  not  from  Spitzbergen  by  Kramp  (1929),  although 
already  known  from  Jan  Meyen  Island  (Broch,  1918).  Calder  (1970)  among  others  recorded 
the  species  from  W  Greenland;  but  I  (Cornelius,  19750)  misquoted  him  as  including  E 
Greenland  in  its  distribution.  I  know  of  no  records  from  E  Greenland. 

HABITAT  [see  also  Geographical  distribution  (p.  45),  the  habitat  notes  under  O.  dichotoma 
and  Cornelius,  19750].  The  species  was  collected  from  the  fins  of  a  spur-dog  (Squalus 
acanthias)  by  Couch  (1844),  who  deduced  that  planula  settlement  must  be  rapid.  Other 
actively  swimming  animals  on  which  O.  geniculata  has  been  recorded  include  a  sea-horse 
(Hippocampus  guttulatus,  by  Zirpolo,  1939,  1940)  and  parasitic  copepods  themselves  on 
fish  (Lernaeocera  spp.  on  Gadus  (cod),  by  Leloup,  193 1/?;  on  Lernaeocem  on  Clupea 
(herring),  by  Debouteville  &  Nunes,  1951,  1952;  on  Dinemoura  on  Cetorhinus  (basking 
shark)  by  Debouteville  &  Nunes).  Long  distance  travel  on  floating  algae  has  also  been 
recorded  (Arnaud,  Arnaud,  Intes  &  Le  Loeuff,  1976).  Recorded  at  100  m  depth  by  Naumov 
(1960).  However,  some  of  these  records  on  fish  might  have  been  of  0.  dichotoma. 

REMARKS.  A  revision  of  this  species  has  appeared  elsewhere  (Cornelius,  19750)  and  the 
present  account  simply  adds  to  that. 

Couch  (1844)  gave  some  interesting  details.  He  stated  the  number  of  tentacles  of  the 
hydranth  to  vary  widely  (1 1-29  on  p.  2,  14-28  on  p.  38).  He  described  medusa  bud  develop- 
ment but  not  medusa  release,  which  was  not  known  till  1847  in  this  species  (Cornelius, 
19750,  1977a,  b;  see  also  p.  1 18).  He  recorded  a  range  of  tissue  colour  in  the  hydroid  from 
'white  to  deep  red'. 

The  nematocysts  were  described  by  Ostman  (1979)  and  by  others  whom  she  cited. 

Campanularia  vermicularis  Van  Beneden,  1866,  was  described  without  illustrations  and 
identification  has  been  thought  impossible  (Vervoort,  19460;  Leloup,  1947).  The  species  was 
based  on  Belgian  material.  Van  Beneden  kept  live  colonies  for  three  weeks.  He  described  the 
species  as  being  more  robust  than  O.  geniculata,  and  as  having  larger  hydrothecae  and  more 
numerous  annulations.  It  was  collected  off  the  brown  alga,  Fucus  vesiculosus  L.  The 
hydranth  was  said  to  have  as  many  as  40^4  tentacles,  a  character  which  might  one  day  help 
to  identify  his  material  (cf.  Couch's  observations,  immediately  above).  Meanwhile  I  follow 
Vervoort  in  treating  the  species  here  but  not  including  it  in  the  synonymy. 

Obelia  spp.  indet. 

1.  Hydroids.  Obelia  longicyatha  Allman  (1877  :  10,  pi.  7,  figs  4-5)  was  recorded  from 
35°  42'  N,  8°  40'  W  (Cap  Spartel,  Tangier)  and  the  Gulf  of  Cadiz  by  Billard  (1907  :  168,  as 
Clytia  longicyatha),  but  the  identity  of  the  species  remains  unclear.  I  have  discussed  it 
elsewhere  (Cornelius,  19750:264).  See  also  the  note  on  Campanularia  vermicularis  Van 
Beneden,  1866  (immediately  above). 

2.  Medusae.  Some  species  from  the  eastern  North  Atlantic  cannot  be  assigned  to  a  hydroid. 
Obelia  adriatica  Neppi  (1912  :  726-727,  pi.  3,  fig.  8,  8a)  was  described  from  Adriatic 
material;  and  Obeliopsis  fabredomergui  Le  Danois  (1913:110)  was  based  on  material  from 
the  Little  Minch,  NW  Scotland.  It  seems  most  unlikely  that  either  is  valid.  O.  plana  Sars 
(1835  :  28,  pi.  5,  fig.  13,  as  Thaumantias)  was  referred  to  O.  jlabellata  hydroid  by  Bedot 
(1910  :  484)  and  some  later  authors,  and  hence  would  fall  in  the  synonymy  of  O.  dichotoma. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  121 

But  there  seems  slim  reason  for  this  identification  (cf.  Cornelius,  19750:271,  footnote). 
Other  pre-1910  medusa  names  in  this  genus  were  listed  by  Bedot  (1901-1925)  and  Mayer 
(1910),  and  are  not  repeated  here.  As  explained  above  (p.  1 18)  most  cannot  be  confidently 
assigned  to  a  hydroid  stage  and  a  new  list  would  serve  little  purpose. 


Taxa  now  referred  to  other  families 

A  few  genera  and  species  recorded  from  the  eastern  North  Atlantic  are  no  longer  referred  to 
the  Campanulariidae.  As  several  have  been  included  in  this  family  in  some  standard  works 
notes  on  them  are  given  here  to  summarize  the  current  opinions.  The  taxa  are  treated  in  date 
order. 

1.  The  genus  Capsularia  Cuvier,  1797,  was  used  by  Gray  (1848:85-87)  to  embrace 
several  British  species  of  Campanulariidae  but  should  be  regarded  a  junior  synonym  of 
Coryne  Gaertner,  in  Pallas,  1774  (discussion  in  Cornelius,  1 9756  :  378). 

2.  Sert ularia  fruticosa  Esper,  71810  [pi.  34,  figs  1-2  (see  note  2,  p.  124);  also  Hammer,  in 
Esper,   1829:162-163  (syn.  Sertularia  laxa  Lamarck,   1816:116;  Laomedea  sauvagii 
Lamouroux,  1816:  206;  both  nom.  nov.  pro  S.  fruticosa)]  was  recorded  from  the  Adriatic  Sea 
by  Marktanner-Turneretscher  (1890  :  205)  as  Campanularia  fruticosa.  The  species  is  now 
referred  to  the  genus  Thyroscyphus  in  the  family  Sertulariidae  (e.g.  Vervoort,  1967;  Millard, 
1975).   It  is  otherwise  unrecorded  from  the  eastern  North  Atlantic  and  Marktanner- 
Turneretscher's  record  may  well  be  wrong.  However,  T.fruticosus  occurs  in  the  Red  Sea  as 
far  north  as  the  Gulf  of  Suez  (Vervoort,  1 967). 

The  exact  date  of  publication  of  the  species  name  is  unusually  difficult  to  determine  and 
the  necessary  bibliographic  work  on  Esper's  important  multi-part  book  has  apparently  not 
been  done.  Precise  dating  would  be  useful  as  the  plates  were  issued  several  at  a  time  around 
1810,  and  they  carry  binominals.  They  thus  predate  the  posthumous  text,  edited  by  Hammer 
(Esper,  1 829),  from  which  this  part  of  the  work  is  often  thought  to  date.  The  plates  predate 
also  several  important  European  works  on  zoophytes  [e.g.  those  by  Lamarck  (1816)  and 
Lamouroux  (1812,  1816,  1821;  Lamouroux  et  al.,  1824)].  Esper's  work  is  rare,  and  few  of 
the  original  wrappers  survive;  but  the  information  on  an  incomplete  set  of  wrappers  in  the 
BMNH  library  gives  hope  that  dating  would  be  simple  if  a  complete  copy  were  located.  See 
also  section  7,  below. 

3.  The  genus  Cymodocea  Lamouroux,  1816:214,  originally  included  two  species,  C. 
simplex  and  C.  ramosa,  both  described  as  new.  C.  simplex  was  based  on  material  from  Great 
Yarmouth  and  'Ireland',  C.  ramosa  on  fragments  from  the  Antilles.  Johnston  (1838) 
commented  on  the  genus  and  on  the  identities  of  the  two  species;  and  Gray  (1848)  dismissed 
them  as  'doubtful  species'  at  the  end  of  the  Campanulariidae.  Lamouroux  (1821)  and 
Lamouroux  et  al.  (1824)  included  additional  species  in  the  genus  but  these  do  not  enter  the 
present  discussion.  Billard  (1909),  who  saw  some  of  Lamouroux'  original  material,  referred 
the   two   species  to  Nemertesia  antennina  (Linnaeus,    1758),   family   Plumulariidae.   I 
designate  C.  ramosa  type  species  of  the  genus  Cymodocea,  the  name  of  which  hence 
becomes  a  junior  subjective  synonym  of  Nemertesia  Lamouroux,  1812.  Billard  was  confi- 
dent that  the  C.  ramosa  material  he  saw  was  type,  but  had  doubts  about  that  of  C.  simplex. 
His  doubts  are  significant  since  in  J.  Fleming's  opinion,  quoted  by  Johnston  (1838),  the 
original  illustrations  of  C.  simplex  might  be  identified  as  an  Obelia  dichotoma  colony  lacking 
hydrothecae.  I  concur  with  Fleming's  opinion;  and  disagree  with  Johnston  who  thought 
simplex  was  a  plumularid.  If  the  illustration  is  actually  of  O.  dichotoma  then  Billard's 
caution  was  justified  and  the  specimen  he  saw  was  not  type.  Almost  all  the  Lamouroux 
collections  were  destroyed  during  the  liberation  of  Caen  on  7  July,  1944,  so  the  C.  simplex 
material  is  no  longer  available  (Redier,  1967).  However,  C.  simplex  is  here  provisionally 
referred  to  O.  dichotoma  (p.  1 1 7). 


122  P.  F.  S.  CORNELIUS 

The  species  Cymodocea  comata  Lamouroux,  1821,  was  recorded  from  the  coast  of  Devon 
by  W.  E.  Leach  (in  Johnston,  1838);  but  the  species  was  referred  to  Nemertesia  ramosa 
(Lamarck,  1 8 1 6)  by  Billard  ( 1 909),  who  saw  type  material. 

4.  Campanularia  intertexta  Couch,  1844  :  41^2,  pi.  11,  fig.  3,  was  based  on  material 
then  in  the  Museum  of  the  Royal  Institution  of  Cornwall,  Truro.  The  material  almost 
certainly    no    longer   exists   (Curator,    Roy.    Inst.    Cornwall,    pers.    comm.).    Johnston 
(1847:109-110)  repeated  Couch's  description   verbatim  but   regarded  the   species  as 
unsoundly  based.  Gray  (1848  :  88)  listed  it  without  comment,  incidentally  assigning  to  it 
Shetland  material  of  Lafoea  dumosa  (Fleming,  1820),  BMNH  reg.  no.  1847.9.24.66.  At 
the  same  time  Gray  proposed  the  new  genus  Conchella  (?lapsus  pro  Couchella),  of  which 
Campanularia  intertexta  is  type  species  by  monotypy.  Hincks  (1868:220)  thought  the 
species   might   be   L.   dumosa,   stating   that   Couch's   description   was   inadequate   for 
identification.  Later  Bedot  (1905  :  157)  referred  intertexta   to  'Coppinia  arcta\  an  invalid 
species  accepted  by  several  nineteenth  century  authors  but  merely  based  on  the  coppinia  or 
reproductive  branch  of  L.  dumosa.  Couch's  illustration  shows  an  unidentifiable  campanu- 
larid  hydroid,  possibly  Orthopyxis  Integra,  growing  on  a  coppinia  of  L.  dumosa.  Thus  the 
type  'series'  was  mixed.  The  name  intertexta  is  here  restricted  solely  to  the  illustrated 
coppinia,  which  becomes  lectotype.  Hence  Campanularia  intertexta  is  to  be  regarded  a 
junior  synoym  of  L.  dumosa;  and  the  genus  name  Conchella  a  junior  synonym  of  Lafoea 
Lamouroux,  1821.  See  also  page  65. 

5.  Campanularia   lacerata   Johnston,    1847:111,   pi.   28,   fig.    3,   is   now   known   as 
Opercularella  lacerata  and  referred  to  the  Campanulinidae  (e.g.  Millard,  1975).  The  species 
has  been  referred  to  a  variety  of  genera  in  the  literature,  including  Capsularia,  Laomedea, 
Wrightia  and  Calycella  (references  in  Hincks,  1868). 

6.  Laomedea  obliqua  Johnston,  1847  :  106-107,  pi.  28,  fig.  1,  based  on  British  material,  is 
today  known  as  Monotheca  obliqua  and  Plumularia  obliqua  by  different  authors  and 
referred  to  the  Plumulariidae  (e.g.  Millard,  1975). 

7.  Campanularia  fruticosa  Sars,  1850  :  138-139,  is  today  referred  to  Lafoea  dumosa 
(Fleming,  1820),  in  the  Lafoeidae  (e.g.  Cornelius,  19756).  It  has  often  been  given  full 
specific  status,  as  L.  fruticosa.  See  also  section  2,  above. 

8.  Campanularia  abietina  Sars,  1850  :  139,  based  on  Norwegian  material,  has  long  been 
known  as  Grammaria  abietina  and  assigned  to  the  family  Lafoeidae  (e.g.  Cornelius,  19756). 

9.  Campanularia  parvula  Hincks,  1853  :  178,  pi.  5a,  was  probably  based  on  material  of 
Calycella  syringa  (Linnaeus,  1767)  lacking  operculae.  Several  authors  have  referred  the 
species  to  Lafoea  Lamouroux,  1821,  but  this  seems  wrong  (references  and  discussion  in 
Cornelius,  19756). 

10.  Campanularia  gracillima  Alder,   1856a:361,  pi.   14,  figs  5-6,  based  on  hydroid 
material  from  NE  England,  has  frequently  been  referred  to  Lafoea,  family  Lafoeidae;  but 
lately  to  the  species  Lafoea  dumosa  (Fleming,  1820)  (e.g.  Cornelius,  19756;  Cornelius  & 
Garfath,  1980). 

11.  Laomedea  acuminata  Alder,  18566  :  441,  pi.  16,  figs  5-8,  based  on  hydroid  material 
from  NE  England,  is  the  hydroid  of  an  Aequorea  sp.  medusa,  family  Aequoreidae.  The  two 
species  recognized  from  British  waters  from  the  medusa  stage  were  described  earlier  so  the 
name  acuminata  should  be  regarded  provisional.  Although  the  medusa  generation  of  the  two 
can  be  separated  the  respective  hydroids  are  morphologically  identical,  so  far  as  is  known. 
Hence  at  present  it  cannot  be  decided  to  which  of  the  'medusa  species'  acuminata  should 
rightly  be  referred  (Russell,  1953). 

12.  The  nominal  species  Laomedea  tenuis  Allman,  1859  :  367-368,  was  once  known  as 
Leptoscyphus  tenuis  (e.g.  Hincks,  1868).  It  is  currently  regarded  as  a  campanulinid,  having 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  123 

been  provisionally  referred  to  Phialella  quadrata  (Forbes,  1848)  by  Stechow  (1923a  :  129) 
and  Rees  (1939:441).  Browne  (1896:479)  summarized  the  mistaken  observations  by 
Allman  who  assigned  the  medusa  of  one  species  to  the  hydroid  of  another.  Campanulina 
tennis  Van  Beneden,  1886  :  174,  176,  pi.  13,  is  a  different  nominal  species,  provisionally 
referred  to  Aequorea  vitrina  Gosse,  1853,  by  Russell  (1953).  Bedot  (1910)  listed  an  earlier 
homonym  of  C.  tenuis. 

13.  Campanularia  fastigiata  Alder,  1860  :  73-74,  pi.  5,  fig.  1,  based  on  Scottish  material, 
was  known  for  some  decades  as  Stegopoma  fastigiatum.  Edwards  (1973)  showed  that 
fastigiatum  was  the  hydroid  stage  of  the  medusa  Modeeria  rotunda  (Quoy  &  Gaimard, 

1827),  the  binominal  of  which  takes  priority.  The  species  is  assigned  to  the  family 
Laodiceidae  (e.g.  Rees  &  Rowe,  1969;  Edwards,  1973;  see  also  Cornelius  &  Garfath,  1980). 

14.  Campanularia  humilis  Hincks,  1866  :  298,  is  now  known  as  Cuspidella  humilis.  Its 
affinities  are  unclear  but  it  was  removed  from  the  Campanulariidae  long  ago  (discussion  in 
Cornelius  &  Garfath,  1 980). 

15.  Clytia  eucophora  Haeckel,  1879  :  168,  was  a  combination  applied  to  the  hydroid  stage 
of  Eucopium  primordiale  Haeckel,  1879  :  168  (sic),  from  Corsica.  As  Mayer  (1910 : 236) 
pointed  out,  the  two  names  are  objective  synonyms  with  the  same  date  of  publication. 
Mayer,  as  first  reviser,  suppressed  eucophora  in  favour  of  primodiale\  but  at  the  same  time 
referred  Haeckel's  species  to  Eucope  picta  Keferstein  &  Ehlers,  1861.  Although  Kramp 
(1961)  omitted  to  treat  E.  picta  there  is  no  doubt  that  it  and  the  Haeckel  species  should  be 
referred  to  the  Phialellidae.  Weismann  (1883  :  158)  introduced  the  combination  Clytia 
eucopophora,  a  lapsus  of  Haeckel's  spelling. 

16.  Campanularia  mutabilis  Ritchie,   1907  :  504,  pi.  23,  figs  3-5,  based  on  Azores 
material,  is  now  known  as  Scandia  mutabilis  and  referred  to  the  Lafoeidae  (e.g.  Millard, 
1975). 

17.  Campanularia  divisa  was  attributed  by  Bassindale  (1941  :  148)  to  Todd  (1906 :  137) 
whom  he  supposed  to  have  reported  material  from  Ilfracombe.  Bassindale  misread  Todd's 
list,  which  included  Campanularia  species  and  Tubularia  indivisa.  The  word  'indivisa'  was 
split  between  the  two  lines  (in/divisa)  and  evidently  read  wrongly  by  Bassindale. 

18.  The  genus  Hincksella  Billard,  1918  :  22,  was  considered  by  Totton  (1930)  and  Ralph 
(1957)  to  be  closely  related  to  the  Campanulariidae,  but  is  now  referred  to  the  Syntheciidae 
(e.g.  by  Millard,  1975). 

19.  The  genus  Billardia  Totton,  1930  :  150  (type  species  B.  novae  zealandiae,  by  original 
designation)  was  based  on  southern  hemisphere  material.  Totton  assigned  Billardia  to  the 
Campanulariidae  but  the  genus  is  similar  to  Hincksella  and  like  it  comes  within  the  scope  of 
the  Syntheciidae  sensu  Millard  (1975),  in  my  opinion.  Blanco  (\961b)  and  Stepanyants 
(1979),  however,  retained  Billardia  in  the  Campanulariidae.  Possibly  a  greater  under- 
standing of  the  reproductive  structures  will  help.  Totton  suspected  that  the  blastostyles  of 
Billardia  were  produced  in  place  of  hydranths,  within  the  hydrothecae,  but  said  his  material 
was  inadequately  preserved  for  him  to  be  certain.  If  his  suspicion  were  confirmed  the 
Syntheciidae  would  be  the  correct  family  for  Billardia. 


Notes 

1  (See  p.  65).  G.  D.  Westendorp  (1813-1868)  apparently  produced  only  two  works 
involving  coelenterates  (Westendorp,  1843,  1853).  Both  were  on  the  zoophytes  of  the 
Belgian  coast.  The  first  was  a  straightforward  taxonomic  account  including  some  new 
genera  and  species,  among  them  Clytia  ryckholtii  (here  referred  to  Orthopyxis  integrd). 
Probably  none  is  valid.  His  second  work,  published  in  1853,  was  remarkable  in  being 
illustrated  by  dried  specimens  mounted  on  sheets.  It  is  rare  but  still  important  since  one  of 


124  P.  F.S.CORNELIUS 

the  included  species  was  described  as  new  in  the  1843  work,  and  the  specimens  used  in 
illustration  might  be  considered  types.  The  1853  work  comprised  a  printed  title  page,  a 
printed  page  giving  a  list  of  species  and  localities,  and  32  hebarium  sheets  each  bearing  one 
species.  There  were  16  hydroids,  14  bryozoans  and  2  sponges.  Examples  of  the  work  were 
seen  by  Neviani  (1903),  Bedot  (1910  :  200-201)  and  Billard  (1914;  also  seen  by  Leloup, 
1947  :  5);  and  another,  imperfect  copy  has  recently  been  found  in  the  British  Museum 
(Natural  History)  (Cornelius,  unpublished  ms  in  BMNH).  I  have  not  seen  an  intact  copy. 

Neviani  listed  the  32  species  but  C.  ryckholtii  was  not  among  them.  The  copy  described  by 
Billard  (1914),  in  Brussels  (Mus.  roy.  Hist,  nat.;  inv.  no.  3440),  similarly  does  not  have  C. 
ryckholtii.  The  BMNH  example  also  does  not  include  that  species.  The  copy  evidently 
came  to  the  (then)  British  Museum  library  about  1867.  Certainly  a  collection  of  specimens 
corresponding  with  Neviani's  list  was  purchased  from  Westendorp  at  that  time,  and  was 
given  the  32  zoological  accession  numbers  1867.5.4.22-24  and  1867.5.7.1-29.  The 
specimens  were  curated  each  under  its  own  species,  and  the  printed  species  list  was  cut  up  to 
provide  labels  which  were  in  most  cases  glued  to  the  herbarium  sheets.  Much,  perhaps  all,  of 
this  material  survives  and  is  still  curated  under  the  various  species;  but  I  have  not  found  the 
title  page  of  this  copy. 

Only  one  type  specimen  of  any  group  is  included  in  this  collection,  that  of  the  hydroid, 
Plumularia  macleodi  Westendorp,  1843  (BMNH  reg.  no.  1867.5.7.7).  It  can  be  considered  a 
syntype  of  P.  macleodi;  and  is  in  fact  a  specimen  of  Kirchenpaueria  pinnata  (Linnaeus, 
1758).  P.  macleodi  is  thus  a  junior  synonym  of  K.  pinnata.  Billard  (1914),  working  on  the 
corresponding  Brussels  syntype  specimen,  reached  the  same  conclusion. 

2  (See  p.  121).  Some  evidence  was  presented  by  Sherborn  (1922  :  1,  entry  under  Esper; 
Sherborn,  1926  :  2528,  entries  under  fruticosa,  Laomedea  and  fruticosa,  Sertularia)  that  at 
least  some  of  the  plates  of  Esper's  (1829)  'Die  Pflanzen-Thiere'  appeared  before  1816, 
possibly  around  1810.  Since  the  plates  bear  binominals,  and  might  predate  the  synoptic 
works  of  Lamouroux  (1812  onwards)  and  Lamarck  (1816  onwards),  correct  dating  is 
important.  However,  it  seems  that  bibliographers  have  not  yet  dated  the  plates  and 
Sherborn's  (1926)  provisional  date  of  1810  for  the  plate  of  Sertularia  fruticosa  Esper  is  the 
best  compromise.  Certainly  the  plate  of  fruticosa  predated  the  works  of  Lamarck  ( 1 8 1 6)  and 
Lamouroux  (1816),  which  referred  to  Esper's  plate  as  being  already  published.  Indeed,  they 
proposed  new  names  in  place  of  fruticosa.  The  three  volume  Esper  work  was  issued  in  parts, 
some  after  Esper's  death  in  1810;  and  the  few  copies  I  have  seen  have  been  collated  with  the 
undoubtedly  earlier  Sertularia  plates  intermingled  with  the  text.  But  the  partial  set  of 
wrappers  with  the  BMNH  copy  gives  much  detailed  information,  and  a  full  set  might 
enable  all  the  plates  of  this  scarce  work  to  be  dated  accurately. 

3  (See  p.  1 12).  Dating  of  the  two  works  here  listed  as  Peron  &  Lesueur,  1810a,  718106,  has 
caused  confusion.  A  clarification  is  desirable  since  in  them  many  genus  names  of  medusae 
were  introduced  which  are  still  used.  The  works  comprised  the  earliest  serious  systematic 
treatment  of  medusae,  and  these  authors  finally  disbanded  the  genus  Medusa  Linnaeus, 
1758.  The  first  of  the  two  works  was  one  of  a  series  of  journal  papers  written  by  the  two 
authors,  and  comprised  their  taxonomic  treatment  of  medusae.  Several  other  papers  in  the 
series  also  dealt  with  marine  animals  but  are  not  important  here.  They  have  been  listed  by 
Goy  (1980).  The  second  work  (Peron  &  Lesueur,  718106),  a  book  entitled  Histoire  generale 
des  meduses,  brought  the  series  of  papers  together  each  forming  a  chapter  of  the  book.  The 
book  version  was  repaginated  from  1  on,  and  was  presumably  published  after  the  series  of 
papers  (although  it  might  conceivably  have  been  issued  in  parts  as  the  papers  appeared). 

The  publication  date  of  the  taxonomic  paper  (Peron  &  Lesueur,  1810#)  has  been  taken  as 

1809  by  most  authors  and  this  date  might  be  inferred  from  the  title  page  of  the  volume  of  the 
journal  in  which  it  appeared.  But  there  is  good  evidence  that  it  did  not  appear  until  January, 

1810  (Sherborn,  1914;  1929  :  4455,  entry  under  Obelia\  Cornelius,  1975a;  Goy,  1980).  The 
book,  the  Histoire  generale  des  meduses,  has  been  almost  entirely  overlooked.  It  was  dated 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  125 

1809  in  the  British  museum  catalogue  of  printed  books.  But  J.  Goy  (pers.  comm.)  and  I 
concur  that  the  book  version  almost  certainly  would  have  followed  the  series  of  papers,  and 
so  would  have  appeared  at  least  as  late  as  1810  (unless  the  book  version  were  issued  in  parts). 

Hence  the  many  names  introduced  by  Peron  &  Lesueur  almost  certainly  date  from  the 
1810^  paper,  published  January,  1810.  Apart  from  minor  heading  changes  the  book  version 
(71810&)  was  apparently  printed  from  the  same  'blocks'  as  the  series  of  papers.  For 
taxonomic  purposes  the  two  versions  differ  only  in  date. 

The  numerous  plates  that  were  to  have  accompanied  the  account  of  the  medusae  (Peron  & 
Lesueur,  1810#)  were  prepared  but  not  published  immediately  (cf.  Cornelius,  \915a  :  253, 
footnote;  1977a:49,  footnote).  But  most  of  the  illustrations  showing  Peron  &  Lesueur's 
'new'  species  were  brought  together  in  an  extremely  rare  work  usually  attributed  to  Lesueur 
alone  (Peron  &  Lesueur,  71815).  This  work  was  cited  by  Haeckel  (1879,  in  several  of  his 
synonymies,  as  the  Planches  inedites)  and  Totton  (1965  :  p.  45,  pi.  8)  among  others  but 
almost  all  twentieth  century  medusa  workers  have  overlooked  it.  Fewer  than  half  a  dozen 
copies  are  known  (Totton,  1965;  Goy,  1980).  Goy  (1980)  cited  evidence  from  the  P.-v. 
Seanc.  Acad.  Sci.  Paris,  1795-1831  (issue  covering  14  August  1815)  5  :  532,  that  1815  is  the 
correct  date;  and  not  71811  as  given  in  the  British  museum  catalogue  of  printed  books 
(187  :  1 1 1)  under  Peron  alone.  She  concluded  that  the  bulk  of  the  illustrations,  showing  most 
of  the  nominal  species  newly  described  by  Peron  &  Lesueur  (1810(2),  were  copied  and 
published  by  a  variety  of  contemporary  compilers  in  their  own  works  and  so  made  public. 
They  included  such  famous  names  as  de  Blainville,  Cuvier,  Milne  Edwards,  Lamarck  and 
Lesson  (references  in  Goy,  1980).  The  original  Lesueur  drawings  are  preserved  in  the 
Museum  at  Le  Havre. 

Although  the  bulk  of  the  rare  work  (Peron  &  Lesueur,  71815)  comprised  illustrations  of 
medusae  exquisitely  drawn  by  Lesueur,  the  title  page  leaves  no  doubt  that  the  authorship 
should  be  ascribed  jointly  to  Peron  &  Lesueur  (Mrs  A.  Datta,  pers.  comm.).  Peron  had  died 
in  1810  and  it  can  be  inferred  that  Lesueur  wished  him  still  to  be  senior  author,  as  he  had 
been  of  the  lengthy  text  (Peron  &  Lesueur,  1 8 1  Oa,  7 1 8 1 06)  of  the  work. 


Acknowledgements 

I  am  indebted  to  Professor  W.  Vervoort,  Rijksmuseum  van  Natuurlijke  Historic,  Leiden,  for 
generous  and  detailed  discussion  on  the  Campanulariidae;  and  through  him  to  the  Stichting 
Jan  Joost  ter  Pelkwijk  Fonds  for  financial  assistance  while  on  a  visit  both  to  the  Leiden 
Museum  and  the  Institut  voor  Taxonomische  Zoologie,  Zoologisch  Museum,  Universiteit 
van  Amsterdam,  where  Dr  R.  W.  M.  van  Soest  was  my  kind  host.  Dr  J.  M.  Paul,  Zoology 
Department,  University  of  Oxford,  helped  trace  literature  about  Laomedea  neglecta.  I  am 
grateful  also  to  Dr  R.  G.  Hughes,  Westfield  College,  University  of  London,  for  new  records  of 
Clytia  paulensis  and  to  Dr  R.  B.  Williams  of  Tring  for  helpful  discussions.  Dr  A.  G.  Long, 
Hancock  Museum,  Newcastle  upon  Tyne,  kindly  provided  facilities  while  I  worked  on  the 
Joshua  Alder  collection.  Lastly  I  am  grateful  to  Miss  L.  J.  Wadmore  for  preparing  the 
illustrations  of  medusae  and  to  R.  H.  Harris  for  histological  assitance.  Other  acknowledge- 
ments are  made  in  the  text. 


Addendum 

On  15  August  1981  I  found  Laomedea  angulata  (p.  98)  common  on  the  eel  grass  bed  at 
Studland,  Dorset.  The  previous  record  from  the  British  mainland  was  dated  1906;  and  from 
Dorset  was  1890,  also  at  Studland.  Whether  or  not  the  species  had  disappeared  from  that 
locality  in  the  meantime  might  be  hard  to  tell.  On  15  September  1981  I  found  the  same 
species  abundant  on  eel  grass  near  Misery  Point,  R  Yealm,  near  Plymouth.  At  this  locality 
too  the  status  of  the  species  during  the  last  several  decades  was  unclear. 


126  P.  F.  S.CORNELIUS 

References 

Agassiz,  L.  1862.  Hydroidae.  Contributions  to  the  natural  history  of  the  United  States  of  America. 

Second  Monograph,  4  :  1 8 1-380.  Boston. 
Alcock,  A.  1892.  A  case  of  commensalism  between  a  gymnoblastic  anthomedusoid  (Stylactis  minoi) 

and  a  scorpaenoid  fish  (Minous  inermis).  Ann.  Mag.  nat.  Hist.  (6)  10  :  207-2 14. 
Alder,  J.  1856a.  A  notice  of  some  new  genera  and  species  of  British  hydroid  zoophytes.  Ann.  Mag.  nat. 

Hist.  (2)  18  :  353-362. 

1 8566.  Descriptions  of  three  new  British  zoophytes.  Ann.  Mag.  nat.  Hist.  (2)18:  439-44 1 . 

1857.  A  catalogue  of  the  zoophytes  of  Northumberland  and  Durham.  Trans.  Tyneside  Nat.  Fid 

Cl.  3  :  93-162.  (Although  dated  1856  on  the  wrapper,  a  footnote  on  page  87  of  a  previous  paper  is 

dated  29  January,  1857.) 

1860.  Descriptions  of  a  zoophyte  and  two  species  of  Echinodermata  new  to  Britain.  Ann.  Mag. 


nat.  Hist.  (3)5:  73-75. 
Alexander,  W.  B.  1932.  The  natural  history  of  the  Firth  of  Tay.  Trans.  Proc.  Perthsh.  Soc.  nat.  Sci. 

9  :  35-42. 

Allman,  G.  J.  1859a.  Notes  on  the  hydroid  zoophytes.  Ann.  Mag.  nat.  Hist.  (3)4  :  137-144. 
18596.  Notes  on  the  hydroid  zoophytes.  Ann.  Mag.  nat.  Hist.  (3)4  :  367-370. 

1864a.  On  the  construction  and  limitation  of  genera  among  the  Hydroida.  Ann.  Mag.  nat.  Hist. 

(3)  13  :  345-380. 

18646.  Report  on  the  present  state  of  our  knowledge  of  the  reproductive  system  in  the  Hydroida. 

Rep.  Br.  Ass.  Advmt  Sci.  (1863)(1) :  351-426. 

1871.  A  monograph  of  the  gymnoblastic  or  tubularian  hydroids.  I.  The  Hydroida  in  general. 

London. 

1876a.  Descriptions  of  some  new  species  of  Hydroida  from  Kerguelen's  Island.  Ann.  Mag.  nat. 

Hist.  (4)  17:  113-115. 

1 8766.  Diagnoses  of  new  genera  and  species  of  Hydroida.  J.  Linn.  Soc.  (Zool.)  12:25 1-284. 

1887.  Report  on  the  Hydroida  collected  during  the  exploration  of  the  Gulf  Stream  by  L.  F.  de 

Pourtales,  Assistant,  United  States  Coast  Survey.  Mem.  Mus.  comp.  Zool.  Harv.  5  (2) :  1-66. 

1888.  Report  on  the  Hydroida  dredged  by  H.M.S.  Challenger  during  the  years  1873-1876.  Part 


II.  The  Tubularinae,  Corymorphinae,  Campanularinae,  Sertularinae,  and  Thalamophora.  Rept. 

sclent.  Results  Voy.  Challenger '23  :  l-90  +  i-lxix. 
Annandale,  N.  1915.  Fauna  of  the  Chilka  Lake.  The  coelenterates  of  the  lake,  with  an  account  of  the 

Actiniaria  of  brackish  water  in  the  Gangetic  delta.  Mem.  Indian  Mus.  5  :  65-1 14. 
Aria,  M.  N.  &  Brinckmann-Voss,  A.  1980.  Hydromedusae  of  British  Columbia  and  Puget  Sound.  Can. 

Bull.  Fish,  aquat.  Sci.  204  :  1-192,  i-viii. 
Arnaud,  F.,  Arnaud,  P.  M.,  Intes,  A.  &  Le  Loeuff,  P.  1976.  Transport  d'invertebres  benthiques  entre 

PAfrique  du  Sud  et  Sainte  Helene  par  les  laminaires  (Phaeophyceae).  Bull.  Mus.  Hist.  nat.  Paris  (3) 

384  (Ecol.  gen.  30) :  49-55. 
Babic,    K.    1912.    Dimorphismus   der  Gonangien   bei    Laomedea   angulata   Hincks.    Zool.   Anz. 

39  : 457-460. 
Bale,  W.  M.  1888.  On  some  new  and  rare  Hydroida  in  the  Australian  Museum  collection.  Proc.  Linn. 

Soc.  N.S.W.  (2)  3:  745-749. 

1914.  Further  notes  on  Australian  hydroids.  III.  Proc.  R.  Soc.  Viet.  (N.S.)  27  :  72-93. 

Bassindale,  R.  1941.  Studies  on  the  biology  of  the  Bristol  Channel.  IV.  The  invertebrate  fauna  of  the 

southern  shores  of  the  Bristol  Channel  and  Severn  Estuary.  Proc.  Bristol  Nat.  Soc.  (4)  9  :  143-201 . 
Baster,  J.  1762.  Opuscula  subseciva,  observationes  miscellaneas  de  animalculis  et  plantis.  Volume  I. 

Haarlem,  Holland. 
Bedot,  M.  1901.  Materiaux  pour  servir  a  1'histoire  des  hydroi'des.  Ire  periode.  Revue  suisse  Zool. 

9:379-515. 

1905.  Materiaux  pour  servir  a  Phistoire  des  hydroi'des.  2me  periode  (1821  a  1850).  Revue  suisse 

Zool.  13:  1-183. 

1910.  Materiaux  pour  servir  a  Phistoire  des  hydroi'des.  3me  periode  (1851  a  1871).  Revue  suisse 

Zool.  18:  189-490. 

1912.  Materiaux  pour  servir  a  Phistoire  des  hydroi'des.  4me  periode  (1872  a  1880).  Revue  suisse 

Zool.  20:213-469. 

1916.  Materiaux  pour  servir  a  Phistoire  des  hydroi'des.  5me  periode  (1881  a  1890).  Revue  suisse 


Zool.  24:  1-349. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  127 

—  1918.  Materiaux  pour  servir  a  1'histoire  des  hydroi'des.  6me  periode  (1819  a  1900).  Revue  suisse 
Zool.  26  (suppl.) :  1-376. 

1925.  Materiaux  pour  servir  a  1'histoire  des  hydroi'des.  7me  periode  (1901  a  1910).  Revue  suisse 


Zool.  32  (suppl.) :  1-657. 
Behner,  A.  1914.  Beitragzur  Kenntnisder  Hydromedusen.  Z.  wiss.  Zool.  Ill  :  381-427. 
Beloussov,  L.  V.  1961.  Vital  observations  on  cell  displacements  in  the  hydroid  polyp  Obelia  flexuosa. 

Dokl.  Akad.  Nauk.  SSSR.  136  : 1490-1 493. 
1973.  Growth  and  morphogenesis  of  some  marine  Hydrozoa  according  to  histological  data  and 

time-lapse  studies.  Publs  Seto.  mar.  biol.  Lab.  20  :  3 1 5-366. 
Beneden,  P.-J.  van  1843.  Memoire  sur  les  campanulaires  de  la  cote  d'Ostende,  considered  sous  le 

rapport    physiologique,    embryogenique    et    zoologique.    (Extrait).    Annls    Sci.    nat.    (Zool.)    (2) 

20  :  350-369. 
1844.   Memoire  sur  les  campanulaires  de   la  cote  d'Ostende,  consideres  sous  le  rapport 

physiologique,  embryogenique  et  zoologique.  Nouv.  Mem.  Acad.  R.  Sci.  Bruxelles,  17  (5) :  1-42. 
1866.  Recherches  sur  la  faune  littorale  de  Belgique  (Polypes).  Mem.  Acad.  r.  Sci.  Lett.  Belg. 


36(2):  1-207. 
Bergh,  R.  S.  1 879.  Studien  iiber  die  erste  Entwicklung  des  Eies  von  Gonothvraea  loveni  (Allm.).  Morph. 

Jb.  5:  22-61. 
Berrill,  N.  J.  1949.  The  polymorphic  transformations  of  Obelia.  Q.  Jl  microsc.  Sci.  90  :  235-264. 

1950.  Growth  and  form  in  calyptoblastic  hydroids.  II.  Polymorphism  within  the  Campanu- 

lariidae.  /.  Morph.  87  :  1-26. 

Betencourt,  A.  1888.  Les  hydraires  du  Pas-de-Calais.  Bull,  scient.  Fr.  Belg.  19  :  201-2 14. 
Billard,  A.  1902.  Les  hydroi'des  de  la  Bai  de  la  Hougue.  Bull.  Mus.  Hist.  nat.  Paris  8  :  53 1-536. 

1904a.  Hydroi'des  recoltes  par  M.  Ch.  Gravier  dans  le  Golfe  de  Tadjourah.  Bull.  Mus.  Hist.  nat. 

Paris  10:  480-485. 

1904/7.  Contribution  a  1'etude  des  hydroi'des.  (Multiplication,  regeneration,  greffes,  variations). 

Annls  Sci.  nat.  (Zool.)  (8)  20  :  1-25 1 . 

1906.  Mission  des  pecheries  de  la  cote  occidentale  d'Afrique.  III.  Hydroi'des.  Act.  Soc.  linn. 

Bordeaux,  56  : 69-76. 

1907.  Hydroi'des.  Exped.  scient.  Travailleur-  Talisman  8  :  153-243. 

1909.  Revision  des  especes  types  d'hydro'i'des  di  la  collection  Lamouroux  conservee  a  1'Institut 

Botanique  de  Caen.  Annls  Sci.  nat.  (Zool.)  (9)  9  :  307-336. 

1912.  Hydroi'des  de  Roscoff.  Archs  Zool.  exp.  gen.  51  :  459-478. 

1914.  Note  sur  les  hydroi'des  de  la  collection  Westendorp  du  Musee  Royal  d'histoire  naturelle  de 

Belgique.  Ann.  Soc.  r.  zool.  malac.  Belg.  48  :  27-30. 
1917.  Note  sur  quelques  especes  d'hydro'i'des  libres.  Bull.  Mus.  Hist.  nat.  Paris,  23  :  539-546. 

1918.  Notes  sur  quelques  especes  d'hydro'i'des  de  1'expedition  du  'Siboga'.  Archs  zool.  exp.  gen. 

51  (notes  et  revue  2) :  21-27. 

1922.  Note  sur  deux  especes  d'hydro'i'des  du  littoral  d'Ostende.  Ann.  Soc.  r.  zool.  malac.  Belg. 

52:  135-139. 

1926.  Rapport  sur  les  Hydroi'des  [of  the  Cambridge  Expedition  to  the  Suez  Canal].  Trans,  zool. 

Soc.Lond.  22:85-104. 

1927.  Les  hydroi'des  de  la  cote  Atlantique  de  France.  C.  r.  Congr.  Socs  sav.  Paris.  Sect.  Sci. 

(1926):  326-346. 

1928.  Clytia  johnstoni  Alder,  Campanularia  raridentata  Alder  et  Thaumantias  inconspicua 

Forbes.  Bull.  Mus.  nat.  Hist.  nat.  Paris  34  :  456^157. 

193  la.  Hydroi'des  de  Mauritanie.  Bull.  Mus.  nat.  Hist.  nat.  Paris  (2) 3  :  673-678. 

193  \b.  Hydroi'des  des  cotes  de  Syrie  (Missions  Gruvel).  Bibltque  Faune.  Colon.fr.  3  :  389-395. 

1934.  Note  sur  quelques  hydroi'des  du  Maroc.  Bull.  Soc.  zool.  France  59  :  227-231. 

1936.  Les  fonds  de  peche  pres  d'Alexandrie.  Hydroidea.  Notes  Mem.  Dir.  Rech.  Pech.,  Le  Caire 

13:  1-11. 

1938.  Note  sure  une  espece  de  campanularides  (Clytia  gravieri  Billard).  Bull.  Mus.  Hist.  nat. 


Paris  (2)  10: 429-432. 
Blanco,  O.  M.  1976a.  Estudio  critico  sobre  las  especies  del  genro  "Silicularia"  Meyen.  1834.  Revta 

Mus.  La  Plata,  N.S.  (Zool.)  9:21 7-241 . 
1967^.  Contribucion  al  conocimiento  de  los  hydrozoarios  Argentines.  Revta  Mus.  La  Plata,  N.S. 

(Zool.)  9:  243-297. 

Bohm,  R.  1878.  Helgolander  Leptomedusen.  Jena  Z.  Naturw.  12  :  68-203. 
Botten-Hansen,  P.  &  Petersen,  S.  1870.  Norsk  Bog-Fortegnelse.  1848-1865.  Oslo. 


128  P.  F.  S.  CORNELIUS 

Bougis,  P.  1963.  Cours  d'oceanographie  biologique.  Fascicule  1.  Hydromeduses.  Caracteres  generaux. 

classification.  Villefranche. 
Brattegard,  T.  1966.  The  natural  history  of  the  Hardangerfjord.  7.  Horizontal  distribution  of  the  fauna 

of  rocky  shores.  Sarsia,  22  :  1-54. 
Breemen,  P.  J.  Van  1905.  Plankton  van  Noord-  en  Zuiderzee.   Tijdschr.  ned.  dierk.   Vereen.  (2) 

9 : 145-324. 
Brinckmann-Voss,  A.  1970.  Anthomedusae/Athecatae  (Hydrozoa,  Cnidaria)  of  the  Mediterranean. 

Part  I.  Capitata.  Fauna  Flora  Golfo  Napoli,  39  :  1-96. 
Broch,  H.  1905.  Nordsee-Hydroiden  von  dem  norwegischen  Fischereidampfer  'Michael  Sars'  in  den 

Jahren  1903-1904  gesammelt,  nebst  Bemerkungen  iiber  die  Systematik  der  Tecaphoren  Hydroiden. 

Bergens  Mm.  Arb.  ( 1 905)  (6) :  1-26. 

1910.  Die  Hydroiden  der  arktischen  Meere.  Fauna  arct.  5  :  127-248. 

1912a.  Coelenteres  du  fond.  In  D uc  d 'Orleans  Campagne  Arctique  de  1907.  Brussels,  pp.  1-29. 

\9\2b.  Hydroidenuntersuchungen  III.  Vergleichende  Studien  an  adriatischen  Hydroiden.  K. 

norske  Vidensk.  Selsk.  Skr.  (1911)(l):l-65. 

1913.  Hydroida  from  the  "Michael  Sars"  North  Atlantic  deep-sea  expedition  1910.  Rep.  scient. 

Results  Michael  Sars  N.  Atlant.  deep  sea  Exped.  3(1):  1-18. 

1918.  Hydroida  (Part  II).  Dan.  Ingolf  Exped.  5  (7) :  1-206. 

1928.  Hydrozoa  I.  Tierwelt  N.-  u.  Ostsee,  3  (b) .  1-100. 

1929.  Hydroidenuntersuchungen  VI.  Studien  an  Silicularia  Meyen.  Meddr  zool.  Mus.,  Oslo 

(1929)20:  1-30. 

1933.  Zur  Kenntnis  der  adriatischen  Hydroidenfauna  von  Split.  Skr.  norske  VidenskAkad. 


mat.-nat.Kl.(\933)4:  1-115. 
Brooks,  W.  K.  1888«.  On  a  new  method  of  multiplication  in  hydroids.  Johns  Hopk.  Univ.  Circ. 

7  : 29-30. 
18886.  The  life  history  of  Epenthesis  mccradvi  (n.sp.).  Stud.  biol.  Lab.  Johns  Hopkins  Univ. 

4:  147-162. 
Browne,  E.  T.  1896.  On  British  hydroids  and  medusae.  Proc.  zool.  Soc.  Lond.  (1896) :  459-500. 

1897.  On  British  medusae.  Proc.  zool.  Soc.  Lond.  (1897) :  816-835. 

1900.  The  fauna  and  flora  of  Valencia  harbour  on  the  west  coast  of  Ireland.  Part  I.  The  pelgaic 

fauna.  II.  Report  on  the  medusae  (1 895-98).  Proc.  R.  Ir.  Acad.  (3)  5  :  694-736. 

1904.  Hydromedusae  with  a  revision  of  the  Williadae  and  Petasidae.  Fauna  Geogr.  Maldive 


Laccadive  Archipel.  2  :  722-749. 
Browne,  E.  T.  &  Vallentin,  R.  1904.  On  the  marine  fauna  of  the  Isles  of  Scilly  J.  R.  Instn  Cornwall, 

16:  120-132. 
Bruce,  J.  R.  1948.  Additions  to  faunal  records,  1941^6.  Rep.  mar.  Biol.  Stn  Port  Erin  (1945^7). 

58-60 : 39-58. 
Bruce,  J.  R.,  Colman,  J.  S.  &  Jones,  N.  S.  1963.  Marine  fauna  of  the  Isle  of  Man.  L.M.B.C.  Mem.  typ. 

Br.  mar.  PL  Anim.  36  :  i-ix  +  lr-307. 

Buchanan,  J.  B.  1957.  The  hydroid  fauna  of  the  Gold  Coast.  Revue  Zool.  Bot.  afr.  56  :  349-372. 
Calder,  D.  R.  1970.  Thecate  hydroids  from  the  shelf  waters  of  northern  Canada.  J.  Fish.  Res.  Bd, 

Canada  27  :  1501-1547. 
1975.  Biotic  census  of  Cape  Cod  Bay :  hydroids.  Biol.  Bull.  mar.  biol.  Lab.,   Woods  Hole, 

149:287-315. 

1976.  The  zonation  of  hydroids  along  salinity  gradients  in  South  Carolina  estuaries.  In  Mackie, 


G.  O.  (editor),  Coelenterate  ecology  and  behavior  (Third  international  symposium  on  coelenterate 

biology),  New  York.  Pp.  165-174. 

Calkins,  G.  N.  1 899.  Some  hydroids  from  Puget  Sound.  Proc.  Boston  Soc.  nat.  Hist.  28  :  333-367. 
Carus,  J.  V.  1884-1893.  Prodromus  faunae  mediterraneae.  Stuttgart.  2  volumes.  (Vol.  1,  pars  I, 

Colenterata,  Echinodermata,  Vermes,  1884;  pp.  1-282.) 
Chas  Brinquez,  J.  C.  &  Rodriguez  Babio,  C.    1977.  Fauna  marina  de  Galicia.  Contribucion  al 

conocimiento  de  los  hidropolipos  del  literal  gallego.  Monogr.   Univ.  Santiago  de  Compostela, 

39:  1^3. 
Chopin,  A.  1894.  A  visit  toCumbrae:  The  veteran  naturalist:  The  Millport  Biological  station.  Trans,  a. 

Rep.  Manchr  micr.  Soc.  (1 894) :  43-54. 

Christiansen,  B.  O.  1972.  The  hydroid  fauna  of  the  Oslo  Fiord  in  Norway.  Norw.  J.  Zool.  20  :  279-3 10. 
Chumley,  J.  1918.  The  fauna  of  the  Clyde  Sea  area,  being  an  attempt  to  record  the  zoological  results 

obtained  b  y  the  late  Sir  John  Murra  v  and  his  assistants  on  board  the  S.  Y.  ' 'Medusa ' '  during  the  rears 

1884  to  1892.  Glasgow. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  129 

Clarke,  S.  F.  1875.  Descriptions  of  new  and  rare  species  of  hydroids  from  the  New  England  Coast. 

Trans.  Conn.  Acad.  Arts  Sci.  3  :  58-66. 
1876.  The  hydroids  of  the  Pacific  coast  of  the  United  States,  south  of  Vancouver  Island.  With  a 

report  upon  those  in  the  Museum  of  Yale  College.  Trans.  Conn.  Acad.  Arts  Sci.  3  :  249-264. 
1877.  Report  on  the  hydroids  collected  on  the  coast  of  Alaska  and  the  Aleutian  Islands,  by  W.  H. 

Dall,  U.S.  Coast  Survey,  and  party,  from  1871  to  1874  inclusive.  Proc.  Acad.  nat.  Sci.  Philad.  (1876) 

28  :  209-238.  (A  note  on  p.  ii  of  the  volume  states  unequivocally  that  the  publication  date  of  this 

paper  was  2  January,  1877.) 
1879.  Report  on  the  Hydroida  collected  during  the  exploration  of  the  Gulf  Stream  and  Gulf  of 

Mexico  by  Alexander  Agassiz,  1877-78.  Bull.  Mus.  comp.  Zool.  Harv.  5  :  239-252. 

1907.  Reports  on  the  scientific  results  of  the  expedition  to  the  eastern  tropical  Pacific,  in  charge  of 


Alexander  Agassiz,  by  the  U.S.  Fish  Commission  steamer  'Albatross',  from  October,  1904,  to  March, 

1905,  Lieut. -Commander  L.  M.  Garrett,  U.S.N.,  commanding.  VIII.  The  hydroids.  Mem.  Mus. 

comp.  Zool.  Harv.  35  (1) :  1-18. 

Congdon,  E.  D.  1907.  The  hydroids  of  Bermuda.  Proc.  Am.  Acad.  Arts  Sci.  42  :  461^85. 
Cornelius,  P.  F.  S.  1975a.  The  hydroid  species  ofObelia  (Coelenterata,  Hydrozoa:  Campanulariidae), 

with  notes  on  the  medusa  stage.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  28  :  249-293. 
19756.  A  revision  of  the  species  of  Lafoeidae  and  Haleciidae  (Coelenterata:  Hydroida)  recorded 

from  Britain  and  nearby  seas.  Bull.  Br.  Mus.  nat.  hist.  (Zool.)  28  :  373^26. 
1977 a.  The  linking  of  polyp  and  medusa  stages  in  Obelia  and  other  coelenterates.  Biol.  J.  Linn. 

Soc.  9  : 45-57. 

1977 b.  The  gradual  discovery  of  the  relation  between  medusa  and  polyp  in  Obelia  and  other 


coelenterates.  Microscopy  53  '.  172-174. 

—  1978.  The  genus  names  Calicella  Hincks  and  Calvcella  Hincks  (Coelenterata:  Hydrozoa).  Bull.  Br. 
Mus.  nat.  Hist.  (Zool.)  33  :  233-234. 

—  1979.  A  revision  of  the  species  of  Sertulariidae  (Coelenterata:  Hydroida)  recorded  from  Britain 
and  nearby  seas.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  34  :  243-32 1 . 

1981.  Clytia  Lamouroux,  1812,  Laomedea  Lamouroux,  1812,  and  Campanularia  Lamarck, 


1816  (Coelenterata,  Hydroida):  proposed  designations  of  type  species  by  use  of  the  plenary  powers, 
and  comments  on  related  genera.  Z.N.(S.)  2326.  Bull.  zool.  Norn.  38  :  208-220. 
—  (unpublished).  Notes  on  the  BMNH  copy  of  a  rare  work  by  Westendorp,  1853.  (MS  in  BMNH 
library.) 

&  Garfath,  J.  B.  1980.  The  coelenterate  taxa  of  Joshua  Alder.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


39:273-291. 

Couch,  R.  Q.    1842.  An  essay  on  the  zoophytes  of  Cornwall.  Rep.  R.  Cornwall  polvtech.  Soc. 
(1840:27-91. 

1844.  A  Cornish  fauna;  being  a  compendium  of  the  natural  history  of  the  county.  Part  3, 

containing  the  zoophytes  and  calcareous  corallines.  Truro. 
Coughtrey,  M.    1875.   Notes  on  the  New  Zealand  Hydroideae.    Trans.   Proc.   N.Z.   Inst.   (1874) 

7:281-293. 

Crawford,  J.  H.  1895.  The  hydroids  of  St.  Andrews  Bay.  Ann.  Mag.  nat.  Hist.  (6)  16  :  256-262. 
Crawshay,  L.  R.  1912.  On  the  fauna  of  the  outer  western  area  of  the  English  Channel.  J.  mar.  biol.  Ass. 

U.K.  9  : 292-393. 
Crothers,  J.  H.  (Ed.).  1966.  Dale  Fort  marine  fauna.  Second  edition.  Fid  Std.  2  (suppl.) :  i-xviv  + 

1-169. 
Crowell,  S.  1957.  Differential  responses  of  growth  zones  to  nutritive  level,  age,  and  temperature  in  the 

colonial  hydroid  Campanularia.  J.  exp.  Biol.  134  :  63-90. 
1961.  Developmental  problems  in  Campanularia.  In  Lenhoff,  H.  M.  &  Loomis,  W.  F.  (editors) 

The  biology  of  Hydra  and  of  some  other  coelenterates:  1961  Miami,  pp.  297-3 16. 

&  Wyttenbach,  C.  R.  1957.  Factors  affecting  terminal  growth  in  the  hydroid  Campanularia.  Biol. 


Bull.  mar.  biol.  Lab.,  Woods  Hole  113  :  233-244. 
Cunha,  A.  X.  da  1944.  Hidropolipos  das  costas  de  Portugal.  Mems  Estud.  Mus.  zool.  Univ.  Coimbra 

161  :  1-101. 
1950.  Nova  contribuicjio  para  o  estudo  dos  hidropolipos  das  costas  de  Portugal.  Archos  Mus. 

Bocage,2\  :  121-144. 
Cuvier,  G.  L.  C.  F.  D.  1797.  Tableau  elementaire  de  I'histoire  naturelle  des  animaux.  Paris.  (Dating 

follows  J.  Typogr.  Biblphique(\191)  1  :  81-82.) 
Dana,  J.  D.  1846,  1849.  United  States  Exploring  Expedition  during  the  years  1838,  1839,  1840,  1841, 


130  P.  F.S.CORNELIUS 

1842.  Under  the  command  of  Charles  Wilkes,  U.S.N.  Zoophytes.  Philadelphia.  Vol.  1,  text  (1846); 

vol.  2,  atlas  ( 1 849).  (Dating  after  Haskell,  1 942.) 
Darwin,  C.  R.  1 860.  Journal  of  researches  into  the  natural  history  and  geology  of  the  countries  visited 

during  the  voyage  of  H. M.S.  Beagle  round  the  world,  under  the  command  ofCapt.  Fitz  Roy,  R.N.  2nd 

edition.  London. 
Debouteville,  C.  D.  &  Nunes,  L.  P.  1951.  Hydraires  epizoi'ques  sur  les  copepodes  parasites.  Vie  Milieu 

2:42\-432. 

1952.  Hydraires  epizoi'ques  sur  les  copepodes  parasites.  Archos  Mus.  Bocage  23:  1-11. 

Deevey,  E.  S.  1954.  Hydroids  of  the  Gulf  of  Mexico.  In  Galtsoff,  P.  S.,  Gulf  of  Mexico.  Its  origin, 

waters,  and  marine  life.  Fishery  Bull.  Fish.  Wildl.  Serv.  U.S.  55  :  267-272. 
den  Hartog,  C.  1970.  The  sea-grasses  of  the  world.  Verh.  K.  ned.  Akad.  Wet.  59  :  1-275. 
Desor,  M.  E.  1849.  Lettre  sur  la  generation  medusipare  des  polypes  hydraires.  Annls  Sci.  nat.  (3) 

12:204-217. 
Du  Plessis,  G.  1871.  Evolution  medusipare  de  Clvtia  (Campanularia)  volubilis.  Bull.  Soc.  vaud.  Sci. 

nat.ll  :  167-170. 
Edwards,  C.  1973.  The  medusa  Modeeria  rotunda  and  its  hydroid  Stegopoma  fastigiatum,  with  a 

review  of  Stegopoma  and  Stegolaria.  J.  mar.  biol.  Ass.  U.K.  53  :  573-600. 
Ehrenberg,  C.  G.  1834#.  Beitra'ge  zur  physiologischen  Kenntniss  der  Corallenthiere  im  allgemeinen, 

und  besonders  des  rothen  Meeres,  nebst  einem  Versuche  zur  physiologischen  Systematik  derselben. 

Phys.  Math.  Abh.  K.  Akad.  Wiss.  Berlin  (1832)  1  :  225-380.  (Reprinted  as  Ehrenberg,  18346.) 
18346.   Die  Corallenthiere  des  rothen  Meeres,  physiologisch   untersucht  und  systematisch 

verzeichnet.  Berlin.  (Reprint  of  Ehrenberg,  1 843a,  in  book  form,  paginated  1-1 56.) 
Ellis,  J.  1 755.  An  essay  towards  a  natural  history  of  the  corallines,  and  other  marine  productions  of  the 

like  kind,  commonly  found  on  the  coasts  of  Great  Britain  and  Ireland.   To  which  is  added  the 

description  of  a  large  marine  polype  taken  near  the  North  Pole  by  the  whale-fishers,  in  the  summer 

1753.  London. 
1756a.  'Account  of  the  vesicles  on  corallines.'  [Running  title  at  top  of  page.]  Gentleman's  Mag. 

26  : 288-290. 

17566.  Essai  sur  I'histoire  naturelle  des  corallines,  et  d'autres  productions  marines  du  meme 


genre,  qu'on  trouve  communement  sur  les  cotes  de  la  Grande- Bretagne  et  d'Irlande;  auquel  on  a 
joint  une  description  d'un  grand  polype  de  mer,  pris  aupres  du  Pole  Arctique,  par  des  pecheurs  de 
baleine,  pendant  I'ete  de  1 753.  The  Hague.  Translated  by  P.  de  Hondt. 

1756c.  Natuurlyke  historic  van  de  koraal-gewassen,  en  andere  dergelyke  zee-lighamen,  die  men 


gemeenelyk  vind  op  de  kusten  van  Groot-Brittanien  en  lerland:  benevens  eene  beschryving  van  een 
grooten  zee-polyp,  in  den  zomer  van  't  jaar  1753  by  den  Noord-Pool  door  de  walvis-vangers 
gevonden.  The  Hague.  Translated  by  J.  Tak. 

—  1767.   Versuch  einer  Natur-Geschichte  der  Corall-Arten  und  anderer  dergleichen  Mer-Korper, 
welche  gemeiniglich  an  den  Kusten  von  Gro/3-Britanien  und  Inland  gefunden  werden;  nebst  der 
Beschreibung  eines  grossen  Buschel-Polypen,  welcher  in  dem  Eis-Mere  gefangen  worden.  Aus  dem 
Englischen  und  Franzosischen  ubersezt,  und  mil  Anmerkungen,  auch  einem  Anhange  fiinf  hieher 
gehoriger  Abhandlungen  der  Herren  Schlosser,  Baster  und  Ellis  begleitet  von  D.  J.  G.  Kriiniz. 
Niirnberg.  Translated  by  D.  J.  G.  Kriiniz. 

—  1 768.  An  account  of  the  Actinia  sociata,  or  clustered  animal-flower,  lately  found  on  the  sea-coasts 
of  new-ceded  Islands.  Phil.  Trans.  R.  Soc.  (1 767)  57  :  428^37. 

&  Solander,  D.  C.  1 786.  The  natural  history  of  many  curious  and  uncommon  zoophytes,  collected 


from  various  parts  of  the  globe.  London.  Edited  by  M.  Watt. 
Elmhirst,  R.  1925.  Lunar  periodicity  in  Obelia.  Nature,  Lond.  116  :  358-359. 
Eschscholtz,  F.  1829.  System  der  Acalephen.  Berlin. 
Esper,  E.  J.  C.  71810.  (There  is  strong  evidence  that  some  of  the  plates  of  Esper  (1829)  were  published 

in  1810,  but  I  have  not  traced  a  perfect  copy.  See  note  2  (p.  1 24)  and  next  reference.) 
1829.  Die  Pflanzen-Thiere.  Volume  3,  pp.  145-283.  Edited  by  F.  L.  Hammer.  Niirnberg.  (See 

also  previous  reference.) 
Evans,  F.  O.  1978.  The  marine  fauna  of  the  Cullercoats  district.  6.  Coelenterata  and  Ctenophora.  Rep. 

Dove  mar.  Lab.  (3)  19  :  1-165. 
Faulkner,  G.  H.  1929.  The  early  prophases  of  the  first  oocyte  division  as  seen  in  life,  in  Obelia 

geniculata.  Q.  Jl  microsc.  Sci.  73  :  225-242. 

Faure,  C.  1 965.  Sur  la  distinction  des  deux  especes  de  Laomedea.  Cah.  Biol.  mar.  6:41 9-426. 
Fey,  A.  1969.  Peuplements  sessiles  de  1'archipel  de  Glenan.  I.  Inventaire :  hydraires.  Vie  Milieu  (B) 

20:387-413. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  131 

Fleming,  J.  1 820.  Observations  on  the  natural  history  of  the  Sertularia  gelatinosa  of  Pallas.  Edinb.  phil. 

J.  2  :  82-89. 

1828.  A  history  of  British  animals.  Edinburgh. 

Forbes,  E.  1848.  A  monograph  of  the  British  naked-eyed  medusae:  with  figures  of  all  the  species. 

London. 
Fraipont,  J.  1880.  Recherches  sur  1'organisation  histologique  et  le  developpement  de  la  Campanularia 

angulata.  Contribution  a  1'histoire  de  1'origine  du  testicule  et  de  1'ovaire.  Archs  Zoo/,  exp.  gen. 

8 : 433-466. 
Fraser,  C.  M.  1913.  Hydroids  from  Nova  Scotia.  Bull.  Victoria  meml  Mus.  1  :  157-186. 

1918.  Hydroids  of  eastern  Canada.  Rep.  Dept.  Naval  Serv.  Ottawal  (suppl.) :  329-367. 

1 944.  Hydroids  of  the  Atlantic  coast  of  North  America.  Toronto. 

1946.  Distribution  and  relationship  in  American  hydroids.  Toronto. 

Freycinet,  L.  de  (editor)  1 824.  Voyage  autour  du  monde  entrepris  par  ordre  du  Roi.  Zoologie  [by  Quoy, 

J.  R.  C.  &  Gaimard,  J.  P.].  Paris.  (The  hydroid  descriptions  were  written  by  J.  V.  F.  Lamouroux.) 
Garcia  Corralles,  P.,  Inchaurbe,  A.  A.  &  Mora,  D.  G.  1978.  Contribucion  al  conocimiento  de  los 

hidrozoos  de  las  costas  espanolas.  Parte  I  :  Halecidos,  campanularidos  y  plumularidos.  Boln  Inst. 

esp.  Oceanogr.  4  (253) :  3-73. 
Gegenbaur,  C.  1854.  Zur  Lehre  vom  Generationswechsel  und  der  Fortpflanzung  bei  Medusen  und 

Polvpen.  Wiirzburg.  (Paginated  1-68;  reprint  of  article  with  same  title  in  Verh.  phvs.-med.  Ges. 

Wurzb.4:  154-221.) 

1856.  Versuch  eines  Systemes  der  Medusen,  mil  Beschreibung  neuer  oder  wenig  gekannter 

Formen;  zugleich  ein  Beitrag  zur  Kenntniss  der  Fauna  des  Mittelmeeres.  Z.  wiss.  Zoo/.  (1857) 
8 : 202-273. 

Giard,  A.  1898.  Sur  1'ethologie  du  Campanularia  caliculata  Hincks.  (Stolonisation  et  allogonie).  C.  r. 

Seanc.  Soc.  Biol.  ( 10)  5  :  1 7-20. 

Godeaux,  J.  1941.  Sur  la  croissance  d'un  hydrozoaire.  C.  r.  Ass.  franc..  Avanc.  Sci.  63(1939) :  707-710. 
Goette,  A.  1907.  Vergleichende  Entwicklungsgeschichte  der  Geschlechtsindividuen  der  Hydropolypen. 

Z.  wiss.  Zoo/.  87  :  1-335. 
Gosse,  P.  H.  1853.  A  naturalist's  rambles  on  the  Devonshire  coast.  London. 

1855-1856.  A  manual  of  marine  zoology  for  the  British  Isles.  London.  2  volumes.  (Volume  1, 

'Zoophytes',  1855,  is  cited  here.) 

Gow,  C.  &  Millard,  N.  A.  H.  1975.  Two  new  species  of  campanularian  hydroids  from  South  Africa. 

Ann.  S.  Afr.  Mus.  67  :  1-6. 
Goy,  J.  1980.  Lesmedusesde  Francois  Peron  et  Charles-Alexandre  Lesueur(  1775-1 810  et  1778-1846) 

revelees  par  les  velins  de  Lesueur.  Bull,  trimest.  Soc.  geol.  Normandie  Amis  Mus.  du  Havre 

67  (2):  65-78,  pis  1-27. 
Gravier-Bonnet,  N.  1979.  Hydraires  semi-profonds  de  Madagascar,  (Coelenterata  Hydrozoa),  etude 

systematique  et  ecologique.  Zoo/.  Vehr.  Leiden  169  :  1-76. 
Gray,  J.  E.  1848.  List  of  the  specimens  of  British  animals  in  the  collection  of  the  British  Museum.  Part 

1.  Cenlroniae  or  radiated  animals.  London. 
Gronovius,  L.-T.  1760.  Observationes  de  animalculus  aliquot  marinae  aquae  innatantibus  atque  in 

littoribus  Belgicis  obviis.  Acta  helv.  4  :  35-40.  [The  author's  name  does  not  appear  in  the  paper, 

which  was  ascribed  to  L.-T.  Gronovius  by  Bedot  ( 1 90 1  :  396).] 
Gudger,  E.  W.  1928.  Association  between  sessile  colonial  hydroids  and  fish.  Ann.  Mag.  nat.  Hist.  (10) 

1  :  17-48. 

1937.   A  glut   herring,   Pomolobus  aestivalis,   with   an   attached  colonial   hydroid,   Obelia 

commensuralis.  Am.  Mus.  Novit.  945  :  1-6. 

Haeckel,  E.  1879.  Das  System  der  Medusen.  Erster  Theil  einer  Monographie  der  Medusen.  Denkschr. 

med.-naturw.  Ges.  Jena,  1  :  1-360. 
Hale,  L.  J.  1964.  Cell  movements,  cell  division  and  growth  in  the  hydroid  Clvtia  johnstoni.  J.  Embryol. 

exp.Morph.  12:517-538. 

1973a.  The  pattern  of  growth  of  Clytia  johnstoni.  J.  Embryol.  exp.  Morph.  29  :  283-309. 

19736.  Morphogenetic  properties  of  the  parts  of  the  colony  of  Clvtia  johnstoni.  J.  Embryol.  exp. 

Morph.  30  :  773-789. 
Hammett,  F.  S.  1943.  The  role  of  the  amino  acid  and  nucleic  acid  components  in  developmental 

growth.  Part  one.  The  growth  of  an  Obelia  hydranth.  Chapter  one.  Description  of  Obelia  and  its 

growth.  Growth  7  :  33 1-339. 

Hamond,  R.  1957.  Notes  on  the  Hydrozoa  of  the  Norfolk  coast.  J.  Linn.  Soc.  (Zool.)  43  :  294-324. 
1 963.  Further  notes  on  the  Hydrozoa  of  the  Norfolk  coast.  Ann.  Mag.  nat.  Hist.  (13)6  :  659-670. 


132  P.  F.  S.  CORNELIUS 

&  Williams,  R.  B.  1977.  The  Ctenophora,  Scyphozoa  and  Anthozoa  of  Norfolk,  with  additional 

notes  on  the  Hydrozoa.  Trans.  Norfolk  Norwich  Nat.  Soc.  24  :  58-74. 
Hargitt,  C.  W.  1909.  New  and  little  known  hydroids  of  Woods  Hole.  Biol.  Bull.  mar.  biol.  Lab.,  Woods 

Hole,  17  :  369-385. 

—  1924.  Hydroids  of  the  Philippine  Islands.  Philipp.  J.  Sci.  24  :  467-507. 
Hartlaub,  C.  1897.  Die  Hydromedusen  helgolands.  Zweiter  Bericht.  Wiss.  Meeresunters.  Helgol.  (N.F.) 

2  (Heft  1):  449-536. 
1901.  Hydroiden  aus  dem  Stillen  Ocean.  Ergebnisse  einer  Reise  nach  dem  Pacific  (Schauinsland 

1896-97).  Zoo/.  Jb.  (Syst.)  14  :  349-379. 

1905.  Die  Hydroiden  der  magelhaensischen  Region  und  chilenischen  Kiiste.  Zoo/.  Jb.  (suppl. 


6): 497-7 14. 
Haskell,  D.  C.   1942.   The  United  States  Exploring  Expedition,  1838-1842,  and  its  publications 

1844-1874.  New  York. 
Heath,  H.  1910.  The  association  of  a  fish  with  a  hydroid.  Biol.  Bull.  mar.  biol.  Lab.,  Woods  Hole, 

19 :  73-78. 

Heller,  C.  1868.  Die  Zoophvten  und  Echinodermen  des  adriatischen  Meeres.  Vienna. 
Hickson,  S.  J.  &  Gravely,  F.  H.  1907.  Hydroid  zoophytes.  Nat.  Antarct.  Exped.  1901-1904  Nat.  Hist.  3 

(Colenterata  II) :  1-34. 
Hincks,  T.  1852.  Notes  on  the  reproduction  of  the  Campanulariadae;  with  a  description  of  a  new 

species  ofLaomedea.  Ann.  Mag.  nat.  Hist.  (2)  10  :  8 1-87. 
1853.  Further  notes  on  British  zoophytes,  with  descriptions  of  new  species.  Ann.  Mag.  nat.  Hist. 

(2)11:  178-185. 

1861fl.  On  new  Australian  Hydrozoa.  Ann.  Mag.  nat.  Hist.  (3)7  :  279-281. 

1861&-1862.  A  catalogue  of  the  zoophytes  of  south  Devon  and  south  Cornwall.  Ann.  Mag.  nat. 

Hist.  (3)  8  (1861):  152-161,251-262,  290-297,  360-366;  9  (1862):  22-30. 
1863.  On  some  new  British  hydroids.  Ann.  Mag.  nat.  Hist.  (3)  11  :  45-47  [plus  pi.  9  from  volume 

10,1862]. 

1866.  On  new  British  Hydroida.  Ann.  Mag.  nat.  Hist.  (3)  18  :  269-299. 

1868.  A  history  of  the  British  hydroid  zoophytes.  2  volumes.  London. 

1871.  Supplement  to  a  "Catalogue  of  the  zoophytes  of  south  Devon  and  South  Cornwall,"  with 

descriptions  of  new  species.  Ann.  Mag.  nat  Hist.  (4)8  :  73-83. 

1872.  Contributions  to  the  history  of  the  Hydroida.  Ann.  Mag.  nat.  Hist.  (4)  10  :  385-395. 

1889.  On  the  Polyzoa  and  Hydroida  of  the  Mergui  Archipelago  collected  for  the  Trustees  of  the 


Indian  Museum,  Calcutta,  by  Dr  J.  Anderson,  F.R.S.,  Superintendent  of  the  Museum.  J.  Linn.  Soc. 

21  :  121-135. 
Hirohito.  1969.  Some  hydroids  of  the  Amakusa  Islands.  Publ.  Biol.  Lab.  Imp.  Household,  Tokyo 

(1969)  (9):  1-32. 
1977.  Five  hydroid  species  from  the  Gulf  of  Aqaba,  Red  Sea.  Publ.  Biol.  Lab.  Imp.  Household, 

Tokyo (1977) (11):  1-26. 
Hoeven,    J.    van    der,    1862.    Eenige    aanteekeningen    over    Martinus    Slabber's    Natuurkundige 

Verlustigingen;  benevens  opgave  der  systematische  namen  van  de  daarin  afgebeelde  diersoorten. 

Versl.  Meded.  K.  Akad.  wet.  Amst.  (Afd.  Nat.)  14  :  270-285. 
Houttuyn,  M.  1770.  Natuurlyke  Historic  ofuitvoerige  beschryving  der  Dieren,  Planten  en  Mineraalen, 

volgens  het  Samenstel  van  den  Heer  Linnaeus.  Eerste  Deels,  veertiende  Stuk.  Amsterdam. 
Houvenaghel-Crevecoeur,  N.  1973.  Sur  le  mode  de  fixation  de  la  planule  de  H ydrall mania  falcata  L. 

(hydroi'de  thecate,  Sertulariidae).  C.  r.  hebd.  Seanc.  Acad.  Sci.,  Paris  (D)  276  :  28 1 3-28 1 5. 
Hughes,  R.  G.  1975.  The  distribution  of  epizoites  on  the  hydroid  Nemertesia  antennina  (L.).  J.  mar. 

biol.  Ass.  U.K.  55  :  275-294. 

1980.  Current  induced  variations  in  the  growth  and  morphology  of  hydroids.  In  Tardent,  P.  & 

Tardent,  R.  (eds)  Developmental  and  cellular  biology  ofcoelenterates.  Amsterdam;  pp.  179-184. 

Hummelinck,  P.  W.  1930.  Beitragezur  Kenntnis  hollandischer  Hydroiden.  I.  Bemerkungen  iibereinige 
Campanuliniden  und  Campanulariiden  vom  Vangdam  und  Nieuwediep.  Tijdschr.  ned.  dierk. 
Vereen.  (3)2(1) :  28^12. 

1936.  Hydropoliepen.  Flora  Fauna  Zuiderzee  (Suppl.) :  41-64. 

Jaderholm,  E.  1904a.  Mitteilungen  ueber  einige  von  der  Schwedischen  Antartic-Expedition 
1901-1903  eingesammelte  Hydroiden.  ArchsZool.  exp.  gen.  (4)  3  (Notes  et  revue) :  I-XIV. 

1904/?.  Aussereuropaische  Hydroiden   im  schwedischen  Reichsmuseum.  Ark.   Zoo/.  (1903) 

1  : 259-3 12. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  133 

1909.  Northern  and  arctic  invertebrates  in  the  collection  of  the  Swedish  State  Museum.  IV. 

Hydroiden.  K.  svenska  VetenskAkad.  Handl.  45  (1) :  1-124. 

Jagerskiold,  L.  A.  1971.  A  survey  of  the  marine  bethonic  macro-fauna  along  the  Swedish  west  coast 

1921-1938.  Ada R.  Soc.  sclent,  litt.  gothoburg.  (Zool.)6  :  1-146. 
Johnston,  G.   1836.  A  catalogue  of  the  zoophytes  of  Berwickshire.  Hist.  Berwicksh.  Nat.  Club, 

1  :  107-108.  (Dating  follows  a  manuscript  note  by  C.  D.  Sherborn  in  the  BMNH  copy,  drawing 

attention  to  the  inclusion  of  the  annual  Presidential  Addresses  in  the  journal.  That  in  the  part 

including  pp.  107-108  was  delivered  in  September,  1836;  but  there  is  no  further  proof  that  the  part 

was  issued  before  1 837.) 

1 838.  A  history  of  the  British  zoophytes.  Dublin,  Edinburgh  &  London. 

1 847.  A  history  of  the  British  zoophytes.  2nd  edition.  2  volumes.  London. 

Kawaguti,  S.  1966.  Electron  microscopy  on  the  Hydrozoa,  Orthopyx'^  caliculata.  Biol.  J.  Okayama 

Univ.  12:93-103. 
Keferstein,  W.  &  Ehlers,  E.  1861.  Zoologische  Beitrdge  gesammelt  im  Winter  1859/60  in  Neapel  und 

Messina.  Leipzig. 
Keller,  C.    1883.   Untersuchungen  iiber  neue  Medusen  aus  dem  rothen  Meere.   Z.   wiss.   Zool. 

38:621-670. 
Knight,  D.  P.  1965.  Behavioural  aspects  of  emergence  in  the  hydroid  of  Campanularia  jlexuosa 

(Hincks).  Nature,  Lond.  206  :  1 170-1 171. 

1970.  Sclerotization  of  the  perisarc  of  the  calyptoblastic  hydroid,  Laomedea  jlexuosa.  \.  The 

identification  and  localization  of  dopamine  in  the  hydroid.  Tissue  Cell 2  :  467-477. 

1971.  Sclerotization  of  the  perisarc  of  the  calyptoblastic  hydroid,  Laomedea  Jlexuosa.  2. 


Histochemical  demonstration  of  phenol  oxidase  and  attempted  demonstration  of  peroxidase.  Tissue 

Cell  3:  57-64. 
Knight-Jones,  E.  W.  &  Jones,  W.  C.  1956.  The  fauna  of  rocks  at  various  depths  off  Bardsey.  Rep. 

Bardsey  Birds  Fid  Obs.  (1955)3:  23-30. 
Kramp,  P.  L.  1919.  Medusae.  Part  I.  Leptomedusae.  Dan.  IngolfExped.  5  (8) :  1-1 12. 

1929.  Marine  Hydrozoa.  Zoology  Faroes,  1(1  :  5) :  1-59. 

1933.  Occasional  notes  on  Coelenterata.  II.  Vidensk.  Meddrdansk.  naturh.  Foren.  94  :  237-247. 

1935.   Polypdyr  (Coelenterata).   I.   Ferskvandspolypper  og  Goplepolypper.   Danm.   Fauna, 

41  :  1-208. 

1938.  Marine  Hydrozoa.  a.  Hydroida.  Zoology  Iceland,  2  (5a) :  1-82. 

1943.  The  zoology  of  east  Greenland.  Hydroida.  Meddr.  Gronland,  121  (11):  1-52. 

1955.  The  medusae  of  the  tropical  west  coast  of  Africa.  Atlantide  Rep.  5  :  239-324. 

1957.  Some  Mediterranean  hydromedusae  collected  by  A.  K.  Totton  in  1954  and  1956.  Vidensk. 


Meddrdansk.  naturh.  Foren.  119  :  1 15-128. 

—  1959.  The  hydromedusae  of  the  Atlantic  Ocean  and  adjacent  waters.  Dana  Rep.  46  :  1-283. 

—  196 1 .  Synopsis  of  the  medusae  of  the  world.  J.  mar.  biol.  Ass.  U.K.  40  :  1-469. 

1965.  The  hydromedusae  of  the  Pacific  and  Indian  Oceans.  [Section  1.]  Dana  Rep.  63  :  1-162. 


Kruse,  D.  N.   1959.  Parasites  of  the  commercial  shrimps,  Pennaeus  aztecus  Ives,  P.  duorarum 

Burkenroad  and  P.  setiferus  (Linnaeus).  Tulane  Stud.  Zool.  7  :  123-144. 
Kubota,  S.  1978.  The  life-history  of  Clytia  edwardsi  (Hydrozoa;  Campanulariidae)  in  Hokkaido, 

Japan.  J.  Fac.  Sci.  Hokkaido  Univ.  (6)  21:31 7-354. 
Ktthl,    H.    1967.    Die    Hydromedusen   der   Emsmiindung.    Veroff.    Inst.    Meeresforsch.    Bremerh. 

10  : 239-246. 
Kuhn,  A.  1913.  Entwicklungsgeschichte  und  Verwandtschaftsbeziehungen  der  Hydrozen.  I  Teil  :  Die 

Hydroiden.  Ergebn.  Fortschr.  Zool.  4(1-2) :  1-284. 

Lamarck,  J.  B.  P.  A.  de  1 8 1 6.  Histoire  naturelle  des  animaux sans  vertebres.  Volume  2.  Paris. 
1836.  Histoire  naturelle  des  animaux  sans  vertebres.  2nd  edition,  edited  by  Deshayes,  G.  P.  & 

Edwards,  H.  M.  Volume  2,  Histoire  des  polypes. 
Lamouroux,  J.  V.  F.  1812.  Extrait  d'un  memoire  sur  la  classification  des  polypiers  coralligenes  non 

entierement  pierreux.  Nouv.  Bull.  Sci.  Soc.  philom.  Paris,  3  :  181-188. 

1816.  Histoire  des  polypiers  coralligenes  flexibles,  vulgairement  nommes  zoophytes.  Caen. 

1 82 1 .  Exposition  methodique  des  genres  de  I'ordre  des  polypiers.  Paris. 

,  Bory  de  Saint-Vincent,  J.  B.  G.  M.  &  Deslongchamps,  E.  1824,  Histoire  naturelle  des  zoophytes, 

ou  animaux  rayonnes,  faisant  suite  a  I'histoire  naturelle  des  vers,  de  Bruguiere.  In:  Encyclopedic 

methodique  (suppl.).  Paris. 
Lang,  A.  1886.  Gastroblasta  rajfaelei.  Eine  durch  eine  Art  unvollstandiger  Theilung  entstehende 

Medusen-Kolonie.  Jena  Z  Naturw.  19  (N.P.  12) :  735-763. 


134  P.  F.  S.  CORNELIUS 

I  .a rw  ood.  G.  P.  &  Rosen,  B.  R.  (editors)  1979.  Biology  and  systematics  of  colonial  organisms.  Syst. 

Ass.  spec.  vol.  1 1 . 

Laverack,  M.  S.  &  Blackler,  M.  1974.  Fauna  and  flora  of  St.  Andrews  Bay.  Edinburgh  and  London. 
Le  Danois,  E.  1913.  Note  sur  trois  nouvelles  meduses  et  liste  des  coelenteres  du  plankton  recueillis 

a  bord  du  Pourquoi-Pas?  dans  sa  croisiere  dans  les  mers  du  Nord.  Bull.  Mus.  Hist.  Nat.  Paris 

19(1913):  110-113. 
Leloup,  E.  1930a.  Campanularia  macrotheca  nov.  sp.  Annls  Soc.  r.  zool.  Belg.  (1929)60  :  101-102. 

19306.  Deux  cas  d'epibiose  de  1'hydropolype  Campanularia  johnstoni  Alder.  Bull.  Mus.  roy. 

Hist.  nat.  Belg.  6  (\9):  1-5. 

193  la.  Trois  nouvelles  especes  d'hydropolypes.  Bull.  Mus.  r.  Hist.  nat.  Belg.  (2)  2  :  1-73. 

19316.  Un  cas  d'epibiose  de  1'hydropolype,  Laomedea  geniculata  (Linne).  Bull.  Mus.  r.  Hist.  nat. 

Belg.  7(24):  1-3. 
1933.  Contribution  a  la  connaissance  des  hydropolypes  de  la  cote  des  Pays-Bas.  Bull.  Mus.  r.  Hist. 

nat.  Belg.  9  (45):  1-30. 

1935.  Hydraires  calyptoblastiques  des  Indes  Occidentales.  Mem.  Mus.  r.  Hist.  nat.  Belg.  (2) 

2:  1-73. 

1939.  Hydropolypes  marins  et  dulcicoles  du  Congo  Beige.  Revue  Zool.  Bot.  afr.  32  :  418-423. 

1940.  Hydropolypes  provenant  des  croisieres  du  Prince  Albert  ler  de  Monaco.  Result.  Camp. 

sclent.  Prince  Albert  1 104  :  1-38. 

1947.  Les  coelenteres  de  la  faune  Beige.  Leur  bibliographic  et  leur  distribution.  Mem.  Mus.  r.  Hist. 


nat.  Belg.  107  :  1-73. 

1952.  Coelenteres.  Faune  Belg.,  pp.  1-283. 


Lendenfeld,  R.  von  1883a.  Eine  ephemere  Eucopide.  Zool.  Anz.  6  :  186-189. 

18836.  Uber  Coelenteraten  der  Siidsee.  IV.  Mittheilung.  Eucopella  Campanularia  nov.  gen.  Z. 

wiss.  Zool.  38  :  497-583. 

1 885.  Addenda  to  the  Australian  Hydromedusae.  Proc.  Linn.  Soc.  N.S.  W.  ( 1 884)  (1)9:  908-924. 


Leslie,  G.  &  Herdman,  W.  A.  1881.  The  invertebrate  fauna  of  the  Firth  of  Forth.  Edinburgh. 

Lesson,  R.-P.  1 843.  Histoire  naturelles  des  zoophytes.  Paris. 

Leuckart,   R.    1856.    Beitrage  zur   Kenntniss  der   Medusenfauna   von   Nizza.   Arch.   Naturgesch. 

22(1):  1-40. 
Levinsen,  G.  M.  R.  1893.  Meduser,  Ctenophorer  og  Hydroider  fra  Grgmlands  Vestkyst,  tilligemed 

Bemaerkninger  om  Hydroidernes  Systematik.   Vidensk.  Meddr  dansk  naturh.  Foren.  (1892)  (5) 

4:  143-212,215-220. 
Linko,  A.  K.   1911.  Hydraires  (Hydroidea).  Volume  I.  Haleciidae,  Lafoeidae,  Bonneviellidae  et 

Campanulariidae.  Fauna  Rossii,  pp.  1-250. 
Linnaeus,  C.  17 '58.  Sy 'sterna  naturae.  1  Oth  edition.  Holmiae. 

1767.  Systema  naturae.  12th  edition.  Tom  I,  Pars  II.  Holmiae. 

Lloyd,  R.  1907.  Nudiclava  monocanthi,  the  type  of  a  new  genus  of  hydroids  parasitic  on  fish.  Rec. 

Indian  Mus.  I  :  28 1-294. 
Lo  Bianco,  S.  1909.  Notizie  biologiche  riguardanti  specialmente  il  periodo  di  maturita  sessuale  degli 

animali  del  golfo  di  Napoli.  Mitt.  zool.  Stat.  Neapel  19:51 3-763. 

Macartney,  J.  1810.  Observations  upon  luminous  animals.  Phil.  Trans.  R.  Soc.  100  :  258-293. 
Macgillivray,  J.  1842.  Catalogue  of  the  marine  zoophytes  of  the  neighbourhood  of  Aberdeen.  Ann. 

Mag.  nat.  Hist.  (1)9  :  462^69. 
Maitland,  R.  T.  1876.  Determinatie  der  dieren,  beschreven  en  afgebeeld  in  de  Werken  van  Job  Baster 

en  Martinus  Slabber.  Tijdschr.  ned.  dierk.  Vereen.  2  :  7-15. 

1 897.  Prodrome  de  la  faune  des  Pays-Bas  et  de  la  Belgique  fla  mande.  Leiden. 

Mammen,  T.  A.  1965.  On  a  collection  of  hydroids  from  south  India.  II.  Suborder  Thecata  (excluding 

family  Plumulariidae).  J.  mar.  biol.  Ass.  India  1 :  1-57. 
Manea,  V.  1972.  Contribujii  la  studiul  hidroidelor  (Coelenterata)  din  Marea  Neagra.  Studii  Cere.  Biol. 

(Zool.)  24: 409-4 17. 

Marine  Biological  Association,  1957.  Plymouth  marine  fauna.  3rd  edition.  Plymouth,  England. 
Marktanner-Turneretscher,  G.  1890.  Die  Hydroiden  des  k.  k.  naturhistorischen  Hofmuseums.  Annln 

naturh.  Mus.  Wien,  5  :  195-286. 
Mathiesen,  O.  1928.  Hydroids  from  northern  Norway,  with  a  list  of  unreported  specimens  in  the 

Tromsjar  Museum.  TromspMus.  Aarsh.  (1926)49  (4) :  1-40. 
Mayer,  A.  G.   1900.  Some  medusae  from  the  Tortugas,  Florida.  Bull.  Mus.  comp.  Zool.  Harv. 

37:  13-82. 
1910.  Medusae  of  the  world.  Volume  II.  The  hydromedusae.  Washington. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  1 35 

McCrady,  J.  1857.  Gymnopthalmata  of  Charleston  Harbor.  Proc.  Elliott  Soc.  nat.  Hist.  1  :  103-221. 
Metschnikoff,  E.  1886a.  Medusologische  Mittheilungen.  Arbor,  zool.  Inst.  Univ.  Wiend  :  237-266. 

1 886/j.  Embryologische  Studien  an  Medusen.  2  vols,  text  &  atlas.  Vienna. 

Meyen,  F.  J.  F.  1834.  Uber  das  Leuchten  des  Meeres  und  Beschreibung  einiger  Polypen  und  anderer 

nieder  Thiere.  Nova  Acta  Acad.  Caesar.  Leop.  Carol.  16  (suppl.  1) :  125-2 16. 
Millard,  N..A.  H.  1957.  The  Hydrozoa  of  False  Bay,  South  Africa.  Ann.  S.  Afr.  Mus.  43  :  173-243. 
1958.  Hydrozoa  from  the  coasts  of  Natal  and  Portugese  East  Africa.  Part  I.  Calyptoblastea.  Ann. 

S.  Afr.  Mus.  44:  165-226. 

1966.  The  Hydrozoa  of  the  south  and  west  coasts  of  South  Africa.  Part  III.  The  Gymnoblastea 

and  small  families  of  Calyptoblastea.  Ann.  S.  Afr.  Mus.  48  :  427-487. 

1973.  Auto-epizoism  in  South  African  hydroids.  Publs  Seto  mar.  biol.  Lab.  20  :  23-34. 

1975.  Monograph  of  the  Hydroida  of  southern  Africa.  Ann.  S.  Afr.  Mus.  68  :  1-513. 

1977.  The  South  African  Museum's  Meiring  Naude  cruises.  Part  3.  Hydroida.  Ann.  S.  Afr.  Mus. 

73:  105-131. 
1978.  The  geographical  distribution  of  southern  african  hydroids.  Ann  S.  Afr.  Mus.  74  :  1 59-200. 

&  Bouillon,  J.    1973.  Hydroides  from  the  Seychelles.  Annls  Mus.  r.  Afr.  cent.  (8°,  Zool.) 

206:  1-106. 

Miller,  M.  C.  1961.  Distribution  and  food  of  the  nudibranchiate  mollusca  of  the  south  of  the  Isle  of 

Man.  J.  anim.  Ecol.  30  :  95-1 16. 
Miller,   R.   L.    1973.   The  role  of  the  gonomedusa  and  gonangium  in  the  sexual  reproduction 

(fertilization)  of  the  Hydrozoa.  Publs  Seto  mar.  biol.  Lab.  20  :  367-400. 
Morri,  C.  \919a.  Quelques  observations  sur  les  hydroides  de  la  Lagune  d'Orbetello.  Rapp.  P.-v.  Reun. 

Commn  int.  Explor.  scient.  Mer  Mediterr.  25-26  :  1 19-120. 

1979/7.  Contribute  alia  conoscenza  degli  idrozoi  lagunari  Italiani:  idropolipi  di  alcuni  laghi 

costieri  Mediotirrenici.  Annali  Mus.  civ.  Stor.  nat.  Geneva  (1978)  82  :  163-171. 

Mulder,  J.  F.  &  Trebilcock,  R.  E.  1914.  Victorian  Hydroida.  With  description  of  new  species.  Geelong 

Nat.  6:  6-15. 
Miiller,  O.  F.  1 776.  Zoologiae  Danicae prodromus,  seu  Animalium  Daniae  et  Norvegiae  indigenarum 

characteres,  nomina  et  synonyma  imprimis popularium.  Pp.  xxxii  +  282.  Havniae. 
Murdock,  G.  R.  1976.  Hydroid  skeletons  and  fluid  flow.  In  Mackie,  G.  O.  (editor),  Coelenterate 

ecology  and  behaviour  (Third  international  symposium  on  coelenterate  biology),  New  York.  Pp. 

33-40. 
Naumov,  D.   V.    1960.  Gidroidy  i  gidromeduzy  morskikh,  solonovatovodnykh  i  presnovodnykh 

basseinov  S.S.S.R.  Fauna  S.S.S.R.  70  :  1-626. 
1969.  Hydroids  and  hydromedusae  of  the  U.S.S.R.  Fauna  S.S.S.R.  70  :  1-660.  Israel  Program  for 

Scientific  Translations,  cat.  no.  5 108. 

&  Stepanyants,  S.  D.  1972.  Marine  invertebrates  from  Adelie  Land,  collected  by  the  Xllth  and 


XVth  French  Antarctic  Expeditions.  3.-Hydroida.  Tethys(Supp\.)4  :  25-60. 
Neppi,  V.  1912.  Adriatische  Hydromedusen.  Sber.  Akad.  Wiss.  Wien  121  (1) :  709-734. 
Neviani,  A.  1903.  Intorno  ad  una  rara  pubblicazione  di  G.  D.  Westendorp.  Boll.  Soc.  zool.  ital.  (2) 

4  : 96-97. 
Nishihira,  M.  1968.  Distribution  pattern  of  Hydrozoa  on  the  broad-leaved  eelgrass  and  narrow-leaved 

eelgrass.  Bull.  mar.  biol.  Stn  Asamushi  13  :  125-138. 

Nobre,  A.  1931.  Contribul'coes  para  o  estudo  dos  coelenterados  de  Portugal.  Porto. 
Norman,  A.  M.  1869.  Shetland  final  dredging  report.  Part  II.  On  the  Crustacea,  Tunicata,  Polyzoa, 

Echinodermata,  Actinozoa,  Hydrozoa,  and  Porifera.  Rep.  Brit.  Ass.  Advmt  Sci.  (1868)  (1) :  247-336. 
Nutting,  C.  C.  1899.  Hydroida  from  Alaska  and  Puget  Sound.  Proc.  U.S.  natn.  Mus.  21  :  741-753. 
190 la.  Papers  from  the  Harriman  Alaska  Expedition.  XXI.  The  hydroids.  Proc.  Wash.  Acad.  Sci. 

3:  157-216. 

1901/7.  The  hydroids  of  the  Woods  Hole  region.  Bull.  U.S.  Fish  Commn  (1899) :  325-386. 

1915.  American  hydroids.  Part  III.  The  Campanularidae  and  the  Bonneviellidae.  Spec.  Bull.  U.S. 


natn.  Mus.,  pp.  1-126. 
Oken,  L.  1815.  Okens  Lehrbuch  der  Naturgeschichte.  Dritte  Theil.  Zoologie.  Jena. 
Orton,  J.  H.  1920.  Sea-temperature,  breeding  and  distribution  in  marine  animals.  J.  mar.  biol.  Ass. 
•   U.K.  12  :  339-366. 
Ostman,  C.  1979.  Two  types  of  nematocysts  in  Campanulariidae  (Cnidaria,  Hydrozoa)  studied  by  light 

and  scanning  electron  microscopy.  Zoologica  Scr.  8  :  5-12. 
Pallas,  P.  S.  1 766.  Elenchus  zoophytorum.  The  Hague. 
1774.  Spicilegia  zoologica.  Tom.  I,  fasc.  10  :  1-42.  Berlin. 


136  P.  F.  S.  CORNELIUS 

Parker,  H.  W.  1939.  Turtles  stranded  on  the  British  coast.  Proc.  Linn.  Soc.  Lond.  151  :  127-129. 
Patriti,  G.  1970.  Catalogue  des  cnidaires  et  ctenaires  des  cotes  atlantiques  marocaines.  Rabat. 
Pennington,  A.  S.  1885.  British  zoophytes:  an  introduction  to  the  Hydroida,  Aclinozoa,  and  Polyzoa 

found  in  Great  Britain,  Ireland,  and  the  Channel  Islands.  London. 
Peron,  F.  &  Lesueur,  C.-A.  1810a.  Tableau  des  caracteres  generiques  et  specifiques  de  toutes  les 

especesde  meduses  connuesjusqu'ace  jour.  Annls  Mus.  Hist.  nat.  Paris  (1809)  14  :  325-366.  (Dating 

of  this  publication  follows  Sherborn,  1914.) 
718106.  Histoire  generate  des  meduses.  Paris.  (Dating  of  this  work  is  discussed  in  note  3, 

p  124). 

71815.  Voyage  de  decouvertes  aux  terres  australes.  Histoire  naturelle.  Histoire  generale 


et  particuliere  de  tous  les  animaux  qui  composent  la  famille  des  meduses,  et  de  quelques  autres 

radiaires  mollasses.  Paris.  (Dating  of  this  work  is  discussed  in  note  3,  p.  124). 
Philbert,  M.  1935a.  Le  phenomene  de  stolonisation  chez  trois  especes  d'hydraires  fixees  sur  des 

posidoniesen  Mediterranee.  Bull.  Inst.  oceanogr.  Monaco 663  :  1-8. 

19356.  Les  hydraires  de  la  region  malouine.  Bull.  Inst.  oceanogr.  Monaco  673  :  1-36. 

Picard,  J.  1949.  Sur  la  presence  en  Mediterranee  de  Clytia  noliformis  (McCrady).  Bull.  Mus.  Hist.  nat. 

Marseille  9:  184-190. 
1950.  Observations  sur  les  hydraires  recoltes  aux  Martigues  dans  le  canal  de  Carente.  Vie  Milieu 

1:51-52. 

1 95 1  a.  Note  sur  les  hydraires  littoraux  de  Banyuls-sur-Mer.  Vie  Milieu  2  :  338-349. 

195 16.  Hydraires  littoraux  du  Senegal  recoltes  par  H.  Sourie  aux  environs  de  Dakar.  Bull.  Inst.fr. 

Afr.  noire  13:  109-115. 
1955.  Hydraires  des  environs  de  Castiglione  (Algerie).  Bull.  Stat.  Aquic.  Peche  Castiglione  (NS) 

7:  177-199. 
1958a.  Notes  sur  une  collection  d'hydroides  provenant  des  cotes  Mediterraneennes  d'Israel.  Bull. 

Sea  Fish.  Res.  Stn  Israel  15  :  1-3. 

19586.  Origines  et  affmites  de  la  faune  d'hydropolypes  (gymnoblastes  et  calyptoblastes)  et 


d'hydromeduses  (anthomeduses  et  leptomeduses)  de  la  Mediterranee.  Rapp.  P.-v.  Reun.  Commn  int. 

Explor.  scient.  Mer  Mediterr.  (NS)  14  :  187-199. 

Pictet,  C.  1 893.  Etude  sur  les  hydraires  de  la  Baie  d'Amboine.  Revue  suisse  Zool.  1  :  1-64. 
Pieper,  F.  W.  1844.  Erganzungen  zu  "Heller's  Zoophyten  etc.  des  adriatischen  Meeres".  Zool.  Anz. 

7:  148-152,164-169,185-188,216-221. 

Poche,  F.  19 14.  Das  System  der  Coelenterata.  Arch.  Naturgesch.  80  (Abt.  A,  Heft  5) :  47-128. 
Prenant,    M.    &    Bobin,    G.    1956.    Bryozoaires.    Premiere    partie.    Entoproctes,    phylactolemes, 

ctenostomes.  Faune  Fr.  60  :  1-398. 
Quoy,  J.  R.  C.  &  Gaimard,  J.  P.  1827.  Observations  zoologiques  faites  a  bord  de  V Astrolabe,  en  mai 

1 826,  dans  le  detroit  de  Gibraltar.  Annls  Sci.  nat  (Zool.)  (1 )  10:5-21,1 72-193, 225-239. 
Ralph,  P.  M.    1956.  Variation  in  Obelia  geniculata  (Linnaeus,   1758)  and  Silicularia  bilabiata 

(Coughtrey,  1875)  (Hydroida,  F.  Campanulariidae).  Trans.  R.  Soc.  N.Z.  84  :  279-296. 
1957.  New  Zealand  thecate  hydroids.  Part  I.  Campanulariidae  and  Campanulinidae.  Trans.  R. 

Soc.  N.Z.  84:  81 1-854. 

&  Thomson,  H.  G.  1968.  Seasonal  changes  in  growth  in  the  erect  stem  of  Obelia  geniculata  in 


Wellington  Harbour,  New  Zealand.  Zool.  Publs  Victoria  Univ.  Wellington  44  :  1-21. 
Rankin,  J.  1901.  Hydroida.  In  Elliot,  G.  F.  S.,  Laurie,  M.  &  Murdoch,  J.  B.  (editors).  Fauna,  flora  & 

geology  of  the  Clyde  area;  Glasgow  [British  Association  for  the  Advancement  of  Science];  pp. 

369-371. 

Rasmussen,  E.  1973.  Systematics  and  ecology  in  the  Isefjord  marine  fauna.  Ophelia,  11  :  1-495. 
Ray,  J.  1 724.  Synopsis  methodica  stirpium  Britannicarum.  Edn.  3.  London. 
Redier,  L.  1967.  Revision  de  la  collection  du  Museum  des  hydraires  de  Lamouroux.  Bull.  Mus.  natn 

Hist.  nat.  Paris  (2)  39  :  381^10. 
Rees,  W.  J.  1939.  A  revision  of  the  genus  Campanulina  van  Beneden,  1847.  Ann.  Mag.  nat.  Hist.  (11) 

3  : 433^47. 
&  Rowe,  M.  1969.  Hydroids  of  the  Swedish  west  coast.  Ada.  R.  Soc.  Scient.  litt.  gothoburg.  (Zool.) 

3  :  1-24. 

&  Thursfield,  S.  1965.  The  hydroid  collections  of  James  Ritchie.  Proc.  R.  Soc.  Edinb.  (B) 


69  : 34-220. 

—  &  White,  E.  1966.  New  records  and  fauna  list  of  hydroids  from  the  Azores.  Ann.  Mag.  nat.  Hist. 
(13)9:271-284. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  137 

Riedl,  R.  1959.  Die  Hydroiden  des  Golfes  von  Neapel  und  ihr  Anteil  an  der  Fauna  unterseeischer 

Hohlen.  Publ.  Staz.  zool.  NapoliSO  (suppl.) :  591-755. 
Rioja  y  Martin,  J.  1906.  Datos  para  el  conocimiento  de  la  fauna  marina  de  Espana.  Boln  R.  Soc.  esp. 

Hist.  not.  6:  275-281. 
Ritchie,  J.  1907.  On  collections  of  the  Cape  Verde  Islands  marine  fauna,  made  by  Cyril  Crossland, 

M.A.  (Cantab.),  B.Sc.  (Lond.),  F.Z.S.,  of  St.  Andrews  University,  July  to  September,  1904.  The 

hydroids.  Proc.  zool.  Soc.  Lond.  (1907) :  488-514. 

1910-1911.  Contribution  to  our  knowledge  of  the  hydroid  fauna  of  the  west  of  Scotland.  Ann. 

Scot.  nat.  Hist.  76  (1910) :  220-225;  77  (191 1) :  29-34;  79  (191 1) :  158-164;  80  (191 1) :  217-225. 

Robins,  M.  W.  1969.  The  marine  flora  and  fauna  of  the  Isles  of  Scilly.  Cnidiaria  and  Ctenophora.  /. 

nat.  Hist.  3:  329-343. 
Robson,  J.  H.   1914.  Catalogue  of  the  Hydrozoa  of  the  north-east  coast  (Northumberland  and 

Durham).  Rep.  Dove  mar.  Lab.  (NS)3  :  87-103. 
Roosen-Runge,  E.  C.  1970.  Life  cycle  of  the  hydromedusa  Phialidium  gregarium  (A.  Agassiz,  1862)  in 

the  laboratory.  Biol.  Bull.  mar.  biol.  Lab.,  Woods  Hole,  139  :  203-22 1 . 
Rossi,  L.  1950.  Celenterati  del  Golfo  di  Rapallo.  Boll.  Musei  Zool.  Anat.  comp.  R.  Univ.  Torino 

2:  193-235. 

197 1 .  Guid  a  cnidari  e  ctenofori  della  fauna  Italiana.  Quad.  civ.  Sta.  idrobiol.  Milano  2  :  5-101 . 

Russell,  F.  S.  1953.  The  medusae  of  the  British  Isles.  Anthomedusae,  Leptomedusae,  Limnomedusae, 

Tr  achy  medusae  and  Narcomedusae.  Cambridge. 
Sars,  M.  1835.  Beskrivelser  og  Jagttagelser  over  nogle  maerkelige  eller  nye  i  havet  ved  den  Bergenske 

Kyst  levende  Dyr  af  Polypernes,  Acalephernes,  Radiaternes,  Annelidernes  og  Molluskernes  Classer, 

med  en  kort  Oversigt  over  de  hidtil  af  Forfatteren  sammesteds  fundne  Arter  og  deres  Forekommen. 

Bergen. 

1846.  Fauna  littoralis  Norvegiae.  Erstes  Heft.  I.  Ueber  die  Fortpflanzungsweise  einiger  Polypen 

(syncoryna — Podocoryna — Perigonimus — Cytaeis).  Christiania. 

1850.  Beretning  om  en  i  Sommeren  1849  foretagen  zoologisk  Reise  i  Lofoten  og  Finmarken. 


Nyt.  Mag.  Naturvid.  6  :  121-211.  [The  date  of  this  paper  has  been  given  as  1851  (e.g.  by  Bedot,  1910; 
Vervoort,  1972),  and  that  date  appears  on  the  title  page  of  the  volume.  However,  Botten-Hansen  & 
Petersen  (1870  :  209)  and  Sherborn  (1922  :  14)  dated  the  paper  1850.  The  paper  was  reprinted  as  a 
book  differing  only  in  being  paginated  1-9 1 ,  published  'Christiana',  1 850.] 

1857.  Bidrag  til  Kundskaben  om  Middelhavets  Littoral-Fauna,  Reisebemaerkninger  fra  Italien. 


Nyt.  Mag.  Naturvid.  9  :  1 10-164.  (This  article  formed  part  of  a  book  of  identical  title  published  in 

Oslo  (Christiania)  in  the  same  year,  paginated  1-55.) 
Savage,  S.  1945.  A  catalogue  of  the  Linnaean  herbarium.  London. 
Schach,  H.   1935.  Kausal  Analyse  der  Entstehung  des  polysiphonen  Stockes  von  Campanularia 

verticillata  L.  Arch.  Entw-Mech.  Org.  Berlin  132  :  615-647. 
Schmidt,  H.-E.  &  Benovic,  A.  1977.  Notes  on  hydromedusae  (Cnidaria)  from  the  Adriatic  Sea.  J.  mar. 

biol.  Ass.  U.K.  51 :  635-640. 
Shaw,  G.  R.  &  Nodder,  F.  P.  1975.  The  naturalist's  miscellany.  Vol.  6.  London.  (Plate  196,  cited  here 

under  Clytia  hemisphaerica,  was  probably  published  in  1795  according  to  Sherborn,  C.  D.  1895  'On 

the  dates  of  Shaw  and  Nodder's  "Naturalist's  Miscellany".'  Ann.  Mag.  nat.  Hist.  (6)  15  :  375-376.) 
Sherborn,  C.  D.  1914.  An  attempt  at  a  fixation  of  the  dates  of  issue  of  the  parts  of  the  publications  of 

the  Musee  d'Histoire  Naturelle  of  Paris,  1802-1850.  Ann.  Mag.  nat.  Hist.  (8)  13  :  365-368. 

1922.  Index  animalium  . . .  Part  I.  Introduction,  bibliography  and  index  A-Aff.  London. 

1926.  Index  animalium  .  . .  Part  X.  Index  Eurystomus-funereus.  London. 

1929.  Index  animalium  . . .  Part  XVIII.  Index  o-nigrum-pallens.  London. 

Sigerfoos,  C.  P.   1893.  Note  on  the  formation  of  blastostyle  buds  on  the  reproductive  organs  of 

Epenthesis  mccradyi.  Johns  Hopk.  Univ.  Circ.  12  :  106. 
Slabber,  M.  1769-1778.  Natuurkundige  verlustigingen,  behelzende  microscopise  waarneemingen  van 

in-  en  uitlandse  water-  en  land-dieren.  Haarlem,  Holland.  (For  bibliographic  details  see  Cornelius, 

19750:289.) 
1775-1781.  Physicalische  Belustigungen  oder  microscopische  Wahrnehmungen  von  drey  und 

vierzigin-  und  auslandischen  Wasser-  und  Landthierchen.  Translated  by  P.  L.  St.  Miiller.  Niirnberg. 
Splettstosser,  W.  1924.  Beitrage  zur  Kenntnis  der  Gattung  Laomedea  (sensu  Broch).  Zool.  Jb.  (Syst.) 

48  :  367^32. 
Stebbing,  A.  R.  D.  1976.  The  effects  of  low  metal  levels  on  a  clonal  hydroid.  J.  mar.  biol.  Ass.  U.K. 

56  :  977-994. 


138  P.  F.  S.  CORNELIUS 

1979.  An  experimental  approach  to  the  determinants  of  biological  water  quality.  Phil.  Trans.  R. 


Soc.  (8)286:465^181. 

198 1 .  The  kinetics  of  growth  control  in  a  colonial  hydroid.  J.  mar.  biol.  Ass.  U.K.  61  :  35-63. 


Stechow,  E.  1913.  Hydroidpolypen  der  japanischen  Ostkiiste.  II  Teil:  Campanularidae,  Halecidae, 
Lafoeidae,  Campanulinidae  und  Sertularidae,  nebst  Erganzungen  zu  den  Athecata  und 
Plumularidae.  Abh.  bayer.  Akad.  Wiss.  suppl.  3  (2) :  1-162. 

1914.  Zur  Kenntnis  neuer  oder  seltener  Hydroidpolypen,  meist  Campanulariden,  aus  Amerika 

und  Norwegen.  Zoo/.  Anz.  45  :  120-136. 

1919a.  Zur  Kenntnis  der  Hydroidenfauna  des  Mittelmeeres,  Amerikas  und  anderer  Gebiete, 

nebst  Angaben  iibereinige  kirchenpauer'sche  Typen  von  Plumulariden.  Zoo/.  Jb.,  Syst.  42  :  1-172. 

19 190.  Neue  Ergebnisse  auf  dem  Gebiete  der  Hydroidenforschung.  Mtinchener  Med.  Wochenschr. 


(1919)30:852-853. 

1919c.  Neue  Ergebnisse  auf  dem  Gebiete  der  Hydroidenforschung.  Sber.  Ges.  Morph.  Physiol. 


Munch.  31  : 9-45. 

—  192 la.  Neue  Genera  und  Species  von  Hydrozoen  und  anderen  Evertebraten.  Arch.  Naturgesch. 
87  (Abt.  A.  Heft  3) :  248-265. 

—  192 10.  Symbiosen  zwichsen  Isopoden  und  Hydroiden.  Zoo/.  Anz.  53  :  221-223. 

—  1923a.  Zur  Kenntnis  der  Hydroidenfauna  des  Mittelmeeres,  Amerikas  und  anderer  Gebiete.  II 
Teil.  Zoo/.  Jb.  (Syst.)  47  :  29-270. 

—  19230.  Die  Hydroidenfauna  der  japanischen  Region.  J.  Coll.  Sci.  imp.  Univ.  Tokyo,  44  (8) :  1-23. 

—  1923c.  Neue  Hydroiden  der  Deutschen  Tiefsee-Expedition,  nebst  Bemerkungen  iiber  einige  andre 
Formen.  Zoo/.  Anz.  56  :  1-20. 

—  1923d.  Uber  Hydroiden  der  Deutschen  Tiefsee-Expedition,  nebst  Bermerkungen  iiber  einige 
andre  Formen.  Zoo/.  Anz.  56  :  97-1 19. 

—  1925.  Hydroiden  der  Deutschen  Tiefsee-Expedition.  Wiss.  Ergebn.  dt.  Tiefsee- Exped.  "Valdivia 
17  :  383-546. 

—  1927.  Die  Hydroidenfauna  der  Ostsee.  Zoo/.  Anz.  70  :  304-3 13. 

1932.  Neue  Hydroiden  aus  dem  Mittelmeer  und  dem  Pazifischen  Ozean,  nebst  Bemerkungen 


iibereinige  wenig bekannte  Formen.  Zoo/.  Anz.  100  :  81-92. 
Stefani,  R.  1956.  II  problema  dell'allogonia  negli  Idrozoi.  1.  Osservazioni  biologiche  ed  istologiche  in 
una  popolazione  di  Campanularia  caliculata  Hincks  del  Golfo  di  Cagliari.  Archo  zoo/,  ital. 
41 : 337-368. 

1959.  Sulla  variabilita  ecologica  di  un  Idrozoo  (Campanularia  caliculata  Hincks).  Boll.  Zoo/. 

26  (2):  115-1 20. 

Stepanyants,  S.  D.  1979.  Gidroidy  vod  antarktikh  i  subantarktikh.  Issled.  Fauny  Morei  22  (30): 

1-200. 
Stephens,  J.  1905.    A  list  of  Irish  Coelenterata,  including  the  Ctenophora  Proc.  R.  Ir.  Acad.  (B) 

25  : 25-92. 

Teissier,  G.  1965.  Inventaire  de  lafaune  marine  de  Roscoff.  Cnidaires-ctenaires.  Roscoff,  France. 
Thiel,  M.  E.   1935.  Zur  Kenntnis  der  Hydromedusenfauna  des  Schwarzen  Meeres.  Zoo/.  Anz. 

Ill :  161-174. 
Thornely,  L.  R.  1899.  The  hydroid  zoophytes  collected  by  Dr  Willey  in  the  southern  seas.  In:  Willey, 

A.,  1898-1902,  Zoological  results  based  on  material  from  New  Britain,  NewGuinea,  Lovalty  Islands 

and  elsewhere.  Collected  during  1895,  1986  and  1897.4  :  45 1-457.  Cambridge. 

1904.  Report  on  the  Hydroida  collected  by  Professor  Herdman,  at  Ceylon,  in  1902.  Rep.  Govt 

Ceylon  Pearl  Oyster  Fish.  GulfManaar,  Suppl.  Rep.  8  :  107-126. 

Todd,  R.  A.  1906.  Marine  zoology.  In  Page,  W.  (editor)  The  Victoria  history  of  the  counties  of 

England:  a  history  of  Devonshire  in  five  volumes.  Volume  1,  pp.  1 3 1-1 58.  London. 
Torrey,  H.  B.  1904.  The  hydroids  of  the  San  Diego  region.  Univ.  Calif.  Publs  Zoo/.  2  :  1-43. 
Toth,  S.  E.  1969.  Aging  and  regression  in  the  colonial  marine  hydroid  Campanularia  flexuosa  with 

special  reference  to  senescence  in  hydroids.  Int.  Rev.  gen.  exp.  Zoo/.  4  :  49-79. 
Totton,  A.  K.  1930.  Coelenterata.  Part  V.  Hydroida.  Nat.  Hist.  Rep.  Br.  antarct.  Terra  Nova  Exped. 

5:  131-252. 

1965.  A  synopsis  of  the  Siphonophora.  London. 

Trebilcock,  R.  E.  1928.  Notes  on  New  Zealand  Hydroida.  Proc.  R.  Soc.  Viet.  41  :  1-3 1 . 

Tutin,  T.  G.  1942.  Zostera  L.  J.  Ecol.  30  :  2 1 7-226. 

Vanhoffen,    E.     1910.    Die    Hydroiden    der    Deutschen    Sudpolar-Expedition     1901-1903.    Dt. 

Sudpol.- Exped.  11  (=  Zool.  3) :  269-340. 
Vannucci,  M.  1955.  On  the  newly  liberated  medusa  ofObelia  hyalina  Clarke,  1 879.  Dusenia  6  :  55-60. 


N.E.  ATLANTIC  CAMPANULARIID  HYDROZOANS  139 

Vatova,  A.  1928.  Compendio  della  Flora  e  Fauna  del  Mare  Adriatico  presso  Rovigno.  Memorie  R. 

Com.  talassogr.  ital.  143  :  1-614. 
Verrill,  A.  E.  1873.  Results  of  recent  dredging  expeditions  on  the  coast  of  New  England.  Am.  J.  Sci.  (3) 

5:  1-16. 
Vervoort,  W.  19460.  Hydrozoa  (CI).  A.  Hydropolypen.  Fauna  Ned.  14  :  1-336. 

19466.  Hydroiden  uit  de  Waddenzee.  Archs  neerl.  Zool.  7  :  334-352. 

1949.  Notes  on  a  small  collection  of  hydroids  from  Jersey  (Channel  Islands).  Zool.  Meded. 

Leiden,  30:  133-162. 

1959.  The  Hydroida  of  the  tropical  west  coast  of  Africa.  Atlantide  Rep.  5:21 1-325. 

1966.  Bathyal  and  abyssal  hydroids.  Galathea  Rep.  8  :  97-174. 

1967.  The  Hydroida  and  Chondrophora  of  the  Israel  South  Red  Sea  Expedition,  1962.  Bull.  Sea 

Fish.  Res.  Stn  Israel,  43  :  1 8-54. 

1968.  Report  on  a  collection  of  Hydroida  from  the  Caribbean  region,  including  an  annotated 


checklist  of  Caribbean  hydroids.  Zool.  Verh.  Leiden92  :  1-124. 

1972.  Hydroids  from  the  Theta,  Vema  and  Yelcho  cruises  of  the  Lamont-Doherty  Geological 


Observatory.  Zool.  Verh.  Leiden  120  :  1-247. 
Warburton,  F.  E.  1953.  Antagonism  between  different  species  of  hydroids  on  the  same  shell.  Ecology 

34:  193-194. 
Warren,  E.   1908.  On  a  collection  of  hydroids,  mostly  from  the  Natal  coast.  Ann.  Natal  Mus. 

1  : 269-355. 

Weismann,  A.  1 883.  Die  Entstehung der  Sexualzellen  bei  den  Hvdromedusen.  Jena. 
West,   D.    L.   &   Renshaw,   R.   W.    1970.   The  life  cycle  of  Clytia  attenuata  (Calyptoblastea  : 

Campanulariidae).  Mar.  Biol.  Berlin  1 :  332-339. 
Westendorp,  G.  D.  1843.  Recherches  sur  les  polypiers  flexibles  de  la  Belgique  el  particulierement  des 

environs  d'Ostende.  Bruges,  pp.  1-48  +  pi.  1.  (Published  for  the  Societe  Medico-Chirurgicale  de 

Bruges  by  F.  de  Pachtere.  Reprinted  from  the  Annales  of  the  Societe,  vol.  4.  I  have  not  seen  the 

Annales  version.) 

1853.  Polypiers  flexibles  de  la  Belgique.   Collection  de  bryozoaires,  sertulaires,  /lustres  el 

spongiaires  qu'on  rencontre  en  Belgique  et  particulierement  aux  environs  d'Ostende.  Coutrai, 
Belgium.  [Not  seen.  These  details  are  from  Bedot(  19 10).  See  note  l,p.  123.] 

Will,  J.  G.  F.  1 844.  Horae  tergestinae  oder  Beschreibung  und  Anatomie  der  im  Herbste  1843  bei  Triest 

beobachteten  Akalephen.  Leipzig. 

Winsor,  M.  P.  1976.  Starfish,  jellyfish,  and  the  order  of  life.  New  Haven,  U.S.A. 
Winther,  G.    1879.   Fortegnelse  over  de  i  Danmark  og  dels  nordlige  Bilande  fundne  Hydroide 

Zoophyter.  Naturh.  Tidsskr.  (3)  12  :  223-278. 
Wood,  G.  W.  1901.  The  hydroid  zoophytes  of  the  Isle  of  Man,  with  a  notice  of  species  not  hitherto 

reported  from  the  district.  Lioar  Manninagh  2  :  12-2 1 . 

Wright,  T.  S.  1858.  Observations  on  British  zoophytes.  Proc.  R.  phys.  Soc.  Edinb.  1  :  447-455. 
1859.  Observations  on  British  zoophytes.  Edinb.  newphil.  J.  (N.S.)  9  :  106-1 14. 

1862.  On  the  reproduction  of  Thaumantias  inconspicua.  Q.  J.  microsc.  Sci.  (N.S.)  2  : 221-222, 

308. 

1 863.  Observations  on  British  zoophytes.  Q.  J.  microsc.  Sci.  (N.S.)  3  :  45-52. 


Wulfert,  J.  1902.  Die  Embryonentwicklung  von  Gonothyraea  loveni  Allm.  Z.  wiss.  Zool.  71  :  296-327. 
Wyttenbach,  C.  R.  1968.  The  dynamics  of  stolon  elongation  in  the  hydroid,  Campanularia  flexuosa.  J. 
exp.  Zool.  167:333-351. 

1969.  Genetic  variations  in  the  mode  of  stolon  growth  in  the  hydroid,  Campanularia  flexuosa. 

Biol.  Bull.  mar.  biol.  Lab.,  Woods  Hole  137  :  547-556. 

,  Crowell,  S.  &  Suddith,  R.  L.  1973.  Variations  in  the  mode  of  stolon  growth  among  different 

genera  of  colonial  hydroids,  and  their  evolutionary  implications.  J.  Morph.  139  :  363-375. 

Yamada,  M.  1959.  Hydroid  fauna  of  Japanese  and  its  adjacent  waters.  Publs  Akkeshi  mar.  biol.  Stn 
9:  1-101. 

1965.  Marine  hydroids  from  Greece.  Publs  Seto  mar.  biol.  Lab.  12  :  359-362. 

Zirpolo,  G.  1939.  Caso  di  epibiosi  di  Obelia  geniculata  su  Hippocampus  guttulatus.  Annuar.  Mus. 
zool.  Univ.  Napoli  (N.S.)  7  (8) :  1-8. 

1940.  Nuovo  caso  di  associazione  di  idroidi  e  pesci  con  revisione  critica  dei  casi  gia  noti.  Boll. 

Soc.  nat.  Napoli  ( 1938-1 939)  50  :  127-139. 

Manuscript  accepted  for  publication  24  July  1981. 


Index 

Valid  names  are  in  Roman  type,  new  names  in  bold  and  invalid  names  in  italics.  New  combinations  are 
indicated  by  an  asterisk  (*).  Main  page  references  are  in  bold.  References  to  substrates  are  not  included. 


abietina,  Campanularia  &  Grammaria  1 22 

acuminata,  Campanulina  74 

acuminata,  Laomedea  122 

acuta,  Campanularia  74, 81 

adelungi,  Obelia  1 1 4 

adriatica,  Obelia  1 20 

Aequorea  122 

vitrina  123 
affinis,  Clytia  7 1 
affinis,  Eucope  11,  73,  74 
africana,  Campanularia  60 
africana,  Orthopyxis  39, 60 
agas,  Campanularia  54 
Agastra  48,  57-58 

caliculata  6 1 

mira  6 1 , 67 

rubra6l,63,61 
aha,  Campanularia  53, 54 
alternata,  Clytia  84,  85, 86 
alternata,  Eucope  &  Obelia,  1 14 
andersoni,  Obelia  1 14 
angulata,  Campanularia  &  Eulaomedea  98 
angulata,  Laomedea  41,  42,  43,  44,  98-102, 

104,  105;  Fig.  17,  p.  99 
angulosa,  Obelia  1 14 
antennina,  Nemertesia  121 
arborescens,  Clytia  1 13,  1 14, 117 
arcta,  Coppinia  122 
armata,  Oceania  8 1 
arruensis,  Obelia  1 14 
articulata,  Eucope  1 14 
asymmetrica,  Orthopyxis  6 1 , 67 
atlantica,  Hypanthea  50 
atlantica,  Silicularia  50 
attenuata,  Campanularia  73-74, 79, 81 
attenuata,  Clytia  40, 42,  8 1 
attenuata,  Obelia  1 14 
australis,  Obelia  1 14 
austrogeorgiae,  Obelia  1 13, 117 


bicophora,  Clytia  (Trochopyxis)  73, 80 

bicophorum,  Phialidium  74, 80 

bicuspidata,  Obelia  90, 113,  1 14 

bidentata,  Obelia  40,  43,  44,  85,  96,  97,  1 10, 

111,113-117 
bifurca,  Obelia  11 4,  115 
bifurcata,  Obelia  1 14 
bilabiata,  Campanularia  58,  59, 60 
bilabiata,  Eucopella  58 
bilabiata,  Silicularia  60 
Billardia  123 
novaezealandiae  123 


biserialis,  Obelia  1 14 

bistriata,  Laomedea  84,  85, 86 

borealis,  Campanularia  6 1 , 66 

borealis,  Obelia  1 14 

brachycaulis,  Campanularia  53,  55-56,  74, 82 

brasiliensis,  Campanularia  1 14 

braziliensis,  Obelia  1 14 

breviscyphia,  Campanularia  60-6 1 , 66 

brochi,  Laomedea  47,  107, 110 

brochi,  Paralaomedea  47 

buskiana,  Thaumantias  74 

buskianum,  Phialidium  74 


calceolifera,  Campanularia  102 

calceolifera,  Eulaomedea  98,  102 

calceolifera,  Laomedea  41,  42,  43,  44,  47,  98, 

100,  101,  102-105;  Fig.  18,  p.  103 
calceolifera,  f.  of  Laomedea  angulata  100 
calceolifera,  Obelia  102 
caliculata,  Agastra  6  1 
caliculata,  Campanularia  51,  58,  60-61,  64, 

65-66 

caliculata,  Eucopella  58,61 
caliculata,  Orthopyxis  48,  6  1  ,  63,  65-66 
Calycellal22 

syringa  55,  57,  78,  122 
calyculata,  Campanularia  51,61 
Camp  alar  ia  97 

conferta  102-105 
campanella,  Medusa  73 
Campanula  51 

Campanularia,  Eucopella  58,  60,  61  ,  63,  67 
Campanularia  4  1  ,  47,  48,  49,  50,  51-52,  57,  70, 
71,97,112 

abietina  122 

acuta  74,  81 

africana  60 

agas  54 


angulata  98 

attenuata!3,14,19,Sl 

bilabiata  58,  59,  60 

borealis  6  1  ,  66 

brachycaulis  53,  55,  56,  74,  82 

brasiliensis  1  1  4 

breviscyphia  60,  6  1  ,  66 

calceolifera  102 

caliculata  5  1  ,  58,  60,  6  1  ,  64,  65-^6 

calyculata  51,61 

caulini  1  14 

cavolinii  1  14 

cheloniae\\4,  117,  118,119 


INDEX 


141 


compressa  60,  6 1 ,  64, 66-67,  73 

confer  ta  100,  104 

coruscans  1 14 

crenata  52-53,  59,  60;  Fig.  2,  p.  55 

crenata  f.  intermedia  59 

decipiens  107 

devisa  123 

edwardsill,  81 

elongata  107,110 

everta5\,58,6Q 

fastigiata  123 

flabellatall4 

flemingii95,96,\\3 

flexuosa  105 

fragilis  107, 109,110 

fruticosa  121,  122 

gegenbauri!3,8Q 

gelatinosa  95 

geniculata  sens.  Lister  9 1 

gigantea  73,  75,81 

gracilis  sens.  Allman  60,  61,  66;  sens.  Meyen 

50;sens.  Sars  74,  77,81 
gracillima  122 
gravieri%4,85 
groenlandica  52,  55,  56, 57 
hicksoni  54 

hincksii  5 1 , 53-55,  80,  82;  Fig.  3,  p.  55 
humilis  123 
hummelincki  82 
inconspicua  73 
Integra  52,60 
integriformis  6 1 ,  64, 66-67 
intermedia  58,60 
inter  text  a  65 ,  122 
johnstoni  71 ,  73,  75,  77, 79-80 
kincaidi  9 1 
lacerata  122 
7flevis54,60,61,65 
lennoxensis  58, 60 
macrotheca  53,54 
magnifica  52,53 
maior  1 1 4 
morgansi  5 1 
mutabilis  123 
neglecta  107 
noliformis  80 
obliqua  84 
obtusidens  1 14 
parvula  122 
paulensis  88 
pelagica  73, 78-79 
platycarpa  6 1 
prolifera  1 14 
ptychocyathus  80 
pulcratheca  5 1 
rara  53 

raridentata  73,  74, 75,  76, 80,  8 1 , 90 
retroflexa  5 1 
60 


senulatall,8Q 
speciosa  52,  53 
speciosus  52 
spinulosa  1 14 


tulipifera  70 

tulpifera  70 

vermicularis  120 

verticellata  67 

verticillata67,69 

villafrancensis  74, 81 

volubiliformis  73, 80 

volubilis  48,  5 1 ,  54,  55-57,  69,  70,  73,  76,  77, 

79;  Fig.  4,  p.  55 

CAMPANULARIIDAE  47^19 
CAMPANULARIINAE  48-49, 50-51, 69-70 
campanulata,  Eucope  7 1 ,  73,  74 
Campanulata  5 1 

verticillata  68 
Campanulina  acuminata  74 

tennis  123 
Capsularia  121,  122 

Integra  6 1 

/aev/5  6 1 , 65 
carnea,  Podocoryne  43 
caulini,  Campanularia  1 14 
cavolinii,  Campanularia  1 14 
cheloniae,  Campanularia  1 14,  1 17,  1 18,  1 19 
chinensis,  Obelia  1 14 
Clytea  70 

vicophora  73 
Clythia  57,  70 
C/y//z/ajohnstoni  74 

poterium  6 1 , 66 
Clytia  41 , 42, 47, 48, 49,  57,  69, 70-72 

affinis  1  \ 

alter nata  84,85,86 

arborescens  1 13,  1 14, 117 

attenuata  40,42,81 

bicophora  73,80 

compressa  14, 82 

cylindrica  42,  73, 80 

discoida*  42, 72-73;  Fig.  8,  p.  72 

edwardsi  8 1 

ed  wards  ia  43 

elsaeoswaldae  80 

eucophora,  eucopophora  123 

flavidula  73,  74,  76,  77,  80, 81 

folleata  9 1 

foxi  84, 86 

gardineri  9 1 

geniculata  84,  86 

gigantea  40,  74,  77 

gracilis  42,  71,91 

gravieriS5,86 

hemisphaerica  40,  41,  42,  43,  45,  70-71, 
73-82,  86,  87,  89,  90,  91,  92,  94;  Fig.  9, 
p.  75 

hendersonae  84 


142 

hummelincki  4 1 , 42, 44, 69, 82-83,  86; 
Fig.  10,  p.  82 

islandica*  42, 84;  Fig.  11,  p.  84 

johnstoni  69,  70,  73,  74,  80 

kincaidi  9 1 

teem  55, 57, 73 

tanguida  8 1 

linearis*  42, 84-86;  Fig.  12,  p.  85 

longicyatha  120 

longiiheca  1 1 4 

mccradyi*  42, 87-88;  Fig.  13,  p.  87 

mo///s55,56,57,80 

«o///brw/.s73,80,  81 

obeliformis  13,15, &\ 

obliqua85,86 

olivacea  67 

Clytia  (Orthopyxis)  poten'MW  57,  58,60,61,66 
Clytia  paulensis  42,  43,  44,  76,  85,  86,  88-91, 
113,  115,  117;  Fig.  14,  p.  89 

pelagica  73 

pentata42,9l 

Clytia  (Platypyxis)  70, 71,  80 
Clytia  posterior  6 1 

poterium61,63 

rijckholtii  6 1 , 65 

ryckholtii  6 1,65,  123-124 

sarsi41,74,78 

serrata  85 

serrulata  73,  74,  76 

simplex  1?>,&\ 

striata  86 

Clytia  (Trochopvxis)  70 
Clytia  ulvae  88, 91 

undulata  60, 6 1 , 65 

unijlora  74 

urnigera  64-65,  73,  78 

vicophora  73 

villafrancensis  74 

viridicans  74 

volubilis73,74,76,77,78 

volubis  43 

CLYTIINAE  38, 48-49,  50, 69-70 

clytioides,  Orthopyxis  50,  118-119 
clytioides,  Silicularia  50,  1 18-1 19 
clytioides,  Tubularia  50,  1 14,  1 17, 118 
Cmpanularia  5 1 

comata,  Cymodocea  &  Nemertesia  121,  122 
commensuralis,  Obelia  1 14,  1 17, 118 
commissuralis,  Obelia  1 14,  1 17, 119 
compressa,  Campanularia  60, 6 1 , 64, 66-67,  73 
compressa,  Clytia  74, 82 
compressa,  Orthopyxis  6 1 , 66 
ConcHella  122 

intertexta  122 

conferta,  Campalaria  102-103,  104-105 
conferta,  Campanularia  100,  104 
conferta,  Laomedea41,  102-103,  104 
congdoni,  Laomedea&  Obelia  1 14, 119 


INDEX 


convexa,  Thaumantias  73 

Coppinia  arcta  \  22 

corona,  Obelia  1 14 

coruscans,  Campanularia  1 14 

Coryne65,  122 

Couchella\2\ 

coughtreyi,  Obelia  1 14 

crenata,  Campanularia  52-53,  59, 60;  Fig.  2, 

p.  55 

crenata,  Eucopella  52,  58 
crenata,  Orthopyxis  40, 42,  52,  58-60;  Fig.  5, 

p.  59 

Cuspidella  humilis  123 
cylindrica,  Clytia  42,  73, 80 
cylindrica,  Platypyxis  73 
cymbaloidea,  Epenthesis  73,  74 
cymbaloidea.  Medusa  73 
Cymodocea  1 2 1 

comata  122 

ramosa  1 2 1 

simplex  114,  117,121 


decipiens,  Campanularia  107 
decipiens,  Laomedea  107,  109, 110 
delicata,  Orthopyxis  58,  59, 60 
diaphana,  Eucope  &  Thaumantias  1 14 
dichotoma,  Obelia  40,  41,  43,  45,  73,  96,  100, 

101,   102,   104,   112,   113,  114,  117-119, 

120,121 

dichotoma,  Sertularia  \  1 7 
dischotoma,  Obelia  1 14,  1 17 
discoida,*  Clytia  42, 72-73;  Fig.  8,  p.  72 
discoida,  Oceania  72 
discoidum,  Phialidium  73 
divaricata,  Laomedea  1 1 4 
divisa,  Campanularia  123 
dubia,  Obelia  114 
dubia,  Thaumantias  73 
dumosa,  Lafoea  47, 65,  1 22 


echinata,  Hydractinia  43 
edwardsi,  Campanularia  73, 81 
edwardsi,  Clytia  8 1 
edwardsia,  Clytia  43 
elongata,  Campanularia  107, 110 
elsaeoswaldae,  Clytia  80 
Epenthesis  70, 71 

cymbaloidea  73,  74 

mccradyi  87 

equilateralis,  Obelia  1 14 
Eucalix  48-49, 50-51 

morgansi  5 1 

retroflexus  5 1 
Eucampanularia  47-48,  5 1 , 52, 97 

groenlandica  52 

integra  52 


INDEX 


143 


speciosa  52 
verticillata  52 
volubilis52 
Eucope  70,  71,  1  12 
affinis  71,73,74 
alternata  1  1  4 
articulata  1  1  4 
campanulata  7  1  , 
diaphana  1  14 
exigua  73 
fusiform  is  1  14 
gemmifera  73 
pamsitica  \  1  4 
pictall,  123 
polygena  1  14 
polystyla  1  1 
primordiale  123 
pyriformis  1  1  4 
thaumantias  74 
thaumahtoides  71 
variabilis  73 


73,  74 


73 


bilabiata  58 

caliculata  58,61 

campanularia  58,  60,  6  1  ,  63,  67 

crenata52,58 

eucophora  (&  eucopophora),  Clytia  123 
Eucopium  primordiale  123 
Eulaomeda  97 

flexuosa  105,  117 
Eulaomedea  47,  97-98,  107 

angulata  98 

calceolifera98,  102 

flexuosa  98,  107,  1  17 

pseudodichotoma  1  1  1 
everta,  Campanularia  51,58,  60 
<?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) 
Chance ,  change  &  challenge 

Two  multi-author  volumes  from  one  of  the  foremost  scientific  institutions  in  the  world. 
General  Editor:  P.  H.  Greenwood 

The  Evolving  Earth 

Editor:  L.  R.  M.  Cocks 

The  Evolving  Biosphere 

Editor:  P.  L.  Forey 

In  the  first  volume,  The  Evolving  Earth,  twenty  scientists  have  been  asked  to  review 
the  present  state  of  knowledge  in  their  particular  field,  ranging  from  the  origin  of 
the  Earth,  through  ocean  sediments  and  soils  to  continental  drift  and  palaeogeography. 

In  the  companion  volume,  The  Evolving  Biosphere,  museum  scientists  have  chosen 
an  evolutionary  concept — speciation,  coevolution,  biogeography  etc.  and  related 
this  to  the  group  of  animals  or  plants  in  which  they  are  specialising.  Thus  beetles 
and  birds  exemplify  sympatric  and  allopatric  speciation,  butterflies  mimicry  and 
certain  fishes  explosive  evolution. 

In  both  volumes  the  text  is  supplemented  by  over  one  hundred  specially-commissioned 
pieces  of  two-colour  artwork. 

These  two  books  will  be  invaluable  to  all  sixth-form  and  undergraduate  biology  and 
geology  students. 

The  Evolving  Earth:  276x219  mm,  280pp,  138  line  illustrations,  42  halftones 
The  Evolving  Biosphere:  276x219  mm,  approx.  320pp,  133  line  illustrations 
Published:  May  1981 

Co-published  by  the  British  Museum  (Natural  History),  London  and  Cambridge 
University  Press,  Cambridge. 


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 


1982 


Bulletin  of  the 

British  Museum  (Natural  History) 


Miscellanea 


Zoology  series    Vol  42  No  3    27  May  1982 


The  Bulletin  of  the  British  Museum  (Natural  History),  instituted  in  1949,  is  issued  in  four 
scientific  series,  Botany,  Entomology,  Geology  (incorporating  Mineralogy)  and  Zoology,  and 
an  Historical  series. 

Papers  in  the  Bulletin  are  primarily  the  results  of  research  carried  out  on  the  unique  and 
ever-growing  collections  of  the  Museum,  both  by  the  scientific  staff  of  the  Museum  and  by 
specialists  from  elsewhere  who  make  use  of  the  Museum's  resources.  Many  of  the  papers  are 
works  of  reference  that  will  remain  indispensable  for  years  to  come. 

Parts  are  published  at  irregular  intervals  as  they  become  ready,  each  is  complete  in  itself, 
available  separately,  and  individually  priced.  Volumes  contain  about  300  pages  and  several 
volumes  may  appear  within  a  calendar  year.  Subscriptions  may  be  placed  for  one  or  more  of 
the  series  on  either  an  Annual  or  Per  Volume  basis.  Prices  vary  according  to  the  contents  of 
the  individual  parts.  Orders  and  enquiries  should  be  sent  to: 

Publications  Sales, 

British  Museum  (Natural  History), 
Cromwell  Road, 

London  SW7  5BD, 
England. 


World  List  abbreviation:  Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


©  Trustees  of  the  British  Museum  (Natural  History),  1982 


The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 
Keeper  of  Zoology     :    Dr  J.  G.  Sheals 
Editor  of  Bulletin      :    Dr  C.  R.  Curds 
Assistant  Editor         :    Mr  C.  G.  Ogden 


ISSN  0007-1498  Zoology  series 

Vol  42  No  3  pp  149-225 
British  Museum  (Natural  History) 
Cromwell  Road 
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 


^ 

U 

«»» 

s-orssaa-s-***- 

-  — 

^—            {^^           C^J 

vooo>m>o  —  r-oo'oooo  —  OONTT 

I/I   (N    CN|                                 l~~    1  — 

Q 

^™            ^^           ^^ 

rx|—  i«NO(^)TfTt>orr>ON-^--^too^OO 

oo  t~-  O                  10  m 

C/5 

•* 

r<N  ON                       —  — 

—  m                      — 

£ 

ON  ON         TT 

ON  VO                             O 

(U 

^J     ^^ 

•  fN  i/~>  O                ^         ^O         <N  ON  |n  >/">  r<^ 

OO    Tt                                    >O    Tf 

DA 

C 

T  *?  °f 

"]>ri4Tri40?0fTrTr?rTl<Tri4<Nr? 

7  7  °f           77 

CO 

06 

oo  ^r 

^^    ^^     —     —                            ^^                fXj                fS)     f^.     —     fVj    fsl 

^f  ^"                   rs  ^^ 

8 

^ 

—  —          <N 

CM  r-                      — 

g. 

CO 

c 

»n 

B 

CO 

CO 

. 

CU 

2 

—  NO  r~ 

2^!H  —  ^^^^P^^r^oN^?^ 

r~-  o                  <N  rsi 

(N 

fN  OO                                 — 

Efl 

o 
Z 

^s  ON  ON 

ON  ON  ON  ON  ON  ON  ON  ON  ON  ON  *O  ON  ON  ON  ON 

ON  ON  t    •                        ON  ON 

| 

TTTfin 

r-          NOONOONOO<N</->(~-vO<N»/~>fN'<3- 

ON                 —  ON  ^"  00  00 

1 

U 

2 

m  r-  vo 

ON         roOfN«0«0—  .fNfNOOOOO 

Tf               OOOO 

C/3 

Q 

O  O  m 

t~-         t~-rr)i/~)^1'ONOO^O>O(Nt~~-  —  —  — 

ON                CO  en  f*"i  r  — 

C/5 

C/5 

<N 

—          —                 —          t~-  -T  —  '/-,  — 

^f  r*^  r*^  ^-  ON 

'J£ 

ON 

U 

tN 

'Q 

CN 

ON  (N           r~ 

r-                  ON 

t/5 
jS 

C 

„ 

00 
CO 

g 

.     ON 
—  IO  OO 

\    \    \ 

O  fO  1^ 
m  10 

7     7  T  ^f  "?  T  T  *?  °f  7  'V  7  ^  T 

**?      ^  ir  ^  ^  7 

ro                ON  ON  I"--  ON  CO 

^ 

"b 

rn 

—  —         <N 

t  —                      ^^ 

£ 

u 

a 

— 

c 

03 

m 

•^ 

c 

c 

o 

S 

CU 

CO 

5j 

^> 

*n 

^^         ^^  ^^                ^^         rn         ^"  ON  0*4  ^f  <N 

ro  co  <N  c^i  rs 

Q 

—  —         <N 

oo                   — 

c 
o 

o 

o  o  o 

O         O  0  O  O  0  O  O  O  r-  O  O  NO  O 

O                ON  O  O  O  O 

Z 

5 

"b 

a 

!> 

ON 

0. 

a 

t«j 

>^ 

>/"> 

"Q 

_o 

^H     <O     OO 

(N         O  ^J"  ^  ^  "  ••  t  —  —  "  ^O  NO  NO  ON         ^f 

rt              vo  v~>  w~>  m  —  • 

s 

o 

SO 

r«->         r-j  ON  —         <N 

^g  ro  <N  <N  rn 

k. 

3C 

o 

-s: 

c_ 

CO  X) 

o  -5 

si 

> 

^ 

u. 

2 

.0 

^ 

**                                       i. 

M 

—  c                             o  *- 

U 

_  •a  E             fe    *|  •§  x 

jS                  SB 

g 

"SSg          J3J2-2      C'£12 

00                 O  ^^   CJ    I- 

u 

OU  ^~*   5^^                        *-»    ^_^      UN      f^      TO      IS   *  ^ 

C                 *7j    fj    5S    QJ 

E 

3 

^     TO    fi                  C  .i-    ^    OJQ  iu    o    !>s 

—  -"     •             O     Cd  *O     r" 

C/5 

™"*  ^^  r"  (•        ^  ^       C-  «  i  ^^  r? 

QJ  .  i—     ._-    ^^    >*~*^ 

CO 

C/5 

"_y       -^      "    "i^    ^^    *""*               ^      QJ      ?r         u      W 

TO    ^            c/5     r*    O    prt 

u 

**••  *v      O 

&                    OJD  _*™|     ^^   ^*   O  ""    O  "^ 

*-*    <U     ^  *  ^^   "^    ^^  ,  _^ 

s 

k> 

V 

E'-s 

E  2 

1  1  1|  1  '5  1|  g>  |  S  |  g 

£i  >2  g  s|.§ 

^ 

£ 
- 

Charact 

^g 

00^ 

c 

1>    U 

i  '•=  cx-=  •?•=  —  —  'c  -S  2  2  p 

__    2    l>J=-£-J-'COcO'JQ.flja>w" 
^  J—    CjD.D,D.OO>Oi--t.-  '>? 

Ouj     aCCCggUiUxx^ 

ZUc/5<<<aaoaZOuJUj2 

O"^*J2<u5_oc 

_C-        CH        ^»     UH        -^     "^     "^             , 

"T^J   —     CQ    ^j     O     ^     C3   77^ 
•-    3    >    C  'Q,   3    C    03 
2>O-<C/5C/5<h- 

U 


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. 


References 

Bell,  F.  J.  1893.  Catalogue  of  the  British  Echinoderms  in  the  British  Museum  (Natural  History). 

London  xvii  +  202  pp.,  16  pis. 
1917.  Echinoderma.  1.  Actinogonidiata.  Brit.  Antarct.  (Terra  Nova)  Exped.  1910.  Zool.  4  (1): 

1-10, 2  pis. 
Bernasconi,  I.  1 94 1 .  Dos  nuevas  especies  argentinas  de  "'Luidia'".  Physis,  B.  Aires  19  :  117-118. 

1942.  Los  Asteroideos  sudamericanos  de  la  familia  "Luidiidae".  Physis,  B.  Aires  19  :  252-253. 

1943.  Los  Asteroideos  sudamericanos  de  la  familia  Luidiidae.  An.  Mus.  argent,  B.  Aires  41  :  1-20, 

5  pis. 
1958.  Equinoideos  y  asteroideos  de  la  collection  del  Institute  Oceanografico  de  la  Universidad  de 

San  Pablo.  2.  Bol.  Inst.  oceanogr.  S.  Paulo  1 :  1 19-149, 4  pis. 

1960.  Los  equinodermos  de  la  campana  1958  del  buque  oceanografico  Capitdn  Cdnepa  A.R.A. 


Adas  Trab.  I  Congr.  sudamer.  Zool.,  1959.  2  :  2 1-32,  3  pis. 
Blake,  D.  B.   1973.  Ossicle  morphology  of  some  recent  asteroids  and  description  of  some  west 

American  fossil  asteroids.  Univ.  Calif.  Publs  geol.  Sci.  104  :  1-59,  3  figs,  19  pis. 
1982.  Somasteroidea,  Asteroidea  and  the  affinities  of  Luidia  (Platasterias)  latiradiaia  (Gray), 

1 840,  Palaeontology  25  :  1 67-1 9 1 . 
Boone,  L.  1933.  Coelenterata,  Echinodermata  and  Mollusca.  Bull  Vanderbilt  mar.  Mus.  4  :  1-217,  133 

pis. 


182  A.M.CLARK 

Cadenat,  J.   1941.  Les  Echinodermes  de  la  cote  occidentale  d'Afrique.  Description  d'une  asterie 

nouvelle  de  la  region  du  Cap  Blanc.  Ann.  Soc.  Sci  nat.  Charente  Inf.  N.S.  3  :  53-67,  3  figs. 
Carrera-Rodriguez,  C.  J.  &  Tommasi,  L.  R.  1977.  Asteroidea  de  la  plataforma  continental  de  Rio 

Grande  do  Sul  (Brasil).  Bolm  Inst.  oceanogr.  S.  Paulo  26  (1 ):  5 1-1 30,  33  figs. 
Cherbonnier,  G.  1959.  Echinodermes  de  la  Guyane  franchise.  2.  Bull.  Mus.  natn.  Hist.  nat.  Paris  31  : 

168-1 72,  figs  3, 4. 
1963.  Echinodermes  des  cotes  du  Cameroun  recoltes  par  A.  Crosnier.  Bull.  Mus.  natn.  Hist.  nat. 

Paris  35:  179-193,3  figs. 
Clark,  A.  H.  1945.  A  new  starfish  of  the  genus  Luidia  from  the  coast  of  Georgia.  J.  Wash.  Acad.  Sci. 

35:  19-21. 
1954.  Echinoderms  (other  than  holothurians)  of  the  Gulf  of  Mexico.  Bull.  U.S.  Fish.  Commn  55  : 

373-379. 
Clark,  A.  M.  1952.  Some  echinoderms  from  South  Africa.  Trans.  R.  Soc.  S.  Afr.  33  :  193-221,  3  figs, 

Ipl. 
1953.  Notes  on  asteroids  in  the  British  Museum  (Natural  History).  3.  Luidia.  4.  Tosia  and 

Pentagonaster.  Bull.  Br.  Mus.  nat.  Hist.  Zool.  1  :  379-412,  1 5  figs,  8  pis. 

1955.  Echinodermata  of  the  Gold  Coast.  Jl  W.  Afr.  Sci.  Assn  1  :  16-56, 23  figs,  1  pi. 

1977.  The  South  African  Museum's  Meiring  Naude  cruises.  4.  Echinoderms.  Annls  S.  Afr.  Mus. 


73:  133-147. 
Clark,  A.  M.  &  Court  man-Stock,  J.  1976.  The  echinoderms  of  southern  Africa.  Publs  Br.  Mus.  nat. 

Hist.  No.  776, 277  pp.,  276  figs. 
Clark,  H.  L.  1923.  The  echinoderm  fauna  of  South  Africa.  Annls  S.  Afr.  Mus.  13  :  221^35,  4  figs,  6 

pis. 
1933.  A  handbook  of  the  littoral  echinoderms  of  Porto  Rico  and  the  other  West  Indian  islands. 

Scient.  Surv.  P.  Rico  16  :  1-147,  7  pis. 

1941.  Reports  on  the  scientific  results  of  the  Atlantis  Expeditions  to  the  West  Indies.  The 


echinoderms.  Mem.  Soc.  cubana  Hist.  nat.  15  :  1-54,  10  pis. 
Doderlein,    L.    1920.    Die   Asteriden   der  Siboga-Expedition.   2.    Die  Gattung  Luidia   und   ihre 

stammesgeschichte.  Siboga  Exped.  46b  :  193-291,  5  figs,  3  pis. 
Downey,  M.  E.  1973.  Starfishes  from  the  Caribbean  and  the  Gulf  of  Mexico.  Smithson.  Contr.  Zool. 

No.  126:  1-1 58, 48  pis. 
Duben,  M.  W.  von  (&  Koren,  J.)  1845.  Norriges  Hafs-fauna.  Ofvers.  K.  VetenskAkad  Forh.  1844  : 

110-116. 
1846.    Ofversigt    af    Skandinaviens    Echinodermer.    K.    svenska     VetenskAkad.    Handl. 

1844:  229-328, 6  pis. 
Engel,  H.,  Croes,  A.  F.  &  Schroevers,  G.  F.   1960.  Les  Asterides  recoltees  par  1'expedition  du 

navire-ecole  beige  Mercator,   1935-36.  A.  Les  Asterides  recoltees  sur  les  cotes  orientales  de 

1'Amerique  entre  la  Floride  et  la  Venezuela.  (Engel  &  Schroevers).  B.  Les  Asterides  de  la  cote 

occidentale  d'Afrique.  (Engel  &  Croes).  Bull.  Inst.  roy.  Sci.  nat.  Belg.  36  (54) :  1-10;  10-15, 6  pis. 
Fell,  H.  B.  1963.  The  phylogeny  of  sea-stars.  Phil.  Trans.  246B  :  38 1^35, 2  pis. 
Fisher,  W.  K.  1911.  Asteroidea  of  the  North  Pacific  and  adjacent  waters.  Bull.  U.S.  natn.  Mus.  76  (1) : 

1-419,  122  pis. 

-  1940.  Asteroidea.  'Discovery'  Rep.  20  :  69-316, 23  pis. 

Forbes,  E.  1839.  On  the  Asteriadae  of  the  Irish  Sea.  Mem.  Wernerian  nat.  Hist.  Soc.  8  :  1 13-130, 
2  pis. 

1841.  A  history  ofbritish  starfishes  and  other  animals  of  the  class  Echinodermata.  xx  +  270  pp., 

figs.  London. 

Gray,  I.  E.,  Downey,  M.  E.  &  Cerame-Vivas,  M.  J.  1968.  Seastars  of  North  Carolina.  Fishery  Bull. 

Fish  Wildl.  Serv.  U.S.  67  :  127-163, 40  figs. 
Gray,  J.  E.  1840.  A  synopsis  of  the  genera  and  species  of  the  class  Hypostoma  (Asterias,  Linn.)  Ann. 

Mag.  nat.  Hist.  6  :  1 75-1 84;  275-290. 

Jangoux,  M.  1975.  Note  sur  le  genre  Tethyaster  Sladen  (Echinodermata,  Asteroidea).  Revue  zool.  afr. 
89  (4):  76 1-768, 4  figs. 

-  1978.  Les  etoiles  de  mer  (Echinodermata:  Asteroidea)  recoltees  par  le  M/V  Calamar  au  large  des 
Guyanes  et  du  Venezuela.  Bull.  zool.  Mus.  Univ.  Amsterdam  6(13):  93-101 ,  3  figs. 

John,  D.  D.  &  Clark,  A.  M.  1954.  The  "Rosaura"  Expedition.  3.  Echinodermata.  Bull.  Br.  Mus.  nat. 

Hist.  Zool.  2  (6) :  139-162,  12  figs,  1  pi. 
Koehler,  R.  1895.  Notes  Echinologiques.  Revue  biol.  N.  Fr.  7  :  317-342,  1  pi. 


NOTES  ON  ATLANTIC  ASTEROIDEA  2.  LUIDIIDAE  183 

—  1909.  Echinodermes  provenant  des  campagnes  du  yacht  Princesse- Alice.  Result.  Camp,  scient 
Prince  Albert  34  :  1-3 1 7,  32  pis. 

1911.  Mission  Gruvel  sur  la  cote  occidentale  d'Afrique.  Echinodermes.  Annls  Inst.  oceanogr. 


Monaco  2  (5):  1-25,  3  pis. 

—  1914.  Asteroidea,  Ophiuroidea  et  Echinoidea.  Beitr.  Kennt.  Meeresfauna  Westafr.  1  (2) :  129-303, 
12  pis. 

—  1921.  Echinodermes.  Faune  Fr.  1  :  1-210,  153  figs. 

1923.  Asteries  et  Ophiures  recueillis  par  1'expedition  antarctique  suedoise  1901-03.  Further  zool. 


Results  Swed.  Antarct.  Exped.  I  (1) :  1-145,  1  fig.,  15  pis. 
Lamarck,  J.  B.  P.  A.  de  1816.  Histoire  naturelle  des  animaux  sans  vertebres.  Ed.  1.  2:  522-568:  3  : 

1-59;  60-76.  Paris. 

Linnaeus,  C.  1758.  Systema  Naturae.  Ed.  10. 1  :  824  pp.  Holmiae. 

Ludwig,  H.  1 897.  Die  Seesterne  des  Mittelmeeres.  Fauna  Flora  Golf.  Neapel  24  :  1-49 1 ,  1 2  pis. 
Liitken,  C.  1859.  Bidrag  til  Kundskab  om  de  ved  Kysterne  af  Mellem-  og  Syd-America  levende  arter  af 

Sostjerner.  Vidensk.  Meddrdansk.  naturh.  Foren.  1859  :  25-96. 
Mcknight,  D.  G.   1977.  Classification  of  recent  paxillosid  sea-stars  (Asterozoa:  Echinodermata). 

NZOIRec.  3(12) :  1 13-1 19,  figs. 
Madsen,  F.  J.  1950.  The  echinoderms  collected  by  the  Atlantide  Expedition,  1945-46.  1.  Asteroidea. 

Atlantide  Rep.  1  :  167-222,  1 1  figs,  3  pis. 
Marenzeller,   E.   von    1893.   Veroffentlichungen  der  Commission  fur  Enforschung  des  ostlichen 

Mittelmeeres.  Sber.  Akad.  Wiss.  Wien  Math,  naturw.  Kl.  102  :  1-5. 
Mortensen,  T.  1925.  Echinodermes  du  Maroc  et  de  Mauritanie.  Bull.  Soc.  Sci.  nat.  Maroc  5  :  178-187, 

I  pi. 

1 927.  Handbook  of  the  Echinoderms  of  the  British  Isles,  ix  +  47 1  pp.,  269  figs.  London. 

1933a.  Echinoderms  of  South  Africa  (Asteroidea  and  Ophiuroidea).   Vidensk.  Meddr  dansk 

naturh.  Foren.  93  :  2 1 5^400, 9 1  figs,  1 2  pis. 

19336.  The  echinoderms  of  St  Helena.  Vidensk.  Meddr  dansk  naturh.  Foren.  93  : 401-472,  29 


figs,  3  pis. 
Miiller,  J.  &  Troschel,  F.  H.  1840.  Mber.  Akad.  Wiss.  Berlin  1840  :  100-106. 

1842.  System  der  Aster iden.  1 .  Asteriae.  2.  Ophiuridae.  xx  +  1 34  pp.,  12  pis.  Braunschweig. 

Nataf,  G.  &  Cherbonnier,  G.  1973.  Les  asterides  d'Afrique  occidentale,  utilisation  du  microscope 

electronique  a  balayage  pour  etude  systematique  des  Luidia.  Bull.  Mus.  natn.  Hist.  nat.  Paris  (Zool) 

No.  81  :  69-101, 9  pis. 

Pennant,  T.  1 777.  British  Zoology.  Ed.  4. 4  :  1-1 54, 93  pis.  London. 
Perrier,  E.  1 869.  Recherches  sur  les  Pedicellaires  et  les  Ambulacres  des  Asteries  et  les  Oursins.  1 88  pp., 

2  pis.  Paris  [Also  in  Annls  Sci.  nat.  12[1869] :  197-304;  13[1870] :  1-81.] 
1875.  Revision  de  la  collection  de  Stellerides  du  Museum  d'Histoire  Naturelle  de  Paris.  384  pp. 

Paris.  [Also  in  Archs  Zool.  exp.  gen.  4[1875] :  265^50;  5[1876] :  1-104;  209-309.] 

1881.  Description  sommaire  des  especes  nouvelles  d'Asteries.  Bull.  Mus.  comp.  Zool.  Harv. 


9:  1-31. 

—  1882.  In:  Milne  Edwards,  A.  Rapport  sur  les  traveaux  de  la  Commission  chargee  par  M.  le 
ministre  de  1'Instruction  publique  d'etudier  la  faune  sous-marine  dans  le  grandes  profundeurs  de  la 
Mediterranee  et  de  1'Atlantique.  (Echinodermes).  Arch.  Missions  sci.  litt:  Rapp.  et  Instr.  ser  3,  9  : 
46^9,  fig. 

—  1884.  Memoire  sur  les  Etoiles  de  Mer  recueillies  dans  la  Mer  des  Antilles  et  le  Golfe  de  Mexique. 
Nouv.  Archs  Mus.  Hist.  nat.  Paris  (2)6  :  127-276,  10  pis. 

1894.  Stellerides.  Exped.  scient.  Travailleur-  Talisman.  431  pp.,  26  pis.  Paris. 


Philippi,  R.  1837.  Ueber  die  mit  Asterias  aurantiaca  verwandten  und  verwechselten  Asterian  der 

sicilianischen  Kiiste.  Arch.  Naturgesch.  3  :  193-194. 
Sars,  M.  1835.  Radiater.  Beskrivelser  og  Jagttagelser  over  nogle  mdrkelige  eller  nye  i  Havel  ved  den 

Bergenske  Kyst  levende  Dyr.  pp.  39^6.  Bergen. 
Say,  T.  1 825.  On  the  species  of  the  Linnaean  genus  Asterias  inhabiting  the  coast  of  the  United  States.  J. 

Acad.  nat.  Sci.  Philad.  5  :  141-154. 
Sibuet,  M.  1975.  Asterides  abyssales  de  1'Atlantique  sud.  Bull.  Mus.  natn.  Hist.  nat.  Paris  (Zool.)  No. 

199:  28 1-296,  2  figs,  pi. 

Sladen,  W.  P.  1889.  Asteroidea.  Rep.  scient.  Results  Voy.  Challenger  Zool  30  :  1-935,  1 18  pis. 
Spencer,  W.  K.  &  Wright,  C.  W.  1966.  Asterozoans.  In:  Moore,  R.  C.  [Ed.]  Treatise  on  invertebrate 

paleontology.  U.  Echinodermata  3(1):  4-107,  89  figs.  Geological  Society  of  America  Inc.:  University 

of  Kansas  Press. 


184  A.M.CLARK 

Studer,  T.    1884.   Verzeichniss  der  wahrend  der  Reise  S.M.S.   Gazelle  urn  die  Erde    1874-76 

gessammelten  Asteriden  und  Euryaliden.  Phys.  Abh.  K.  Akad.  Wiss.  Berlin  [1883,  1884]  3  :  1-64, 

5  pis. 
Sussbach,  S.  &  Breckner,  A.  191 1.  Die  Seeigel,  Seesterne  und  Schlangensterne  der  Nord-  und  Ostsee. 

Wiss.  Meeresunters.  KielN.S.  12  :  167-300,  3  pis. 
Tattersall,  W.  M.  &  Sheppard,  E.  M.  1934.  Observations  on  the  Bipinnaria  of  the  asteroid  genus 

Luidia.  James  Johnstone  Memorial  Volume:  Lanes  Sea-Fisheries  Laboratory,  Liverpool,  pp.  35-61, 

8  figs. 
Tommasi,  L.  R.  1958.  Os  equinodermas  do  literal  de  Sao  Paulo.  2.  Contrcoes  avuls.  Inst.  oceanogr.  S. 

Paulo  No.  2:  1-2 7, 6  pis. 
1970.  Lista  dos  asteroides  recentes  do  Brasil.  Contrcoes  Inst.  oceanogr.  Univ.  S.  Paulo  No.  18  : 

1-6 1,60  figs. 

1974.  Equinodermes  do  Brasil.  3.  P ubl.  Inst.  oceanogr.  Univ.  S.  Paulo  No.  369  :  1-1 5,  12  figs. 


Tortonese,  E.  1 965.  Echinodermata.  Fauna  d'ltalia.  6  :  xv  +  422,  1 86  figs.  Bologna. 

Ummels,  F.  1963.  Asteroids  from  the  Netherlands  Antilles  and  other  Caribbean  localities.  Stud.  Fauna 

Curacao  15  :  72-101 , 3  figs,  9  pis.  [Also  in  Natuurwet.  Stud.  Suriname  No.  30.] 
Ursin,  E.  1960.  A  quantitative  investigation  of  the  echinoderm  fauna  of  the  central  North  Sea.  Medd. 

Danmarks  Fisk.  Havundersog.  N.S.  2  (24) :  1-204, 96  figs. 
Verrill,  A.  E.  1885.  Results  of  the  explorations  made  by  the  steamer  Albatross  off  the  northern  coast  of 

the  United  States  in  1883.  Rep.  U.S.  Fish.  Commn  1883  :  503-543,  pis  13-19. 
1899.  Revision  of  certain  genera  and  species  of  starfishes  with  descriptions  of  new  forms.  Trans. 

Conn.  Acad.  Arts  Sci.  1899  :  145-234,  pis  24-30. 

1915.  Report  on  the  starfishes  of  the  West  Indies,  Florida  and  Brazil.  Bull.  Labs  nat.  Hist.  St.  Univ. 


/a  7(1):  1-232, 29  pis. 
Walenkamp,  J.  H.  C.  1976.  The  asteroids  of  the  coastal  waters  of  Surinam.  Zoo/.  Verh.  Leiden  No. 

147:  1-91,  16  figs,  18  pis. 
1979.  Asteroidea  (Echinodermata)  from  the  Guyana  shelf.  Zoo/.  Verh.  Leiden  No.  170  :  1-97,  31 

figs,  20  pis. 
Zoppi  de  Roa,  E.  1967.  Contribucion  al  estudio  de  los  Equinodermos  de  Venezuela.  Acta  biol.  venez. 

5:  267-333, 29  figs. 

Manuscript  accepted  for  publication  25  June  1 98 1 


New  and  little  known  species  of  Oncaeidae 
(Cyclopoida)  from  the  Northeastern  Atlantic 

S.  J.  Malt 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London 
SW7  5BD 


Introduction 

A  large  number  of  new  species  and  genera  belonging  to  the  Oncaeidae  have  been  described 
in  recent  years.  Many  common  species  have  been  re-examined  and  recognized  as  polytypic, 
comprising  two  or  more  varieties  (Tanaka,  1960;  Moulton,  1973;  Ferrari,  1975;  Boxshall, 
1977)  or  two  or  more  species  (Heron,  1977).  Now  that  some  of  the  better  known  species  have 
been  redescribed  in  greater  detail  it  is  obvious  that  many  of  the  newly  described  species  are 
also  in  need  of  more  exact  description.  Taxonomic  study  of  oncaeids  has  become  a  very 
laborious  process,  demanding  the  recognition  of  relatively  small  morphological  differences 
in  these  microscopic  animals,  half  the  known  species  of  which  are  less  than  O6  mm  in  body 
length. 

Most  records  of  oncaeids  are  from  the  epipelagic  zone,  but  the  more  recent  discovery  of 
several  species  extending  down  into  the  deep  bathypelagic  zone  (below  2000  m)  indicates 
that  previous  records  reflect  the  limitations  of  the  sampling  programmes  rather  than  any  true 
restrictions  in  their  depth  range. 

A  new  species,  Oncaea  heronae,  and  two  little  known  species  belonging  to  the  genus 
Oncaea  are  here  described.  The  genera  Conaea,  Epicalymma  and  Myctospictosum  are 
synonymized  with  Oncaea.  New  locality  records  are  presented  for  six  species  and  two 
previously  undescribed  males  and  two  juveniles  of  known  species  are  also  described. 

Materials  and  methods 

The  material  described  below  was  donated  to  the  British  Museum  (Natural  History)  by  the 
Institute  of  Oceanographic  Sciences  (Wormley)  and  the  Marine  Biological  Association 
(Plymouth).  It  was  collected  in  April  1977  at  station  9541  (20°  N  21°  W)  in  the  NE  Atlantic 
during  cruise  82  of  the  R.R.S.  Discovery,  and  in  September  1979  at  the  mouth  of  the  English 
Channel  (48°  N  7°30'  W)  during  a  cruise  of  the  R.V.  Sarsia.  The  Discovery  material  was 
collected  by  a  80  jn  mesh  diatom  net  attached  to  the  RMT  1  +  8  net  system.  Sarsia  material 
was  taken  using  a  pump  system  with  80  n  and  200  //  mesh  filters. 

Material  was  preserved  in  10%  sea  water  formalin,  stained  in  chlorazol  black,  dissected  in 
lactophenol  and  mounted  in  polyvinyl  lactophenol.  Drawings  were  made  with  the  aid  of  a 
camera  lucida.  Specimens  were  measured  using  an  eyepiece  micrometer,  total  body  length 
being  the  distance  from  the  tip  of  the  rostrum  to  the  apex  of  the  caudal  rami. 

Descriptions  of  species 
Oncaea  tregoubo vi  Shmeleva,  1968 
O.  tregoubovi  Shmeleva,  1968  :  1784-1785,  figs  1-12. 

DESCRIPTION  Female.  Ratio  of  prosome  and  urosome  lengths  1  -8  :  1 .  Prosome  elongate,  oval 
in  dorsal  aspect  (Fig.  la).  Mean  body  length  of  3  specimens  0'34  mm  (range  0'31  to 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  42  (3) :  1 85-205  Issued  27  May  1 982 


186  S.  J.  MALT 

0*37  mm).  Third  prosome  segment  without  a  dorsal  projection.  Head  (first  prosome  division) 
1*3  times  longer  than  wide.  Proportional  lengths  of  ursome  and  caudal  rami  8  :  57  : 
5:4:  13  :  13  (Fig.  Ib).  Genital  apparatus  located  on  dorsal  surface  anterior  to  midpoint  of 
genital  segment;  each  area  armed  with  a  setule.  Maximum  width  of  genital  segment  at  level 
of  genital  apparatus;  length  to  width  ratio  1  '75  :  1 .  Caudal  rami  twice  as  long  as  wide. 

First  antenna  6-segmented  (Fig.  Ic);  armature:  1-2,  II-7,  III-3,  IV-3,  V-2,  VI-5  (some 
elements  may  be  missing);  ratio  of  segment  lengths  10  :  13  :  46  :  13:8:  10.  Second  antenna 
3 -segmented  (Fig.  Id);  first  segment  bearing  1  long  pinnate  seta  distally;  second  segment 
bearing  a  row  of  denticles  along  internal  surface;  terminal  segment  longer  than  second  and 
bearing  5  curved  spines  and  2  curved  setae.  Mandible  bearing  5  elements  (Fig.  le):  a  stout 
seta  on  external  surface,  2  broad  blades,  1  bearing  a  row  of  setules  along  internal  edge,  and 
the  other  unarmed,  and  2  setae,  1  long  and  hirsute,  the  other  small  and  unarmed.  First 
maxilla  bilobed  (Fig.  1  f);  bearing  1  seta  on  internal  surface;  2  setae  on  internal  lobe  and  2 
setae,  a  setose  seta  and  a  spine  on  external  lobe.  Second  maxilla  2-segmented  (Fig.  Ig); 
second  segment  produced  distally  as  an  elongate,  curved,  bilaterally  spinulose  claw;  also 
having  an  external  pinnate  seta  and  an  internal  bilaterally  spinulose  element.  Maxilliped 
4-segmented  (Fig.  1  h);  first  segment  unarmed;  internal  surface  of  second  segment  with  2 
spines,  distal  spine  bidentate;  third  segment  reduced;  terminal  segment  produced  as  a  long 
claw  and  armed  with  a  row  of  setules  on  concave  surface,  an  internal  basal  spine  and  an 
external  basal  setule. 

Endopodites  and  exopodites  of  natatory  legs  3-segmented  (Figs  2a-d);  external  spines  of 
exopodites  bilaterally  serrate;  armature  of  natatory  legs  as  follows: 

Coxa         Basis  Endopodite  Exopodite 

Legl  0-0  1-1  0-1;  0-1;  0,1, 5  I-0;I-1;  III,  1,4 

Leg2  0-0  1-0  0-1; 0-2;  II,  1, 3  1-0;  !*-!*;  Ill,  1, 5 

Leg  3  0-0  1-0  0-1;  0-2;  II,  1, 2  1-0;  I-1;II,1, 5 

Leg  4  0-0  1-0  0-1;  0-2;  II,  1,1  I-0;I-1;II,1, 5 

*  =  segment  missing  from  figured  specimen  (Fig.  2b),  present  in  other  material. 

Terminal  spines  of  exopodites  longer  than  terminal  segments  bearing  them  but  terminal 
spines  of  endopodites  shorter  than  terminal  segments;  each  terminal  spine  bordered  by  a 
serrate  membrane  unilaterally;  endopodites  of  legs  1-3  terminate  in  conical  projections;  all 
setae  on  legs  plumose.  Fifth  leg  comprising  a  small  cylindrical  segment  bearing  a  single 
terminal  seta  on  body  surface  near  leg  (Fig.  1  i). 

MATERIAL  EXAMINED.  399  from  sample  9541.24,  2980-3560  m.  20°  N  21°  W.  BM(NH) 
1981.114. 

REMARKS.  This  species  was  identified  by  the  following  combination  of  characters:  the 
relative  lengths  of  the  segments  of  the  first  and  second  antennae,  the  armature  of  the 
maxilliped  and  natatory  legs,  and  the  relative  dimensions  of  the  urosome  segments.  Some 
minor  differences  were  found  between  the  present  material  and  the  original  material  figured 
by  Shmeleva  (1968).  Most  notably  Shmeleva  (1968  :  Fig.  9)  shows  the  fourth  natatory  leg 
with  the  internal  margin  of  the  third  endopodite  segment  unarmed,  whereas  the  present 
specimens  have  a  single  seta.  The  absence  of  this  seta  is  atypical  for  Oncaea  species.  It  may 
be  significant  that  Shmeleva  does  not  mention  this  character  in  the  text  of  the  description, 
and  the  original  drawing  may  be  inaccurate  or  based  on  a  specimen  with  incomplete 
armature.  This  species  has  only  been  recorded  once  before,  from  the  southern  Adriatic 
where  it  was  taken  between  200  and  300  metres  depth. 

Oncaea  ivlevi  Shmeleva,  1966 

O.  ivlevi  Shmeleva,  1966  :  932-933,  Plate  I,  figs  1-1 1 . 

DESCRIPTION.  Female.  Ratio  of  prosome  and  urosome  lengths  1-7:1  (Fig.  3a).  Body  length  of 

3  specimens  0*33  mm.  Third  prosome  segment  without  a  dorsal  projection  (Fig.  3b).  Head 


NEW  AND  LITTLE  KNOWN  SPECIES  OF  ONCAEIDAE  187 

1  •  1  times  longer  than  wide.  Proportional  lengths  of  urosome  segments  and  caudal  rami 
8  :  50  :  7:6:18:11  (Fig.  3c).  Genital  apparatus  situated  anterior  to  midpoint  of  dorsal 
surface  of  genital  segment;  each  area  armed  with  a  setule.  Maximum  width  of  genital 
segment  at  the  level  of  genital  apparatus;  length  to  width  ratio  2:1.  Genital  segment  not 
markedly  swollen.  Caudal  rami  nearly  twice  as  long  as  wide,  curving  outwards  and  each 
bearing  a  stout  spine  at  external  corner. 

First  antenna  armature  similar  to  O.  tregoubovi  (Fig.  3d);  ratio  of  segment  lengths  19  :  19  : 
35  :  12  \  6  :  9.  Second  antenna  armature  similar  to  O.  tregoubovi  except  that  first  segment 
armed  with  a  row  of  setules  on  internal  surface,  and  second  segment  with  a  row  of  spinules; 
terminal  segment  equal  in  length  to  second  segment  (Fig.  3e).  Mandible  and  first 
maxilla  lost  during  dissection.  Second  maxilla  like  that  of  0.  tregoubovi  (Fig.  30-  Maxilliped 
4-segmented  (Fig.  3g);  first  segment  unarmed;  internal  surface  of  second  segment  with  2 
spines,  longer  distal  spine  spinulose;  second  segment  also  bearing  a  row  of  denticles  along 
internal  surface;  third  segment  reduced;  terminal  claw  bearing  a  row  of  spinules  along 
concave  surface  and  an  internal  basal  spine. 

Armature  of  natatory  legs  similar  to  O.  tregoubovi  (Figs  3h-k),  but  terminal  segment  of 
second  endopodite  bearing  1  spine  on  external  surface  instead  of  2;  endopodites  2,  3  and  4 
tipped  with  relatively  small  conical  projections;  terminal  spines  of  exopodites  1  and  2 
slightly  longer  than  terminal  segments;  those  of  exopodites  3  and  4  slightly  shorter;  all 
terminal  spines  of  endopodites  shorter  than  terminal  segments  and  bilaterally  flanged. 
Fifth  leg  comprising  a  small  cylindrical  free  segment  bearing  2  setae,  outer  double  length  of 
inner;  and  a  seta  on  body  surface  near  the  leg. 

Male.  Ratio  of  prosome  and  urosome  lengths  1-6  :  1  (Fig  4a  and  b).  Mean  body  length  of  6 
specimens  0-33  mm  (range  0'28  to  0'35  mm).  Head  I'l  times  longer  than  wide.  Proportional 
lengths  of  urosome  segments  and  caudal  rami  7  :  60  :  2  :  3  :  2  :  13  :  13.  Genital  lappets 
produced  into  small  postero-lateral  processes  (Fig.  4c).  Genital  segment  with  length  to  width 
ratio  2-1  :  1 .  Caudal  rami  twice  as  long  as  wide,  as  in  female  possessing  a  stout  spine  on  each 
ramus. 

Mouthparts  similar  to  those  of  female  except  first  antenna  and  maxilliped.  Three  distal 
segments  of  the  first  antenna  fused.  Maxilliped  3 -segmented  (Fig.  4d);  first  segment 
unarmed;  second  segment  bearing  2  short  spines  and  a  row  of  setules  on  internal  surface; 
terminal  claw  bearing  spinules  on  concave  surface  and  a  stout  external  basal  spine.  Legs  1-5 
similar  to  those  of  female. 

MATERIAL  EXAMINED.  399  and  6dtf  from  sample  F166-F170,  8(MO  m,  48°  N  7°30'  W. 
BM(NH) 99  1981. 115-116,^  1981. 117-1 19  and  1981. 124. 

REMARKS.  This  species  was  identified  by  the  following  combination  of  characters:  the 
armature  of  the  natatory  legs,  the  relative  dimensions  of  the  urosome  segments,  and  the 
presence  of  the  stout  caudal  spine.  The  present  specimens  differ  from  Shmeleva's 
descriptions  in  a  few  minor  details:  in  the  possession  of  3  terminal  setae  in  association  with 
the  spine  on  the  caudal  rami  rather  than  2,  the  possession  of  an  extra  row  of  spinules  on  both 
the  distal  spine  of  the  second  segment  of  the  female  maxilliped  and  the  terminal  claw.  There 
are  also  small  differences  in  the  lengths  of  the  terminal  exopodite  spines.  This  species  has 
also  been  recorded  from  the  southern  Adriatic  where  it  was  taken  between  50  and  100  metres 
depth  (Shmeleva,  1966)  and  from  the  Atlantic  at  15  stations  from  9°  S  25°  W  to  9°  S  45°  W 
between  10  and  2000  metres  depth  (Shmeleva,  1969). 

Oncaea  hispida  (Heron,  1977)  Comb.  nov. 
Conaea  hispida  Heron,  1977  :  90-95,  Figs  33h-j,  34a-k. 

DESCRIPTION.  Female.  Ratio  of  prosome  and  urosome  lengths  1-75  :  1  (Fig.  5a).  Body  length 
of  1  specimen  0'56  mm.  Third  prosome  segment  without  dorsal  projection.  Proportional 
lengths  of  urosome  segments  and  caudal  rami  8:48:7:7:  18:  12  (Fig.  5b).  Genital 
apparatus  located  on  dorsal  surface  anterior  to  midpoint  of  genital  segment,  each  area  armed 


188  S.  J.  MALT 

with  a  setule.  Maximum  width  of  genital  segment  at  level  of  genital  apparatus;  length  to 
width  ratio  1'6  :  1.  Caudal  rami  1*3  times  as  long  as  wide.  Appendages  similar  to  those 
described  by  Heron  with  only  minor  differences  apparent  (Figs  5c-h  and  6a-d). 

Male.  Ratio  of  prosome  and  urosome  lengths  2  :  1  (Fig.  6e).  Body  length  of  1  specimen 
0'55  mm.  Head  1-2  times  longer  than  wide.  Proportional  lengths  of  urosome  segments  and 
caudal  rami  10:51  :3:3:3:  18:  12  (Fig..  6f).  Genital  lappets  produced  into  small  posterior 
processes.  Genital  segment  with  length  to  width  ratio  1'5  :  1.  Caudal  rami  approximately  as 
long  as  wide. 

Mouthparts  similar  to  those  of  female  except  first  antenna  and  maxilliped.  The  3  distal 
segments  of  first  antenna  fused.  Maxilliped  3-segmented  (Fig.  6g);  first  segment  unarmed; 
second  segment  bearing  2  rows  of  dentiform  processes  on  internal  surface;  terminal  claw 
unarmed.  Legs  1-5  similar  to  those  of  female. 

Fifth  Copepodid  (female).  Ratio  of  prosome  and  urosome  lengths  2  :  1  (Fig  6h).  Body 
length  of  1  specimen  O46  mm.  Head  approximately  as  long  as  wide.  Urosome  4-segmented 
(Fig.  6h).  Proportional  lengths  of  urosome  segments  and  caudal  rami  10  :  45  :  6  :  29  :  10. 
Genital  segment  without  visible  genital  apparatus,  length  to  width  ratio  1'5  :  1.  Caudal  rami 
1-2  times  as  long  as  wide.  Appendages  similar  to  those  of  adult  female  except  that  proximal 
seta  on  second  segment  of  juvenile  maxilliped  is  pinnate  (Fig.  6j). 

MATERIAL  EXAMINED.  19,  Irf,  19  juvenile  from  sample  9541.24,  3980-3960  m,  20°  N  21°  W. 
BM(NH)1981.120-122. 

REMARKS.  This  species  is  here  transferred  from  the  genus  Conaea  to  Oncaea  (for  discussion 
see  page  191),  and  was  identified  by  the  following  combination  of  characters:  the  length  of 
the  terminal  segment  of  the  second  antenna  and  its  armature,  the  armature  of  the  maxilliped 
and  the  length  of  the  third  segment  of  the  fourth  endopodite.  There  are  minor  differences 
between  the  present  material  and  Heron's  description  of  the  female  in  the  ratio  of  urosome 
segment  lengths,  in  the  proximal  spine  of  the  second  segment  of  the  maxilliped  which  lacks 
the  spinules  in  the  adult  female  (but  not  in  the  juvenile),  and  the  fifth  leg  has  2,  rather  than  1, 
tubercles  each  bearing  at  seta.  These  differences  are  considered  to  represent  geographical 
variation  within  the  species,  as  the  only  previous  records  of  this  species  are  from  the  antarctic 
zone  of  the  southwest  Pacific  between  1000  and  2000  metres  depth.  The  male  and  fifth 
copepodid  have  not  previously  been  described. 


Oncaea  heronae  sp.  nov. 

DESCRIPTION.  Female.  Ratio  of  prosome  and  urosome  lengths  1-5  :  1.  Prosome  elongate 
and  oval  in  dorsal  aspect  (Fig.  7a).  Mean  body  length  of  4  specimens  0'33  mm  (range  0'3 1  to 
0'35  mm).  Third  prosome  segment  without  a  dorsal  projection.  Head  I'l  times  longer  than 
wide.  Proportional  lengths  of  urosome  segments  and  caudal  rami  9:50:7:7:  13:  14  (Fig. 
7b).  Genital  apparatus  located  on  dorsal  surface  anterior  to  midpoint  of  genital  segment. 
Maximum  width  of  genital  segment  at  level  of  genital  apparatus;  length  to  width  ratio 
1  -2  :  1 .  Caudal  rami  twice  as  long  as  wide. 

First  antenna  6-segmented  (Fig.  7c);  armature:  1-3,  II-6,  III-3,  IV-1,  V-2,  VI-5  (some 
elements  may  be  missing);  ratio  of  segment  lengths  12  :  22  :  34  :  13  :  7  :  12.  Second  antenna 
3-segmented  (Fig.  7d);  first  segment  bearing  1  long  pinnate  seta  distally;  second  segment 
unarmed;  terminal  segment  equal  to  second  segment  in  length,  bearing  1  curved  spine  and  3 
curved  setae  proximally  and  5  curved  spines  and  1  curved  seta  distally.  Mandible  lost  during 
dissection.  First  maxilla  (Fig.  7e)  and  second  maxilla  (Fig.  70  with  armature  similar  to  O. 
tregoubovi.  Internal  surface  of  second  segment  of  maxilliped  bearing  2  overlapping  rows  of 
setules  and  2  spines;  larger  distal  spine  bilaterally  dentate  (Fig.  7g);  terminal  claw  with 
smooth  concave  surface;  single  internal  basal  spine. 

Armature  of  natatory  legs  similar  to  O.  tregoubovi  except  for  armature  of  terminal 
segments  of  endopodites  2-4: 


NEW  AND  LITTLE  KNOWN  SPECIES  OF  ONCAEIDAE  1  89 

Coxa         Basis  Endopodite  Exopodite 

Legl  0-0  1-1  0-1;  0-1;  0,1, 5  1-0;  I-1;III,  1,4 

Leg  2  0-0  ?-0  0-1;  0-2;  1, 1, 3  I-0;I-1;III,  1, 5 

Leg  3  0-0  1-0  0-1;  0-2;  0,1, 2  1-0;  1-1 ;  II,  I,  5 

Leg  4  0-0  ?-0  0-1;  0-2;  0,1,1  I-0;I-1;  II,  1, 5 

Bilaterally  serrate  spines  on  external  margins  of  exopodites  lack  pronounced  flanges  (Figs 
7h-k);  first  endopodite  with  conical  terminal  projection;  all  terminal  spines  shorter  than 
terminal  segments  except  terminal  spines  of  fourth  exopodite  and  endopodite,  these  being 
longer  than  terminal  segments.  Fifth  leg  comprising  small  free  cylindrical  segment  bearing  2 
terminal  setae,  the  longer,  internal  seta  extending  to  level  of  genital  apertures. 

Male.  Not  known. 

MATERIAL  EXAMINED.  499  from  sample  9541.24,  3980-3960  m,  20°  N  21°W.  BM(NH) 
Holotype  1981.125,  paratypes  1981.126. 

ETYMOLOGY.  This  species  has  been  named  after  Gayle  A.  Heron  of  the  University  of 
Washington,  Seattle,  in  recognition  of  her  work  on  the  taxonomy  of  the  Oncaeidae. 

REMARKS.  The  second,  third  and  fourth  endopodites  of  O.  heronae,  O.  brodskii  Shmeleva, 
1968  and  O.  longipes  Shmeleva,  1968  have  the  same  unusual  armature  described  above,  but 
O.  heronae  can  be  distinguished  by  other  small  differences  in  the  armature  of  the  natatory 
legs,  by  the  2  overlapping  rows  of  setules  on  the  second  segment  of  the  maxilliped,  and  by  the 
structure  of  the  free  segment  of  the  fifth  leg  which  is  elongate  in  O.  longpipes  and  reduced  to 
a  tubercle  in  O.  brodskii,  but  small  and  delimited  from  the  body  segment  in  O.  heronae. 

Oncaea  setosa  Heron,  1977 
O.  setosa  Heron,  1977  :  73,  figs  22a-h. 

DESCRIPTION.  Female.  Mean  body  length  of  4  specimens  0'5  mm  (range  0'42  to  0*58  mm). 
Body  figured  (Figs  8a-c).  Appendages  of  O.  setosa  similar  to  those  described  by  Heron  with 
only  minor  differences  apparent  (Figs  8d-n). 

Male.  Ratio  of  prosome  and  urosome  lengths  1-5:1  (Figs  9a  and  b).  Body  length  of  1 
specimen  0'58  mm.  Head  1-2  times  longer  than  wide.  Proportional  lengths  of  urosome 
segments  and  caudal  rami  12  : 47  :  3  :  3  :  3  :  18  :  14  (Fig.  9c).  Genital  lappets  extending 
postero-laterally  into  acute  points. 

Mouthparts  like  those  of  female,  except  first  antenna  and  maxilliped.  First  antenna 
4-segmented  (Fig.  9d).  Maxilliped  3 -segmented  (Fig.  9e);  second  segment  bearing  2  spines  on 
internal  surface  and  3  groups  of  setules;  terminal  claw  bearing  a  stout  internal  basal  spine. 

First  and  second  natatory  legs  like  those  of  female,  but  terminal  spines  of  third  and  fourth 
exopodites  longer  than  those  of  female,  being  respectively  1-5  times  and  twice  the  length  of 
terminal  segments.  Two  spines,  representing  the  fifth  leg,  borne  on  tubercle  not  clearly 
delimited  from  body. 

Fifth  Copepodid  (female).  Ratio  of  prosome  and  urosome  lengths  3  :  1  (Figs  9f  and  g).  Body 
length  of  1  specimen  0'51  mm.  Head  1-2  times  longer  than  wide.  Urosome  4-segmented 
(Fig.  9h).  Proportional  lengths  of  urosome  segments  and  caudal  rami  12  :  40  :  8  :  24  :  16. 
Mouthparts  similar  to  those  of  adult  female  except  maxilliped,  bearing  a  group  of  setules  on 
internal  surface  of  second  segment  (Fig.  9i).  Natatory  legs  bearing  terminal  spines 
proportionally  longer  than  those  of  adult  female  (Figs  9j  and  k,  lOa  and  b).  (lOj  and  k,  1  la 
and  b). 

MATERIAL  EXAMINED.  499(1  lost),  Irf,  1 9  juvenile  from  sample  954 1.24,  3980-3960  m,  20°  N 
21°  W.BM(NH)  1981. 127-129. 

REMARKS.  This  species  was  identified  by  the  cluster  of  spinules  on  the  second  segment  of  the 
second  antenna,  the  armature  of  the  maxilliped  and  the  relative  dimensions  of  the  urosome 
segments.  The  female  differs  from  Heron's  description  in  some  minor  details:  the  terminal 


190  S.  J.  MALT 

spines  of  the  third  and  fourth  endopodites  are  longer  in  proportion  to  their  terminal 
segments  in  the  present  material.  The  male  and  the  fifth  copepodid  have  not  previously  been 
described.  Heron  reported  O.  setosa  from  2  stations  in  the  antarctic  zone  of  the  southwest 
Pacific  between  1000  and  2000  metres  depth. 

Oncaea  rotunda  Heron,  1977 
O.  rotunda  Heron,  1977  :  77-79,  figs  24o-q,  25a-m. 

DESCRIPTION.  Female,  body  length  of  1  specimen  0-55  mm.  Body  (Figs  lOc-e)  and  second 
antenna  (Fig.  1  Of)  figured. 

MATERIAL  EXAMINED.  19  from  sample  9541.24.  3980-3960  m,  20°  N  21°  W.  BM(NH) 
1981.130. 

REMARKS.  This  specimen  was  assigned  to  O.  rotunda  because  of  its  distinctive  second 
antenna,  which  has  a  very  short  terminal  segment  relative  to  the  second  segment  and  bears 
relatively  short  terminal  spines  and  setae.  Also  the  terminal  segment  of  the  fourth 
endopodite  is  reduced.  Although  somewhat  smaller  than  the  type  material  (mean  length 
0'74  mm),  this  specimen  otherwise  corresponds  closely  to  Heron's  description.  Heron 
reported  O.  rotunda  from  3  stations  in  the  antarctic  zone  of  the  Pacific  between  1000  and 
2000  metres  depth. 

Oncaea  brocha  Heron,  1977 
O.  brocha  Heron,  1977  :  60,  figs  14f-n,  15a-n. 

DESCRIPTION.  Female.  Body  length  of  2  specimens  0'81  and  0'83  mm.  Urosome  (Fig.  lOg), 
second  antenna  (Fig.  lOh),  maxilliped  (Fig.  lOi)  and  second  and  third  natatory  legs  (Figs  lOj 
and  k)  figured. 

MATERIAL  EXAMINED.  299  from  sample  9541.24,  3980-3960.  20°  N  21°W.  BM(NH) 
1981.131. 

REMARKS.  The  specimens  were  identified  by  the  following  combination  of  characters:  the 
relative  lengths  of  the  segments  of  the  first  and  second  antennae,  the  armature  of  the 
maxilliped,  of  the  natatory  and  of  the  fifth  legs.  The  second  antenna  has  an  additional 
terminal  spine  not  figured  by  Heron.  This  may  have  been  lost  from  Heron's  material  or  may 
represent  geographical  variation.  Another  minor  difference  observed  was  the  length  of  the 
terminal  spines  of  the  second  and  third  endopodites  which  are  relatively  longer  in  the  present 
material.  The  refractive  granules  mentioned  by  Heron  are  absent  from  these  specimens,  but 
but  these  may  have  been  artefacts  or  may  represent  the  positions  of  cuticular  pores.  Heron 
reported  O.  brocha  from  1  station  in  the  antarctic  zone  of  the  southwest  Pacific  between 
1000  and  2000  metres  depth. 

Oncaea  schmitti  (Heron,  1977)  Comb.  nov. 
Epicalymma  schmitti  Heron,  1977  :  82-84,  figs  28a-n,  29a-e. 

DESCRIPTION.  Female.  Body  length  2  specimens  0*33  and  0'34  mm.  Body  (Figs  1  la  and  b), 
second  antenna  (Fig.  1 1  c)  and  maxilliped  (Fig.  1 1  d)  figured. 

MATERIAL  EXAMINED.  299  from  sample  9541.24,  3980-3960  m.  20°  N  21°W.  BM(NH) 
1981.132. 

REMARKS.  This  species  is  here  transferred  from  the  genus  Epicalymma  to  Oncaea  (for 
discussion  see  page  191).  Specimens  are  clearly  identifiable  as  O.  schmitti  by  the  armature 
and  relative  segment  lengths  of  the  second  antenna,  the  armature  of  the  maxilliped  and  the 
dorsal  projections  of  the  caudal  rami.  However,  the  specimens  are  so  thinly  chitinized  that 
the  genital  segment  had  partially  collapsed  during  capture  or  preservation.  Therefore,  the 
drawing  of  the  urosome  may  not  represent  the  morphology  of  the  living  animal.  This  species 


NEW  AND  LITTLE  KNOWN  SPECIES  OF  ONCAEIDAE  1 9  1 

has  also  been  recorded  from  the  antarctic  zone  of  the  southwest  Pacific  where  it  was  taken 
between  1000  and  2000  metres  depth. 

Oncaea  umbonata  (Heron,  1977)  Comb.  nov. 
Epicalymma  umbonata  Heron,  1977  :  84-86,  figs  29f-p,  30a-c. 

DESCRIPTION.  Female.  Body  length  of  1  damaged  specimen  0*62  mm,  without  caudal  rami. 
Body  (Figs  1 1  e  and  0,  second  antenna  (Fig.  1 1  g)  maxilliped  (Fig.  1 1  h)  figured. 

MATERIAL  EXAMINED.  \g  from  sample  9541.24,  3980-3960  m.  20°  N  21°  W.  BM(NH) 
1981.133. 

REMARKS.  This  species  is  here  transferred  from  the  genus  Epicalymma  to  Oncaea  (for 
discussion  see  below).  Although  the  specimen  lacks  its  caudal  rami  it  was  assigned  to  O. 
umbonata  on  the  basis  of  its  body  size  and  maxilliped  armature.  There  are  also  differences  of 
the  armature  of  the  second  antenna  between  this  and  the  specimens  recorded  as  O.  schmitti, 
the  two  species  are  otherwise  very  close.  The  natatory  legs  and  other  mouthparts  correspond 
closely  to  Heron's  original  description.  This  species  has  also  been  recorded  from  the 
antarctic  zone  of  the  southwest  Pacific  where  it  was  taken  between  1000  and  2000  metres 
depth. 


Phylogenetic  relationships  within  the  genus  Oncaea 

Giesbrecht  (1892)  designated  the  following  character  states  as  typical  of  the  genus 
Conaea:  the  elongate  terminal  segment  of  the  second  antenna  with  very  large,  hook-tipped 
spines;  endopodite  of  the  fourth  leg  shorter  than  the  exopodite,  third  segment  is  reduced 
(shorter  than  the  first  or  second  segment);  the  fifth  leg  reduced  to  a  single  plumose  seta. 
However  Heron  (1977)  was  able  to  demonstrate  the  presence  of  a  minute  spinule 
accompanying  the  seta  of  the  fifth  leg  of  O.  gracilis  Dana,  1852  (=syn.  C.  rapax),  the  type 
species.  Heron  also  described  2  new  species  of  Conaea,  C.  succurva  and  C.  hispida.  Both  of 
these  have  a  seta  on  a  small  prominence  representing  leg  5,  and  a  reduced  endopodite  of  the 
fourth  leg,  but  the  third  endopodite  segment  exceeds  the  first  segment  in  length.  These  2 
species  also  lack  the  hooked  tips  to  the  terminal  spines  of  the  second  antenna.  Oncaea 
expressa  Gordejeva,  1973  shares  these  and  other  character  states  but  differs  in  the 
morphology  of  the  natatory  legs  (Table  1).  No  less  than  27  other  species  of  Oncaea  also  show 
1  or  more  of  these  Conaea-\ike  character  states  (Table  2)  The  2  species  of  Epicalymma 
which  Heron  (1977)  described  also  exhibit  the  following  character  states:  elongate  terminal 
segment  of  the  second  antenna  with  very  large  hook-tipped  spines,  endopodite  of  the  fourth 
leg  shorter  than  the  exopodite,  third  segment  reduced  (but  longer  than  the  first  or  second 
segment),  fifth  leg  with  a  single  terminal  seta. 

A  continuous  gradation  of  character  states  within  Oncaea,  Conaea  and  Epicalymma  is 
thus  becoming  apparent  as  more  species  of  Oncaea  are  discovered.  The  armature  of  the 
third  exopodite  segments  of  the  swimming  legs  also  illustrates  this.  Most  Oncaea  species 
have  an  external  spine  formula  of  III,  III,  II,  II  but  there  is-considerable  variation  within  the 
genus:  II,  III,  III,  I  (O.  ancora  Gordejeva,  1973);  III,  III,  III,  II  (O.  minor  Shmeleva,  1979  and 
O.  parobscura  Shmeleva,  1979);  III,  II,  II,  I  (O.  expressa  Gordejeva.  1973);  III,  II,  II,  II  (O. 
brodskii  Shmeleva,  1968  and  O.  longipes  Shmeleva,  1968);  II,  III,  II,  I  (O.  exigua  Farran, 
1908);  II,  II,  I,  I  (O.  atlantica  Shmeleva,  1967  and  O.  vodjanitskii  Shmeleva  and  Delalo, 
1969;  II,  III,  II,  II  (O.  zernovi  Shmeleva,  1966  and  O.  mollicula  Gordejeva,  1975).  The 
typical  formula  for  Conaea  is  II,  III,  II,  I  and  for  Epicalymma  II,  III,  III,  I.  Such  variation 
represents  a  continuum  which  cannot,  in  my  opinion,  justifiably  be  subdivided  into  2  or 
more  genera  as  it  is  at  present.  It  is  probable  that  Conaea-\\ke  and  Epicalymma-like 
character  states  have  been  derived  by  reduction  and  loss  of  armature  elements  from  ancestors 
within  the  genus  Oncaea.  Separation  of  the  Conaea  and  Epicalymma  species  would  leave 


192  S.  J.  MALT 

Table  1     Comparison  of  morphology  of  natatory  legs  ofOncaea  expressa.  O.  succurva  and  O.  hispida. 


Legl 


Leg  2 


Leg  3 


Leg  4 


O.  expressa 

Exopod.  spine 

Exopod.  spine 

Exopod.  spine 

Exopod.  spine 

Gordejeva,  1973 

2  x  length 

2  x  length 

2-  5  x  length 

>  2x  length 

term.  segm. 

term.  segm. 

term.  segm. 

term.  segm. 

Endopod.  spine 

Endopod.  spine 

Endopod.  spine 

Endopod.  spine 

shorter  than 

1-5  x  length 

>4x  length 

>  4  x  length 

term.  segm. 

term.  segm. 

term,  segm 

term.  segm. 

O.  succurva 

Exopod.  spine 

Exopod.  spine 

Exopod.  spine 

Exopod.  spine 

(Heron,  1977) 

2  x  length 

>2  x  length 

2-5  x  length 

>2'5  x  length 

comb.  nov. 

term.  segm. 

term.  segm. 

term.  segm. 

term.  segm. 

(syn.  Conaea 

Endopod.  spine 

Endopod.  spine 

Endopod.  spine 

Endopod.  spine 

succurva) 

same  length 

1-5  x  length 

2-  5  x  length 

3  x  length 

term.  segm. 

term.  segm. 

term.  segm. 

term.  segm. 

O.  hispida 

Exopod.  spine 

Exopod.  spine 

Exopod.  spine 

Exopod.  spine 

(Heron,  1977) 

2  x  length 

>2x  length 

2-5  x  length 

>  3  x  length 

comb.  nov. 

term.  segm. 

term.  segm. 

term.  segm. 

term.  segm. 

(syn  Conaea 

Endopod.  spine 

Endopod.  spine 

Endopod.  spine 

Endopod.  spine 

hispida) 

same  length 

1  -5  x  length 

2  x  length 

>4  x  length 

term.  segm. 

term.  segm. 

term.  segm. 

term.  segm. 

Oncaea  as  a  paraphyletic  group.  In  order  to  retain  Oncaea  as  a  monophyletic  group  (sensu 
Hennig,  1966)  it  is  proposed  that  Conaea  and  Epicalymma  be  synonymized  with  Oncaea 
and  that  the  species  of  Conaea  and  Epicalymma  be  transferred  to  Oncaea,  C.  gracilis,  C. 
succurva,  and  C.  hispida  becoming  O.  gracilis,  O.  succurva  and  O.  hispida,  also  E.  schmitti 
and  E.  umbonata  becoming  O.  schmitti  and  O.  umbonata  respectively. 

Kazatchenko  and  Andreev  (1977)  described  Myctospictosum  as  a  genus  of  uncertain 
taxonomic  position.  A  single  male  of  M.  philippinensis  was  found  on  the  gills  ofMyctophum 
spinosum  in  the  western  Pacific.  It  is  clear  from  the  figures  (Kazatchenko  and  Andreev, 
1977  :  Figs  9  and  10)  that  this  is  an  oncaeid.  It  has  the  body  shape,  the  4-segmented  first 
antenna  and  the  3-segmented  second  antenna  of  a  typical  Oncaea  male.  The  other 
mouthparts  have  been  interpreted  differently  by  the  authors  but  are  clearly  those  of  a  male 
Oncaea.  The  natatory  legs  are  also  typical  of  the  genus.  It  is  proposed,  therefore,  that 
Myctospictosum  should  be  synonymized  with  Oncaea.  The  armature  of  the  natatory  legs  and 
the  relative  lengths  of  the  posterior  segments  separate  O.  philippinensis  comb.  nov.  from  any 
of  the  known  males  of  the  genus  Oncaea. 

Oncaeids  are  often  found  clinging  by  their  maxillipeds  to  other  organisms  and  debris  in 
plankton  samples,  there  is,  therefore,  no  reason  to  suppose  that  O.  philippinensis  is  parasitic, 
it  is  probably  a  free-living  planktivore  like  other  species  of  the  genus. 

The  generic  diagnosis  ofOncaea  is  ammended  as  follows: 

ONCAEA  Philippi,  1843 

Oncaea  Philippi,  1843  :  63. 

AntariaDana,  1846:229. 

Conaea  Giesbrecht,  1891  :  477. 

Epicalymma  Heron,  1977  :  82. 

Myctospictosum  Kazatchenko  and  Andreev,  1977  :  47. 

DIAGNOSIS.  Cyclopoid  shape.  Rostral  area  thickened  with  rounded  posteroventral  margin. 
Female  urosome  5-segmented,  male  6-segmented.  First  antenna  6-segmented  in  female, 


NEW  AND  LITTLE  KNOWN  SPECIES  OF  ONCAEIDAE  193 

Table  2     Conaea-like  character  states  exhibited  by  species  ofOncaea. 


c  rj 

u  .«  o 

r;  CO  O 

SP  C  «      .  u    Q  —    C  M-5    p 


?s  ^       U^  -*-^  <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 


R.  W.  INGLE 


°  - 

c  </> 


•s     ± 


It 


U    00 

>   1 


o.  a 

—  «  (N    I 


N  = 

'r'N 


•a 


°~  E  ° 


~^  _>>  u.  £> 

00  -C    ^-  2 

C    oo  O  S 

i  o  =5  -g  &  i 


E  o 


S  -t"? 
S"iS 

™"       '    O 

+  o  c 
TT   a   S  m 

«=>  g  5  + 

O  i:  -i^O 


X      1^1  + 


rs 

u 

— 
« 


g 

'S  s 


•S  ?  5b 


—  1    <N  —  O 


•2.8 


+  o-g; 


J3    O  ' 


4>    "•  Q, 


I 


ce 
ara 


ral  spin 
pace  d 


I    ' 


ft  J  J 

•s       °  S. 


u  i: 
E  JS 

II 
^^ 


gt 


00 

-^ 

«      ^ 

TT 

vO 

V    00 

2  £ 

00 

§ 

a 

O.        <Nr 

— 

+ 

JO    C 

f-  J3 

c 

o 

+ 

— 

<N     1    <N 

m 

°' 

—     1 

1  —   »/"!  V"i 

—  — 

(j    ^ 

i 

J2   i   + 

oo  < 

u 

^_^ 

M 

__ 

o  X 

c 

N 

y,       f*> 

+ 

~  JS 

'5. 

O       ' 

U 

Oil 

c   oo 

00 

c 

+ 

a 

a     *s 

<N    +  <N 

•<t 

- 

-   + 

7 

r-  w>  f> 

'5.J 

'5. 

i 

1   i    i 

jd 

1 

•j?  ^ 

c 

'* 

Abdomen  posterio-lateral  spines: 

2  setae  on  1st  abdominal  segment: 

Setae  on  2nd  abdominal  segment: 

Dorso-lateral  proc.  on  3rd  abd.  sgm 
Setal  formula  basis  1st  maxilliped: 

No.  setae  basis  of  2nd  maxilliped: 

Setal  formula  endopod  2nd  maxillip 

Setal  formula  endopod  maxillule: 
Telson  dorsal  spine  +  or  -: 

megalopa 

No.  abdominal  segments  (incl.  telso 
No.  pairs  of  pleopods  (incl.  uropods 
No.  setae  on  each  terminal  pleopod: 

Frontal  region  of  carapace-submedi 
-rostrum 

Carapace,  each  protogastric  region  i 

orbital  region  with: 

cardiac  region  with: 
Spines  on  abdominal  segments: 
Processes  or  spine,  1  st  ped.  sgmt  anl 

c        u 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB       2  1 3 

Provisional  key  to  the  larvae  of  N.E.  Atlantic  Oxyrhyncha 

Zoeae 

Carapace  lateral  spines  present 2 

Carapace  lateral  spines  absent 5 

Carapace  with  a  group  of  5  lateral  spines  on  each  side 

t  Dory  nchusthomsoni  Thomson,  (Williamson,  1960). 

Carapace  with  one  lateral  spine  on  each  side 3 

Telson  medio-lateral  margin  strongly  concave  or  medially  cleft;  middle  pair  of  medio-lateral 
setae  longest;  each  telson  furca  with  one  dorsal  and  one  (rarely  2)  lateral  spines;  3rd 
abdominal  segment  with  dorso-lateral  process 4 

Telson  medio-lateral  margin  not  concave  or  cleft;  medio-lateral  setae  of  equal  length;  each 
telson  furca  without  a  dorsal  spine,  one  lateral  spine  present;  3rd  abdominal  segment 
without  dorso-lateral  process .  .  .  .  Rochinia  carpenteriThomson,  (Ingle,  1979). 

Carapace  dorsal  spine  much  longer  than  carapace  length;  abdominal  posterio-lateral  process 
very  long;  2nd  maxilliped  basis  with  4  setae;  each  furca  of  telson  with  one  lateral  spine 
Hyas  coarctatus  (Leach),  H.  araneus  (Linnaeus),  (Christiansen,  1973). 

Carapace  dorsal  spine,  at  the  most,  slightly  longer  than  carapace  length;  abdominal 
posterio-lateral  processes  short;  2nd  maxilliped  basis  with  3  setae;  each  furca  of  telson 
with  2  lateral  spines Majasquinado(Herbst),(Lebour,  1927,  1928). 

Carapace  rostral  spine  long;  each  telson  furca  with  a  dorsal  spine 

Eurynome  aspera  (Pennant),  (Lebour,  1 928).  E.  spinosa  Hailstone,  (see  Table  2,  footnote). 

Carapace  rostral  spine  small  or  absent;  each  telson  furca  without  a  dorsal  spine      .       .  6 

Telson  medio-lateral  margin  strongly  cleft,  marginal  setae  in  2  groups;  maxillule  endopod* 

with  0  +  3-4  setae;  abdominal  posterio-lateral  spines  long 7 

Telson  medio-lateral  margin  not  cleft,  marginal  setae  not  in  2  groups;  maxillule  endopod 

with  1  +  6  setae;  abdominal  posterio-lateral  spines  short 

tPfaa  annaia(Latreille),  (Ingle  &  Clark,  1980). 

Lateral  spine  of  each  telson  furca  very  long  (about  \  length  of  furca  and  exceeding  ^  maximum 

width  of  telson) Achaeus  cranchii  Leach,  (Bocquet,  1954). 

Lateral  spine  of  each  telson  furca  short,  never  reaching  |  length  of  furca  ....  8 

Lateral  spine  of  each  telson  furca  large;  middle  medio-lateral  seta  of  telson  not  longest; 
maxillule  endopod  with  0  +  4  setae 

Inachus  dorsettensis  (Pennant),  /.  p/ialangium  (Fabricius),  /.  leptochirus  Leach,  (Clark,  1980). 
Lateral  spine  of  each  telson  furca  small;  middle  medio-lateral  seta  of  telson  longest; 
maxillule  endopod  with  0  +  3  setae 

Macropodia  rostrata  (Linnaeus),  M .  tenuirostris  (Leach),  (see  p.  2 1 1 ),  M .  deflexa  Forest  (Lebour, 

1928). 

Megalops 

Abdomen  composed  of  5  segments  (excluding  telson);  4  pairs  of  pleopods,  terminal  pair  with 

8  setae  on  exopod 2 

Abdomen  composed  of  6  segments  (excluding  telson);  5  pairs  of  pleopods,  terminal  pair  with 

3-5  setae  on  exopod 4 

Dorsal  margins  of  abdominal  segments  without  spinules;  carapace  submedian  spines  not 

developed 

Macropodia  rostrata  (Linnaeus),  M.  tenuirostris  (Leach)  (see  p.  2 1 1 ),  M.  deflexa  Forest  (Lebour, 

1928). 
Dorsal  margins  of  abdominal  segments  with  small  spinules;  carapace  submedian  spines  well 

developed 3 

Orbital  spines  present 

Inachus  dorsettensis  (Pennant),  /.  phalangium  (Fabricius),  /.  leptochirus  Leach,  (Clark,  1 980). 
Orbital  spines  absent Achaeus  cranchii  Leach,  (Bocquet,  1954). 


*Not  known  for  Achaeus  cranchii;  tHeegaard's  (1963)  description  of  zoea  I  of  Pisa  tetraodon  differs  considerably 
from  that  of  P.  armata  (see  Table  2,  footnote)  and  is  excluded  from  this  present  key. 


214  R.  W.  INGLE 

4  Carapace  submedian  spines  present 

Hyas  coarctatus  (Leach),  H.  araneus  (Linnaeus),  (Christiansen,  1973). 

-  Carapace  submedian  spines  absent 5 

5  First  peduncular  segment  of  antenna  without  a  distal  process;  exopod  of  each  terminal 

pleopod  with  3  setae 

•  Eurynome  aspera  (Pennant),  (Lebour,  1928),  E.  spinosa  Hailstone,  (see  Table  2,  footnote). 
First  pedunular  segment  of  antenna  with  an  obtuse  process  or  spine;  exopod  of  each 
terminal  pleopod  with  5  setae 6 

6  A  stout  cardiac  spine  on  carapace     .       .       .           Rochinia  carpenteri  (Thomson),  (Ingle,  1979). 
Without  a  cardiac  spine  on  carapace 7 

7  Rostral  spine  well  developed;  each  protogastric  region  with  a  carina;  exopods  of  lst-4th 

pereiopods  with  10  setae Pisa  armata  (Latreille),  (Ingle  &  Clark,  1 980). 

-  Rostral  spine  minute;  each  protogastric  region  with  (at  the  most)  a  tubercle;  exopods  of 

1  st-4th  pereiopods  with  8  setae     .       .       .         A/a/a  squinado  (Herbst),  (Lebour,  1 927,  1 928). 

Acknowledgements 

I  wish  to  thank  Dr  R.  B.  Manning  for  sponsoring  my  visit  to  Tunisia  under  the  Smithsonian 
Institution  Foreign  Currency  Program;  Dr  D.  I.  Williamson  for  his  assistance  in  obtaining 
ovigerous  crabs  from  the  Isle  of  Man  region,  and  Mr  A.  D.  M.  Mattacola  and  crew  of  the 
Sepia  for  their  help  in  collecting  material  in  Plymouth  waters. 

References 

Barnard,  K.  H.  1950.  Descriptive  Catalogue  of  South  African  Decapod  Crustacea  (Crabs  and  Shrimps). 

Ann.  S.  Afr.  Mus.  38  :  837  pp. 
Bocquet,  C.  1954.  Developpment  larvaire  d'Achaeus  cranchii  Leach  (Decapode  Oxyrhynque).  Bull. 

Soc.  zool.  Fr.  79  :  50-56. 
Bourdillon-Casanova,  L.   1960.  Le  meroplancton  du  Golfe  de  Marseille:  les  larves  de  crustaces 

decapodes.  Red  Trav.  Stn  mar.  Endoume3Q  :  1-286. 

Cano,  G.  1 893.  Sviluppo  e  Morfologia  degli  Oxyrhynchi.  Mitt.  zool.  Stn  Neapel  10  :  527-583. 
Christiansen,  M.  E.  1969.  Marine  invertebrates  of  Scandinavia,  No.  2.  Crustacea  Decapoda  Brachyura. 

Universitetsforlaget,  Oslo,  1-143  pp. 
1973.  The  complete  larval  development  of  Hyas  araneus  (Linnaeus)  and  Hyas  coarctactus  (Leach 

(Decapoda,  Brachyura,  Majidae)  reared  in  the  laboratory.  Norw.  J.  Zool.  21  :  63-89. 
Clark,  P.  F.   1980.  British  spider  crabs  of  the  genus  Inaehus;  a  morphological  study  of  larval 

development.  M.Sc  Modern  Taxonomy  (C.N.A.A.)  Thesis.  Polytechnic  of  Central  London,  140  pp. 

London. 
Heegaard,  P  1963.  Decapod  larvae  from  the  Gulf  of  Napoli  hatched  in  captivity.  Vidensk.  Meddr 

dansk  naturh.  Foren.  125  :  449^493. 
Ingle,  R.  W.   1979.  The  larval  development  of  the  spider  crab  Rochinia  carpenteri  (Thomson) 

[Oxyrhyncha:  Majidae]  with  a  review  of  majid  subfamilial  larval  features.  Bull.  Br.  Mus.  nat.  Hist. 

(Zool.)  37:  47-66. 

&  Clark,  P.  F.  1 977.  A  laboratory  module  for  rearing  crab  larvae.  Crustaceana  32  :  220-222. 

1980.  The  larval  and  post-larval  development  of  Gibb's  spider  crab.  Pisa  armata  (Latreille) 

[family  Majidae:  subfamily  Pisinae],  reared  in  the  laboratory.  /.  nat.  Hist.  14  :  723-735. 
Ingle,  R.  W.  &  Manning,  R.  B.  (in  press).  Infraspecific  variation,  synonomy  and  distribution  of  the 

spider  crab,  Macropodia  rostrata  (Linnaeus).  Colloquium  Crustacea  Decapoda  Mediterranean  May 

1979  Ancona. 
Lebour,  M.  V.  1927.  Studies  of  the  Plymouth  Brachyura.  I.  The  rearing  of  crabs  in  captivity,  with  a 

description  of  the  larval  stages  of  Inaehus  dorsettensis.  Macropodia  longirostris  and  Maia  squinado. 

J.  mar.  biol.  Ass.  U.K.  14  :  795-820. 

1928.  The  larval  stages  of  the  Plymouth  Brachyura.  Proc.  zool.  Soc.  Lond.  2  :  473-560. 

Lo  Bianco,  S.  1904.  Pelagische  Tiefseefischeri  der  "Maja"  in  der  umgebung  von  Capri.  Beitr.  Kennt. 

Meeres  und  Seiner  Bewohner  1  :  1-91  Jena. 
Manning,  R.  B.  &  Holthius,  L.  B.  1981.  West  African  Brachyuran  Crabs  (Crustacea:  Decapoda). 

Smithson.  Contr.  Zool.  306  :  1-379. 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB       2 1  5 

Monod,    T.    1932.    Brachyuran    Maroccana    2e    Partie:    Dromiidea,    Oxystomata,    Oxyrhyncha, 

Brachyrhyncha(excl.  Pinnotheridae).  Bull.  Soc.  Sci.  nat.  Maroc  12  :  7-8. 
Rice,  A.  L.  1980.  Crab  zoeal  morphology  and  its  bearing  on  the  classification  of  the  Brachyura.  Trans. 

zool.  Soc.  Lond.  35  :  271^424. 
&  Ingle,  R.  W.  1975.  The  larval  development  of  Carcinus  maenas  (L.)  and  C.  mediterraneus 

Czerniavsky  (Crustacea,  Brachyura,  Portunidae)  reared  in  the  laboratory.  Bull.  Br.  Mus.  nat.  Hist. 

(Zool.)  28:  101-119. 
Steedman,  H.  F.  1976.  General  and  applied  data  on  formaldehyde  fixation  and  preservation  of  marine 

zookplankton  (pp.  103-171)  In:  Steedman,  H.  F.  (Edit.).  Zooplankton  fixation  and  preservation. 

Monographs  on  oceanographic  methodology  4  :  350  pp.  UNESCO  Press,  Paris. 
Stuxberg,  A.  1874.  Karcinologiska  iaktagelser.  Ofvers.  K.  VetenskAkad.  Forh.  Stockh.  9  :  3-23. 
Thompson,  J.  V.  1836.  Of  the  double  metamorphosis  in  Macropodia  phalangium,  or  spider  crab,  with 

proofs  of  the  larvae  being  zoea  in  Gegarcinus  hydrodomus.  Thelpusa  erythropus.  Eriphia  carribaea. 

and  Grapsus  pelagicus.  Ent.  Mag.  3  :  370-375. 
Williamson,  D.  I.  1960.  A  remarkable  zoea,  attributed  to  the  Majidae  (Decapoda,  Brachyura).  Ann. 

Mag.  nat.  Hist.  (13)3  :  141-144. 
Zariquiey  Alvarez,  R.  1968.  Crustaceos  Decapodos  Ibericos.  Investigacion  pesq.  32  :  i-ix  +  5 10  pp. 

Manuscript  accepted  for  publication  3  July  1981 


216 


R.  W.  INGLE 


Fig.  1  Macropodia  rostrata  (L.):  a,  b  zoea  I  &  II  respectively  from  lateral  aspects;  c  zoea  II 
carapace  and  eyes  from  frontal  aspect;  d  megalopa  from  dorsal  aspect  and  e  lateral  aspect  of 
carapace  and  eye;  f  dorsal  and  g  lateral  aspect  of  megalopal  abdomen;  h  first  and  i  fourth  left 
megalopal  pleopods;  carapace  of  the  first  crab  from  j-Isle  of  man  cf.  with  k-Tunisian  waters. 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB        2 1 7 


1  002' 


Fig.  2  Macropodia  rostrata  (L.):  zoea  I,  a  antennule  and  antenna,  b  mandible;  zoea  II,  c 
antennule  and  antenna,  d  mandible;  megalopa,  e  antennule,  f  antenna,  g  mandible;  first  crab,  h 
antennule  &  antenna,  i  mandible. 


218 


R.  W.  INGLE 


Fig.  3    Macropodia  rostrata  (L.):  maxillule  of-  a  zoea  I;  b  zoea  II;  c  megalopa;  d  first  crab. 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB         2  1 9 


Fig.  4    Macropodia  rostrata  (L.):  Maxilla  of-  a  zoea  I;  b  zoea  II;  c  megalopa;  d  first  crab. 


220 


R.  W.  INGLE 


Fig.  5    Macropodia  rostrata  (L.):  a  &  b  1st  &  2nd  maxillipeds  of  zoea  I;  c  &  d  1st  &  2nd 

maxillipeds  of  zoea  II. 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB       22  1 


Fig.  6    Macropodia  rostrata  (L.):  abdominal  segments  and  telson  a  &  c  lateral,  b  &  d  dorsal 
aspects  of-  a,  b  zoea  I;  c,  d  zoea  II;  e  spines  on  right  medio-lateral  margin  of  telson  of  zoea  I. 


222 


R.  W.  INGLE 


Fig.  7    Macropodia  rostrata  (L.):  1  st  (a,  c)  &  2nd  (b,  d)  maxillipeds  a,  b  megalopa;  c,  d  first  crab. 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB       223 


Fig.  8    Macropodia  rostrata  (L.):  3rd  maxilliped-  a  megalopa;  b  first  crab;  cheliped-  c  megalopa; 

d  first  crab. 


224 


R.  W.  INGLE 


Fig.  9    Macropodia  rostrata  (L.):  a  2nd  pereiopod  of  megalopa;  b,  c  2nd  &  3rd  pereiopods  of  1st 

crab. 


LARVAL  AND  POST-LARVAL  DEVELOPMENT  OF  THE  SLENDER-LEGGED  SPIDER  CRAB         225 


Fig.  10    Macropodia  rostrata  (L.):  a-c  3rd-5th  pereiopods  of  megalopa;  d  &  e  4th  &  5th 

pereiopods  of  first  crab. 


British  Museum  (Natural  History) 
British  Marine  Amphipoda:  Gammaridea 

by  R.  J.  Lincoln 

658pp    2,300  figures    4to  hard  bound 
ISBN  0  565  00818        £50.00 

Amphipods  are  both  numerous  and  diverse  in  numbers  of  genera  and  species  in 
British  coastal  waters,  but  in  the  absence  of  any  form  of  modern  systematic  synopsis 
or  key  this  group  of  crustaceans  has  acquired  the  reputation  of  being  notoriously 
difficult  to  identify.  This  monograph,  which  is  the  first  comprehensive  and  illustrated 
text  on  British  gammaridean  amphipods  to  be  published  in  more  than  a  century, 
should  go  a  long  way  towards  solving  the  problem. 

The  systematic  section  of  the  book  contains  descriptions  and  figures  of  all  271  species 
of  marine  and  brackish  water  amphipods,  in  123  genera  and  36  families,  recorded 
from  British  coasts  and  the  adjacent  continental  shelf  to  a  depth  of  200  metres.  Keys 
are  provided  at  all  levels,  as  well  as  relevant  synonymies  and  diagnoses  of  genera  and 
families.  The  text  is  illustrated  with  about  2,300  separate  figures  which  have  been 
drawn  by  the  author  from  Museum  and  other  material,  in  many  cases  with  reference 
to  type  specimens. 

The  work  has  been  carefully  edited  to  bring  corresponding  descriptions,  keys  and 
figures  into  close  proximity  within  the  text.  The  systematic  section  is  supported  by 
chapters  dealing  with  morphology,  systematics,  geographical  distribution,  biology 
and  ecology,  the  latter  being  presented  in  the  form  of  an  annotated  subject  index  of 
research  literature.  Finally,  there  is  an  extensive  bibliography  of  about  1,200 
references  that  includes  most  of  the  British  marine  amphipod  literature  published  to 
date. 

Publications  Sales, 

British  Museum  (Natural  History), 

Cromwell  Road, 

London  SW7  5BD, 

England. 


Titles  to  be  published  in  Volume  42 


The  tick  collection  (Acarina:  Ixodoidea)  of 

the  Hon.  Nathaniel  Charles  Rothschild  deposited  in  the  Nuttall  and 

general  collections  of  the  British  Museum  (Natural  History). 

By  James  E.  Kekans 

Hydroids  and  medusae  of  the  family  Campanulariidae  recorded 
from  the  eastern  North  Atlantic,  with  a  world  synopsis  of  genera. 
By  P.  F.  S.  Cornelius 

Miscellanea 
Miscellanea 


Printed  by  Henry  Ling  Ltd,  Dorchester 


902 


British  Museum  (Natural  History) 


Miscellanea 


Zoology  series    Vol  42  No  4    24  June  1982 


The  Bulletin  of  the  British  Museum  (Natural  History),  instituted  in  1949,  is  issued  in  four 
scientific  series,  Botany,  Entomology,  Geology  (incorporating  Mineralogy)  and  Zoology,  and 
an  Historical  series. 

Papers  in  the  Bulletin  are  primarily  the  results  of  research  carried  out  on  the  unique  and 
ever-growing  collections  of  the  Museum,  both  by  the  scientific  staff  of  the  Museum  and  by 
specialists  from  elsewhere  who  make  use  of  the  Museum's  resources.  Many  of  the  papers  are 
works  of  reference  that  will  remain  indispensable  for  years  to  come. 

Parts  are  published  at  irregular  intervals  as  they  become  ready,  each  is  complete  in  itself, 
available  separately,  and  individually  priced.  Volumes  contain  about  300  pages  and  several 
volumes  may  appear  within  a  calendar  year.  Subscriptions  may  be  placed  for  one  or  more  of 
the  series  on  either  an  Annual  or  Per  Volume  basis.  Prices  vary  according  to  the  contents  of 
the  individual  parts.  Orders  and  enquiries  should  be  sent  to: 


Publications  Sales, 

British  Museum  (Natural  History), 
Cromwell  Road, 

London  SW7  5BD, 
England 


World  List  abbreviation:  Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


©  Trustees  of  the  British  Museum  (Natural  History),  1982 


The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 
Keeper  of  Zoology     :    Dr  J.  G.  Sheals 
Editor  of  Bulletin       :    Dr  C.  R.  Curds 
Assistant  Editor         :    Mr"C.  G.  Ogden 


ISSN  0007-1498  Zoology  series 

Vol  42  No  4  pp  227-335 
British  Museum  (Natural  History) 
Cromwell  Road 
London  SW7  5BD  Issued  24  June  1982 


.*,    LIBRARY 


^ 


a*0; 


Miscellanea 

Contents 

New  species  of  marine  nematodes  from  Loch  Ewe,  Scotland.  By  H.  M.  Platt  and 
Z.N.Zhang       .............. 

The  larval  development  of  Crangon  crangon  (Fabr.  1795)  (Crustacea:  Decapoda). 
By  A.  R.Gurney      ............ 

A  revision  of  the  spider  genus  Cocalodes  with  a  description  of  a  new  related  genus 
(Araneae:  Salticidae).  By  F.  R.  Wanless       ........ 

Anatomy  and  evolution  of  the  jaws  in  the  semiplotine  carps  with  a  review 
of  the  Genus  Cyprinion  Heckel,  1  843  (Teleostei:  Cyprinidae).  By  G.  J.  Howes 


221 
247 
263 
299 


A*^<?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 

o 

3 

<S23           <^<S 

c^ccc               ooScoc 
D222             D.2D2.0 
8       tt       8       o       E                 tt                tt       8       fi 

*~ 

•  —        {<3*J*Jti"^        ^        o        fO.'S        " 

§ 

1  s  i  i  •§  i  5  |  8  •§  i 

s 

^uiSSSa5uflQ(iiSv3 

13 

C    u 

is 

<-.          |         «-.          |                   e-                             e-        c-         + 

(5    <» 

H 

•o 

D          JL> 

a"«0   2 

1           1          +'          1                                                     + 

r*     ** 

c  B  c 

+          1           1                      1 

O    rt    CO 

S 

C/i 

•0"3 

ag 

^M                  IO                                                                     CO                 *^                                           OO                 t  —  • 

es-       e^-       —  i       p—       t>-       ^"       ^o       ^       s>"       r~»       r^ 

-H                                      —       —                            r^ 
o 

CO 

C 

oo 

I                       V                               1                                                          1                                                                                                                                                  . 

—    _j_ 

3 

o 

CO 

.a  u 

*                                                *        *                  * 

.C     00    ~!r 

iO         v^          ^^          vO          *^~>         fN          C^          ^O         CO          *^         C?N 

&&- 

O                         0 

U  « 

*                                                                  *            *                         * 

s 

CO  "*•? 

t^         *O         ro         *O         ^D        OJ         OO        ^"         *^         f*^        oo 

^ 

"5  -^ 

ooooooooooo 

s: 

35 

0 

o 

|3 

j= 

^) 

aj= 

u  «-> 

GO 

0    00 

(N         C?\         t~~        TT         OO                    ~*                     ^         <^> 

3 

*^    4> 

^—                                 ^^                                 ^^                                 ^^ 

c 

—  _ 

00 

a> 

s 

CO 

m-rj-mt^mfNooO^ooo 
~*Tt<Nm'O(Nl/^rOI^<Nt^ 

_>, 

*© 

c 

V) 

o 

jj 

X 

O"O            O«o"o            O"O*O            3            O^D 

1 

C/3 

u- 

0 

~ 

^- 

w  c 

I1 

<N           (N                                                                                   2 

T           ~T 

OOT3 
tS    <L> 

Table  I  Tabular 

00 

U 

'G 

_           „,          (vj          r~           ^          Sd2.rr>          "" 

g-^-j       in       ^       _^       ON      ovc"""1       °Na""f'^ 

S^gON.^U           0         X.         —    3-0'        —    ^  T3           C 

Q  —   •t-    uT  ^    C  '5    cS^r'S^    O^    u2S    OQ    S^> 
s     •  S  &  $  Q  £»  O       9  <B  s  s  5  S  (i  ^  3k  «l  ^  C  Q 

^    O    Ji  '~C  1?    V          D  .^    u    5«>    U  *-i    ^  "ts    1>    2  j™   C    "    S    rv 

^calculated  from  I 
+'vacuoles  arrang 

NEW  SPECIES  OF  MARINE  NEMATODES 


239 


Fig.  6  Pictorial  key  to  the  species  of  Ceramonema:  (a)  C.  attenuatum  9  (after  Cobb,  1920);  (b)  C. 
carinatum  d  (after  Wieser,  1959);  (c)  C.  chitwoodi  9  (after  De  Coninck,  1942);  (d)  C.filipjevi  d 
(after  De  Coninck,  1942);  (e)  C.  pisanum  <3  (after  Gerlach,  1952);  (0  C.  pisanum  juvenile  9  (after 
Gerlach,  1952);  (g)  C.  rectum  9  (after  Gerlach,  1957);  (h)  C.  reticulatum  d  (after  Chitwood,  1936); 
(i)  C.  salsicum  d  (after  Gerlach,  1956);  (j)  C.  sculpturatum  juvenile  (after  Chitwood,  1936);  (k)  C. 
undulatum  9  (after  De  Coninck,  1 942);  (1)  C.  yunfengi  sp.  nov. 


DIMENSIONS. 
Holotype  rf,  : 

Allotype  9,  : 


-  115     M  550 
15    23     23     19 

-  125  455  680    930/rni;  a  = 

17     25     32     23 


=  7-4;c  =  3-7;  V  =  49°/o 


cT2:  L  =  860  //m;     a  =  30;      b  =  7'5;  c  =  3'7 

a  =  35;      b  =  7'3;  c  =  3'5 

a  =  29*;     b  =  7'3;  c  =  4-2 

a  =  25*;     b  =  7'5;  c  =  3'5 
4  and  d"5  were  slightly  flattened. 


240 


H.  M.  PLATT  &  Z.  N.  ZHANG 


Fig.  7  Acantholaimus  ewensis:  (a)  whole  body  of  d1,;  (b)  head  of  d1,;  (c)  head  of  9,  showing  surface 
structures  only;  (d)  head  of  9,  showing  buccal  cavity  structure.  Bar  scales:  a=100wm; 
b-d=  10//m. 

DESCRIPTION.  Cuticle  punctated  and  laterally  differentiated.  Punctation  extends  anterior  to 
amphids  (Fig.  7c)  and  in  the  anterior  oesophageal  region  the  lateral  punctations  are  irregular. 
Further  posteriorly,  the  punctations  become  smaller  and  arranged  in  transverse  rows.  In  the 
middle  of  the  body,  the  transverse  rows  of  punctations  can  be  difficult  to  distinguish.  Lateral 
differentiation  consisting  of  two  longitudinal  rows  of  larger  punctations  begins  level  with  the 
middle  of  the  posterior  oesophageal  bulb  and  ends  at  about  65%  of  the  tail  length.  The  lateral 
rows  are  about  1  -5  /zm  apart  and  there  is  a  space  between  these  rows  and  the  transverse  rows 
of  punctations  (Fig.  8f).  There  are  four  sublateral  rows  of  hypodermal  pores,  which  are  larger 


NEW  SPECIES  OF  MARINE  NEMATODES 


241 


anteriorly  (Figs  8d-f).  Long  somatic  setae  are  present  throughout  the  length  of  the  body  and 
arranged  more  or  less  in  four  sublateral  rows.  In  the  oesophageal  region,  the  length  of  the 
somatic  setae  increases  from  15-20//m  at  the  anterior  to  3  5-40 //m  at  the  posterior.  In  the 
middle  of  the  body,  the  longer  setae  are  about  45  //m  long  but  decrease  in  length  again  to 
about  30-40  //m  in  the  anal  region  and  10-20  //m  in  the  tail. 

At  the  anterior  extremity  there  is  a  circle  of  six  2*5-3 /zm  stout  cephalic  sensilla, 
presumably  the  R2  sensilla.  Just  behind  are  four  30-33  //m  R3  cephalic  setae  which,  in  some 
specimens,  appear  to  have  a  joint  at  about  two-thirds  of  the  length  (Fig.  7b).  Amphids 
8'5-9'5//m  wide  (45-50%  c.d.),  circular  with  a  well  cuticularised  boarder  and  a  ventrally 
directed  posterior  inflection. 

Buccal  cavity  rather  complex  but  basically  it  is  conical  with  a  distinct  dorsal  and  two 
subventral  teeth  and  anteriorly  bears  twelve  rugae.  Oesophagus  has  characteristic  plasmatic 
interruptions  (Fig.  7d)  throughout  its  length  and  posteriorly  there  is  a  distinct  bulb, 
28  x  18  //m.  Tail  elongated  (11-12  a.b.d.),  gradually  tapering  but  with  a  slightly  bulbous  tip 
and  a  pointed  spinneret  (Fig.  8c). 


Fig.  8  Acantholaimus  ewensis:  (a)  tail  of  d1,;  (b)  copulatory  apparatus  of  d,;  (c)  tail  tip  of  $,;  (d)  d, 
cuticle  ornamentation  about  40  //m  from  anterior;  (e)  d,  cuticle  ornamentation  level  with  the 
posterior  of  the  oesophagus;  (f)  d,  cuticle  ornamentation  at  middle  of  body.  Bar  scales: 
a  =  30  //m;  b-f  =  1 0  //m. 


242 


H.  M.  PLATT  &  Z.  N.  ZHANG 

7A/ 


a 


Fig.  9  /J/zips  paraornata:  (a)  anterior  region  of  d1,;  (b)  head  of  d1,;  (c)  head  of  another  cf;  (d)  tail  of 
d1,;  (e)  copulatory  apparatus  of  d1,;  (fHO  cuticle  patterns  at  positions  indicated  in  Fig.  10.  Bar 
scales:  a,d  =  30  ^m;  b,c,e-l  =  10  /am. 


Spicules  paired,  equal,  26-28  ^m  long  and  with  a  characteristic  ventral  apophysis. 


Gubernaculum  absent.  In  d1,,  a  feint  cuticularised  structure  was  observed  lateral  to  the 
spicule  (Fig.  8b):  it  was  not  detected  in  the  other  specimens.  There  is  a  single  stout  ventral 
precloacal  seta,  5-6  //m  long.  Testis  single  and  mature  specimens  contain  large  pear-shaped 
sperm. 

Ovary  paired,  opposed  and  reflexed.  In  the  female  studied,  there  were  sperm  present  in  the 
oviduct  and  a  large  egg  in  each  uterus  containing  a  developing  larva. 

DIFFERENTIAL  DIAGNOSIS.  Acantholaimus  ewensis  sp.  nov.  can  be  distinguished  from 
Acantholaimus  species  with  two  lateral  longitudinal  rows  of  larger  punctations,  A.  poly- 


NEW  SPECIES  OF  MARINE  NEMATODES 


243 


Fig.  10    Rhips  paraornata:  whole  body  of  rf,  showing  positions  of  cuticle  pattern  drawings  in  Fig. 

9f-l.  Bar  scale  =100 //m. 

dentatus  Gerlach,  195 1  and  A.  calathus  Gerlach,  Schrage  &  Riemann,  1979,  on  the  length  of 
the  R3  cephalic  setae,  somatic  setae  and  tail.  A.  polydentatus  has  shorter  cephalic  setae 
(20-22  urn,  1-5  h.d.  vs.  30-33 /zm;  2'0  h.d.),  shorter  somatic  setae  (30 /zm,  0'75  c.d.  vs. 
45  um,  2-0  c.d.)  and  a  shorter  tail  (8  a.b.d.vs.  11-12  a.b.d.).  A  calathus  has  shorter  cephalic 
setae  ( 1 2  um,  1  -0  h.d.),  shorter  setae  (27  um,  1  -0  c.d.)  and  longer  tail  ( 1 7  a.b.d.). 

DISCUSSION.  The  genus  Acantholaimus  Allgen,  1933  has  been  fully  revised  recently  by 
Gerlach,  Schrage  &  Riemann  (1979)  who  described  seven  new  forms  and  provided  a  key  to 
the  twelve  known  species.  A.  ewensis  fits  into  the  first  part  of  their  key  as  follows: 

1  (4)       Cuticle  laterally  with  two  longitudinal  rows  of  larger  punctations      .       .  2 

2  (3)       Amphid  situated  close  behind  the  cephalic  setae 21 

T  (2")      Tail  8  anal  diameters  long  A.  polydentatus  Gerlach,  1951. 

1"  (2')      Tail  Tl-12  anal  diameters  long  A  ewensis  sp.  nov. 

3  (2)       Amphid  situated  half  the  head  diameter  behind  the  cephalic  seta.  Tail  1 7  anal 

diameters  long.  A.  calathus  Gerlach,  Schrage  &  Riemann,  1979 

4  (1)       Cuticle  with  limited  or  no  lateral  differentiation,  without  longitudinal  rows  of 

punctations 5 


Rhips  paraornata  sp.  nov. 
Figs  9- 10 

MATERIAL  STUDIED.  Holotype:  rf,  BM(NH)  1 98 1 .4. 19. 

Allotype:  9,  BM(NH)  1981.4.20.  Paratypes:  nine  males  and  two  females;  rf6  BM(NH) 

1 98 1 .4.2 1 ,  remainder  in  collection  of  Z.N.Z. 


244  H.  M.  PLATT  &  Z.  N.  ZHANG 

DIMENSIONS. 

Holotyperf,:  -  180  M  990  1 1050m;a  =  48;b  =  6-l;c  =  9-6;S  =  7 

11     21  23    22 

Allotypeg,:  -  179  M  938  10800m;a  =  37;b  =  6-0;c  =  7-6;  V  =  54% 

10    26  29    20 

rf2:L=  10850m;  a  =  38;  b  =  6'l;  c=8'7 

dy.L=13300m;  a  =  50;  b  =  6'5;  c=  9*3 

rf4:L=11300m;  a  =  48;  b  =  6'5;  c=ll'9 

rf5:L=12600m;  a  =  44;  b  =  6'4;  c=   8'5 

d"6:L=12600m;  a  =  47;  b  =  6*9;  c=ll'0 

d"7:L=  9200m;  a  =  42;  b  =  5'3;  c=   8-7 

d-8:L=12800m;  a  =  53;  b  =  6'7;  c=10'0 

cT9:L=12750m;  a  =  53;  b  =  7'4;  c=13'0 

<*w'.L=  11600m;  a  =  50;  b  =  6'9;  c=9'0 

92:L=10950m;  a  =  44;  b  =  6'6;  c=  9'0 

93:L=  9800m;  a  =  36;  b  =  6'5;  c=   7-3 

DESCRIPTION.  The  cuticle  is  conspicuously  annulated  and  heterogeneously  ornamented 
laterally.  In  the  anterior  third  of  the  oesophagus,  from  about  the  point  where  the  body 
characteristically  narrows  (Fig.  9a)  each  annule  bears  a  row  of  long  and  a  row  of  smaller 
round  punctations  (Fig.  90,  the  latter  being  difficult  to  distinguish  and  liable  to  be  over- 
looked. In  the  posterior  part  of  the  oesophagus,  the  punctations  are  smaller  and  appear  to  be 
partly  linked  diagonally  (Fig.  9g),  giving  a  reticulated  appearance.  The  regular  nature  of  the 
ornamentation  breaks  down  posterior  to  the  oesophagus  and  lateral  differentiation  of  large 
dumb-bell-shaped  punctations  begins  (Fig.  9h).  Just  posterior  to  the  mid-point  of  the  body, 
the  pattern  of  the  lateral  differentiation  reverses  and,  at  the  point  of  change,  the  dumb-bell- 
shaped  lateral  punctation  is  represented  by  a  single  round  dot  (Fig.  9j).  Lateral  differentiation 
ends  just  anterior  to  the  anus  and  transverse  rows  of  discrete  punctations  are  present  on  the 
tail  (Fig.  91). 

Short  4-6  0m  sublateral  somatic  setae  are  present  at  infrequent  intervals  throughout  the 
body  and  at  a  third  of  the  oesophagus  length,  there  are  four  longer  somatic  setae  (Fig.  9a). 
The  head  bears  six  setose  R,  sensilla.  The  six  2-3  0m  R2  sensilla  are  situated  just  anterior  to 
the  four  shorter  (1*5-2  0m),  R3  sensilla.  The  elongated  first  body  annule  forms  a  cephalic 
shield  and  bears  six  triangular  flap-like  extensions  anteriorly  which  alternate  with  the 
R2  +  R3  sensilla  (Fig.  9b,  c).  The  shield  is  irregularly  punctated  and  bears  the  large  transverse 
amphids.  The  amphid  has  strongly  cuticularised  borders  and  is  1 1  0m  wide,  about  0*85  c.b.d. 
The  buccal  cavity  is  surrounded  anteriorly  by  rugae  which  protrude  beyond  the  lips.  There  is 
a  large  pointed  dorsal  tooth  and  two  smaller  subventral  teeth.  The  oesophagus  widens 
posteriorly  to  a  weak  bulb.  The  tail  is  conical  and  has  an  unstriated  end  cone. 

The  male  copulatory  apparatus  is  complex,  consisting  of  two  long  double-jointed  spicules, 
a  paired  gubernaculum  and  two  lateral  pieces.  In  the  holotype,  the  posterior  part  of  the 
spicule  measures  40  0m  as  the  arc  (or  29  0m  as  the  chord)  and  the  anterior  part  measures 
38  0m  as  the  arc  (or  35  0m  as  the  chord).  In  five  other  males,  the  total  arc  length  of  the 
spicule  is  70-96  0m.  The  lateral  pieces  are  about  22  0m  long  and  the  gubernaculum  18  0m. 
The  cloacal  opening  is  surrounded  by  irregular  cuticular  excrescences  and  anteriorly  there  is 
a  small  ventral  spine.  The  ventral  part  of  the  precloacal  body  rings  are  more  thickly 
cuticularised:  they  gradually  reduce  in  thickness  anterior  to  the  level  of  the  proximal  end  of 
the  spicules  (Fig.  9d).  There  is  a  single  testis  situated  to  the  right  of  the  gut.  The  female  has 
two  opposed,  reflexed  ovaries. 

DIFFERENTIAL  DIAGNOSIS.  Rhips  paraornata  sp.  nov.  can  be  distinguished  from  the  only 
other  valid  species  in  the  genus,  R.  ornata  Cobb,  1920,  by  having  slightly  shorter  R2  cephalic 
setae,  wider  amphids,  spicules  with  each  part  about  the  same  length  and  possibly  a 
differently  shaped  lateral  differentiation. 


NEW  SPECIES  OF  MARINE  NEMATODES  245 

DISCUSSION.  This  is  only  the  fifth  time  that  valid  specimens  of  this  genus  have  been  reported. 
Timm(  1961)  described  a  species/?,  longicauda  from  the  Bay  of  Bengal,  but  the  description  is 
very  poor  and  based  only  on  a  single  immature  female:  it  must  be  considered  a  dubious 
species. 

The  original  description  of  the  type  and  only  other  species,  R.  ornata,  was  by  Cobb  (1920): 
the  specimens  from  Loch  Ewe  are  clearly  similar  to  this  species  from  Florida.  The  species 
was  found  again  by  Wieser  &  Hopper  (1967)  who  provide  a  brief  redescription  and  figure  the 
head.  Allgen  (1932)  found  what  is  certainly  a  male  Rhips  species  from  Campbell  Island  and 
considered  it  conspecific  with  Cobb's  species  and  Gerlach  (1957)  found  R.  ornata  in  Brazil 
but  did  not  describe  it. 

Cobb  (1920)  described  the  unusual  triangular  scale-like  cephalic  flaps  but  only  saw  the 
four  sublateral  ones:  Wieser  &  Hopper  (1967)  correctly  observed  all  six.  The  four  long 
cervical  setae,  located  at  about  one-third  the  oesophagus  length  in  R.  paraornata,  were  not 
reported  in  R.  ornata  but  they  are  fine  and  may  be  lost  on  handling  so  that  their  absence  in  R. 
ornata  cannot  be  assumed.  The  four  sublateral  cephalic  flaps  were  positioned  'just  in  front  of 
the  ends  of  the  amphids'  in  R.  ornata  according  to  Cobb  (1920)  and  as  figured  by  Wieser  & 
Hopper  (1967),  whilst  the  ends  of  the  amphids  in  R.  paraornata  extend  further  beyond  the 
flaps.  Both  Cobb  (1920)  and  Wieser  &  Hopper  (1967)  describe  the  lateral  differentiation  in 
the  posterior  portion  of  the  body  as  being  'V-shaped  but  unfortunately  provide  no 
illustrations  of  the  cuticle  patterns.  If  the  lateral  differentiation  is  similar  in  both  species, 
then  we  would  not  have  chosen  to  describe  the  lateral  differentiation  as  'V'-shaped.  It  is 
possible,  then,  that  the  cuticle  patterns  are  different.  Finally,  the  spicules  ofR.  ornata  have 
an  anterior  part  which  is  1-5  times  the  length  of  the  posterior  part  whilst  in  R.  paraornata  the 
lengths  are  almost  the  same,  measured  as  the  arc  in  both  cases.  Therefore,  although  there  are 
many  points  of  similarity  in  overall  dimensions  and  general  anatomy,  there  are  sufficient 
points  of  difference  in  relation  to  the  specific  sizes  of  cephalic  setae,  amphids  and  spicules 
and  possibly  the  form  of  the  cuticle  lateral  differentiation  to  substantiate  the  creation  of  a 
new  species  for  the  specimens  from  Loch  Ewe. 

Species  of  the  genus  Rhips  seem  to  be  very  closely  related  to  Actinonema,  particularly 
through  the  excellent  redescription  of  the  common  species  A.  pachydermatum  Cobb,  1920 
by  Lorenzen  (1972):  both  have  similar  amphids,  cuticle  patterns  and  the  six  triangular 
extensions  to  the  cephalic  shield,  'Kopfpanzer',  although  the  flaps  in  A.  pachydermatum  are 
not  as  conspicuous  as  those  in  R.  paraornatum  and  may  have  been  overlooked  by  earlier 
workers.  However,  Actinonema  does  not  have  large  double-jointed  spicules  and,  according 
to  Lorenzen  (1972),  between  those  structures  which  he  terms  spicules,  but  which  resemble 
the  lateral  pieces  of  Rhips,  there  lies  a  single  thin  cuticularised  tube  which  Lorenzen  (1972) 
interprets  as  the  cuticularised  terminal  portion  of  the  vas  deferens.  We  have  studied  several 
specimens  of  A.  pachydermatum,  sympatric  with  R.  paraornata,  and  find  the  cuticularised 
tube  very  difficult  to  distinguish.  However,  in  some  specimens  it  is  just  possible  to  make  out 
two  tubes:  if  so,  they  cannot  be  vas  deferens  since  A.  pachydermatum  is  monorchic.  This 
problem  is  of  systematic  importance  since  Rhips  and  Actinomena,  together  with  a  number  of 
other  genera  including  Euchromadora,  are  grouped  together  in  the  subfamily  Euchro- 
madorinae.  This  subfamily  was  erected  by  Gerlach  &  Riemann  (1973)  without  explanation 
but  presumably  because  of  the  presence  of  the  'L'-shaped  lateral  pieces  in  addition  to 
spicules  and  gubernaculum.  For  Actinonema  to  fit  into  this  group,  which  seems  reasonable 
through  its  similarity  to  Rhips,  the  spicules  of  Actinonema  must  be  considered  homologous 
with  the  lateral  pieces  of  Rhips  and  either  the  cuticularised  tube(s?)  are  vestigial  spicules  or 
spicules  are  absent.  For  the  moment,  this  problem  must  remain  open  for  further  study. 

Acknowledgements 

We  would  like  to  thank  Dr  A.  D.  Mclntyre  of  the  D.A.F.S.  Marine  Laboratory,  Aberdeen  for 
providing  the  opportunity  for  one  of  us  (Z.N.Z.)  to  spend  two  years  studying  benthos  at  that 
institute:  we  named  one  of  our  new  species  in  his  honour.  We  would  also  like  to  thank  the 


246  H.  M.  PLATT  &  Z.  N.  ZHANG 

staff  of  the  benthos  section  for  help,  encouragement  and  the  provision  of  specimens,  in 
particular  Dr  A.  Eleflheriou  and  Mr  D.  J.  Murison. 

References 

Allgen,  C.  1932.  Weitere  Beitrage  zur  Kenntnis  der  marinen  Nematodenfauna  der  Campbell-insel.  Nyt. 
Mag.  Naturvid.  70  :  97-198. 

-  1933.  Freilebende  Nematoden  aus  dem  Trondhjemsfjord.  Capita  zool.  4  :  1-162. 

Andrassy,  I.  1976.  Evolution  as  a  basis  for  the  systematization  of  nematodes.  288pp.  Pitman  — 

London,  San  Francisco,  Melbourne. 
Boucher,  G.  1975.  Nematodes  des  sables  fins  infralittoraux  de  la  Pierre  Noire  (Manche  occidentale).  I. 

Desmodorida.  Bull.  Mus.  natn.  Hist.  nat.  Paris  285  :  101-128. 
Chitwood,  B.  G.    1936.  Some  marine  nematodes  from  North  Carolina.  Proc.  helminth.  Soc.  Wash. 

3:  1-16. 
Cobb,  N.  A.  1920.  One  hundred  new  nemas  (type  species  of  100  new  genera).  Contrib.  to  a  Science  of 

Nematology9  :  217-343. 
De  Coninck,  L.  A.  1942.  Sur  quelques  especes  nouvelles  de  Nematodes  libres  (Ceramonematinae  Cobb, 

1933),  avec  quelques  remarques  de  systematique.  Bull.  Mus.  r.  Hist.  nat.  Belg.  18:  1-37. 
Filipjev,  I.  N.  1918.  Free-living  marine  nematodes  of  the  Sevastopol  area.  Trudy  osob.  zool.  Lab. 

Sevastop.  biol.  Sta.  4  :  1-350  (in  Russian). 
Gerlach,  S.  A.   1951.  Drei  bemerkenswerte  neue  Nematoden  aus  der  Kieler  Bucht.  Zool.  Anz. 

147  :  37^3. 

-  1952.  Nematoden  aus  dem  Kiistengrundwasser.  Abh.  math,  -naturw.  Kl.  Akad.  Wiss.  Mainz 
6:315-372. 

-  1956.  Diagnosen  neuer  Nematoden  aus  der  Kieler  Bucht.  Kieler  Meeresforsch.  12  :  85-109. 
1957.  Die  Nematodenfauna  des  Sandstrandes  an  der  Ku'ste  von  Mittelbrasilien  (Brasilianische 


Meeres-Nematoden  IV).  Mitt.  zool.  Mus.  berl.  33  :  41 

19630.  Freilebende  Meeresnematoden  von  den  Maldiven  II.  Kieler  Meeresforsch.  14  :  67-103. 
19636.  Robbea  tenax  sp.  n.,  ein  merkwiirdiger  mariner  Nematode  von  den  Maldiven.  Int.  Revue 
ges.  Hydrobiol.  Hydrogr.  48  :  1  53-1  58. 

&  Riemann,  F.  1973.  The  Bremerhaven  checklist  of  aquatic  nematodes.  A  catalogue  of  Nematoda 
Adenophorea  excluding  the  Dorylaimida.  Verojf.Inst.  Meeresforsch.  Bremerh.  Suppl.  4  :  1-404. 
-,  Schrage,  M.  &  Riemann,  F.  1979.  Die  GattungAcantholaimus  (Nematoda,  Chromadoridae),  und 


Beobachtungen  iiber  einen  mutmasslichen  Transportmechanismus  fur  Spermien  bie  A.  calathus  sp. 

n.  Veroff.Inst.  Meeresforsch.  Bremerh.  18  :  35-67. 
Haspeslagh,  G.  1973.  Superfamille  des  Ceramonematoidea  (Cobb,  1933)  (Nematoda),  evolution  et 

systematique.  Annls  Soc.  r.  zool.  Belg.  102  :  235-25 1 . 
Hopper,  B.  E.  &  Cefalu,  R.  C.  1973.  Free-living  marine  nematodes  from  Biscayne  Bay,  Florida  V. 

Stilbonematinae:  contributions  to  the  taxonomy  and  morphology  of  the  genus  Eubostrichus  Greef 

and  related  genera.  Trans.  Am.  microsc.  Soc.  94  :  578-59 1 . 
Inglis,  W.  G.  1968.  Interstitial  nematodes  from  St.  Vincent's  Bay.  New-Caledonia.  Editions  Fondat. 

Singer- Polignac,  Paris,  19672:  29-74. 
Lorenzen,  S.  1972.  Die  Nematodenfauna  im  Verklappungsgebiet  fur  Industrieabwasser  nordwestlich 

von  Helgoland  II.  Desmodorida  und  Chromadorida.  Zool.  Anz.  187  :  283-302. 
1981.  Entwurf  eines  phylogenetischen  Systems  der  freilebenden  Nematoden.    Veroff.  Inst. 

Meeresforsch.  Bremerh.  Suppl.  7  :  1-472. 
Mclntyre,  A.  D.  &  Eleftheriou,  A.  1968.  The  bottom  fauna  of  a  flatfish  nursery  ground.  J.  mar.  biol. 

Ass.  U.K.  48:  113-142. 
&  Murison,  D.  J.  1973.  The  meiofauna  of  a  flatfish  nursery  ground.  J.  mar.  biol.  Ass.  U.K. 

53:93-118. 
Platt,  H.  M.  1973.  Freeliving  marine  nematodes  from  Strangford  Lough,  Northern  Ireland.  Cah.  Biol. 

mar.  14:295-321. 

Steele,  J.  H.  &  Baird,  I.  E.  1968.  Production  ecology  of  a  sandy  beach.  Limnol.  Oceanogr.  13  :  14-25. 
Timm,  R.  W.  1961 .  The  marine  nematodes  of  the  Bay  of  Bengal.  Proc.  Pakist.  Acad.  Sci.  1  :  1-88. 
Vitiello,  P.  1974.  Nouvelles  especes  de  Desmodorida  (Nematoda)  des  cotes  de  Provence.  Tethys 

5:  137-146. 
Wieser,  W.  1959.  Free-living  nematodes  and  other  small  invertebrates  ofPuget  Sound  beaches.  1 79  pp. 

University  of  Washington  Press,  Seattle. 
&  Hopper,  B.  Marine  nematodes  of  the  east  coast  of  North  America.  Bull.  Mus.  comp.  Zool.  Harv. 

135  : 239-244. 

Manuscript  accepted  for  publication  14  September  1981 


The  larval  development  of  Crangon  cmngon 
(Fabr.  1795)  (Crustacea:  Decapoda) 

A.  R.  Gurney 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London 
SW7  5BD 

Introduction 

The  first  larval  stage  of  Crangon  crangon  (Fabr.,  1795)  was  recorded  by  Du  Cane  in  1839 
and  since  then  descriptions  of  some  of  the  subsequent  larval  stages  have  been  added.  In 
1890  Ehrenbaum  described  five  larval  stages  but  suggested,  as  they  were  taken  from 
plankton  samples,  that  his  series  may  not  have  been  complete.  Williamson  (1901) 
reared  the  first  three  stages  in  the  laboratory,  and  selected  examples  from  tow  net 
plankton  samples  concluding  that  there  were  probably  five  larval  stages  in  all. 
Experimental  evidence  on  the  development  of  larvae  under  non-circadian  light/dark 
cycles  given  by  Dalley  (1980),  also  suggested  there  were  five  larval  stages.  The  present 
study  is  based  on  specimens  reared  in  the  laboratory,  and  describes  six  larval  stages,  one 
more  than  those  previously  recorded. 

Materials  and  method 

Ovigerous  Crangon  crangon  (Fabr.,  1795)  were  trawled  in  the  Sound,  Plymouth,  in 
March,  1980.  Rearing  techniques  follow  those  of  Fincham,  1977.  Representatives  of 
each  stage  are  deposited  in  the  Crustacea  collections  of  the  British  Museum  (Natural 
History),  registration  numbers  1981  :  394^00. 


Description  of  larval  stages 

Key  characters  are  printed  in  italic  type',  many  setal  counts  have  been  omitted  but  these 
are  recorded  in  Table  1 . 

ZOEA  1  (Fig.  1 )  mean  size  2  mm 
Head  (Figs  la,  b):  eyes  are  sessile. 
Carapace  (Figs  la,  b):  without  spines;  rostrum  short. 

Antenna  1   (Fig.   Ic):  single  peduncle  segment  bearing  external  flagellar  segment  with 
three  aesthetascs  distally  plus  one  seta;  terminal  setose  spine  on  peduncle  segment. 
Antenna  2  (Fig.  Id):  expedite  broad  and  flat  with  ten  setae,  all  plumose  except  the  penulti- 
mate, subapical  medial  spine  absent;  endopodite  with  terminal  spinous  seta;  spine  on 
distal  edge  of  peduncle  segment  at  base  of  endopodite. 
Mandibles  (Fig.  le):  symmetrical. 

Maxilla  1  (Fig.  If):  coxa  with  seven  spines;  basis  with  five  spines;  endopod  with  five 
setae  plus  one  small  spine. 

Maxilla  2  (Figs  Ig,  h):  endopod  coxa  bilobed  on  inner  margin,  proximal  lobe  with  six  setae, 
distal  lobe  with  four  setae;  basis  one  with  four  setae;  basis  two  with  four  setae, 
endopodite  with  four  lobes  on  inner  margin,  fourth  proximal  lobe  with  three  setae,  third 
sub  proximal  lobe  with  two  setae,  second  subdistal  lobe  with  one  seta,  distal  lobe  with  two 
setae;  exopodite  with  five  setae. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  42  (4) :  247-262  Issued  24  June  1 982 


248  A.  R.  GURNEY 

Maxillipeds  1-3  (Figs  li-1):  with  natatory  exopodites. 

Pereiopod  1  (Fig.  1  m):  rudimentary,  biramous. 

Pereiopods  2-4  (Figs  1  m):  rudimentary,  uniramous. 

Pereiopod  5:  absent. 

Abdomen  (figs  la,  b):  5  somites,  6th  somite  continuous  with  telson. 

Telson  (Fig.  In):  fans  out  distally,  distal  margin  with  7+7  terminal  plumose  spines. 

ZOEA  2  (Fig.  2)  mean  size  2 -4  mm 

Head  (Figs  2a,  b):  eyes  'stalked'. 

Antenna  1  (Fig.  2c):  peduncle  two  segmented;  internal  distal  margin  of  second  segment 

extended. 

Pereiopod  1  (Fig.  2m):  1- segmented  endopod  with  one  apical  seta;  natatory  exopodite. 

Telson  (Fig.  2o):  distal  margin  with  8  +  8  terminal  plumose  spines. 

ZOEA  3  (Fig.  3)  mean  size  3'0  mm 

Antenna  1  (Fig.  3c):  external  jlagellum  with  four  distal  aesthetascs,  one  narrower  than 

others  and  one  distal  seta. 

Pereiopod  1  (Fig.  3k):  2-segmented  endopodite. 

Telson  (Fig.  3n):  divided  from  somite  6  by  suture;  exopod  and  endopod  of  uropod  with 

fringing  plumose  setae. 

ZOEA  4  (Figs  4  &  5)  mean  size  3'5  mm 

Antenna  1  (Fig.  4c):  rudimentary  stylocerite. 

Periopod  1  (Fig.  5d):  3 -segmented  endopodite. 

Abdomen  (Fig.  4b):  rudimentary  buds  of  pleopods  on  somites  1-5. 

ZOEA  5  (Figs  6  &  7)  mean  size  4- 1  mm 

Antenna  1  (Fig.  6c):  external  jlagellum  segment  with  4  wide  distal  aesthetascs  and  one 

narrow  sub-distal  aesthetasc  plus  one  setule. 

Pereiopod  1  (Fig.  7d):  4-segmented  endopodite. 

Pereiopods  2-3  (Fig.  7e):  uniramous  1 -segmented. 

Pereiopod  4  (Fig.  7f):  uniramous  4-segmented. 

Pereiopod  5  (Fig.  7g):  uniramous  5-segmented. 

ZOEA  6  (Figs  8  &  9)  mean  size  4*2  mm 
Pereiopod  1  (Fig.  9d):  5-segmented  endopodite. 
Pereiopods  2-5  (Figs  9e-f):  uniramous  6-segmented. 

POST  LARVA  1  (Figs  10-12)  mean  size  4-4  mm 

Carapace  (Figs  lOa,  b):  rostrum  short;  one  dorso-medial  tooth. 

Antenna  1  (Fig.  lOc):  3-segmented  peduncle;  external  flagellum  2-segmented  with  four 

distal  and  two  sub-distal  aesthetascs,  internal  flagellum  3-segmented;  stylocerite  with 

terminal  plumose  setae. 

Antenna  2  (Fig.  lOd):  exopodite  with  19  plumose  setae;  endopodite  multisegmented. 

Mandibles  (Fig.  10e):  symmetrical. 

Maxilla  1  (Fig.   10f):  coxa  with  5  spines;  basis  with  9  spines  plus  one  plumose  seta; 

endopod  with  one  spine. 

Maxilla  2  (Fig.  lOg):  endopod  reduced;  exopod  >22  plumose  setae. 

Maxilliped  1  (Fig.  11  a):  endopod  reduced  with  one  plumose  seta;  exopod  with  3  short 

plumose  setae  distally  plus  one  seta,  one  plumose  seta  proximally;  epipodite  present. 

Maxilliped  2  (Figs,  lib,  c):  endopod  5-segmented,  exopod  with  reduced  setae. 

Maxilliped  3   (Fig.    lid):   endopod  3-segmented;  exopod  with  very  reduced  plumose 

setae. 

Pereiopod  1  (Figs  lie,  0:  exopod  reduced;  endopod  4-segmented,  propodus  and  dactylus 

subchelate. 

Pereiopod  2  (Fig.  1  Ig):  5-segmented,  chelate;  epipodite  present. 

Pereiopod  3  (Figs  1 1  h,  i):  7-segmented. 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 


249 


Table  1     Morphological  comparison  of  larval  stages  1-6  and  post  larva.  (v  =  visible;  d  =  developed; 
a  =  absent;  p  =  present;  r  =  rudimentary;  b  =  biramous;  re  =  reduced) 


ZOEA  STAGE 

1 

2 

3 

4 

5 

6 

PL 

MEAN  SIZE 

2*0  mm 

2*4  mm 

3-0  mm 

3'5  mm 

4'1  mm 

4-2  mm 

4-4  mm 

ANTENNA  1 

No.  of  aesthetascs. 

3 

3 

4 

4 

5 

6 

6 

Stylocerite. 

a 

a 

a 

V 

V 

V 

d 

No.  of  segments. 

1 

2 

2 

2 

2 

2 

3 

ANTENNA  2 

Presence  of  subapical  medial 

spine. 

a 

P 

P 

P 

P 

P 

P 

Exopodite;  no.  of  plumose  setae 

incl.  fine  setae  immediately 

after  spine. 

10 

11 

12-13 

13-14 

13-14 

14-16 

19 

MAXILLA  1 

No.  of  setae  on  coxa; 

7 

7 

7 

7 

8 

8 

1 

basis; 

5 

7 

8 

8 

9 

9 

10 

endopod. 

5  +  1 

5  +  1 

5  +  1 

5  +  1 

5+1 

5  +  1 

4+1 

MAXILLA  2 

No.  of  endite  setae  on  endopodite 

lobes  1,2,3,  4; 

2  1.2.3 

2.1.2.3 

2.1.2.3 

2.1.2.3 

2.1.2.3 

2.1.2.3 

re 

basis  1  ; 

4 

4 

4 

4 

4 

4 

re 

basis  2; 

4 

4 

4 

4 

4 

4 

re 

coxa  (prox.  &  distal). 

6  +  4 

7+4 

7  +  4 

7  +  4 

7+4 

7+4 

re 

No.  of  plumose  setae  on  exopodite. 

5 

10 

10-13 

13-15 

15-19 

20-22 

>22 

MAXILLIPED  1 

No.  of  setae  on  endite  of  coxa; 

4 

5 

5 

5 

5 

5 

re 

basis. 

11 

13 

14 

14 

14 

14 

re 

No.  of  plumose  setae  on  exopodite. 

4 

5 

5 

5 

5 

5 

re 

PEREIOPOD  1 

No.  of  segments  in  endopodite. 

r/b 

1 

2 

3 

4 

5 

4 

No.  of  setae  on  endopodite. 

a 

1 

2 

V 

V 

V 

- 

Presence  of  natatory  exopodite. 

a 

P 

P 

P 

P 

P 

re 

PEREIOPOD  2 

No.  of  segments. 

r 

r 

r 

r 

r 

6 

5 

No.  of  setae. 

- 

- 

- 

1 

1 

- 

PEREIOPOD  3 

No.  of  segments. 

r 

r 

r 

r 

r 

6 

7 

No.  of  setae. 

- 

- 

- 

- 

1 

1 

- 

PEREIOPOD  4 

No.  of  segments. 

r 

r 

r 

r 

4 

6 

7 

PEREIOPOD  5 

No.  of  segments. 

a 

r 

r 

r 

5 

6 

7 

ABDOMEN 

Somite  6  —  distal  suture  present. 

a 

a 

P 

P 

P 

P 

P 

TELSON 

No.  of  segments. 

14 

16 

16 

16 

16 

16 

10 

UROPODS 

Presence. 

a 

a 

P 

P 

P 

P 

P 

250  A.R.  GURNEY 

Pereiopod  4  (Figs  1  Ij,  k):  7-segmented. 

Pereiopod  5  (Figs  1 11,  m):  7-segmented. 

Pleopod   1   (Fig.   12a):  inner  margin  of  exopod  with  6  plumose  setae,  2  apical  setae, 

external  margin  with  4  plumose  setae. 

Pleopods  2-4   (Figs    12b-d):   exopod   inner  margin   6   plumose  setae,  2   apical   setae, 

external  margin  with  5  plumose  setae. 

Pleopod  5  (Fig.  12e):  exopod  inner  margin  with  5  plumose  setae,  2  apical  setae,  external 

margin  with  4  plumose  setae. 

Telson  (Fig.  120:  5  +  5  spines;  exopod  of  uropod  with  one  spine  next  to  subapical  medial 

spine. 


Discussion 

The  larval  stages  of  Crangon  crangon  (Fabr.,  1795)  have  not  previously  been  described 
solely  from  laboratory  reared  material.  Williamson  (1901)  figured  his  larval  series  from 
the  first  three  stages  reared  in  the  laboratory,  supplemented  by  larval  stages  four  and 
five  and  the  post  larva  taken  from  plankton  samples.  All  earlier  records  indicate  five 
larval  stages.  In  the  present  work  six  larval  stages  are  produced  consistently  under 
laboratory  conditions.  The  insertion  of  an  additional  larval  stage  prolonging  development 
is  a  feature  shared  by  other  carideans  (Fincham,  1977).  In  planktonic  larvae  delaying  the 
onset  of  metamorphosis  until  conditions  are  favourable  provides  the  maximum  chance 
of  survival  for  the  mainly  benthic  juveniles. 

Minor  differences  related  to  the  insertion  of  an  additional  larval  stage  become 
apparent  when  comparisons  to  Williamson's  (1901)  work  are  made.  The  numbers  of 
aesthetascs  recorded  in  this  paper  accord  with  Williamson  up  to  stage  four  when  he 
then  records  6  (4  distal  and  2  subdistal)  compared  with  5  aesthetascs  (4  distal  and  1 
subdistal)  for  stage  5;  7  (4  distal  and  3  subdistal)  in  first  post  larva  compared  with  6  (4 
distal  and  2  subdistal)  in  post  larva  recorded  here.  Maxilliped  1  showed  variation  in  the 
early  stages.  Williamson  records  10  spines  on  the  basis  in  stage  1,  12  in  stage  2,  13  in 
stage  3  compared  with  11  in  stage  1,  13  in  stage  2  and  14  in  stage  3  recorded  here. 
Pereiopods  2-5  in  the  last  two  larval  stages  differed  in  segmentation.  Williamson 
showed  stage  4  pereiopod  2  as  1 -segmented,  pereiopod  3  as  3-segmented  and 
pereiopods  4  and  5  as  2-segmented;  stage  5  pereiopods  2-5  as  7-segmented. 
Recorded  in  this  paper  stage  5  pereiopod  2  is  rudimentary,  pereiopod  3  is  1 -segmented, 
pereiopod  4  is  4-segmented  and  pereiopod  5  is  5-segmented;  stage  6  pereiopods  2-5 
are  6-segmented.  These  differences  are  almost  certainly  related  to  the  insertion  of  the 
extra  larval  stage  after  stage  2  prior  to  metamorphosis. 


References 

Dalley,  R.  1980.  The  survival  and  development  of  the  shrimp  Crangon  crangon  (L.)  reared  in  the 
laboratory  under  non-circadian  light-dark  cycles.  J.  exp.  mar.  Biol.  Ecol.  47  :  101-112. 

Du  Cane,  C.  1 839.  Metamorphosis  of  Crustacea.  Ann.  Mag.  nat.  Hist.  2  :  1 78-1 8 1 . 

Ehrebaum,  E.  1890.  Zur  naturgeschichte  von  Crangon  vulgaris  Fabr.  In:  Studien  tiber  Bau, 
Entwichklung,  Lebenweise  und  Fanguerhdltnisse  des  Nordsee-Granat  im  Auftrage  der  Sektion  fur 
Kusten-und  Hochseefischerei  des  Deutschen  Fischerei-  Vereins.  Moeser,  Berlin.  124  pp. 

Fincham,  A.  A.  1977.  Larval  development  of  British  prawns  and  shrimps  (Crustacea:  Decapods: 
Natantia).  1.  Laboratory  methods  and  a  review  of  Palaemon  (Palaeander)  elegans  Rathke,  1837. 
Bull.  Br.  Mus.  nat.  Hist.  (Zool.)32(l) :  1-28. 

Williamson,  H.  C.  1901.  On  the  larval  stages  of  decapod  Crustacea — the  shrimp  (Crangon  vulgaris, 
Fabr.)  Rep.  Fishery  Bd  Scotl.  19  :  92-1 19. 

Manuscript  accepted  for  publication  28  September  1 98 1 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 

b     /- \  c  /    I  d 


251 
h 


Fig.  1     Zoea  1:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e)  mandibles;  (f) 
maxilla  1 ;  (g)  maxilla  2;  (h)  armature  of  maxilla  2;  (i)  maxilliped  1 ;  (j)  armature  of  maxilliped  1 ; 
(k)  maxilliped  2;  (1)  maxilliped  3;  (m)  pereiopods  1-5;  (n)  telson. 
Bar  scales:  a,  b  =  0'35  mm;  c-n  =  0'  1  mm. 


Fig.  2    Zoea  2:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e)  mandibles;  (0 
maxilla  1;  (g)  maxilla  2;  (h)  armature  of  maxilla  2;  (i)  maxilliped  1;  (j)  armature  of  maxilliped  1; 
(k)  maxilliped  2;  (1)  maxilliped  3;  (m)  pereiopod  1;  (n)  pereiopods  2-5;  (o)  telson;  (p)  detail  of 
spine,  3rd  from  right,  on  telson. 
Bar  scales:  a,  b  =  0'3  mm;  c-p  =  0'  1  mm. 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 


n 


Fig.  3    Zoea  3:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e)  mandibles;  (0 
maxilla  1;  (g)  maxilla  2;  (h)  maxilliped  1;  (i)  armature  of  maxilliped  1;  (j)  maxilliped  2;  (k) 
maxilliped  3;  (1)  pereiopod  1 ;  (m)  pereiopods  2-5;  (n)  telson. 
Bar  scales:  a,  b  =  0-2  mm;  c-g  =  0 1  mm. 


254 


A.  R.  GURNEY 


Fig.  4    Zoea  4:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e)  mandibles;  (0 
maxilla  1 ;  (g)  maxilla  2. 
Bar  scales:  a,  b  =  0-3  mm;  c-g  =  0- 1  mm. 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 


255 


Fig.  5    Zoea  4:  (a)  maxilliped   1;  (b)  maxilliped  2;  (c)  maxilliped  3;  (d)  pereiopod   1;  (e) 
pereiopods  2  &  3;  (f)  pereiopods  4  &  5;  (g)  telson. 
Bar  scales:  a-g  =  0'  1  mm. 


256 


A.  R.  GURNEY 


Fig.  6    Zoea  5:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e)  mandibles;  (0 
maxilla  1 ;  (g)  maxilla  2. 
Bar  scales:  a,  b  =  0'3  mm;  c-g  =  0- 1  mm. 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 


257 


Fig.  7    Zoea  5:  (a)  maxilliped   1;  (b)  maxilliped  2;  (c)  maxilliped  3;  (d)  pereiopod   1;  (e) 
pereiopods  2  &  3;  (f)  pereiopods  4  &  5;  (g)  telson. 
Bar  scales:  a-g  =  0- 1  mm. 


258 


A.  R.  GURNEY 


' 


Fig.  8    Zoea  6:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e)  mandibles;  (f) 
maxilla  1 ;  (g)  maxilla  2. 
Bar  scales:  a,  b  =  0*3  mm;  c-g  =  0- 1  mm. 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 


259 


Fig.  9    Zoea  6:  (a)  maxilliped   1;  (b)  maxilliped  2;  (c)  maxilliped  3;  (d)  pereiopod   1;  (e) 
pereiopods  2  &  3;  (0  pereiopods  4  &  5;  (g)  telson. 
Bar  scales:  a-g  =  0'  1  mm. 


260 


A.  R.  GURNEY 


Fig.  10     Post  Larva  1:  (a)  dorsal  view;  (b)  lateral  view;  (c)  antenna  1;  (d)  antenna  2;  (e) 
mandibles;  (0  maxilla  1 ;  (g)  maxilla  2. 
Bar  scales:  a  =  0*3 5  mm;b  =  0-4  mm;c-g  =  0-l  mm. 


LARVAL  DEVELOPMENT  OF  CRANGON  CRANGON 


261 


Fig.  11     Post  Larva  1:  (a)  maxilliped  1;  (b)  maxilliped  2;  (c)  detail  of  endopod  of  maxilliped  2; 
(d)  maxilliped  3;  (e)  pereiopod  1;  (0  detail  of  pereiopod  1;  (g)  pereiopod  2;  (h)  pereiopod  3;  (i) 
detail  of  terminal  segment  pereiopod  3;  (j)  pereiopod  4;  (k)  detail  of  terminal  segment 
pereiopod  4;  (1)  pereiopod  5;  (m)  detail  of  terminal  segment  pereiopod  5. 
Bar  scales:  a-m  =  0- 1  mm. 


262 


a 


A.  R.  GURNEY 

be  d 


Fig.  12     Post  Larva  1:  (a)  pleopod  1;  (b)  pleopod  2;  (c)  pleopod  3;  (d)  pleopod  4;  (e)  pleopod  5; 
(0  telson. 
Bar  scales:  a-f  =  0'  1  mm. 


A  revision  of  the  spider  genus  Cocalodes  with  a 
description  of  a  new  related  genus  (Araneae: 
Salticidae) 

F.  R.  Wanless 

Department  of  Zoology,  British  Museum  (Natural  History)  Cromwell  Road,  London 
SW7  5BD. 


Introduction 

The  genus  Cocalodes  Pocock,  1897,  known  from  Amboina,  Seram,  Halmahera  and 
New  Guinea,  is  comprised  of  12  species,  six  of  which  are  described  here  as  new. 
Petrunkevitch  (1928)  placed  Cocalodes  in  the  subfamily  Boethinae,  but  as  far  as  I  am 
aware  it  shows  no  close  affinities  with  the  nominate  genus  Boethus  or  related  genera 
(sensu  Wanless,  1981).  It  is,  however,  closely  allied  to  Allococalodes  gen.  n.,  proposed 
here  for  two  new  taxa  from  New  Guinea. 

Both  genera  are  unique  in  that  the  majority  of  males  possess  a  well  developed  median  horn 
arising  from  a  sclerite  between  the  lower  basal  margins  of  the  chelicerae  (Figs  8  A,  2 1 B).  This 
structure  has  not  been  described  in  Cocalodes  by  earlier  authors,  for  in  the  majority  of 
species  it  does  not  protrude  beyond  the  edge  of  the  clypeus  and  is  readily  overlooked.  The 
horn  is  not  homologous  with  the  paired  horns  ofPadilla  Peckham  &  Peckham,  which  arise 
from  the  basal  region  of  the  anterior  surface  of  the  chelicerae.  Neither  is  it  homologous  with 
the  horn  of  Thorellia  Keyserling  which  arises  from  the  clypeus. 

The  male  palps  of  both  Cocalodes  and  Allococalodes  are  also  of  interest  in  possessing  a 
functional  conductor  and  median  apophysis  which  seldom  occur  in  other  Salticidae.  In 
Allococalodes  the  functional  conductor  and  median  apophysis  are  lobe-like  (Fig.  16E)  and 
relatively  simple  when  compared  with  the  elaborate  fan-like  functional  conductor  and  bifid 
median  apophysis  characteristic  of  Cocalodes  (Fig.  6G).  The  less  complex  structures  of 
Allococalodes  are  possibly  vestigial  or  germinal  in  respect  of  those  of  Cocalodes,  but  for  the 
present  it  is  a  matter  for  conjecture  as  the  affinities  of  both  genera  are  uncertain. 

The  standard  abbreviations  and  measurements  are  those  used  by  Wanless  (1978), 
but  for  the  leg  spination  the  system  is  that  used  by  Platnick  and  Shadab  (1975). 

Genus  COCALODES  Pocock 

Cocalodes  Pocock,  1897:627.  Type  species  Cocalodes  leptopus  Pocock,  by  original  designation. 
Simon,  1901:400,  403^405.  Waterhouse,  1902:80.  Petrunkevitch,  1928:181.  Neave,  1939, 
1 :  778.  Roewer,  1954  :  936.  Bonnet,  1956  :  1 172. 

DEFINITION.  Small  or  large  spiders  ranging  from  about  4*5  to  11 -Omm  in  length.  Most 
species  elongate  and  narrow  with  long  slender  legs,  the  abdomen  usually  marked  with 
characteristic  dark  lateral  bands;  chelicerae  robust,  in  males  usually  elongate  and 
porrect  with  a  median  horn  which  sometimes  protrudes  beyond  the  clypeus;  not  hirsute, 
fringes  lacking. 

Carapace:  longer  than  broad,  moderately  high,  widest  at  about  level  of  coxae  II-III; 
fovea  long,  weakly  sulciform,  positioned  more  or  less  midway  between  posterior 
margins  of  posterior  lateral  eyes.  Eyes:  with  black  surrounds  except  anterior  medians; 
posterior  medians  and  posterior  laterals  on  moderately  well  developed  tubercles; 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  42(4):  263-298  Issued  24  June  1 982 


264  F.  R.  WANLESS 

arranged  in  three  transverse  rows,  comprised  of  anterior  medians  (AM)  and  anterior 
laterals  (AL),  posterior  medians  (PM),  and  posterior  laterals  (PL);  anteriors  more  or 
less  contiguous  with  apices  procurved  in  frontal  view  and  recurved  in  dorsal  view; 
anterior  medians  largest;  anterior  laterals  greater  than  half  diameter  of  anterior  medians; 
posterior  medians  relatively  large,  positioned  closer  to  and  on  or  slightly  outside  optical  axis 
of  anterior  lateral  eyes;  posterior  laterals  as  large  or  slightly  smaller  than  anterior  laterals  and 
set  inside  lateral  margins  of  carapace  when  viewed  from  above;  quadrangle  formed  by 
posterior  median  and  posterior  lateral  eyes  broader  than  long  and  wider  behind;  entire 
quadrangle,  measured  from  between  bases  of  anterior  medians  to  posterior  margins  of 
posterior  laterals,  occupying  between  47  and  60  per  cent  of  carapace  length.  Clypeus: 
between  18  and  42  per  cent  of  diameter  of  anterior  median  eyes.  Chelicerae:  robust,  slightly 
diverging  and  inclined  anteriorly  in  females;  in  males  usually  elongate,  diverging  and  porrect 
with  median  horn  of  variable  length  which  arises  from  between  the  lower  margins  of  the 
cheliceral  bases  (Fig.  2 1 B);  pro-  and  retromargins  usually  with  three  teeth.  Maxillae:  long 
and  diverging  with  rounded  apices.  Labium:  oblong,  generally  less  than  half  maxillae 
length.  Sternum:  more  or  less  as  in  Figs  3B,  14G;  slightly  elongate  scutiform. 
Coxae:  I  and  II  generally  larger  than  HI-IV.  Pedicel:  short.  Abdomen:  long  slender 
and  tapering;  spinnerets  moderately  long,  posteriors  longest  with  long  apical  articles, 
anteriors  robust,  slightly  longer  than  more  slender  medians;  spiracle  a  transverse  slit 
just  in  front  of  anterior  spinnerets;  tracheal  system  not  examined;  position  of  colulus 
indicated  by  scant  group  of  hairs;  anal  tubercle  cone-shaped.  Legs:  long  and  slender;  spines 
strong  and  numerous;  claws  pectinate;  tufts  present;  scopulae  absent.  Female  palps:  long  and 
slender  with  apical  claw.  Male  palps:  moderately  complex,  intra-  and  generally  inter- 
specifically  distinct.  Femora  long  and  bowed;  patellae  long;  tibiae  long  with  broad  retro- 
lateral  or  ventral  apophysis;  cymbium  with  hair  tuft  protecting  embolic  tip  (Fig.  2H),  distal 
finger-like  extension,  swelling  on  retrolateral  basal  margin  and  usually  a  group  of  peg-like 
spines  on  dorsal  basal  margin  (Figs  1  A;  ISA);  embolus  (e)  very  long  and  slender,  sometimes 
thread-like,  resting  distally  in  well  developed  membraneous  fan-like  functional  conductoi  r 
(c),  the  tip  of  which  ends  in  a  sclerotized  spur;  a  bifid  median  apophysis  arises  from  a  trans- 
lucent pleated  membrane  on  the  tegulum  (Figs  1  B;  4H);  tegulum  (t)  irregular  in  form  with 
seminal  ducts;  median  hematodocha  in  form  of  a  membraneous  sac  between  tegulum  and 
subtegulum  only  evident  in  fully  expanded  palps;  subtegulum  (st)  a  sclerotized  ring  at  distal 
end  of  basal  hematodocha  (bh).  Epigynes:  of  various  forms;  median  septum  sometimes 
present  and  occasionally  bearing  lateral  pouches;  introductory  ducts  long,  simply  looped  or 
convoluted,  of  variable  width,  sometimes  narrow  (Fig.  8G,  H),  or  sac-like  (Figs  4F,  G; 
6F)  or  resembling  spermathecae  (Fig.  141,  J);  spermathecae  of  various  shapes  with  leaf-like 
fertilization  ducts;  spherical  objects  frequently  present  in  interstitial  spaces  (Fig.  2 1C-F). 

AFFINITIES.  The  general  habitus,  presence  of  a  median  cheliceral  horn,  functional  conductor 
and  median  apophysis  suggest  that  Cocalodes  and  Allococalodes  are  closely  related,  even 
sister  groups.  Both  genera  are  also  probably  allied  to  Holcolaetis  Simon,  from  Africa  and 
Sonoita  Peckham  &  Peckham  from  South  Africa,  as  the  male  papal  organs  evidently  possess 
an  homologous  conductor  and  median  apophysis. 

DIAGNOSIS.  Males  of  Cocalodes  are  readily  distinguished  from  other  Oriental  salticids  by  the 
distinctive  palpal  organs.  Females  are  separated  with  more  difficulty  by  the  geographical 
distribution,  relatively  large  posterior  median  eyes,  by  the  structure  of  the  epigynes  and  by 
having  three  teeth  on  the  posterior  margin  of  the  chelicerae.  A  more  useful  diagnosis  or 
assessment  of  affinities  cannot  be  given  until  supposedly  related  genera  have  been  revised. 

REMARKS.  1.  The  sclerite  from  which  the  median  horn  arises  in  Cocalodes  and 
Allococalodes  has  not  been  routinely  examined  in  previous  revisions  and  not  even  in 
all  females  of  Cocalodes  in  the  present  study,  as  the  chelicerae  have  to  be  displaced  or 
detatched  from  the  carapace  to  expose  it.  A  cursory  examination  of  several  Salticidae 
suggests  that  the  sclerite  is  present  in  all  members  of  the  family.  It  is  usually  seen  as 


SPIDER  GENUS  COCALODES 


265 


ma 


B 


Fig.  1  Cocalodes  papuanus  Simon.  Expanded  d1  palp:  A,  retrolateral;  B,  prolateral. 
Abbreviations:  bh,  basal  hematodocha;  c,  functional  conductor;  e,  embolus;  ma,  median 
apophysis;  p,  peg-like  cymbial  spines;  st,  subtegulum;  t,  tegulum. 


266  F.  R.  WANLESS 

an  indistinct  elongate  plate  embedded  in  tissue  between  the  bases  of  the  lower 
margins  of  the  chelicerae.  In  females  of  Cocalodes  thoracicus  Szombathy  and  in 
males  and  females  of  Holcolaetis  it  forms  a  peg-like  horn  similar  to  that  of  C. 
papuanus  Simon  (Fig.  2 IB).  This  does  not,  however,  necessarily  support  a  relation- 
ship between  Holcolaetis  and  Cocalodes  as  our  knowledge  of  the  development  of 
this  sclerite  is  inadequate. 

2.  The  spherical  objects  found  clustered  or  irregularly  distributed  in  the  interstitial 
spaces  of  the  epigynes  of  most  female  Cocalodes  vary  in  diameter  from  8  to  20  //. 
Sometimes  they  can  be  seen  through  the  cuticle  of  intact  epigynes  (Fig.  14H),  but  they 
are  most  readily  observed  in  epigynes  which  have  been  cleared  in  lactic  acid  (Figs 
20F;  21C-F).  When  examined  by  transmitted  light  microscopy  most  spheres  appear 
to  have  a  refractile  centre,  but  with  interferance  microscopy,  the  centres  are  trans- 
formed into  surface  depressions  which  resemble  a  lunar  crater  with  indistinct  lines 
radiating  outwards  from  the  raised  margins  t(Fig.  2 IF).  Some  spheres  appear  to  have 
collapsed  and  look  ragged  with  irregular  and  illdefmed  craters. 

These  objects,  which  are  not  known  to  occur  in  other  Salticidae,  resemble  the 
unidentified  spheres  described  by  Forster  (1980)  from  the  epigyne  of  a  Gamaso- 
morpha  species  (Family  Oonopidae).  A  concensus  of  opinion  (Forster,  1980)  favoured  a 
sporozoan  infection,  but  this  seems  unlikely  in  the  present  case  as  the  spheres  lack  the 
rod-like  structures  illustrated  by  Forster,  furthermore  their  presence  in  almost  every  female 
of  a  wide  range  of  species  suggests  they  are  a  natural  feature  of  these  epigynes,  which  could  be 
associated  with  spermatogenisis,  oviposition  or  even  plugging.  Clearly,  they  require  further 
investigation. 

Lists  of  species  in  the  genus  Cocalodes  Pocock,  1897 

Cocalodes  cygnatus  sp.  n. 

C.  expers  sp.  n. 

C.  innotabilis  sp.  n. 

C.  leptopus  Pocock,  1897 

C.  longicornis  sp.  n. 

C.  longipes  (Thorell,  1881) 

C.  macellus  (Thorell,  1878) 

C.  papuanus  Simon,  1900 

C.  platnicki  sp.  n. 

C.  protervus  (Thorell,  1881) 

C.  turgidus  sp.  n. 

C.  thoracicus  Szombathy,  1915 

The  species  of  Cocalodes  could  on  the  basis  of  the  development  of  the  epigynal 
septum  be  divided  into  two  groups.  However,  these  groups  have  not  been  proposed  as 
there  does  not  appear  to  be  any  correlation  with  characters  of  the  male  palpal  organs. 
This  may  be  a  natural  phenomenon  or  the  result  of  incorrectly  matching  males  and 
females.  Matching  the  sexes  has  been  difficult  in  the  present  study  and  where  doubts 
have  existed,  these  are  mentioned  in  the  species  descriptions.  As  is  often  the  case, 
additional  material  should  resolve  many  of  these  problems. 

Key  to  species  of  Cocalodes 

Males  (the  males  of  expers,  protervus  and  turgidus  are  unknown) 

1  Dorsal  prong  of  median  apophysis  broad  and  distally  truncate  (Fig.  2C,  F);  chelicerae  with 

dorsal  cluster  of  stout  bristles  (Fig.  2  A) papuanus  Simon  (p.  267) 

-      Dorsal  prong  of  median  apophysis  otherwise;  chelicerae  lacking  stout  bristles 

2  Peg-like  cymbial  spines  present  (Figs  4C;  ISA) 3 


SPIDER  GENUS  COCALODES  267 

Peg-like  cymbial  spines  absent 8 

3  Conductor  tip  cygniform  (Fig.  6G) cygnatussp.  n.  (p.  273) 

-  Conductor  tip  otherwise 4 

4  Dorsal  and  ventral  prongs  of  median  apophysis  more  or  less  equal  in  length  (Fig. 

IOC,  D) thoracicus  Szombathy  (p.  280) 

-  Dorsal  prong  of  median  apophysis  much  longer  than  ventral  prong           ....  5 

5  Dorsal  prong  of  median  apophysis  finger-like  or  scimitar-shaped  (Figs  1 3C,  E;  1 5D,  E)  6 
Dorsal  prong  of  median  apophysis  acuminate  (Figs  4C;  7H,  I) 7 

6  Conductor  tip  forming  a  triangular  plate  (Fig.  13G);  dorsal  prong  of  median  apophysis 

scimitar-shaped  (Fig.  13 E) longipes  (Thorell)  (p.  284) 

Conductor  tip  otherwise  (Fig.  1 5F);  dorsal  prong  of  median  apophysis  finger-like 

(Fig.  15D,  E) innotabilis  sp.  n.  (p.  288) 

7  Conductor  tip  slender  with  minute  barb  (Fig.  7G,  I)      ....      macellus  (Thorell)  (p.  275) 
Conductor  tip  robust,  barb  lacking  (Fig.  4H) leptopus  Pocock  (p.  270) 

8  Dorsal  prong  of  median  apophysis  relatively  slender  (Fig.   14D,  E);  cheliceral  horn 

not  protruding  beyond  clypeal  margin platnickisp.  n.  (p.  286) 

-  Dorsal  prong  of  median  apophysis  relatively  robust  (Fig.  8B,  C);  cheliceral  horn 

protruding  well  beyond  clypeal  margin  (Fig.  8A,  D)  .       .       .       .        longicornis  sp.  n.  (p.  277) 

Females  (the  female  of  innotabilis  is  unknown) 

1  Epigyne  with  median  septum  

Epigyne  lacking  median  septum 7 

2  Epigyne  with  relatively  large  openings  (Fig.  3D) papuanus  Simon  (p.  267) 

-  Epigyne  with  relatively  small  openings  3 

3  Epigynal  septum  narrow  and  not  extending  to  posterior  margin  (Fig.  7C)         ...  4 

-  Epigynal  septum  broad,  extending  to  posterior  margin 6 

4  Clypeus  white  haired;  epigynal  openings  more  or  less  ovoid  in  outline  (Figs  9B,  7C)  5 
Clypeus  not  white  haired;  epigynal  openings  indistinct  (Fig.  8F) .       .        longicornis  sp.  n.  (p.  277) 

5  Spermathecae  large  and  dark  (Fig.  9 B) protervus  (Thorell)  (p.  279) 

Spermathecae   small   and   pale,   hardly   if  at  all   visible  through   integument  (Fig. 

7C,  D) macellus  (Thorell)  (p.  275) 

6  Epigynal  septum  with  median  bulge  (Fig.  12 B) turgidus  sp.  n.  (p.  283) 

Epigynal  septum  lacking  median  bulge  (Fig.  1 1  B,  C)     .       .        .      thoracicus  Szombathy  (p.  280) 

7  Epigyne  with  small  median  openings  and  broad  posterior  ledge  (Fig.  1 4H)    platnicki  sp.  n.  (p.  286) 

-  Epigyne  otherwise 8 

8  Epigyne  slightly  depressed  with  comma-shaped  introductory  ducts  (Fig.  1 3B)  . 

longipes  (Thorell)  (p.  284) 
Epigyne  otherwise 9 

9  Epigyne  with  deep  median  notch  on  posterior  margin  (Fig.  5C)  .       .       .         expers  sp.  n.  (p.  27 1 ) 
Epigyne  otherwise 10 

10  Epigyne  with  two  notches  on  posterior  margin  (Fig.  6 D)       ....     cygnatussp.  n.  (p.  273) 
Epigyne  without  notches  on  posterior  margin  (Fig.  4E)         .       .       .       .  leptopus  Pocock  (p.  270) 


Cocalodes  papuanus  Simon 
(Figs2A-H;3A-E;  18C,E;  19A,B;21B) 

Cocalodes  papuanus  Simon,  1900  :  32,  rf.  LECTOTYPE  rf  (here  designated)  Irian  Jaya  (MNHN,  Paris) 

[examined].    Simon,    1901:403^04.    Roewer,    1954:936.    Bonnet,    1956:1173.    Proszynski, 

1971  :390. 
Cocalodes    armatissimus    Strand,    1913:122,    d.    LECTOTYPE    <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. 


References 

Alexander,  R.  McN.  1966.  The  function  and  mechanism  of  the  protrusile  upper  jaw  in  two  species  of 

cyprinid  fish.  J.  Zool.,  Land.  149  :  288-296. 
Ballantijn,  C.  M.  1969.  Functional  anatomy  and  movement  co-ordination  of  the  respiratory  pump  of 

the  carp  (Cyprinus  carpio  L.).  J.  exp.  Biol.  50  :  547-567. 
Banister,  K.  E.  &  Bunni,  M.  K.  1980.  A  new  blind  cyprinid  fish  from  Iraq.  Bull.  Br.  Mus.  nat.  Hist. 

(Zool.)  38  (3):  151-158. 


JAW  ANATOMY  OF  SEMIPLOTINE  CARPS  335 

&  Clarke,  M.  A.  1977.  The  freshwater  fishes  of  the  Arabian  Peninsula,  In:  The  scientific  results  of 

the  Oman  flora  and  fauna  survey  1975.  Jl.  Oman  Stud.  1977:  111-154. 
Berg,  L.  S.  1949.  Freshwater  fishes  of  Iran  and  of  neighbouring  countries.  Trudy  zool.  Inst.  Leningr., 

8  (4) :  783-858. 
Bremer,  K.  &  Wanntorp,   H.  G.    1979.   Hierarchy  and  reticulation  in  systematics.  Syst.   Zool. 

28  (4) :  624-627. 
Day,  F.  1980.  On  the  fishes  of  Afghanistan.  Proc.  zool.  Soc.  Lond.  :  224-232. 

1889.  The  fauna  of  British  India,  including  Ceylon  and  Burma.  Fishes  I  London  548  pp. 

Fowler,  H.  W.  1958.  Some  new  taxonomic  names  of  fishlike  vertebrates.  Notul.  nat.  310  :  1-16. 
Hora,  S.  L.  1937.  Notes  on  fishes  in  the  Indian  Museum.  XXIX.  On  a  collection  offish  from  Nepal. 

Rec.  Ind.  Mm.  39  (1) :  43-46. 

1942.  Notes  on  fishes  in  the  Indian  Museum.  XLII.  On  the  systematic  position  of  the  Indian 

species  ofScaphiodon  Heckel.  Rec.  Ind.  Mus.  54  (1) :  1-10. 

&  Gupta,  J.  C.  1940.  On  a  collection  offish  from  Kalimpong  Duars  and  Siliguri  Terai,  Northern 


Bengal.  J.  roy.  Asiat.  Soc.  Bengal 6  (2) :  77-83. 
Howes,  G.  J.    1976.  The  cranial  musculature  and  taxonomy  of  characoid  fishes  of  the  tribes 

Cynodontini  and  Characini.  Bull.  Br.  Mm.  nat.  Hist.  (Zool.)  29  (4) :  201-248. 
1978.  The  anatomy  and  relationships  of  the  cyprinid  fish  Luciobrama  macrocephalus  (Lacepede). 

Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  34  (1) :  1-64. 
1979.  Notes  on  the  anatomy  of  the  cyprinid  fish  Macrochirichthys  macrochirus  with  a  review  of 

the  subfamily  Cultrinae.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  36  (3) :  147-200. 
1980.  The  anatomy,  phylogeny  and  classification  of  bariliine  cyprinid  fishes.  Bull.  Br.  Mus.  nat. 

Hist.  (Zool.)  37  (3):  129-198. 

1981.  Anatomy  and  phylogeny  of  the  Chinese  major  carps  Ctenopharyngodon  Steind.,  1866  and 


Hypophthalmichthys  Blkr,  1860.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  41  (1) :  1-52. 
Karaman,  M.  S.  1969.  Susswasserfische  der  Tiirkei  7.  Revision  der  kleinasiatischen  und  vorder- 

asiatischen  Arten  des  Genus  Capoeta  (Varicorhinus,  partim).  Mitt.  Hamburg  Zool.  Mus.  Inst. 

66:  17-54. 
I. iem,  K.  F.  1980.  Adaptive  significance  of  intra-  and  interspecific  differences  in  the  feeding  repertoires 

ofcichlid  fishes.  Amer.  Zool.  20  :  295-314. 
Matthes,  H.  1963.  A  comparative  study  of  the  feeding  mechanism  of  some  African  Cyprinidae  (Pisces, 

Cypriniformes).  Bijdr.  Dierk.  33  :  3-35. 
Mirza,  M.  R.  1969.  Fishes  of  the  genus  Cyprinion  Heckel  (Cyprinidae,  Osteichthyes)  from  West 

Pakistan.  Pakistan  J.  Zool.  1  (2) :  141-1 50. 
Regan,  C.  T.  1911.  The  classification  of  the  teleostean  fishes  of  the  order  Ostariophysi.  1 .  Cyprinoidea. 

Ann.  Mag.  nat.  Hist.  (8)  8  :  1 3-32. 
Smith,  H.  M.  1931.  Description  of  new  genera  and  species  of  Siamese  fishes.  Proc.  U.S.  natn  Mus.  79 

art  7:  1-48. 

1945.  The  fresh-water  fishes  of  Siam  or  Thailand.  Bull.  U.S.  natn  Mus.  188  :  1-622. 

Takahasi,  N.  1925.  On  the  homology  of  the  cranial  muscles  of  the  cypriniform  fishes.  J.  Morph. 

40:  1-109. 
Taki,  Y.  1974.  New  species  of  the  genus  Scaphognathops.  Cyprinidae,  from  the  Lao  Mekong  river 

system.  Jap.  J.  Ichthyol.  21  (3) :  129-136. 
Cyprinid  fishes  of  the  genera  Onychostoma  and  Scaphiodonichthys  from  upper  Laos,  with 

remarks  on  the  dispersal  of  the  genera  and  their  allies.  Jap.  J.  Ichthyol.  22  (3) :  143-1 50. 
Tarsitano,  S.  &  Hecht,  M.  K.  1980.  A  reconsideration  of  the  reptilian  relationships  of  Archaeopteryx. 

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 

Publications  Sales 
British  Museum  (Natural  History) 

Cromwell  Road 

London  SW7  5BD 

England 


Titles  to  be  published  in  Volume  42 


The  tick  collection  (Acarina:  Ixodoidea)  of 

the  Hon.  Nathaniel  Charles  Rothschild  deposited  in  the  Nuttall  and 

general  collections  of  the  British  Museum  (Natural  History). 

By  James  E.  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 


BOUND