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26NOV1987 

PRESENTED 
GFNEH/U  LIB3AFV 


Bulletin  of  the 

British  Museum  (Natural  History) 


Zoology  series        Vol  52        1987 


British  Museum  (Natural  History) 
London  1987 


Dates  of  publication  of  the  parts 

Nol    .        .        . 29  January  1987 

No  2 •        •  26  February  1987 

No  3 .        .  26  March  1987 

No  4 30  April  1987 

No  5  .        .        . 28  May  1987 

No  6 25  June  1987 

No  7 30  July  1987 

No  8  27  August  1987 


ISSN  0007-1 498 


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


Contents 
Zoology  Volume  52 


No  1       Miscellanea 


A  revision  of  the  genus  Pseudovorticella  Foissner  &  Schiffmann,  1974 
(Ciliophora:  Peritrichida).  By  A.  Warren 1 

The  taxonomic  status  of  the  genera  Pontigulasia,  Lagenodifflugia  and 
Zivkovicia  (Rhizopoda:  Difflugiidae).  By  C.  G.  Ogden        .         .         .         13 

A  revision  of  the  foraminiferal  genus  Adercotryma  Loeblich  & 
Tappan,  with  a  description  of  A.  wrighti  sp.  nov.  from  British  waters. 
By  P.  Bronnimann  &  J.  E.  Whittaker 21 

Hermit  crabs  associated  with  the  bryozoan  Hippoporidra  in  British 

waters.  By  J.  D.  D.  Bishop 29 

The  first  zoea  of  three  Pachygrapsus  species  and  of  Cataleptodius 
floridanus  (Gibbes)  from  Bermuda  and  Mediterranean  (Crustacea: 
Decapoda:  Brachyura).  By  R.  W.  Ingle 31 

A  classification  of  the  phylum  Sipuncula.  By  P.  E.  Gibbs  &  E.  B. 

Cutler 43 

Two  new  species  ofGarra  (Teleostei:  Cyprinidae)  from  the  Arabian 
peninsula.  By  K.  E.  Banister 59 

No  2      A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The 

Paracineta  and  Corynophora  problem.  By  Colin  R.  Curds  .         .         .        71 

No  3       Notes  on  spiders  of  the  family  Salticidae  1 .  The  genera  Spartaeus, 

MintoniaandTaraxella.RyF.R.'Wan\ess 107 

No  4       Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata: 

Parasitidae)  in  the  British  Isles.  By  K.  H.  Hyatt 139 

No  5      The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus 

Aulopyge  Heckel,  1841,  with  an  appraisal  of  the  genus  Barbus  Cuvier 

&  Cloquet,  1816  and  the  subfamily  Cyprininae.  By  Gordon  J.  Howes .       1 65 

No  6       Revision    of   the    genera    Acineria,    Trimyema,    and    Trochiliopsis 

(Protozoa,  Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam      .       197 

No  7      The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae) 
with  a  systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and 
the  descriptions  of  a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L. 
Harrison .       225 

No  8       Notes     on     some     species     of    the     genus     Amathia     (Bryozoa, 

Ctenostomata).  By  P.  J.  Chimonides 307 


Bulletin  of  the 

British  Museum  (Natural  History) 


Miscellanea 


Zoology  series    Vol52    No  1     29  January  1987 


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-crowing  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),  1986 


The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 

Keeper  of  Zoology     :     Mr  J.  F.  Peake 
Editor  of  Bulletin       :     Dr  C.  R.  Curds 
Assistant  Editor  Mr  C.  G.  Ogden 


ISBN  0565  05025  7 

ISSN  0007- 1498  Zo?l°,glTSeT      ,   ™ 

Vol  52  No.  1  pp  1-70 

British  Museum  (Natural  History) 

Cromwell  Road  , 

London  SW7  5BD  Issued  29  January  1987 


Miscellanea 


Contents 


A  revision  of  the  genus  Pseudovorticella  Foissner  &  Schiffmann,   1974  (Ciliophora: 
Peritrichida).  By  A.  Warren 

The   taxonomic   status   of  the   genera   Pontigulasia,   Lagenodifflugia   and   Zivkovicia 
(Rhizopoda:  Difflugiidae).  By  C.  G.  Ogden 

A  revision  of  the  foraminiferal  genus  Adercotryma  Loeblich  &  Tappan,  with  a  description 
of  A.  wrighti  sp.  nov.  from  British  waters.  By  P.  Bronnimann  &  J.  E.  Whittaker     . 

Hermit  crabs  associated  with  the  bryozoan  Hippoporidra  in  British  waters.  By  J.  D.  D. 
Bishop 

The  first  zoea  of  three  Pachygrapsus  species  and  of  Cataleptodius  floridanus  (Gibbes)  from 
Bermuda  and  Mediterranean  (Crustacea:  Decapoda:  Brachyura).  By  R.  W.  Ingle. 

A  classification  of  the  phylum  Sipuncula.  By  P.  E.  Gibbs  &  E.  B.  Cutler    .... 


Page 
1 

13 
21 
29 

31 

43 


Two  new  species  of  Garra  (Teleostei:  Cyprinidae)  from  the  Arabian  peninsula.  By 
K.  E.  Banister 


59 


A  revision  of  the  genus  Pseudovorticella  Foissner  & 
Schiffmann,  1974  (Ciliophora:  Peritrichida) 

A.  Warren 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road, 


London  SW7  5BD 


Introduction 

The  genus  Pseudovorticella  was  erected  by  Foissner  &  Schiffmann  ( 1 974)  to  include  those  peritrichs 
which  are  morphologically  similar  to  Vorticella  but  which  have  a  reticulate  silver  line  system 
with  lines  running  vertically  as  well  as  horizontally.  The  reticulate  pattern  of  silver  lines  under- 
lies a  system  of  pellicular  tubercles  which  covers  the  entire  zooid  surface  except  the  disc  and 
infundibulum. 

Pellicular  tubercles  have  been  studied  by  several  workers  over  the  past  century.  Schroder  (1906) 
showed  that  the  tubercles  of  Pseudovorticella  monilata  are  surface  features,  the  distribution 
of  which  corresponds  to  that  of  the  underlying  striations.  Ultrastructural  studies  by  TEM 
(Kawamura,  1 973)  and  SEM  (Carey  &  Warren,  1 983)  have  confirmed  this  observation.  Kawamura 
(1973)  also  showed  that  each  tubercle  of  P.  monilata  is  a  semisphere,  about  2-0  um  in  diameter,  and 
contains  a  sphere  of  electron  dense  material.  Further  investigations  using  histochemical  staining 
(Faure-Fremiet&Thaureaux,  1 944;  Pratt  &  Rosen,  1983)  and  microanalysis  (Pratt  &  Rosen,  1983) 
indicate  that  the  tubercles  contain  paraglycogen.  The  function  of  the  tubercles  is  not  known 
although  it  has  been  suggested  that  they  may  aid  predator  avoidance  (Spoon,  1975). 

Foissner  &  Schiffmann  (1974)  noted  that  the  silver  line  system  is  particularly  useful  for  species 
diagnosis  in  Pseudovorticella,  and  biometric  analyses  have  been  carried  out  on  several  species 
(Foissner  &  Schiffmann,  1974  &  1975;  Foissner,  1979).  Parameters  which  are  of  particular  taxo- 
nomic  value  include  the  total  number  of  silver  lines  per  zooid  and  dimensions  of  the  grids  formed 
by  the  intersecting  vertical  and  horizontal  lines.  Morphological  features  traditionally  used  in 
vorticellid  taxonomy  are  also  useful  diagnostic  characters  for  the  species  of  Pseudovorticella;  these 
include  the  size  and  shape  of  the  zooid,  the  number  and  position(s)  of  the  contractile  vacuole(s)  and 
the  shape  and  position  of  the  macronucleus  (Noland  &  Finley,  1931;  Foissner,  1979;  Warren, 
1986). 

Sixteen  species  of  Pseudovorticella  are  recognised,  twelve  of  which  originally  belonged  to  the 
genus  Vorticella.  A  key  to  their  identification  is  provided. 


Systematics 

In  the  scheme  adopted  by  the  Committee  on  Systematics  and  Evolution  of  the  Society  of  Proto- 
zoologists  (Levine  et  al.,  1980),  the  taxonomic  position  of  the  genus  Pseudovorticella  was  given  as 
follows: 

Phylum:  Ciliophora  Doflein,  1 90 1 

Class:  Oligohymenophora  de  Puytorac  et  al.,  1974 

Subclass:  Peri  trichia  Stein,  1859 

Order:  Peritrichida  Stein,  1859 

Suborder:  Sessilina  Kahl,  1933 

Family:  Vorticellidae  Ehrenberg,  1838 

Genus:  Pseudovorticella  Foissner  &  Schiffmann,  1974 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  52(1):  1-12  Issued  29  January  1987 


2  A.  WARREN 

Diagnosis 

Solitary  bell-shaped  zooids  borne  upon  a  spirally  contractile  stalk.  In  all  respects  save  one,  the 
body  and  stalk  of  Pseudovorticella  resemble  those  of  Vorticella  from  which  it  cannot  be  differen- 
tiated until  impregnated  with  silver,  which  reveals  a  reticulate  silver  line  pattern  quite  unlike 
that  of  Vorticella  (see  Warren,  1986).  In  addition  to  Vorticella  this  genus  could  be  mistaken  for 
Haplocaulis  in  which  the  stalk  contracts  in  a  zigzag  rather  than  a  helical  manner. 


Key  to  the  species  of  Pseudovorticella 

1  With  endosymbiotic  zoochlorellae 2 

Without  endosymbiotic  zoochlorellae 3 

2  Zooid  about  40  um  long;  macronucleus  C-shaped      ...          P.  zooanthelligera  (Fig.  9b) 
Zooid  75-95  um  long;  macronucleus  J-shaped P.  chlorelligera  (Fig.  la) 

3  Diameter  of  peristomial  lip  less  than  or  equal  to  maximum  body  width 4 

Diameter  of  peristomial  lip  greater  than  maximum  body  width 10 

4  Diameter  of  peristomial  lip  less  than  maximum  body  width 5 

Diameter  of  peristomial  lip  equal  to  maximum  body  width 6 

5  Macronucleus  J-shaped P.  difficilis  (Fig.  1  c  &  d) 

Macronucleus  C-shaped P.  papillata  (Fig.  5c) 

6  Body  length  less  than  x  2  maximum  body  width 7 

Body  length  at  least  x  2  maximum  body  width P.  micata  (Fig.  2b) 

7  One  contractile  vacuole 8 

Two  contractile  vacuoles P.  sphagni  (Fig.  7b) 

8  Macronucleus  lies  vertical  with  respect  to  major  axis  of  zooid 9 

Macronucleus  lies  horizontal  with  respect  to  major  axis  of  zooid         .        .      P.  stilleri  (Fig.  9a) 

9  Zooid  65-80  urn  long  and  with  44-54  transverse  striations .        .        .        .P.  quadrat  a  (Fig.  7a) 
Zooid  35-45  urn  long  and  with  20-33  transverse  striations .         .         .       P.  sauwaldensis  (Fig.  8) 

10  Zooid  with  two  contractile  vacuoles 11 

Zooid  with  one  contractile  vacuole 13 

1 1  Zooid  with  centrally  located  constriction;  scopular  region  rounded     .      P.  margaritata  (Fig.  2a) 
Zooid  without  centrally  located  constriction;  scopular  region  tapers  towards  stalk      .        .        .12 

12  Zooid  40-45  urn  long;  stalk  length  x  16- 18  zooid  length    ....       P.  mollis  (Fig.  3a) 
Zooid  60-70  um  long;  stalk  x  3  zooid  length P.  monilata  (Fig.  3b,  c  &  d) 

13  Diameter  of  peristomial  lip  less  than  body  length 14 

Diameter  of  peristomial  lip  greater  than  body  length P.  punctata  (Fig.  6) 

14  Zooid  less  than  x  3  maximum  body  width 15 

Zooid  length  greater  than  x  3  maximum  body  width ....      P.  mutans  (Fig.  4  c  &  d) 

1 5  Zooid  50-70  urn  long  x  22^48  um  wide;  typically  marine    .         .         .P.  nebulifera  (Fig.  4  a  &  b) 
Zooid  32-50  um  long  x  20  um  wide;  typically  freshwater    .       P.  pseudocampanula  (Fig.  5  a  &  b) 


Description  of  Species 

P.  chlorelligera  (Kahl,  1935)  Jankowski,  1976 

V.  margaritata  f.  chlorelligera  Kahl,  1935 

P.  margaritata  f.  chlorelligera  (Kahl,  1935)  Foissner  &  Schiffmann,  1975 

DIAGNOSIS  (Fig.  la  &  b).  Zooid  inverted  bell-shaped,  78-95  um  longx  50  um  wide;  peristomial  lip  80  um 
diameter;  infundibulum  reaches  half  body  length;  macronucleus  J-shaped;  numerous  endosymbiotic  zoo- 
chlorellae present  in  cytoplasm;  zooid  has  a  total  of  33-53  (mean  47-7)  transverse  striations;  grid  size 


GENUS  PSEUDOVORTICELLA 


Fig.  1.  (a)  P.  chlorelligera  zooid,  bar  —  50  um;  (b)  telotroch,  bar  =  25  um  (after  Foissner  &  Schiffmann, 
1975);  (c)  P.  difficilis  (after  Kahl,  1935);  (d)  P.  difficilis,  bar  =  50um  (after  Foissner  &  Schiffmann, 
1975;  called  P.  difficilis  var.  magnistriata). 


1-3-3-2  um  x  2-0-4-5  um;  zooid  surface  with  15-28  (mean  22)  pellicular  pores  per  100  um2;  telotroch  nearly 
cylindrical  in  shape  and  with  a  prominent  epistomial  membrane  (Fig.  lb). 

HABITAT.  Freshwater. 

P.  difficilis  (Kah\,  1933)  Jankowski,  1976 
V.  difficilis  Kahl,  1933 
P.  difficilis  var.  magnistriata  Foissner  &  Schiffmann,  1974 

DIAGNOSIS  (Fig.  Ic  &  d).  Zooid  60-140  um  long  x  40-70  um  wide;  diameter  of  peristomial  lip  less  than 
maximum  body  width;  infundibulum  reaches  half  body  length;  single  contractile  vacuole  situated  in  upper 
part  of  zooid  close  to  infundibulum;  macronucleus  J-shaped;  39-49  (mean  43-9)  transverse  striations  per 
zooid;  grid  size  3- 1-4-7  um  x  2-7-3-4  jam;  spasmoneme  with  numerous  thecoplasmic  granules. 

HABITAT.  Freshwater  or  marine. 

P.  margaritata  (Fromentel,  1874)  Jankowski,  1976 

DIAGNOSIS  (Fig.  2a).  Zooid  inverted  bell-shaped,  59-70  um  long  x  50  um  wide,  with  a  slight  constriction  in  the 
central  region  and  rounded  at  the  scopular  end;  peristomial  lip  70  um  in  diameter;  two  contractile  vacuoles 
situated  in  anterior  part  of  zooid;  macronucleus  C-shaped  and  situated  in  centre  of  zooid. 

HABITAT.  Freshwater,  particularly  eutrophic  lakes  and  stagnant  water. 
REMARKS.  This  species  has  been  redescribed  by  Kahl  (1935)  and  Stiller  (1971). 

P.  micata  (Kahl,  1933)  nov.  comb. 
V.micata  Kahl,  1933 

DIAGNOSIS  (Fig.  2b).  Zooid  elongate,  65  um  long  x  25  um  wide;  peristomial  lip  25  um  in  diameter;  disc  flat  and 
slightly  elevated  above  peristome;  infundibulum  reaches  one  third  zooid  length;  contractile  vacuole  situated 
in  upper  part  of  zooid  close  to  infundibulum. 

HABITAT.  Marine. 


A.  WARREN 


Fig.  2.     (a)  P.  margaritata  ,  bar  =  25  urn  (composite  from  Kahl,  1935  and  Stiller,  1971);  (b)  P.  micata, 

bar  =  25  um  (after  Kahl,  1935). 


P.  mollis  (Stokes,  1887)  nov.  comb. 
V.  mollis  Stokes,  1887 

DIAGNOSIS  (Fig.  3a).  Zooid  inverted  bell-shaped,  40-45  urn  long  x  25  um  wide;  peristomial  lip  40  um  in 
diameter;  infundibulum  reaches  one  third  body  length;  two  contractile  vacuoles  situated  in  anterior  part  of 
zooid;  stalk  x  16-1 8  zooid  length. 

HABITAT.  Freshwater 

REMARKS.  Although  this  species  was  not  drawn  by  Stokes  (1887),  it  has  been  observed  and  figured  by 
Nenninger(1948). 

P.  monilata  (Tatem,  1870)  Foissner  &  Schiffmann,  1974 

V.  lockwoodii  Stokes  1884 
V.  monilata  Tatem,  1870 

DIAGNOSIS  (Fig.  3b,  c  &  d).  Zooid  inverted  bell-shaped,  45-70  urn  long  x  40-^45  urn  wide;  peristomial  lip  50  um 
in  diameter;  infundibulum  reaches  half  body  length;  two  contractile  vacuoles  situated  in  anterior  part  of 
zooid;  macronucleus  J-shaped;  31-41  (mean  35-3)  transverse  striations  per  zooid;  grid  size  2-5-3-5  umx 
1 -5-2-5  um;  stalk  x3  body  length;  spasmoneme  with  thecoplasmic  granules;  telotroch  cone-shaped  with 
prominent  epistomial  membrane. 

HABITAT.  Freshwater,  often  forming  pseudocolonies;  Pratt  &  Rosen  (1983)  reported  large  numbers  of 
Pseudovorticella  (Vorticella)  monilata  attached  the  Cyanobacterium  Anabaenaflos-aquae. 

P.  mutatis  (Penard,  1922)  Foissner,  1979 
V.  mutans  Penard,  1922 

DIAGNOSIS  (Fig.  4c  &  d).  Zooid  inverted  bell-shaped,  65-95  um  long  x  1 8-25  um  wide;  peristomial  lip  25  um  in 
diameter;  disc  convex;  infundibulum  reaches  half  body  length;  contractile  vacuole  situated  in  upper  half  of 
body  close  to  infundibulum;  macronucleus  J-shaped;  zooid  has  40-^47  (mean  43)  transverse  striations;  grid 


GENUS  PSEUDOVORTICELLA 


Fig.  3.  (a)  P.  mollis,  bar  =  25  ^m  (after  Nenninger,  1948);  (b)  P.  monilata  showing  oral  ciliation  (detail 
from  Patsch,  1974);  (c)  zooid,  bar  =  25  urn;  (d)  telotroch  (after  Foissner,  1979).  G  =  germinal  kinety; 
H  =  haplokinety;  Pl ,  P2,  P3  =  1 , 2,  3,  peniculus;  PO  =  polykinety. 


Fig.  4.     (a)  P.  nebulifera  zooid,  bar  =  25  ^m  (after  Noland  &  Finley,  1 93 1 );  (b)  telotroch  (after  Barlow  & 
Finley,  1976b);  (c)P.  mutans  telotroch;  (d)  zooid,  bar  =  25  urn  (after  Foissner,  1979). 


6  A.  WARREN 

size  1  -4-1  -5  umx  1-5-2-2  um;  stalk  x5  body  length  and  8-0  urn  wide;  spasmoneme  with  thecoplasmic 
granules;  telotroch  with  prominent  epistomial  membrane. 

HABITAT.  Freshwater. 

P.  nebulifera  (Miiller,  1786)  Jankowski,  1976 
V.  nebulifera  Miiller,  1*786 

DIAGNOSIS  (Fig.  4a  &  b).  Zooid  inverted  bell-shaped,  38-78  urn  (mean  60  urn)  long  x  22-48  um  (mean  37  um) 
wide;  slightly  constricted  beneath  peristomial  lip  which  measures  32-66  um  (mean  53  um)  in  diameter; 
single  contractile  vacuole  situated  close  to  infundibulum;  macronucleus  J-shaped;  stalk  50-800  um  (mean 
1 50  um)  long  x  3-5-6-0  um  (mean  4-7  um)  wide;  spasmoneme  with  thecoplasmic  granules;  telotroch  47-75  um 
(mean  60  um)  long;  cyst  37  um  in  diameter. 

HABITAT.  Marine  or  freshwater. 

REMARKS.  Redescribed  by  Noland  &  Finley  (1931);  for  telotroch  and  SEM  studies,  see  Barlow  &  Finley 
(1976a&b). 

P.papillata  (Stiller)  Jankowski,  1976 
V.  microstoma  f.  monilata  Stiller  (see  Stiller,  1971) 

DIAGNOSIS  (Fig.  5c).  Zooid  35-80  um  (mean  55  jam)  long  x  22-50  um  (mean  35  um)  wide,  the  maximum 
body  width  being  the  mid  region  of  the  zooid;  peristomial  lip  12-25  um  (mean  23  urn)  in  diameter;  disc 
convex;  infundibulum  reaches  one  third  body  length;  contractile  vacuole  situated  in  anterior  part  of  zooid; 
macronucleus  C-shaped  and  lies  longitudinally  with  respect  to  major  axis  of  zooid. 

HABITAT.  Freshwater,  particularly  under  conditions  of  high  biochemical  oxygen  demand  (BOD5). 


Fig.  5.    (a)  P.  pseudocampanula  relaxed  zooid,  bar  =  25  um;  (b)  contracted  zooid  (after  Foissner,  1 979); 
(c)  P.papillata,  bar  =  25  um  (after  Stiller,  1971). 


GENUS  PSEUDOVORTICELLA 


Fig.  6.     P.  punctata,  (a)  bar  =  25  urn  (after  Dons,  1918);  (b)  after  Stiller  (1946)  (called  Vorticella 
subconica);  (c)  after  Kahl  (1935)  (called  Vorticella perlata). 


P.  pseudocampanula  Foissner,  1 979 

DIAGNOSIS  (Fig.  5a  &  b).  Zooid  conical/inverted  bell-shaped,  32-50  um  (mean  40  um)  long  x  20  urn  wide; 
peristomial  lip  35  um  in  diameter;  upon  contraction,  peristomial  lip  becomes  puckered  (Fig.  5b);  infundibu- 
lum  reaches  half  body  length;  contractile  vacuole  situated  close  to  infundibulum;  macronucleus  J-shaped; 
zooid  has  44-51  (mean  46-6)  transverse  striations;  grid  size  1-3-2-6  urn  x  1-5-3-0  urn;  stalk  x  7  body  length; 
thecoplasmic  granules  present  on  spasmoneme. 

HABITAT.  Freshwater. 

P.  punctata  (Dons  1918)  nov.  comb. 

V.  punctata  Dons,  1918 

V.  subconica  Stiller,  1946 

P.  subconica  (Stiller,  1946)  Jankowski,  1976 

DIAGNOSIS  (Fig.  6).  Zooid  conical  or  inverted  bell-shaped,  40-50  um  Iongx40um  wide;  peristomial  lip 
50-55  um  in  diameter;  disc  convex;  infundibulum  reaches  one  third  body  length;  contractile  vacuole  situated 
in  upper  part  of  zooid;  macronucleus  J-shaped;  stalk  x  4-5  body  length  and  4-0  um  wide. 

HABITAT.  Marine. 

P.  quadrata  Foissner,  1 979 

DIAGNOSIS  (Fig.  7a).  Zooid  65-80  um  (mean  70  um)  long  x  55  um  wide;  peristomial  lip  60  um  in  diameter; 
infundibulum  reaches  half  body  length;  contractile  vacuole  situated  in  anterior  part  of  zooid;  macronucleus 
J-shaped;  zooid  has  44-54  (mean  48-3)  transverse  striations;  grid  size  1-5-2-8  um  x  1-3  x  2-7  um;  stalk  x  7 
body  length  and  9-0  um  wide;  spasmoneme  with  thecoplasmic  granules. 

HABITAT.  Freshwater. 

P.  sauwaldensis  Foissner  &  Schiffmann,  1979 

DIAGNOSIS  (Fig.  8).  Zooid  shape  variable,  usually  inverted  bell-shaped  35^5  um  long  x  20  um  wide; 
peristomial  lip  20  um  in  diameter  and  3-0  um  thick;  disc  convex;  infundibulum  reaches  half  body  length; 


A.  WARREN 


a  b 

Fig.  7.     (a)  P.  quadrata,  bar  =  25  nm;  (b)  P.  sphagni,  bar  =  25  um  (after  Foissner,  1 979). 


Fig.  8.     P.  sauwaldensis  (a)  normal  zooid,  bar  =  20  um;  (b)  contracted  zooid;  (c)  showing  variability  of 
macronucleus  and  zooid  shape,  bar  =  20  urn  (after  Foissner  &  Schiffmann,  1979). 

contractile  vacuole  situated  in  upper  part  of  zooid  close  to  infundibulum;  macronucleus  vermiform,  variable 
in  shape  and  situated  longitudinally  with  respect  to  major  body  axis;  pellicle  has  20-33  (mean  29)  transverse 
striations;  grid  size  0-9-1-5  um  x  0-7-2-5  um;  stalk  x  1-3  body  length. 

HABITAT.  Freshwater. 


GENUS  PSEUDOVORTICELLA 


Fig.  9.     (a)  P.  stilleri,  bar  =  50  um  (after  Stiller,  1 963);  (b)  P.  zooanthelligera,  bar  =  25  urn  (after  Stiller, 

1968).  EZ  =  endosymbiotic  zoochlorellae. 


P.  sphagni  Foissner,  1979 

DIAGNOSIS  (Fig.  7b).  Zooid  inverted  bell-shaped,  40-50  um  long  x  30  um  wide;  peristomial  lip  30  um  in 
diameter;  infundibulum  reaches  one  third  body  length;  two  contractile  vacuoles  situated  in  anterior  part  of 
zooid;  macronucleus  J-shaped  with  elongate  distal  arm;  zooid  has  34-37  (mean  35-5)  transverse  striations; 
grid  size  1  -6-1  -9  um  x  2-5-2-7  um. 

HABITAT.  Freshwater,  originally  isolated  from  Sphagnum  bogs. 

P.  stilleri  n.  sp. 

V.  campanula  f.  monilata  Stiller,  1963 

DIAGNOSIS  (Fig.  9a).  Zooid  inverted  bell-shaped,  85  um  long  x  80  um  wide;  peristomial  lip  80  um  in  diameter; 
infundibulum  reaches  half  body  length;  macronucleus  C-shaped  and  lies  horizontally  across  centre  of  zooid. 

HABITAT.  Freshwater,  attached  to  the  duckweed  Lemna  minor. 

P.  zooanthelligera  (Stiller,  1968)  nov.  comb. 
V.  zooanthelligera  Stiller,  1968 

DIAGNOSIS  (Fig.  9b).  Zooid  inverted  bell-shaped,  40-42  um  long  x  40  jam  wide;  peristomial  lip  50  um 
in  diameter;  disc  flat;  infundibulum  reaches  one  third  body  length;  macronucleus  C-shaped  and  lies 
longitudinally  in  zooid;  cytoplasm  contains  numerous  endosymbiotic  zoochlorellae;  stalk  x  5  body  length. 

HABITAT.  Freshwater. 


Incertae  sedis 

Pseudovorticella  sp.  (Graham  & -Graham,  1978)  nov.  comb. 
Vorticella  sp.  Graham  &  Graham,  1978 

Graham  &  Graham  (1978)  made  an  ultrastructural  study  of  a  vorticellid  (Vorticella  sp.)  furnished  with 
pellicular  tubercles  and  containing  endosymbiotic  zoochlorellae.  The  presence  of  pellicular  tubercles  suggests 


10  A.  WARREN 

that  this  organism  should  belong  to  the  genus  Pseudovorticella.  However  other  important  diagnostic  features, 
for  example  the  macronucleus,  contractile  vacuole(s)  and  shape  of  the  relaxed  zooid,  were  not  recorded.  Only 
when  such  data  is  available  will  it  be  possible  to  determine  the  exact  status  of  this  organism. 


Acknowledgements 

I  would  like  to  thank  Dr  W.  Foissner  for  his  helpful  criticism  of  the  manuscript. 

References 

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Peu  Connues.  364  pp.  G.  Masson,  Paris. 
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Ostsee,  Lief.  23  (Tiel,  II,  c3);  Leipzig,  pp.  147-183. 
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A.  R.,  Lorn,  J.,  Lynn,  D.,  Merinfield,  E.  G.,  Page,  F.  C.,  Poljansky,  G.,  Sprague ,  V.,  Vavra,  J.  &  Wallace, 

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GENUS  PSEUDOVORTICELLA  1  1 

Pratt,  J.  R.  &  Rosen,  B.  H.  1983.  Association  of  species  of  Vorticella  (Peritrichida)  and  planktonic  algae. 
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Puytorac,  P.  de,  Batisse,  A.,  Bohatier,  J.,  Corliss,  J.  O.,  Deroux,  G.,  Didier,  P.,  Dragesco,  J.,  Fryd-Versavel, 
G.,  Grain,  J.,  Groliere,  C.  A.,  Hovasse,  R.,  Itfode,  F.,  Laval,  M.,  Roque,  M.,  Savoie,  A.  &  Tuffrau,  M.  1974. 
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Clermont-Ferrand).  Compte  Rendu  Hebdomadaire  des  Seances  de  I'Academie  des  Sciences.  Paris  278: 
2799-2802. 

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395^10. 

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hrsg.  von  der  Senckenbergischen  Naturforschenden  Gesellschaft  41:  285-313. 

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bearbeitet  I.  206  pp.  Leipzig. 

Stiller,  J.  1940.  Beitrage  zur  Peritrichenfauna  des  Groben  Ploner  Sees  in  Holstein.  Archiv  fur  Hydrobiologie 
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fauna. Internationale  Revue  der  Gesamten  Hydrobiologie  und  Hydrographie  48:  603-612. 

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Academiae  Scientiarum  Hungaricae  14:  185-21 1. 

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Stokes,  A.  C.  1883.  A  new  vorticellid.  American  Monthly  Microscopical  Journal  4:  208. 

1884.  A  new  infusorien  belonging  to  the  genus  Vorticella.  American  Naturalist  18:  829-830. 

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244-255. 
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194-195. 
Warren,  A.  1986.  A  revision  of  the  genus  Vorticella  (Ciliophora:  Peritrichida).  Bulletin  of  the  British  Museum 

(Natural History).  Zoology  Series  50(1):  1-57. 

Manuscript  accepted  for  publication  5  January  1986 


Index 

Index  of  extant  species;  annotated  list  of  nominal  species. 

P.  anabaenae  (Stiller,  1940)  Jankowski,  1976  was  transferred  to  the  genus  Haptocaulis  by  Stiller  (1971). 

P.  Morelligera(Kah\,  1935)  Jankowski,  1976 2 

P.chlamydophora(PemLTd,  1922)  Jankowski,  1976=  Vorticella  vestita  Stokes,  1 883  (see  Warren,  1986). 

P.  difficilis  (Kahl,  1933)  Jankowski,  1976 3 

P.  difficilis  var.  magnistriata  Foissner  &  Schiffmann,  1974  =  P.  difficilis. 

P.  lima  (Kahl,  1935)  Jankowski,  1976.  This  species  appears  to  have  pellicular  granules  rather  than 

tubercles;  it  should  therefore  remain  in  the  genus  Vorticella  (V.  lima)  until  it  has  been  redescribed. 

P.  margaritata  (Fromenlel,  1874)  Jankowski,  1976 3 

P.  micata  (Kahl,  1933)  nov.  comb 3 

P.  mollis  (Stokes,  1887)  nov.  comb 4 

P.  monilata  (Tatem,  1870)  Foissner  &  Schiffmann,  1974 4 

P.  wwtaAW  (Penard,  1922)  Jankowski,  1976 4 

P.  nebulif era  (Mutter,  1786)  Jankowski,  1976 6 

P.  papillata  (Stiller)  Jankowski,  1976 6 

P.  pelagica  (Gajewskaja,  1933)  Jankowski,  1976  was  transferred  to  the  genus  Haplocaulis  by  Stiller 

(1971). 


1 2  A.  WARREN 

P. perlata (Kahl,  1933)  Jankowski,  1916  =  P.  punctata. 

P.  plicata  (Gourret  &  Roeser,  1886)  Jankowski,  1976  appears  to  be  identical  to  Vorticella  elongata 

Fromentel,  1874  (Warren,  1986). 

P.  pseudocampanula  Foissner,  \979 7 

P.  punctata  (Dons,  1918)  nov.  comb 7 

P.  quadrat  a  Foissner,  1979 7 

P.  sauwaldensis  Foissner  &Schiffmann,  1979 7 

P.  sphagni  Foissner,  1979 9 

P.  stilleri  (Stiller,  1963)  n.  sp 9 

P.  subconica  (Stiller,  1946)  Jankowski,  1976  appears  to  be  identical  to  P.  punctata. 

P.  vestita  (Stokes,  1883)  Jankowski,  1976.  A  membranous  alveolar  covering  overlays  the  pellicle  of  this 

species.  There  is,  however,  no  evidence  of  pellicular  tubercles  or  of  an  underlying  reticulate  pattern  of 

silver  lines.  This  species  should  therefore  remain  in  the  genus  Vorticella  (  V.  vestita)  until  it  has  been 

redescribed. 
P.  voeltzkowi  (Sondheim,  1929)  Jankowski,  1976.  This  species  has  spine-like  projections  on  its  pellicle. 

There  is,  however,  no  evidence  that  it  has  either  pellicular  tubercles  or  a  reticulate  pattern  of  silver 

lines.  It  should  therefore  remain  in  the  genus  Vorticella  (V.  voeltzkowi)  until  a  redescription  is 

available. 
P.  zooanthelligera  (Stiller,  1968)  nov.  comb 9 


The  taxonomic  status  of  the  genera  Pontigulasia, 
Lagenodifflugia  and  Zivkovida  (Rhi/opoda: 
Difflugiidae) 

Colin  G.  Ogden 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7  5BD 

Introduction 

The  significance  of  the  different  structural  elements  utilised  to  provide  an  internal  dividing  wall  in 
the  shell  of  the  genus  Pontigulasia  Rhumbler,  1896  was  employed  by  Ogden  (1983)  to  divide  the 
genus  into  two,  with  the  creation  of  a  new  genus  Zivkovida  Ogden,  1983.  At  the  same  time  a  new 
genus  Lagenodifflugia  was  erected  by  Medioli  &  Scott  (1983)  to  accommodate  Difflugia  vas  Leidy, 
1 874,  a  species  that  he  (Leidy,  1 879)  later  redescribed  as  a  variety  of  Difflugia  pyr  if ormis.  Medioli  & 
Scott  (1983)  considered  their  new  genus  to  be  distinct  from  species  of  Difflugia  because  the  shell  was 
divided  into  a  bulbous  main  part  and  a  neck,  the  two  parts  being  separated  by  an  internal 
diaphragm  pierced  by  a  single,  central,  usually  large  orifice.  In  the  same  work  Medioli  &  Scott 
redefined  the  genus  Pontigulasia,  describing  the  main  characters  as  a  constriction  of  the  neck 
marking  the  position  where  a  perforated  internal  diaphragm  often  extends  across  this  region,  and 
at  times  a  bent  neck  combined  with  the  internal  restriction  simulating  a  spiral  arrangement.  They 
also  stated  that  the  diaphragm  is  seldom  present  in  fossilised  forms.  The  amendment  of  the 
diagnosis  to  contain  a  'bent  neck'  was  made  to  allow  Medioli  &  Scott  to  include  some 
'Lecquereusia-like'  specimens  in  their  description  of  Pontigulasia  compressa,  and  to  suggest  the 
possibility  of  combining  the  genera  Lesquereusia  and  Pontigulasia,  the  former  being  the  senior 
synonym. 

I  am  most  grateful  to  Dr  Drew  Haman  and  Dr  Georges  Merinfeld  for  directing  my  attention  to 
the  nomenclature  problems  posed  by  the  creation  of  these  two  new  genera.  The  present  report  is  an 
attempt  to  clarify  the  status  and  diagnosis  of  the  three  genera  Pontigulasia,  Lagenodifflugia  and 
Zivkovida. 


Taxonomy 

The  taxonomic  problem  caused  by  the  creation  of  two  new  genera  relates  to  the  interpretation  and 
validity  of  the  structures  and  openings  found  associated  with  the  inner  dividing  wall.  There  are 
only  a  few  descriptions  and  figures  of  these  features  around  which  the  diagnoses  have  been  erected. 
One  reason  for  this  is  the  difficulty  of  trying  to  see  inside  the  shell  either  en  face  or  laterally  due 
mainly  to  the  opacity  of  the  mineral  particles  of  which  it  is  constructed.  Several  techniques  have 
been  tried  to  overcome  this,  for  example  immersion  in  clove  oil  or  Canada  balsam  and  deminerali- 
sation  by  hydrofluoric  acid.  Modern  techniques  have  now  obviated  this  problem  and  allow  a  new 
appraisal  of  this  feature  (Ogden,  1983). 

Each  genus  will  be  reassessed  here  on  the  basis  of  the  available  descriptions  and  the  author's 
earlier  observations. 

Pontigulasia  Rhumbler,  1 896 

The  first  author  to  recognise  the  inner  dividing  wall  was  Rhumbler  (1896)  who  erected  the  genus 
Pontigulasia  using  this  feature  as  a  major  diagnostic  character  to  separate  it  from  Difflugia,  and 
described  and  clearly  figured  it  as  an  internal  'schlundbriicke'-throat  bridge.  He  described  three 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  52(1):  13-17  Issued  29  January  1987 


14  C.  G.  OGDEN 

new  species  belonging  to  this  genus,  P.  compressa,  P.  incisa  and  P.  spiralis.  Since  then  Hopkinson 
(1919)  has  redescribed  P.  compressa  under  a  new  name  P.  r humbler i,  the  name  compressa  was 
preoccupied  by  Carter's  (1864)  species,  and  P.  incisa  has  become  a  synonym  off.  elisa  (Penard, 
1 893),  both  the  earlier  descriptions  being  for  species  of  Difflugia.  The  authoritative  date  for  P. 
rhumbleri  is  considered  to  be  Hopkinson's  redescription  (1919),  not  the  note  mentioned  as  an 
addition  to  P.  elisa  on  p.  162  of  Cash  &  Hopkinson  (1909)  where  Hopkinson  suggests  that 
'Rhumbler's  P.  compressa  (which  might  now  be  called  P.  Rhumbleri}\  his  brackets.  Furthermore, 
the  date  of  Rhumbler's  work  is  erroneously  quoted  by  Cash  et  al.  (1919)  and  Loeblich  &  Tappan 
(1964)  as  1895,  which  is  the  date  of  presentation,  whereas  the  publication  date  of  the  volume  was 
1896,  as  listed  in  Penard's  (1902)  bibliography. 

The  internal  bridge  has  been  redescribed  by  Ogden  ( 1 983),  who  examined  the  dividing  wall  in  situ 
using  scanning  electron  microscopy,  and  showed  it  to  be  correctly  termed  a  bridge,  as  it  represented 
a  rather  weak  connection  between  the  two  lateral  walls  of  the  compressed  shell  in  P.  rhumbleri.  It  is 
often  difficult  to  see  by  optical  microscopy  because  the  shells  are  laterally  compressed,  and  in  this 
the  normal  viewing  position  the  bridge  appears  to  be  a  dark  floating  band  situated  centrally  in  the 
neck  region  without  any  apparent  connection  to  the  shell  wall.  This  is  not  surprising,  because  each 
junction  of  the  bridge  with  the  wall  is  about  a  sixth  of  the  shell  diameter  at  this  point. 

Although  Rhumbler  described  three  species,  since  that  time  two  of  these  P.  incisa  and  P.  spiralis 
have  been  considered  to  be  synonyms  (Cash  &  Hopkinson,  1909),  and  it  was  not  until  Leoblich  & 
Tappan  (1964)  that  a  type  species,  P.  rhumbleri,  was  designated. 

DIAGNOSIS.  Shell  pyriform,  sometimes  with  a  constriction  of  the  neck,  either  circular  or  compressed 
in  transverse  section;  composed  mainly  of  agglutinate  mineral  particles  with  some  diatom  frustules 
or  siliceous  plates,  bound  by  a  network  of  organic  cement;  aperture  terminal,  circular;  internally 
the  shell  is  divided  into  two  regions  by  a  narrow  bridge,  made  mainly  of  organic  cement  with  some 
agglutinate  particles,  stretched  between  the  lateral  walls  at  about  one  third  of  the  body  length  from 
the  aperture.  Type  species  P.  rhumbleri  Hopkinson,  1919,  with  four  other  species  (Chardez,  1985), 
P.  compressoidea  Jung,  1 942;  P.  elisa  (Penard,  1 893);  P.  sarrazinensis  Chardez  &  Caspar,  1 984  and 
P.  spiralis  Rhumbler,  1896. 

Lagenodifflugia  Medioli  &  Scott,  1983 

This  genus  was  established  for  the  single  species  Difflugia  vas  Leidy,  1874.  After  his  original 
description  Leidy  later  considered  (1879)  that  this  species  was  a  variety  of  Difflugia  pyriformis, 
from  which  it  differed  by  a  constriction  of  the  neck,  there  was  no  reference  to  an  internal  structure 
associated  with  the  constriction.  Penard  (1902)  transferred  this  species  to  Pontigulasia  and  con- 
sidered it  to  be  a  synonym  of  his  new  species  P.  spectabilis.  It  was  corrected  to  the  valid  binomen  P. 
vas  by  Schoutenden  (1906),  and  spectabilis  has  since  been  considered  a  synonym  of  vas.  As  P.  vas 
was  described  as  having  a  similar  external  constriction  to  spectabilis,  the  assumption  was  that  it 
had  two  openings  bisecting  the  internal  diaphragm.  Notwithstanding  this,  Stump  (1935,  1936  and 
1 943)  in  a  series  of  experiments  with  specimens  he  initially  thought  were  Difflugia  oblonga,  later 
found  that  in  sectioned  shells  there  was  an  internal  division  with  a  single  opening  and  subsequently 
described  them  as  Pontigulasia  vas.  Although  Stump  does  not  clearly  state  that  his  specimens  had  a 
single,  central  opening,  his  diagrams  without  exception  suggest  that  this  interpretation  is  correct. 
That  the  strength  of  the  diaphragm  is  equal  to  that  of  the  shell  wall  is  extrapolated  from  his 
de-mineralised  sections  which  show  the  continuous  nature  of  these  structures. 

Using  the  joint  reports  of  Leidy  and  Stump,  a  composite  description  of  a  species  emerges:  it 
sometimes  has  a  constriction  of  the  neck  that  separates  the  anterior  third  of  the  shell  from  the  main 
body;  at  this  point  an  internal  diaphragm  is  present  which  is  pierced  by  a  single,  central,  circular 
opening. 

Medioli  &  Scott  (1983)  do  not  illustrate  the  internal  division  in  their  specimens  but  describe  it  as 
a  'large  orifice'.  They  presume  that  their  material  is  conspecific  with  that  of  Leidy  and  Stump, 
possibly  because  all  the  specimens  were  collected  in  America.  Fortunately,  a  single  specimen  with 
an  internal  division  was  found  in  material  kindly  left  at  the  British  Museum  (Natural  History)  by 
F.  S.  Medioli  and  helps  to  confirm  their  presumption.  The  specimen  had  the  following  measure- 


GENERA  PONTIGULASIA,  LA GENODIFFLUGIA  AND   ZIVKOVICIA  15 

ments:  174  um  long,  108  um  broad,  diameter  of  aperture  37  urn  and  diameter  of  internal  opening 
27  um.  Furthermore  the  organic  cement  pattern  of  this  specimen  was  typical  of  that  illustrated 
earlier  (Ogden,  1983). 

DIAGNOSIS.  Shell  pyriform,  often  with  a  constriction  of  the  neck,  most  frequently  circular  in 
cross-section  but  sometimes  slightly  compressed,  composed  mainly  of  agglutinate  mineral 
particles  bound  by  an  organic  cement;  aperture  terminal,  circular;  internally  the  shell  is  partitioned 
into  two  regions  by  a  diaphragm  constructed  as  part  of  the  shell  wall  but  having  a  single  central 
orifice.  Type  species  L.  vas  (Leidy,  1 874).  Three  other  species  are  here  attributed  to  the  genus  on  the 
basis  of  having  a  single  opening  in  a  well  constructed  diaphragm:  L.  bryophila  (Penard,  1902)  (see 
Ogden,  1983  for  recent  description  of  this  species);  L.  montana  (Ogden  &  Zivkovic,  1983)  and  L. 
epiouxi  (Chardez  &  Caspar,  1984). 

Zivkovicia  Ogden,  1987  gen.  nov. 

This  genus  was  erected  by  Ogden  (1983)  to  accommodate  those  species  of  Pontigulasia  which  had  a 
diaphragm  with  either  one  or  two  internal  openings.  At  that  time  no  distinction  was  drawn 
between  the  number  of  openings  in  the  diaphragm.  With  the  creation  of  Lagenodifflugia  to 
represent  those  specimens  with  a  single  opening,  Zivkovicia  is  redefined  here  to  represent  those 
species  with  two  openings. 

In  the  earlier  report  (Ogden,  1983),  due  to  an  oversight,  a  type  species  for  Zivkovicia  was  not 
designated,  so  under  Article  1 3(b)  of  the  International  Code  of  Zoological  Nomenclature  the  genus 
is  not  taxonomically  valid.  Nevertheless,  the  name  is  still  available  and  to  avoid  confusion  is  used 
again  here,  with  an  amended  diagnosis. 

Although  Carter's  (1 864)  original  description  ofDiffugia  compressa  did  not  include  a  mention  of 
an  internal  diaphragm,  his  drawings  are  so  precise  that  they  show  the  typical  V-shaped  notch  on 
the  shell  which  represents  the  internal  diaphragm.  This  structure  is  clearly  illustrated  by  Figs  1 8-25 
of  Ogden  (1983).  It  is  therefore  proposed  as  the  type-species  of  the  genus.  The  earlier  discussion 
(p.  14)  relating  to  the  incorrect  synonymy  of  P.  vas  and  P.  spectabilis,  allows  spectabilis  to  be 
available  to  include  the  original  description  of  specimens  with  a  bisected  diaphragm  (Penard,  1902) 
and  the  recent  description  by  Ogden  (1983),  whose  P.  vas  now  becomes  a  synonym  of  Z.  spectabilis. 

DIAGNOSIS.  Shell  pyriform,  often  with  a  distinct  constriction  of  the  neck  region,  either  circular  or 
compressed  in  transverse-section,  composed  mainly  of  agglutinate  mineral  particles  bound  by  an 
organic  cement  matrix;  aperture  terminal,  usually  circular;  internally  the  shell  is  partitioned  into 
two  parts  by  an  extension  of  the  shell  wall  to  form  a  diaphragm  which  is  bisected  by  two  circular 
openings.  Type  species  Z.  compressa  (Carter,  1864),  other  species  Z.  spectabilis  (Penard,  1902)  and 
Z.flexa  (Cash  &  Hopkinson,  1 909).  A  recent  description  of  the  latter  species  can  be  found  in  Ogden 
(1983). 


Discussion 

Associated  problems  of  clearly  identifying  the  internal  openings  were  discussed  previously  (Ogden, 
1 983).  Suffice  to  say  here  that  both  these  openings  and  the  aperture  can  be  sealed  by  an  organic  cyst 
membrane,  and  the  incidence  of  a  single  specimen  with  a  trisected  diaphragm  in  Z.  compressa  (Fig. 
28,  Ogden,  1983)  is  considered  to  be  an  isolated  deformity. 

In  discussing  a  possible  relationship  between  Pontigulasia  Rhumbler,  1896  and  Lesquereusia 
Schlumberger,  1845,  Medioli  &  Scott  (1983)  suggest  that  both  genera  are  characterised  by  a 
constriction  at  the  base  of  the  neck  which  corresponds  to  an  internal  diaphragm,  and  further  state 
that  a  morphological  intergradation  exists  between  the  two  genera.  They  consider  that  the 
remaining  difference  between  the  genera,  that  Lesquereusia  is  constructed  of  siliceous  idiosomes 
whereas  Pontigulasia  is  always  reported  to  be  composed  of  xenosomes,  is  insufficient  to  separate 
these  two  genera. 

It  has  already  been  established  by  Stump  (1936,  1943)  that  P.  vas  would  not  construct  a  shell  or 
reproduce  in  the  absence  of  extraneous  material,  even  in  the  presence  of  abundant  food,  and  that 


16  C.  G.  OGDEN 

such  deprived  animals  commenced  normal  reproductive  activities  when  shell  making  material  was 
reintroduced.  He  concluded  that  P.  vas  was  unable  to  secrete  its  own  shell  material  and  suggested  a 
possible  alternative  that  individuals  might  be  produced  without  a  shell  covering.  No  reports  of 
such  naked  individuals  have  been  recorded  in  the  literature,  but  it  is  well  known  that  some 
agglutinate  species  are  capable  of  constructing  an  organic  shell,  identical  to  that  which  in  the  field 
incorporates  mineral  particles  (Hedley  et  al.,  1976;  Netzel,  1972,  1976).  Furthermore,  the 
deposition  of  siliceous  structures  by  Lesquereusia  spiralis  are  carried  out  in  the  cytoplasm  of  the 
animal  (Harrison  et  al.,  1981),  and  this  species  is  capable  of  constructing  a  shell  in  the  absence  of 
extraneous  material.  It  is  equally  capable  of  incorporating  xenosomes  and  Stump  used  this  ability 
for  creating  'windows'  to  observe  cytoplasmic  activity.  A  new  family  the  Lesquereusiidae  was 
designated  (Ogden,  1979)  to  include  those  members  of  the  Lobosia  which  secrete  their  own 
siliceous  elements,  e.g.  Lesquereusia,  Netzelia  Ogden,  1979,  and  Quadrulella  Cockerell,  1909. 

The  suggestion  by  Medioli  &  Scott  (1983)  of  considering  a  relationship  between  Lesquereusia 
and  Pontigulasia  is  perhaps  best  treated  as  an  indiscretion  on  the  part  of  geologists  venturing  into 
the  alien  field  of  biology.  Phylogenetic  interpretations  should  be  based  on  the  animal  as  a  complete 
organism,  which  in  protozoa  would  include  information  on  cytoplasmic  detail,  movement,  repro- 
duction as  well  as  external  coverings,  especially  when  such  information  exists  in  publication  and 
the  animals  are  easily  collected  from  the  field. 


References 

Carter,  H.  J.  1864.  On  freshwater  Rhizopoda  of  England  and  India;  with  illustrations.  Annals  and  Magazine 

of  Natural  History.  London.  13(3):  18-39. 
Cash,  J.  &  Hopkinson,  J.  1909.  The  British  Freshwater  Rhizopoda  and  Heliozoa.  Vol.  II.  Rhizopoda,  part  2. 

The  Ray  Society,  London.  166  pp. 
Cash,  J.,  Wailes,  G.  H.  &  Hopkinson,  J.  1919.  The  British  Freshwater  Rhizopoda  and  Heliozoa.  Vol.  IV. 

Supplement  to  the  Rhizopoda.  The  Ray  Society,  London.  130  pp. 
Chardez,  D.  1985.  Note  sur  les  genres  Pontigulasia  Rhumbler  et  Zivkovicia  Ogden  (Rhizopoda,  testacea). 

Revue  Vervietoise  d'Histoire  Naturelle  42: 13-16. 
&  Caspar,  Ch.  1984.  Nouveaux  thecamoebiens  aquatiques  du  domain  des  Epioux  (Ardenne,  Belgique). 

Biologische  Jaarb  52:  57-63. 
Harrison,  F.  W.,  Dunkelberger,  D.,  Watabe,  N.  &  Stump,  A.  B.  1981.  Ultrastructure  and  deposition  of  silica  in 

rhizopod  amebae.  In:  T.  L.  Simpson  &  B.  E.  Volcani  eds,  Silicon  and  siliceous  structures  in  biological 

systems.  Springer- Verlag,  New  York.  pp.  281-294. 
Hedley,  R.  H.,  Ogden,  C.  G.  &  Mordan,  N.  J.  1976.  Manganese  in  the  shell  of  Centropyxis  (Rhizopodea: 

Protozoa).  Cell  and  Tissue  Research  171:  543-549. 
Leidy,  J.  1874.  Notice  of  some  new  freshwater  Rhizopods.  Proceeding  of  the  Academy  of  Natural  Sciences  of 

Philadelphia,  ser.  3:  77-79. 
1 879.  Freshwater  Rhizopods  of  North  America.  In:  'United  Stated  Geological  Survey  of  the  Territories', 

Vol.  12.  Washington,  324  pp. 
Loeblich,  A.  R.  &  Tappan,  H.  1964.  Thecamoebians'.  In:  'Treatise  on  Invertebrate  Paleontology.  Part  C. 

Protista  2,  Vol.  1.  C16-C54.  The  Geological  Society  of  America. 
Medioli,  F.  S.  &  Scott,  D.  B.  1983.  Holocene  Arcellacea  (Thecamoebians)  from  eastern  Canada.  Cushman 

Foundation  for  Foraminiferal  Research.  Special  publication  No.  21, 63  pp. 
Netzel,  H.  1972.  Die  Bildung  der  Gehausewand  bei  der  Thekamobe  Centropyxis  discoide  (Rhizopoda, 

Testacea).  Zeitschr if t  fur  Zellforschung  und  Mickroskopische  Anatomic  135: 45-54. 
1976.  Die  Abscheidung  der  Gehausewand  bei  Centropyxis  discoides  (Rhizopoda,  Testacea).  Archivfur 

Protistenkunde  118:  53-91. 
Ogden,  C.  G.  1979.  Siliceous  structures  secreted  by  members  of  the  subclass  Lobosia  (Rhizopoea:  Protozoa). 

Bulletin  of  the  British  Museum  (Natural  History)  (Zoology)  36:  203-207. 
1983.  The  significance  of  the  inner  dividing  wall  in  Pontigulasia  Rhumbler  and  Zivkovicia  gen.  nov. 

(Protozoa:  Rhizopoda).  Protistologica  19: 215-229. 

Penard,  E.  1902.  Faune  Rhizopodique  du  Bassin  du  Leman.  Geneva.  700  pp. 
Rhumbler,  L.  1896.  Beitrage  zur  Kenntnis  der  Rhizopoden.  Zeitschrift  fur  Wissenschaftliche  Zoologie  61: 

38-110. 


GENERA  PONTIGULASIA,  LAGENODIFFLUGIA  AND   ZIVKOVICIA  17 

Stump,  A.  B.  1935.  Observations  on  the  feeding  of  Difflugia,  Pontigulasia  and  Lesquereusia.  The  Biological 

Bulletin  oj  the  Marine  Laboratory •,  Woods  Hole  69:  136-142. 
1936.  The  influence  of  test  materials  on  reproduction  in  Pontigulasia  vas  (Leidy)  Schouteden.  The 

Biological  Bulletin  of  the  Marine  Laboratory,  Woods  Hole  70:  142-147. 
1943.  Mitosis  and  cell  division  in  Pontigulasia  vas  (Leidy)  Schouteden.  Journal  of  the  Elisha  Mitchell 

Scientific  Society  59:  14-22. 

Manuscript  accepted  for  publication  7  July  1986 

Since  this  manuscript  was  submitted  for  publication  the  author  has  become  aware  of  a  paper  by 
Medioli  &  Scott  (1985),  in  which  they  have  designated  type  specimens  for  certain  species.  In  dealing 
with  species  of  Pontigulasia,  Lagenodifflugia  and  Zivkovicia  they  make  assumptions  based  solely 
on  published  descriptions  and  micrographs  of  fossil  specimens,  which  do  not  have  information 
relating  to  the  main  diagnostic  feature  the  internal  dividing  diaphragm.  In  their  original  paper 
(Medioli  &  Scott,  1983),  they  comment  on  never  having  been  able  to  satisfactorily  observe  the 
diaphragm  of  P.  compressa  (Carter,  1864),  and  their  failure  is  again  reiterated  in  Medioli  &  Scott 
(1985),  As  neither  their  figures  nor  plates  illustrate  an  internal  structure  it  suggests  that  they 
have  failed  by  both  optical  and  electron  microscopy  to  see  this  feature  in  any  of  their  specimens. 
They  presume  the  presence  of  a  diaphragm  solely  on  it  corresponding  in  position  to  the  external 
constriction.  Although  they  admit  that  this  external  constriction  is  often  obscured  by  agglutinate 
material. 

Furthermore,  the  specimen  they  selected  as  neotype  of  L.  vas,  from  their  Maritime  Canada 
sample  does  not  even  resemble  the  original  selected  figure  from  Leidy  (1879).  In  fact  they  chose  an 
example  which  was  intermediate  between  Leidy's  figured  specimen,  a  smoothly  agglutinate  form, 
and  their  extreme  cases  of  coarse  agglutination. 

Both  of  Medioli  &  Scott's  (1983, 1985)  papers  concerning  these  genera  must  be  of  minimal  value 
because  of  their  failure  to  describe  or  identify  the  internal  structures  which  typify  these  genera.  Its 
absence  suggests  that  they  have  been  examining  species  of  Difflugia.  Their  insistance  that  the 
diaphragm  is  absent  in  fossil  forms  (p.  35,  1983;  p.  29,  1985)  of  Pontigulasia  compressa  (Carter, 
1 864)  would  indicate  that  they  are  dealing  with  species  of  Difflugia.  The  construction  of  the 
diaphragm  in  this  species  (Ogden,  1 983)  is  so  robust,  being  continuous  and  as  thick  as  the  shell  wall, 
that  even  in  fractured  shells  it  is  the  wall  that  breaks  and  not  the  diaphragm.  Although  often  by 
convenience  such  breaks  occur  at  the  junction  of  the  wall  and  diaphragm.  Notwithstanding  this, 
specimens  clearly  identified  as  Zivkovicia  compressa  (Carter,  1864)  have  been  recovered  from  core 
samples  taken  in  Lake  Ullswater  which  had  complete  diaphragms  (Ogden  &  Ellison,  in  prep.),  and 
in  addition  possessed  the  typical  organic  cement  units,  described  by  Ogden  (1983),  specific  to  this 
genus. 

Medioli,  F.  S.  &  Scott,  D.  B.  1985.  Designation  of  types,  for  one  genus  and  nine  species  of  Arcellaceans 
(Thecamoebians),  with  additional  original  reference  material  for  four  other  species.  Journal  of 
Foraminiferal  Research  15:  24-37. 


A  revision  of  the  foraminiferal  genus  Adercotryma 
Loeblich  &  Tappan,  with  a  description  of  A.  wrightisp. 
nov.  from  British  waters 

P.  Bronnimann 

9G,  Chemin  de  Bedex,  1226  Thonex,  Geneva,  Switzerland 

J.  E.  Whittaker 

Department  of  Palaeontology,  British  Museum  (Natural  History),  Cromwell  Road,  London 
SW7  5BD 

Adercotryma  Loeblich  &  Tappan  (1952)  was  erected  to  accommodate  Lituola  glomerata  Brady 
(1878),  a  species  assigned  subsequently  to  Haplophragmium,  Haplophragmoides  and  Trochammina 
by  various  authors.  A  recent  examination  of  the  type  material  in  the  British  Museum  (Natural 
History),  and  of  specimens  from  other  collections  deposited  there  and  in  the  National  Museum  of 
Ireland,  led  to  the  discovery  that  the  original  definition  was  inadequate  and  to  the  recognition  of  a 
second  species.  The  purpose  of  this  paper  is  to  emend  the  diagnosis  of  Adercotryma,  to  redescribe 
A.  glomeratum  (Brady),  and  to  describe  the  new  species. 

The  generic  diagnosis  below  is  based  on  the  redescription  of  the  type  species,  and  differs  from  the 
original  definition  in  that  it  recognises  the  significance  of  the  asymmetrically  placed  aperture  and 
shows  the  coiling  to  be  trochospiral.  Adercotryma  is  therefore  transferred  from  the  Lituolacea  to 
the  Trochamminacea  and  placed  in  a  new  subfamily  of  the  Trochamminidae.  The  definition 
follows  the  format  adopted  by  Bronnimann  et  al.  (1983)  in  their  reclassification  of  the 
Trochamminacea. 

Superfamily  TROCHAMMINACEA  Schwager,  1877 

Family  TROCHAMMINIDAE  Schwager,  1877 

Subfamily  ADERCOTRYMINAE  nov. 

DEFINITION.  Test  free;  adult,  a  completely  or  almost  completely  involute,  cone-like,  trochospire; 
wall  agglutinated,  imperforate,  single-layered,  aperture  interiomarginal,  single;  without  secondary 
septa  or  infoldings  of  the  umbilical  chamber  walls  or  incomplete  secondary  partitions. 

TYPE  GENUS.  Adercotryma  Loeblich  &  Tappan,  1952. 

REMARKS.  The  Adercotryminae  differs  from  all  other  subfamilies  in  that  its  members  are 
completely  or  almost  completely  involute  on  the  spiral  side.  Bronnimann  et  al.  (1983:  204) 
distinguished  the  Trochamminacea  from  the  Ataxophragmiacea  on  the  ratio  of  spire  height  to 
umbilical  diameter:  the  former  being  always  smaller  than  the  latter  in  the  Trochamminacea.  At 
first  sight,  the  high  cone-shaped  test  of  Adercotryma  does  not  fulfil  this  criterion,  but  since  the 
coiling  is  involute  and  the  proloculus  is  situated  within  the  shell  (see  Figs  3,  6),  the  spire  height 
measured  from  the  proloculus  is  invariably  less  than  the  umbilical  diameter. 

Genus  ADERCOTRYMA  Loeblich  &  Tappan,  1952 

TYPE  SPECIES.  Lituola  glomerata  Brady,  1878.  Recent,  marine;  distribution  apparently  worldwide. 
Lectotype  from  Arctic  waters. 

EMENDED  DEFINITION.  Test  free;  coiling  trochospiral,  adult  an  inverted  cone,  completely  or  almost 
completely  involute  on  both  sides.  Chambers  axially  elongate.  Aperture  single,  interiomarginal, 
umbilical,  symmetrical  with  respect  to  long  axis  of  chamber.  Wall  agglutinated,  single  layered, 
imperforate. 


Bull.  Br.  Mm.  nat.  Hist.  (Zool.)  52(1):  19-28  Issued  29  January  1987 


20 


P.  BRONNIMANN  &  J.  E.  WHITTAKER 


REMARKS.  The  slit-like  aperture  rests  with  its  border  on  the  first  and  on  the  penultimate  chamber  of 
the  final  whorl  (Paratrochammina-type  aperture).  Adercotryma  differs  from  Paratrochammina 
Bronnimann,  1979  (type  species:  P.  madierae  Bronnimann,  1979)  and  all  other  genera  of  the 
Trochammininae  by  its  spirally  involute  enrolment,  axially  elongate  chambers,  symmetrical 
interiomarginal  aperture  (with  respect  to  the  long  axis  of  the  chamber),  and  inverted  cone-like  test. 

Adercotryma  glomeratum  (Brady) 
Figs  1,  2A-F,  3 A,  4A-E,  5A-J,  6A-F 

1878  Lituola  glomerata  Brady:  433,  pi.  20,  figs  la-c. 

1 884  Haplophragmiwn  glomeratum  (Brady);  Brady:  309,  pi.  34,  figs  15-18. 

1910  Haplophragmoides  glomeratum  (Brady)  (sic);  Cushman:  104,  figs  158-161  (after  Brady,  1884). 

1 93 1  Trochammina  glomerata  (Brady);  Wiesner:  1 1 2,  pi.  1 7,  figs  204,  205. 

1952  Adercotryma  glomerata  (Brady)  (sic);  Loeblich  &  Tappan:  141,  figs  1-4. 

1961  Adercotryma  glomerata  glomerata  (Brady);  Saidova:  35,  pi.  10,  fig.  54. 

1961  Adercotryma  glomerata  abyssorum  Saidova  (sic):  36,  pi.  10,  fig.  55. 

1975  Adercotryma  glomerata  antarctica  Saidova  (sic):  75,  pi.  96,  fig.  6. 

MATERIAL.  Extant  material  in  the  Brady  Collection  of  the  British  North  Polar  Expedition 
(1875-1876),  labelled  Lituola  glomerata,  is  as  follows:  Station  A,  off  Tyndall  Glacier,  27fms 
(49m);  F,  between  Walrus  Shoal  and  Victoria  Head,  57fms  (104m);  H,  Franklin  Pierce  Bay, 
13-1 5  fms  (24-28  m);  I,  Allman  Bay,  25  fms  (46m);  J,  Dobbin  Bay,  45^17  fms  (82-86  m);.  K, 
Dobbin  Bay,  1 1 3  fms  (207  m);  N,  off  Hayes  Point,  35  fms  (64  m);  O,  off  Cape  Frazer,  50  fms  (92  m) 
and  P,  off  Cape  Frazer,  80  fms  (146  m).  These  localities  are  from  the  northern  part  of  Baffin  Bay 
and  Smith  Sound  (between  Ellesmere  Island,  NE  Canada,  and  W  Greenland).  All  the  slides 
contain  a  few  specimens  at  least,  and  some  (e.g.  station  G)  as  many  as  50. 


Fig.  1.  Adercotryma  glomeratum  (Brady).  Paralectotype,  1955.10.28.1732.  Interpretative  drawing  of 
specimen  in  Figs  6A-C,  taken  at  the  third  level  of  dissection  (see  explanation  of  Fig.  6C),  showing 
chambers  1  to  9.  Hatched  areas  represent  exposed  walls  of  earliest  chambers,  x  300. 

From  Franklin  Pierce  Bay,  lat.  79°28'N,  station  H,  depth  46  fathoms  (84  m).  British  North  Polar 
Expedition  of  1875-1876,  ex  BMNH  slide  no.  1955.10.28.1731-1780,  labelled  'Lituola  glomerata 
Brady'. 


GENUS  ADERCOTRYMA 


21 


Fig.  2A-F.  Adercotryma  glomeratum  (Brady).  2A-C,  Paralectotype,  1955.10.28.1701.  Interpretative 
drawing  of  specimen  in  Figs  5E-G,  J.  2A,  umbilical  view  showing  the  overlapping  chamber  walls  and 
the  preserved  apertural  slits  of  the  last  three  chambers;  2B,  spiral  view;  2C,  edge  view  showing  aperture 
of  final  chamber,  with  axis  of  coiling  marked  by  line  A.  2D-F,  Paralectotype,  1955.10.28.1700. 
Interpretative  drawing  of  specimen  in  Figs  5A-D.  2D,  umbilical  view  showing  aperture  of  final 
chamber  in  part  masked  by  agglutinated  or  secreted  material;  2E,  spiral  view;  2F,  edge  view  of  inverted 
cone-like  test  with  aperture  of  final  chamber  in  part  masked,  axis  of  coiling  is  indicated  by  line  A.  Both 
x!50. 

Both  from  slide  labelled  ' Lituola  glomerata  Brady'.  British  North  Polar  Expedition  of  1875-1876. 
Cape  Frazer,  lat.  79°45'N,  station  O,  depth  50  fathoms  (92m),  ex  BMNH  slide  no. 
1955.10.28.1700-1731. 


LECTOTYPE.  1955.10.28.1781  (Figs  4A-E).  From  Brady's  syntypic  series,  obtained  from  Station  P, 
off  Cape  Frazer,  Arctic  Canada,  depth  80  fathoms  (146  m).  Believed  to  be  the  specimen  figured  by 
Brady  (1878,  pi.  20,  fig.  Ib). 

DESCRIPTION  (LECTOTYPE).  Test  free;  a  dextral,  tightly  coiled  trochospire,  with  4  chambers  in  the 
final  whorl,  each  gradually  increasing  in  size;  involute  on  spiral  side.  Test  a  short,  broad,  inverted 
cone-like  structure,  flatly  truncated  spirally,  rounded-convex  umbilically,  broadly  rounded  peri- 
pherally and  somewhat  rounded  laterally.  In  edge  view,  3  chambers  seen  on  both  sides.  In  spiral/ 
umbilical  view,  oval-lobate;  umbilical  side  with  a  small,  well-defined  subcircular  and  shallow 
axial  depression.  Adult  chambers  much  elongated  in  axial  (edge)  view,  narrow  radially  and  some- 
what elongate  tangentially,  more  inflated  towards  the  spiral,  than  towards  the  umbilical  side. 
Intercameral  sutures  straight  but  indistinct  spirally;  straight,  distinct,  laterally  and  umbilically. 
Aperture  single,  interiomarginal,  a  narrow  elongate  slit  with  rounded  extremities,  at  umbilical  end 
of  chamber;  symmetrical  with  respect  to  its  long  axis.  Border  of  aperture  rests  on  the  first  and 


22 


P.  BRONNIMANN  &  J.  E.  WHITTAKER 


Fig.  3A  Adercotryma  glomeratum  (Brady).  Paralectotype,  1955. 10.28. 1782.  Section  cut  parallel  to  axis 
of  coiling.  Note  the  thin-walled  proloculus  already  slightly  elongate  in  the  direction  of  the  coiling  axis. 
x205. 

From  Cape  Frazer,  lat.  79°45'N,  station  P,  depth  80  fathoms  (146m).  British  North  Polar 
Expedition  of  1875-1876,  ex  BMNH  slide  no.  1955.10.28.1781-1799,  labelled  'Lituola  glomerata 
Brady'. 

Fig.  3B.  Adercotryma  wrighti  Bronnimann  &  Whittaker  sp.nov.  ZF  4453.  Section  cut  slightly  obliquely 
to  axis  of  coiling.  x250. 

From  south  of  Mull,  W  Scotland,  depth  20  fathoms  (37m).  S.Y.  Runa  station  2,  collected  1913. 
Heron-Allen  &  Earland  Collection  (BMNH),  slide  labelled  ' Haplophragmium  glomeratum  (Brady)'. 


penultimate  chambers  of  final  whorl  (Paratrochammina-type).  Final  chamber  covers  about  half  of 
the  preceding  apertural  slit.  Wall  agglutinated,  imperforate,  coarser  on  truncated,  spiral  side  than 
on  rounded-comvex,  umbilical  side.  Colour,  prior  to  coating  for  SEM  photography,  yellowish- 
brown. 

DIMENSIONS  (LECTOTYPE).  Maximum  spiral/umbilical  diameter  320  um,  minimum  diameter 
270  um,  height  260  um.  Height  of  apertural  slit  c.  12  um. 

PARALECTOTYPES:  3  sinistral  specimens  (1955.10.28.1700-1702)  are  figured  in  Figs  2A-F,  5A-J; 
another  3  (1955.10.28.1732,  1955.10.28.1783  and  1955.10.28.1703),  dissected  out  to  show  various 
aspects  of  the  internal  coiling,  are  figured  in  Figs  1,  6A-F,  whilst  a  further  paralectotype 
(1955.10.28.1782)  has  been  sectioned  and  is  illustrated  in  Fig.  3 A.  These  specimens,  as  with  others 
remaining  in  Brady's  syntypic  series,  vary  considerably  in  their  dimensions,  elongation  of  the 
chambers  in  the  final  whorl,  depression  of  the  sutures,  depth  of  the  umbilicus  and  spiral  aspect.  For 
further  comments,  see  the  figure  explanations  and  Remarks  section  below.  The  maximum  spiral/ 
umbilical  diameter  of  the  figured  paralectotypes  varies  from  230  to  290  um,  the  test  height,  from 
250  to  270  um. 

REMARKS.  Brady's  small  and  enigmatic  species  was  placed  by  authors  in  Lituola,  Haplophragmium, 
Haplophragmoides  and  Trochammina  prior  to  the  erection  of  Adercotryma  by  Loeblich  &  Tappan 
( 1 952).  The  generic  changes  stem  mainly  from  differing  interpretations  of  the  mode  of  coiling  of  the 
curious  cone-shaped  test. 

Although  Brady  (1878)  originally  referred  to  the  test  as  merely  '. . .  spiral  in  arrangement',  his 
subsequent  comparison  (Brady,  1884)  of  the  overall  shape  with  that  of  a'. . .  nautiloid  species,  such 
as  Haplophragmium  latidorsatum,  drawn  out  as  the  umbilici  so  as  to  form  a  test  bearing  some 
resemblance  to  the  oval  Alveolinae',  implied  that  the  coiling  was  planispiral.  He  clearly  was  not 
sure,  however,  as  he  made  much  in  these  two  papers  of  the  unusual  'unsymmetrical  convolutions'. 

It  was  Cushman  (1910)  who  first  described  the  coiling,  without  reservation,  as  planispiral, 
placing  Brady's  species  in  his  new  genus  Haplophragmoides,  an  assignment  which  was  generally 


GENUS  ADERCOTRYMA 


Fig.  4A-E.  Adercotryma  glotneratum  (Brady).  Lectotype,  1955.10.28.1781.  4A-D,  spiral,  edge 
(apertural),  umbilical  and  edge  (antapertural)  views,  x  1 50. 4E,  detail  of  lateral,  open  part  of  aperture, 
x525. 

From  slide  labelled ' Lituola glomeratum  Brady'.  British  North  Polar  Expedition  of  1 875-1 876.  Cape 
Frazer,  lat.  79°45'N,  station  P,  depth  80  fathoms  (146  m),  ex  BMNH  slide  no.  1955.10.28.1781-1799. 


followed  for  over  forty  years.  The  only  exception  was  Wiesner  (1931)  who  placed  glomerata  in 
Trochammina,  although  he  did  not  make  a  detailed  examination  of  its  morphology  and  his  paper 
offers  no  evidence  for  trochospiral  coiling.  A  year  earlier,  however,  Lacroix  (1930)  had  considered 
the  position  of  the  aperture,  ignored  completely  by  Cushman,  to  be  more  in  keeping  with  a 
trochospiral  genus.  In  terms  of  coiling,  Lacroix  considered  Brady's  species  transitional  between 
the  planispiral  Haplophragmoides  and  the  trochospiral  Trochammina,  but  nevertheless  retained  it 
in  the  former  genus. 

In  1952,  Loeblich  &  Tappan  erected  a  new  lituolid  genus  Adercotryma,  with  Lituola  glomerata 
Brady  as  type.  The  name  refers  to  the  apertural  features,  derived  from  two  Greek  words  aderco- 
unseen,  invisible,  and  tryma-  meaning  a  hole  or  aperture.  The  gender  of  the  name  Adercotryma  is 
neuter,  and  the  specific  name  should  be  construed  as  glomeratum,  not  glomerata  as  originally 
written.  Loeblich  &  Tappan  (1952)  distinguished  their  new  genus  from  Haplophragmoides  on  the 
somewhat  asymmetrical,  completely  involute,  rather  than  slightly  evolute  test  which  has  its 
greatest  dimension  in  the  axis  of  coiling,  and  by  its  aperture  which  lies  near  the  umbilicus  of 
one  side,  rather  than  in  the  plane  of  coiling  at  the  periphery.  Of  these  features,  only  two  are 
fundamentally  different  from  those  of  Haplophragmoides:  the  asymmetrical  test  morphology  and 
the  asymmetrical  interiomarginal  apertural  position.  In  no  part  of  their  original  paper,  nor  in  1964, 
did  Loeblich  &  Tappan  discuss  the  curious  asymmetry  of  what  they  obviously  assumed  to  be  a 
planispiral  test.  Adercotryma  was  placed  in  the  Haplophragmoidinae  Maync,  1952  (Lituolidae  de 
Blainville,  1825),  in  which  were  included  both  planispiral  and  streptospiral  forms. 

Even  though  the  test  of  A.  glomeratum  is  involute,  the  external  and  internal  morphology  clearly 


24 


P.  BRONNIMANN  &  J.  E.  WHITTAKER 


Figs  5A-J.  Adercotryma  glomeratum  (Brady).  5A-D,  Paralectotype,  1955.10.28.1700.  Spiral  edge, 
oblique-umbilical  and  umbilical  views,  x  150.  5E-G,  J,  Paralectotype,  1955.10.28.1701,  E-G,  spiral, 
edge  and  umbilical  views,  x  1 50;  5J,  detail  of  interiomarginal  apertures  of  final  and  penultimate 
chambers,  in  umbilical  view,  x475.  5H,  I,  Paralectotype,  1955.10.28.1702.  Spiral  and  edge  views, 
x!50. 

All  specimens  from  slide  labelled  'Lituola  glomerata  Brady'.  British  North  Polar  Expedition  of 
1875-1876.  Cape  Frazer,  lat.  79°45'N,  station  O,  depth  50  fathoms  (92m),  ex  BMNH  slide  no. 
1955.10.28.1700-1731. 


Figs  6A-F.  Adercotryma  glomeratum  (Brady).  6A-C,  Paralectotype,  1955. 10.28. 1 732.  Stereo-pairs  of 
three  stages  of  dissection,  perpendicular  to  axis  of  coiling;  the  involute  spiral  side  has  been  removed. 
The  third  stage  of  dissection  (6C)  has  broken  open  the  earliest  whorl  and  proloculus  (see  Fig.  1,  for 
interpretative  drawing).  6D,  E,  Paralectotype,  1955.10.28.1783.  Stereo-pairs  of  dissected  specimen 
shown  at  two  different  tilts.  Dissection  is  in  plane  virtually  parallel  to  coiling  axis.  6F,  Paralectotype, 
1955.10.28.1703.  Stereo-pair  of  specimen  dissected  perpendicular  to  axis  of  coiling.  All  x  150. 

Figs  6A-C  from  Franklin  Pierce  Bay,  lat.  79°28'N,  station  H,  depth  46  fathoms  (84  m),  ex  BMNH 
slide  no.  1955.10.28.1731-1780.  Figs  6D,  E  from  Cape  Frazer,  lat.  79°45'N,  station  P,  depth  80 
fathoms  (146m),  ex  BMNH  slide  no.  1955.10.28.1781-1799.  Fig.  5F,  same  locality,  station  O,  depth 
50  fathoms  (92m),  ex  BMNH  slide  no.  1955.10.28.1700-1731.  British  North  Polar  Expedition  of 
1875-1876. 


GENUS  ADERCOTRYMA 


25 


26 


P.  BRONNIMANN  &  J.  E.  WHITTAKER 


Figs.  7A-J.  Adercotryma  wrighti  Bronnimann  &  Whittaker  sp.nov.  7A-D,  Holotype,  NMI  no. 
149. 1985.  Spiral,  edge  (apertural),  umbilical  and  edge  (antapertural)  views.  7E,  F,  Paratype,  NMI  no. 
4.1980.  Oblique-umbilical  and  edge  views.  7G-J,  Paratype,  NMI  no.  5.1980.  Spiral,  edge  (apertural), 
edge  (antapertural)  and  umbilical  views.  All  x  175. 

All  from  offDrogheda,  E  Ireland,  depth  16  fathoms  (29m);  ex  slide  no.  34,  labelled  'Dublin:  off 
Drogheda,  16  fms  &  Lambay  Deep,  70  fms  (mixed)',  J.  Wright  Collection,  13-1921,  National  Museum 
of  Ireland. 


indicates  a  trochospiral  mode  of  coiling  (Figs  1-6).  As  well  as  the  asymmetrical  aperture,  the  adult 
test,  when  orientated  with  the  axis  of  coiling  in  vertical  position,  shows  a  truncated,  more  or  less 
flattened  aboral  or  spiral  side,  and  an  obtusely  pointed,  ovoid-rounded  oral  or  umbilical  side.  This 
differentiation  is  typical  of  trochospiral  tests. 

Saidova  (1961 ;  1975)  introduced  two  new  subspecies  of  A.  glomeratum,  namely  A.  g.  abyssorum 
and  A.  g.  antarctica,  respectively.  They  are  figured  together  with  a  typical  A.  g.  glomeratum  also  in 
Saidova  ( 1 975,  pi.  96,  figs  4-6).  The  difference  in  shell  morphology  said  to  characterise  the  two  (test 
size,  chamber  shape  and  elongation)  falls  within  the  range  of  variation  seen  in  our  paralectotypes 
(compare  our  Figs  5B  and  51  with  Saidova's  pi.  96,  figs  6  and  5,  respectively),  whilst  the  type  of 


GENUS  ADERCOTRYMA  27 

agglutinant,  also  used  by  Saidova  (1961)  as  a  distinguishing  feature  of  A.  g.  abyssorum,  is  not 
considered  by  us  to  have  any  taxonomic  validity. 

A.glomeratum  (Brady)  differs  from  A .  wrighti  sp.  nov.  in  having  a  broadly  inverted  cone-like  test 
with  4  axially  elongate  chambers  in  the  final  whorl,  in  the  shape  of  the  adult  chambers,  and  in  the 
apertural  features.  See  also  pp  27,  28  for  further  remarks  on  their  differences.  In  the  material 
studied,  A.  glomeratum  always  has  4  chambers  in  the  final  whorl,  even  the  preceding  whorl  (Figs  1, 
6C)  has  4  chambers.  This  results  in  3  chambers  being  visible  on  either  side  of  the  test  when  seen  in 
edge  view  (aperturally  and  antaperturally). 

A.  glomeratum  is  a  very  wide-ranging  species  both  in  terms  of  latitudinal  and  depth  distribution 
as  noted  by  Saidova  (1975)  and  Culver  &  Buzas  (1985). 

Adercotryma  wrighti  Bronnimann  &  Whittaker  sp.  nov. 
Figs  3B,  7A-J 

1908  Haplophragmium  glomeratum  (Brady);  Millett:  5  (list),  pi.  1,  fig.  6  (non  Lituolaglomerata  Brady,  1878). 
1913  Haplophragmium  glomeratum  (Brady);  Heron- Allen  &  Earland:  46,  pi.  2,  fig.  14. 

DIAGNOSIS.  A  species  of  Adercotryma  with  only  3  chambers  in  the  final  whorl.  In  spiral/umbilical 
view,  test  oval-lobate,  maximum  diameter  often  almost  twice  the  minimum  diameter.  In  edge  view, 
3  chambers  seen  on  apertural  side,  only  2  on  antapertural  side.  Aperture  single,  interiomarginal,  a 
bilobed  narrow,  elongate  slit  without  rounded  extremities  at  umbilical  end  of  chamber. 

NAME.  In  honour  of  Joseph  Wright,  in  whose  collection  from  Dublin  Bay  this  species  was  first 
noticed. 

HOLOTYPE.  National  Museum  of  Ireland  (NMI)  no.  149.1985.  Illustrated  in  spiral,  edge 
(apertural),  umbilical  and  edge  (antapertural)  views  in  Figs  7A-D.  Ex  J.  Wright  Collection,  slide 
34. 

TYPE  LOCALITY.  OffDrogheda,  E  Ireland,  depth  16  fathoms  (30  m). 

DESCRIPTION  (HOLOTYPE).  Test  free;  a  dextral,  tightly  coiled  trochospire,  with  3  chambers  in  the 
final  whorl,  gradually  increasing  in  size;  involute  on  spiral  side.  Test  an  inverted  cone-like  struc- 
ture, truncated  spirally,  rounded-convex  umbilically  and  broadly  rounded  peripherally.  In  edge 
view,  3  chambers  seen  on  apertural  side,  2  on  antapertural  side.  In  spiral/umbilical  view,  oval- 
lobate,  maximum  diameter  almost  twice  the  minimum  diameter;  umbilical  side  with  very  shallow 
and  small  axial  depression.  Adult  chambers  elongate  in  axial  (edge)  view,  less  elongate  in  tangential 
direction,  narrow  in  radial  direction;  inflated  equally  both  spirally  and  umbilically.  Intercameral 
sutures  well  defined  and  slightly  incurved  laterally  and  umbilically,  less  well  defined  and  straight 
spirally.  Aperture  single,  interiomarginal,  a  bilobed  narrow  slit  without  rounded  extremities,  at 
umbilical  end  of  chamber,  symmetrical  with  respect  to  long  chamber  axis.  Border  of  aperture  rests 
on  first  and  on  penultimate  chamber  of  final  whorl  (Paratrochammina-typo).  Wall  agglutinated, 
imperforate,  coarser  on  spiral  side  than  on  umbilical  side.  Colour  of  test,  prior  to  coating  for  SEM 
photography,  orange-brown. 

DIMENSIONS  (HOLOTYPE).  Maximum  spiral/umbilical  diameter  240  um,  minimum  diameter  1 50  um, 
height  220  um. 

PARATYPES.  Two  paratypes  are  figured  herein.  NMI  no.  4. 1980  (Figs  7E,  F)  is  a  sinistral  specimen; 
the  aperture  is  perfectly  preserved  and  shows  the  bilobed,  narrow,  elongate  slit;  the  development  of 
a  central,  triangular  lip-like  projection  of  the  chamber  wall  serving  to  divide  the  aperture  into  two 
virtually  identical  parts.  This  specimen  has  a  maximum  spiral/umbilical  diameter  of  220  um  and 
test  height  of  240  um.  NMI  no.  5.1980,  the  other  illustrated  paratype  (Figs  7G-J),  is  dextrally 
coiled  like  the  holotype.  Its  maximum  spiral/umbilical  diameter  is  200  um,  the  test  height  210  um. 
The  sectioned  specimen  (ZF  4453),  from  the  Heron-Allen  &  Earland  Collection,  off  W  Scotland, 
has  a  maximum  diameter  of  220  um  and  height  of  180  urn;  it  is  figured  in  Fig.  3B. 

REMARKS.  Adercotryma  wrighti  sp.  nov.  is  easily  distinguished  from  A.  glomeratum  (Brady)  by  the 
overall  shape  of  the  test,  only  3  chambers  in  the  final  whorl,  the  shape  of  the  adult  chambers  and  the 


28  P.  BRONNIMANN  &  J.  E.  WHITTAKER 

bilobed  apertural  features  formed  by  the  triangular  lip-like  projection  of  the  chamber  wall.  A. 
glomeratum  always  has  4  chambers  in  the  final  whorl  and  a  test  which  in  spiral/umbilical  aspect  has 
a  maximum  diameter  little  greater  than  the  minimum. 

Both  Millett  (1908)  and  Heron- Allen  &  Earland  (1913)  show  specimens  from  W  Ireland  with 
only  3  chambers  in  the  final  whorl.  We  have  examined  their  collections  and  many  slides  in  the 
Brady  and  the  Norman  Collections  (BMNH),  labelled  Haplophragmium  glomeratum  (Brady), 
from  Scotland,  Ireland  and  N  England  and  all  exclusively  contain  A.  wrighti  rather  than  A. 
glomeratum.  Careful  study  of  specimens  previously  recorded  as  A.  glomeratum  may  extend  the 
present  range  of  A.  wrighti  beyond  the  British  Isles. 

Acknowledgements 

The  Director  of  the  National  Museum  of  Ireland  is  thanked  for  his  permission  to  borrow  and 
photograph  specimens  from  the  Wright  Collection;  Mr  J.  M.  C.  Holmes  facilitated  the  loan.  We 
are  pleased  to  acknowledge  the  technical  skill  of  Mr  R.  L.  Hodgkinson,  British  Museum  (Natural 
History),  in  preparing  the  dissections  and  thin  sections  of  the  Adercotryma  spp.;  Mrs  L.  M. 
McCormick  and  Mr  P.  V.  York  took  the  SEM  and  optical  photographs,  respectively,  whilst  Drs 
C.  G.  Adams  and  M.  K.  Howarth,  also  of  the  same  institution,  kindly  read  the  manuscript  and 
suggested  many  improvements.  The  research  of  P.  Bronnimann  is  in  part  funded  by  the  Fonds 
National  Suisse. 

References 

Blainville,  H.  M.  D.  de  1 825.  Manuel  de  Malacologie  et  de  Conchy  Uologie.  664  pp.,  87  pis.  F.  G.  Levrault,  Paris. 
Brady,  H.  B.  1878.  On  the  reticularian  and  radiolarian  Rhizopoda  (Foraminifera  and  Polycystina)  of  the 

North  Polar  Expedition  of  1875-76.  Ann.  Mag.  nat.  Hist.,  London,  ser.  5, 1: 425-440,  pis  20, 21. 
1884.  Report  on  the  Foraminifera  dredged  by  H.M.S.  Challenger  during  the  years  1873-76.  Rep.  sclent. 

Results  Voy.  Challenger  (Zool.),  London,  9: 1-814,  pis  1-1 15. 
Bronnimann,  P.  1979.  Recent  benthonic  foraminifera  from  Brasil  —  Morphology  and  Ecology;  part  4: 

Trochamminids  from  the  Campos  Shelf  with  description  of  Paratrochammina  n.gen.  Palaeont.  Z., 

Stuttgart,  53:  5-25,  figs  1-10. 
,  Zaninetti,  L.  &  Whittaker,  J.  E.  1983.  On  the  classification  of  the  Trochamminacea  (Foraminiferida).  J. 

foramin.  Res.,  Washington,  13: 202-218,  pis  1-3. 
Culver,  S.  J.  &  Buy  as,  M.  A.  1985.  Distribution  of  Recent  benthic  foraminifera  of  the  North  American  Pacific 

Coast  from  Oregon  to  Alaska.  Smithson.  Contrib.  Mar.  Sci.,  Washington,  26: 1-234,  figs  1-139. 
Cushman,  J.  A.  1910.  A  monograph  of  the  Foraminifera  of  the  North  Pacific  Ocean;  Part  1 ,  Astrorhizidae  and 

Lituolidae.  Bull.  U.S.  natn.  Mus.,  Washington,  71(1):  1-134,  figs  1-203. 
Heron-Allen,  E.  &  Earland,  E.  1913.  The  Foraminifera  of  the  Clare  Island  District,  County  Mayo,  Ireland. 

(Clare  Island  Survey,  part  64.)  Proc.  R.  Ir.  Acad.,  Dublin,  31: 1-188,  pis  1-13. 
Lacroix,  E.  1930.  Les  Lituolides  de  plateau  continental  mediterranean  entre  Saint-Raphael  et  Monaco.  Bull. 

Inst.  oceanogr.  Monaco  549:  547-550,  figs  1-21. 
Loeblich,  A.  R.  Jr  &  Tappan,  H.  1952.  Adercotryma,  a  new  Recent  foraminiferal  genus  from  the  Arctic.  /. 

Wash.  Acad.  Sci.,  42:  141-142,  figs  1-4. 
& 1964.  Protista  2.  Sarcodina  chiefly  'Thecamoebians'  and  Foraminiferida.  In  Moore,  R.  C.  (ed.) 

Treatise  on  Invertebrate  Paleontology,  Part  C,  1:  1-510,  figs  1-399.  University  of  Kansas  Press. 
Maync,  W.  1952.  Critical  taxonomic  study  and  nomenclatural  revision  of  the  Lituolidae  based  upon  the 

prototype  of  the  family,  Lituola  nautiloidea  Lamarck,  1804.  Contrib.  Cushman  Lab.  foramin.  Res.,  Sharon, 

3: 35-56,  pis  9-12. 

Millett,  F.  W.  1908.  The  Recent  Foraminifera  ofGalway.  8  pp,  4  pis.  W.  Brendon  &  Son,  Plymouth. 
Saidova,  Kh.  M.  1961.  Foraminiferal  ecology  and  palaeogeography  of  the  far  eastern  seas  of  the  USSR  and 

northwestern  part  of  the  Pacific  Ocean.  232  pp,  31  pis.  Akad.  Nauk  SSSR,  Inst.  Okeanologii,  Moskow.  (In 

Russian.) 
1975.  Benthonic  foraminifera  of  the  Pacific  Ocean.  3  vols.  875  pp,  116  pis.  Akad.  Nauk  SSSR,  Inst. 

Okeanol.  P.P.  Shirshova,  Moskow.  (In  Russian.) 
Schwager,  C.  1877.  Quadro  del  proposto  sistema  di  classificazione  dei  foraminiferi  con  guscio.  Boll.  R.  com. 

geol.  Ital.,  Florence,  8: 18-27,  pi.  1. 
Wiesner,  H.  1931.  Die  Foraminiferen  der  Deutschen  Siidpolar-Expedition  1901-1903.  Dt.  Sudpol.-Exped., 

Berlin  (Zool.),  20: 49-165,  pis  1-24. 

Manuscript  accepted  for  publication  15  January  1986 


Hermit  crabs  associated  with  the  bryozoan 
Hippoporidra  in  British  waters 

J.  D.  D.  Bishop 

Departments  of  Palaeontology  and  Zoology,  British  Museum  (Natural  History),  Cromwell  Road, 
London  SW7  5BD 

The  cheilostome  bryozoan  Hippoporidra  lusitania  is  not  associated  exclusively  with  the 
hermit  crab  Pagurus  cuanensis  as  some  previous  reports  have  implied.  It  has  been  found 
with  two  other  species  of  pagurid,  Anapagurus  chiroacanthus  and  Cestopagurus  timidus;  its 
occurrence  with  P.  cuanensis  requires  confirmation. 

Species  of  the  cheilostome  bryozoan  genus  Hippoporidra  preferentially  or  exclusively  encrust 
gastropod  shells  inhabited  by  hermit  crabs  (Crustacea:  Anomura:  Paguroidea).  Helicospiral 
growth  of  an  established  bryozoan  colony  may  extend  the  crab's  domicile  well  beyond  the  aperture 
of  the  original  gastropod  shell  (Taylor  &  Cook,  198 1).  The  type-species  of  the  genus,  Hippoporidra 
edax  (Busk),  was  first  described  as  a  fossil  from  the  Coralline  Crag  (Pliocene)  of  eastern  England, 
but  the  name  has  subsequently  been  used  for  living  specimens  from  both  sides  of  the  North 
Atlantic.  However,  material  from  Recent  British  seas  that  had  formerly  been  referred  to  H.  edax 
was  distinguished  as  a  new  species,  H.  lusitania,  by  Taylor  &  Cook  (1981). 

No  details  were  given  by  Taylor  &  Cook  (1981)  of  the  hermit  crabs  with  which  H.  lusitania 
occurs.  A  few  earlier  records  of  Recent  Hippoporidra  edax  from  Britain,  which  may  be  assumed  to 
refer  to  H.  lusitania,  mentioned  Pagurus  cuanensis  Bell  as  the  associated  pagurid.  Thus,  Moore 
(1937)  reported  a  single  colony  of  the  bryozoan  with  P.  cuanensis  collected  off  the  Isle  of  Man.  This 
record  was  repeated  by  Bruce  et  al.  (1963)  and  quoted  by  Cook  (1964).  Eggleston  (1972)  reported 
that  in  Manx  waters  Hippoporidra  was  in  fact  restricted  to  P.  cuanensis,  and  this  apparent  example 
of  extreme  stenotopy  in  a  bryozoan  was  quoted  by  Ryland  (1976).  Hay  ward  &  Ryland  (1979)  gave 
P.  cuanensis  as  the  preferred  species  of  British  Hippoporidra,  and  did  not  name  any  other  pagurid 
with  which  the  bryozoan  was  found. 

The  Bryozoa  collection  of  the  Zoology  Department  of  the  British  Museum  (Natural  History) 
contains  21  colonies  of//,  lusitania,  including  the  type  series  from  Guernsey  and  part  of  the  Manx 
material  studied  by  Eggleston.  The  associated  hermit  crab  is  present  in  only  four  examples,  as 
detailed  in  Table  1 . 

It  is  clear  from  this  that  H.  lusitania  is  not  restricted  to  Pagurus  cuanensis,  even  off  the  Isle  of 
Man.  Indeed,  its  occurrence  with  P.  cuanensis  at  all  requires  confirmation.  P.  cuanensis  reaches  a 
considerably  larger  size  than  either  Anapagurus  chiroacanthus  (Lilljeborg)  or  Cestopagurus  timidus 


Table  1 

Specimen 

H 

W    Hermit  crab 

Isle  of  Man 

D.  Eggleston 

1963.12.30.1 

11-0 

11-5  Anapagurus  chiroacanthus 

Guernsey 

A.  M.  Norman 

1911.  10.1.  1143H 

12-0 

10-0  Anapagurus  chiroacanthus 

(Paratype) 

Guernsey 

A.  M.  Norman 

1911.10.1.11431 

8-5 

8-5  Anapagurus  sp.  (fragment) 

(Paratype) 

Scilly  Isles 

M.  H.  Thurston 

1965.8.18.26 

7-5 

4-5  Cestopagurus  timidus 

H  =  approximate  height  of  gastropod/bryozoan  measured  along  axis,  in  mm. 
W  =  approximate  'body  whorl'  diameter  of  gastropod/bryozoan,  in  mm. 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  52(1):  29-30 


Issued  29  January  1987 


30  J.  D.  D.  BISHOP 

(Roux),  the  carapace  lengths  given  by  Bouvier  (1940)  being  8-12  mm,  4—6  mm  and  4-5  mm  respec- 
tively. All  H.  lusitania  colonies  available  for  study  at  the  BM(NH)  are  relatively  small;  the  largest 
(Plymouth,  T.  Hincks,  1 899.5. 1.1517)  has  a  height  of  c.  17  mm  and  a  'body  whorl'  diameter  of  c. 
14  mm.  It  therefore  seems  probable  from  the  limited  material  available  that  Hippoporidra  lusitania 
may  be  most  commonly  associated  with  relatively  small  hermit  crabs. 


Acknowledgements 

I  wish  to  thank  R.  W.  Ingle  and  P.  L.  Cook  for  help  with  the  pagurids  and  bryozoans  respectively. 

References 

Bouvier,  E.-L.  1940.  Decapodes  marcheurs.  Faune  de  France  37: 1^04. 

Bruce,  J.  R.,  Colman,  J.  S.  &  Jones,  N.  S.  1963.  Marine  fauna  of  the  Isle  of  Man  and  its  surrounding  seas. 

L.M.B.C.  Memoirs  on  Typical  British  Marine  Plants  and  Animals  36:  i-ix  and  1-307. 
Cook,  P.  L.  1964.  Polyzoa  from  West  Africa.  Notes  on  the  genera  Hippoporina  Neviani,  Hippoporella  Canu, 

Cleidochasma  Harmer  and  Hippoporidra  Canu  &  Bassler  (Cheilostomata,  Ascophora).  Bulletin  of  the 

British  Museum  (Natural  History),  Zoology  series  12:  1-35. 
Eggleston,  D.  1972.  Factors  influencing  the  distribution  of  sub-littoral  ectoprocts  off  the  south  of  the  Isle  of 

Man  (Irish  Sea).  Journal  of  Natural  History  6:  247-260. 
Hay  ward,  P.  J.  &  Ryland,  J.  S.  1979.  British  ascophoran  bryozoans.  Synopses  of  the  British  Fauna  (New 

Series)  14:  i-v  and  1-312. 
Moore,  H.  B.  1 937.  Marine  Fauna  of  the  Isle  of  Man.  Proceedings  and  Transactions  of  the  Liverpool  Biological 

Society  50:  1-293. 

Ryland,  J.  S.  1976.  Physiology  and  ecology  of  marine  bryozoans.  Advances  in  Marine  Biology  14: 285-443. 
Taylor,  P.  D.  &  Cook,  P.  L.  1981.  Hippoporidra  edax  (Busk  1859)  and  a  revision  of  some  fossil  and  living 

Hippoporidra  (Bryozoa).  Bulletin  of  the  British  Museum  (Natural  History) ,  Geology  series  35:  243-25 1 . 

Manuscript  submitted  for  publication  6  September  1985 


The  first  zoea  of  three  Pachygrapsus  species  and  of 
Cataleptodius  floridanm  (Gibbes)  from  Bermuda  and 
Mediterranean  (Crustacea:  Decapoda:  Brachyura) 

R.  W.  Ingle 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7  5BD 

Introduction 

Affinities  of  larvae  belonging  to  the  family  Grapsidae  have  been  reviewed  by  Aikawa  (1929),  Wear 
(1970),  Rice  (1980)  and  Wilson  (1980).  Within  the  four  subfamilies  composing  this  family  'the 
larval  development  of  less  than  15%  of  all  the  species  has  been  described'  (Wilson,  1980:  756). 
Many  descriptions  are  insufficient  for  meaningful  comparative  studies  and,  because  of  the 
apparent  difficulty  of  rearing  these  small  zoeae,  a  large  percentage  of  studies  describe  only  the  first 
stage. 

Wilson  (1980)  has  provided  a  useful  and  comprehensive  table  of  seven  comparative  features  of 
the  first  stage  zoeae  of  47  grapsid  species.  To  this  list  may  be  added  the  following  accounts  which 
contain  more  or  less  adequate  details  for  comparative  purposes.  Plagusiinae:  IPlagusia  depressa, 
Rice  &  Williamson,  1977.  Varuninae:  Eriocheir  japonica,  Gaetice  depressus,  Hemigrapsus  longi- 
tarsus,  H .  penicellatus ,  H.  sanguiensis ,  Terada,  1981.  Grapsinae:  Metopograpsus  latifrons,  Kakati, 
1982;  M.  messor,  Rajabai,  1962.  Sesarminae:  Aratus  pisonii,  Hartnoll,  1965;  Chasmagnathus 
convexus,  Saba,  1974;  C.  laevis,  Helograpsushaswellianus,  Green  &  Anderson,  1973;  Metasesarma 
rousseauxi,  Rajabai,  1962;  Sesarma  erythrodactyla,  Green  &  Anderson,  1973;  S.  perracae,  Soh 
Chen  Lam,  1969;  S.  tetragonum,  Rajabai,  1962. 

Within  the  genus  Pachygrapsus  larval  stages  are  known  for  only  three  of  the  fifteen  or  so 
accepted  species  (viz.  P.  marmoratus,  P.  transversus,  P.  crassipes).  Of  these,  the  complete  develop- 
ment has  been  described  for  P.  marmoratus  (Fabricius)  and  P.  crassipes  Randall.  Larval  descrip- 
tions of  P.  marmoratus  are  based,  except  for  the  first  stage,  upon  plankton  collected  material  (see 
Cano,  1892;  Hyman,  1924;  Bourdillon-Casanova,  1960),  but  some  of  Cano's  figures,  also  repro- 
duced by  Hyman,  may  not  even  be  of  a  Pachygrapsus  (see  p.  000).  Laboratory  hatched  first  stage 
zoea  of  P.  transversus  was  described  superficially  by  Lebour  (1944)  and  Rossignol  (1957)  identified 
a  plankton  caught  megalopa  to  this  species.  Villalobos  (1971)  described  the  first  zoeal  stage  of 
P.  crassipes.  This  species  was  laboratory  reared  to  fifth  zoeal  stage  by  Schlotterbeck  (1976)  and  a 
plankton  caught  megalopa  was  tentatively  assigned  to  P.  crassipes  by  Rathbun  (1923). 

During  1973  first  stage  zoeae  of  Pachygrapsus  marmoratus  were  obtained  from  a  laboratory  held 
crab  collected  by  R.  B.  Manning  off  the  coast  of  Tunisia  and  in  1983  the  first  stage  zoeae  of  P. 
gracilis  and  of  P.  transversus  were  hatched  from  crabs  held  by  the  author  in  the  Biological  Station, 
Bermuda.  Although  the  larvae  of  these  species  were  not  reared  beyond  the  first  zoeal  stage  it  would 
seem  desirable  to  give  an  account  of  this  material  to  supplement  meagre  larval  information  at 
present  available  on  this  genus  and  also  to  compare  (see  Table)  the  first  stages  of  these  four 
Pachygrapsus  species.  Opportunity  is  also  taken  to  describe  the  first  stage  zoea  of  the  xanthid 
Cataleptodius  floridanus  (also  hatched  at  Bermuda),  the  larvae  of  which  were  studied  by  Kurata 
(1970)  but  whose  account  was  never  published  (see  Martin  1984:  233,  footnote). 


Materials  and  methods 

The  first  zoea  of  P.  marmoratus  was  hatched  from  a  crab  collected  in  the  canal  leading  from 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  52(1),  31^*1  Issued  29  January  1987 


R.  W.  INGLE 


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FEATURE 

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Carapace,  posterio- 
margins 

Antennule,  aesthete 

Antenna,  spinous  p 

investment 

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Abdomen,  fourth  s< 

Telson,  lateral  spini 

'Schlotterbeck  tabula 
H960)  gives  1-10  mm 
^Lebour  shows  these 
margins  (see  Fig.  3h  i 
specimens  examined. 

FIRST  ZOEA  33 

southern  Punic  Port,  Salammbo,  Tunisia,  18.7.1973.  Ovigerous  Pachygrapsus  gracilis  and  Cata- 
leptodiusfloridanus  were  collected  at  various  localities  from  the  intertidal  mud  flats  at  Ferry  Reach 
and  P.  transversus  from  beneath  stones  at  Whalebone  Bay,  Bermuda,  all  in  September/October 
1983.  The  eggs  hatched  within  2-3  days  of  the  crab's  confinement  in  aerated  aquaria  water  held  at 
20-24°C  and  the  larvae  were  fed  newly  hatched  Anemia  nauplii.  Live  zoeae  were  subsequently 
transported  to  a  rearing  laboratory  at  the  British  Museum  (Natural  History)  in  London  but  none 
survived  to  the  second  stage. 

Measurements  given  are:  T.T.  =  distance  between  tips  of  dorsal  and  rostral  spines;  C.L.= 
carapace  length  from  between  the  eyes  to  the  posterio-lateral  margin.  The  material  has  been 
incorporated  in  the  Collections  of  the  British  Museum  (Natural  History),  accession  number:  1985: 
463^68. 


Descriptions 

Family  GRAPSIDAE  MacLeay,  1838 
Subfamily  GRAPSINAE  MacLeay,  1838 

Pachygrapsus  gracilis  (de  Saussure,  1858) 

Dimensions:  T.T.  0-60-0-70  mm.  C.L.  0-35-0-38  mm. 

Carapace  (Fig.  la):  dorsal  spine  short  and  straight,  stout  proximally,  slightly  more  than  one  third  carapace 

length;  rostral  spine  of  moderate  length  and  stout;  dorso-median  elevation  prominent;  at  least  four  pairs  of 

anterio-median  setules  and  a  pair  of  posterio-median  setules  present;  posterio-lateral  margins  of  carapace 

narrowly  rounded  with  3-4  obtuse  denticles  and  lateral  microscopic  setules  (inset  to  fig.). 

Eyes:  partly  fused  to  carapace. 

Antennule  (Fig.  Ib):  unsegmented  with  four  aesthetascs/setae. 

Antenna  (Fig.  Ic):  spinous  process  as  long  as  rostral  spine  and  with  many  subacute  denticles;  exopod  very 

small,  about  one  ninth  of  spinous  process  length. 

Maxillule  (Fig.  Id):  endopod  2-segmented,  proximal  segment  with  one  seta,  distal  with  one  subterminal  and 

four  terminal  setae;  basial  endite  with  five  spines,  coxal  with  six  spines/setae. 

Maxilla  (Fig.  le):  endopod  stepped  distally,  outer  lobe  broader  than  inner  each  with  two  long  setae;  basial 

endite  incipiently  bilobed  distally,  each  with  four  setae;  coxal  endite  bilobed  distally,  outer  lobe  with  four  and 

inner  with  five  setae  respectively,  seta  on  apex  of  outer  lobe  very  short  almost  a  spine;  scaphognathite  with 

four  plumose  setae  and  a  stout  posterior  process. 

First  maxilliped (Fig.  If):  basis  with  eight  setae  arranged  in  pairs;  endopod  five-segmented  with  1, 2, 1, 2, 4+ 1 

setae;  exopod  incipiently  two-segmented  and  with  four  terminal  plumose  setae. 

Second  maxilliped  (Fig.  Ig):  basis  with  four  setae;  endopod  three-segmented  with  0,  1,  4+1  setae;  exopod 

incipiently  two-segmented  with  four  terminal  plumose  setae. 

Third  maxilliped  and  pereiopods:  not  developed. 

Abdomen  (Fig.  Ih):  composed  of  five  segments  and  a  telson,  somewhat  dorso-ventrally  compressed,  surfaces 

with  microscopic  spinules;  second  and  third  segments  with  a  pair  of  broad  dorso-lateral  processes;  fourth 

segment  laterally  expanded  and  with  a  pair  of  obtuse  lateral  processes  placed  at  a  lower  level  than  the  ones  on 

preceding  segments;  first  segment,  posterio-lateral  margins  truncate,  those  of  other  segments  obtuse  and  of 

third  to  fifth  with  a  very  minute  denticle;  second  to  fifth  segments  each  with  a  pair  of  setules  near  posterio- 

dorsal  margin.  Telson  somewhat  narrowed,  furcae  not  noticeably  directed  outwards,  each  with  numerous 

microscopic  spinules  and  two  very  small  lateral  spinules;  posterior  margin  with  six  equal  plumose  setae. 

Pachygrapsus  transversus  (Gibbes,  1850) 
Pachygrapsus  transversus:  Lebour,  1944:  115,  fig.  5  (zoea  I);  Rossignol,  1957:  89,  fig.  5  (megal.). 

Dimensions:  T.T.  0-60-0-70  mm.  C.L.  0-30-0-35  mm.  Differs  from  P.  gracilis  in  the  following  features. 
Carapace  (Fig.  2a):  dorsal  spine  longer  and  proximally  slightly  stouter,  more  than  one  third  of  carapace 
length;   rostral   spine  noticeably  stouter  proximally;  posterio-lateral  margin  of  carapace  with  very 
inconspicuous  denticles  and  with  microscopic  setules;  only  two  pairs  of  anterio-median  setules  apparent. 
Antenna  (Fig.  2c):  spinous  process  with  numerous  subacute  denticles  developed  distally  as  stout  spine-like 
processes;  exopod  about  one  seventh  of  spinous  process  length. 
Maxillule  (Fig.  2d):  spines/setae  slightly  stouter. 
Maxilla  (Fig.  2e):  basial  endite  noticeably  bilobed  distally. 


34 


R.  W.  INGLE 


Fig.  1.  Pachygrapsus  gracilis  (de  Saussure).  First  zoea.  a,  carapace,  right  lateral  aspect;  b,  antennule;  c, 
antenna;  d,  maxillule;  e,  maxilla;  f,  first  maxilliped;  g,  second  maxilliped;  h,  abdomen  and  telson, 
dorsal  aspect.  Scale  =  0-05  mm. 


FIRST  ZOEA 


35 


Fig.  2.  Pachygrapsus  transversus  (Gibbes).  First  zoea.  a,  carapace,  right  lateral  aspect;  b,  antennule;  c, 
antenna;  d,  maxillule;  e,  maxilla;  f,  first  maxilliped;  g,  second  maxilliped;  h,  abdomen  and  telson, 
dorsal  aspect.  Scale  =  0-05  mm. 


36  R.  W.  INGLE 

Abdomen  (Fig.  2h):  slightly  larger,  posterio-lateral  margins  of  fourth  segment  subacute  and  of  the  other 
segments  (except  first)  more  produced,  denticles  larger.  Telson  slightly  broader,  the  more  posterior  of  the  two 
lateral  spinules  larger;  furcae  slightly  shorter  and  stouter. 

Pachygrapsus  marmoratus  (Fabricius,  1787) 

Pachygrapsus  marmoratus:  Cano,  1892:  S.Tav.III,  figs  IB  (?zoea  III),  ?  1C,  IE,  IF,  ?2c,  2e-f,  ?3c,  4e-f,  5e-f, 
6e-f,7e-f,8e-f  (labelled  as  6  in  fig.),  12e,  13e,  14e,  1 5e(?zoea  IV,  megal.);  Williamson,  1915: 518,  figs  403-^05, 
407-8  (figs  after  Cano);  Hyman,  1924:  2,  PI.  3,  figs  22,  ?23, 25, 26,  ?33,  36a-b,  41, 42, 44, 45, 48, 49,  50,  52,  53, 
54,  56  (figs  after  Cano):  Bourdillon-Casanova,  1960:  188,  fig.  61  (zoea  I);  Paula,  1985:  142,  fig.  3  (zoea  I). 

Dimensions:  T.T.  0-75  mm.  C.L.  0-35  mm. 

Differs  from  P.  gracilis  and  P.  transversus  as  follows. 

Carapace  (Fig.  3a):  dorsal  spine  longer,  exceeding  half  carapace  length;  posterio-lateral  margin  of  carapace 

without  setules,  denticles  very  minute;  anterio-median  setules  not  apparent. 

Antenna  (Fig.  3c):  denticles  on  spinous  process  small  and  numerous  throughout  length  of  process. 

Maxillule  (Fig.  3d):  setules  on  spines/setae  very  long. 

Abdomen  (Fig.  3h):  fourth  segment  not  laterally  expanded  and  without  a  pair  of  obtuse  lateral  processes; 

posterio-lateral  margins  of  third  to  fifth  segments  each  with  a  conspicuous  denticle.  Telson  lateral  spinules  on 

furcae  absent. 

Family  XANTHIDAE  MacLeay,  1838 
Cataleptodiusfloridanus  (Gibbes,  1850) 

Dimensions:  T.T.  1-1  mm.  C.L.  0-43  mm. 

Carapace  (Fig.  4a):  dorsal  spine  long,  distally  curved,  proximally  stout;  rostral  spine  almost  as  long  as  dorsal 

spine  and  with  1  or  2  small  spinules;  lateral  spines  small;  dorso-median  elevation  hardly  developed;  no 

anterio-median  setules  apparent,  a  small  pair  of  posterio-median  setules  present;  posterio-lateral  margin  of 

carapace  with  one  or  two  small  setules. 

Eyes:  partly  fused  to  carapace. 

Antennule  (Fig.  4b):  unsegmented  and  with  four  aesthetascs/setae. 

Antenna  (Fig.  4c):  spinous  process  as  long  as  rostral  spine,  distally  with  many  long  acute  spines;  exopod  small, 

between  one  sixth  and  one  seventh  of  spinous  process  length  and  with  two  small  distal  setules. 

Maxillule  (Fig.  4d):  endopod  two-segmented,  proximal  segment  with  one  distal  seta,  distal  segment  with  five 

setae  (two  subdistal  and  three  distal);  basial  endite  with  five  spines/setae;  coxal  endite  with  seven  setae. 

Maxilla  (Fig.  4e):  endopod  two-lobed,  outer  slightly  stepped,  broader  than  inner  and  with  2  +  2  setae,  inner 

lobe  with  three  setae;  basial  endite  two-lobed,  outer  prominent  and  with  four  setae,  inner  with  five  setae;  coxal 

endite  two-lobed  each  with  four  setae;  scaphognathite  with  four  plumose  setae  and  a  stout  posterior  process. 

First  maxilliped  (Fig.  40:  basis  with  ten  setae  arranged  2,  2,  3,  3,  respectively;  endopod  five-segmented,  with 

3, 2, 1, 2, 4+ 1  setae  respectively;  exopod  incipiently  two-segmented  with  four  terminal  plumose  setae. 

Second maxilliped '(Fig.  4g):  basis  with  four  setae;  endopod  three-segmented,  with  1,1,4+1  setae  respectively; 

exopod  incipiently  two  segmented,  with  four  terminal  plumose  setae. 

Third  maxilliped  and  pereiopods:  not  developed. 

Abdomen  (Fig.  4h):  composed  of  five  segments  and  a  telson;  second  segment  with  a  pair  of  subacute  laterally 

directed  dorso-lateral  processes,  third  segment  with  a  pair  of  small  acute  posteriorly  directed  dorso-lateral 

processes;  posterio-lateral  margins  of  second  segment  acute,  those  of  third  to  fifth  segments  extended  into 

acute  processes;  posterio-dorsal  surface  of  second  to  fifth  segments  each  with  a  small  pair  of  setules  near 

margin;  posterior  margins  of  segments  four  and  five  with  minute  denticles.  Telson  furcae  diverging  slightly, 

each  with  one  long  prominent  dorsal  spine  and  one  long  and  one  smaller  lateral  spine,  furcae  with  minute 

denticles;  posteror  margin  of  telson  with  six  long  setae. 


Remarks 

As  mentioned  earlier,  some  of  the  stages  described  and  figured  by  Cano  (1892)  as  Pachygrapsus 
marmoratus  may  not  belong  to  this  species.  His  figure  depicting  a  first  stage  zoea  (Tav.III,  Fig.  1  A) 
does  not  show  a  dorso-lateral  process  on  the  third  segment  of  the  abdomen  characteristic  of 
Pachygrapsus  zoeae  and  obvious  in  the  present  laboratory  reared  material.  Cano's  figure  IB  clearly 
shows  this  lateral  process  and  although  this  larva  is  depicted  with  four  maxillipedal  exopod  setae  it 


FIRST  ZOEA 


37 


Fig.  3.  Pachygrapsus  marmoratus  (Fabricius).  First  zoea.  a,  carapace,  right  lateral  aspect;  b,  antennule; 
c,  antenna;  d,  maxillule;  e,  maxilla;  f,  first  maxilliped;  g,  second  maxilliped;  h,  abdomen  and  telson, 
dorsal  aspect.  Scale  =  0-05  mm. 


38 


R.  W.  INGLE 


Fig.  4.  Cataleptodius  floridanus  (Gibbes).  First  zoea.  a,  carapace,  left  lateral  aspect;  b,  antennule;  c, 
antenna;  d,  maxillule;  e,  maxilla;  f,  first  maxilliped;  g,  second  maxilliped;  h,  abdomen  and  telson  dorsal 
aspect.  Scale  =  0-05  mm. 


FIRST  ZOEA  39 

is  probably  of  a  later  stage  because  he  figures  incipient  pereiopods  beneath  the  carapace.  Hyman 
(1924)  has  suggested  that  this  is  a  stage  three  zoea  and  was  also  convinced  that  Cano  had  over- 
looked a  fourth  stage  and  that  the  third  stage  described  by  Cano  (purporting  to  be  the  last) 
represented  the  fifth  and  terminal  zoeal  stage  of  this  species.  Costlow  &  Bookhout  (1962)  however, 
maintained  that . . .  'While  some  of  Cano's  (1891)  figures  may  bear  some  slight  inaccuracies,  it  is 
quite  possible  that  the  sequence  and  number  of  larval  stages  which  he  figures  is  correct. . .'  These 
views  were  expressed  in  the  context  of  their  study  of  the  larval  development  ofSesarma  reticulatum 
in  which  there  are  only  three  zoeal  stages  and  it  is  probable  that  P.  marmoratus  passes  through  five 
stages  similar  to  P.  crassipes.  Cano  (1892)  also  assigned  two  megalopal  forms  to  P.  marmoratus. 
The  one  illustrated  in  his  fig.  ID  and  bearing  an  acute  rostral  projection  appears  to  be  of  an 
oxyrhynch  as  it  lacks  dactylar  subterminal  setae  on  the  fifth  pereiopods  characteristic  of  brachy- 
rhynch  megalopas.  Bourdillon-Casanova  (1960)  described  the  first  stage  zoea  of  P.  marmoratus 
from  laboratory  hatched  material  and  illustrated  (Fig.  61)  prominent  denticles  on  the  carapace 
posterio-lateral  margin  as  well  as  two  conspicuous  lateral  setae  (also  shown  by  Paula,  1985,  Fig. 
3,i)  on  each  furca  of  the  telson.  These  two  features  could  not  be  detected  in  specimens  examined 
during  the  present  study.  Lebour's  (1944)  figure  of  the  first  zoea  of  P.  transversus,  also  obtained 
from  Bermudan  laboratory  hatched  crabs,  differs  from  the  present  specimens  in  apparently  having 
two  distal  setae  on  the  antennal  spinous  process,  a  pair  of  conspicuous  curved,  acute  dorso-lateral 
processes  on  the  fourth  abdominal  segment  and  two  equally  developed  lateral  spinules  on  the 
telson  furca.  Her  zoeae  were  also  larger  than  the  present  ones. 

Mid-dorsal  carapace  setules  have  never  been  mentioned  previously  as  occurring  in  Pachy- 
grapsus  zoeae,  and  Gore  &  Scotto  (1982:  518)  suggested  they  may  be  absent  in  grapsinid  zoeae. 
However  in  the  present  study  these  setules  were  found  in  first  stage  zoeae  of  P.  gracilis  and  P. 
transversus  but  were  not  apparent  in  P.  marmoratus.  They  are  somewhat  difficult  to  resolve 
satisfactorily,  even  with  the  aid  of  interference  contrast,  but  appear  to  be  less  numerous  on  the 
anterio-median  region  of  P.  transversus  than  of  P.  gracilis.  In  many  brachyuran  larvae  these  setules 
do  not  appear  until  later  stages. 

Martin  (1984: 232-233)  has  provided  an  excellent  key  to  the  known  xanthid  zoeae  of  the  Western 
Atlantic  and  Gulf  of  Mexico.  The  present  study  confirms  his  assessment  of  the  larvae  of  this  species 
as  belonging  to  his  group  I  of  the  xanthidae  and  they  can  be  assigned  to  C.  floridanus  on  the 
following  combined  features:  (1)  lateral  processes. of  third  to  fifth  abdominal  segments  not 
extending  posteriorly  beyond  half  length  of  following  segment,  (2)  more  than  twenty  spinules  on 
spinous  process  of  antenna,  (3)  abdominal  dorso-lateral  processes  confined  to  second  and  third 
segments,  (4)  telson  with  three  spines  on  each  furca,  (5)  lateral  carapace  spines  present,  (6)  basial 
segment  of  first  maxilliped  endopod  with  three  setae,  (7)  antennal  exopod  very  reduced. 


Acknowledgements 

The  visit  to  the  Bermuda  Biological  Station  was  partly  sponsored  by  an  Exxon  Corporation  Fellowship.  I 
thank  Dr  Wolfgang  Sterrer,  Director  of  the  Biological  Station  and  his  staff  for  assistance  during  my  visit 
and  also  Dr  R.  B.  Manning  for  kindly  presenting  the  P.  marmoratus  zoeae  to  the  BM(NH).  I  also  thank 
Dr  A.  L.  Rice  for  reading  the  manuscript. 


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Rathbun,  M.  J.  1923.  The  Brachyuran  crabs  collected  by  the  U.S.  Fisheries  Steamer  'Albatross'  in  1911, 

chiefly  on  the  West  Coast  of  Mexico.  Bulletin  of  the  American  Museum  of  Natural  History.  New  York. 

48(10):  6 19-637. 
Rice,  A.  L.  1980.  Crab  zoeal  morphology  and  its  bearing  on  the  classification  of  the  Brachyura.  Transactions 

of  the  Zoological  Society  of  London  35: 271—424. 
Rice,  A.  L.  &  Williamson,  D.  1. 1977.  Planktonic  stages  of  Crustacea  Malacostraca  from  Atlantic  Seamounts. 

Meteor  Forschungsergebnisse.  Berlin  &  Stuttgart.  D26: 28-64. 
Rossignol,  M.  1957.  II.  Crustaces  Decapodes  Marines  de  la  Region  de  Pointe-Noire.  In:  Collignon,  J., 

Rossignol,  M.  &  Roux,  Ch.  Mollusques,  Crustaces,  Poissons  Marines  des  cotes  d'A.E.F.  en  collection  au 

Centre  d'Oceanographie  de  1'Institut  d'Etudes  Centrafricaines  de  Pointe-Noire.  Ministere  de  la  France 

d'Outre-Mer.  Paris,  pages  71-136. 
Saba,  M.  1974.  On  the  larval  development  of  Chasmagnathus  convexus  de  Haan  (Grapsinae).  Researches  on 

Crustacea.  The  Carcinological  Society  of  Japan.  Tokyo.  6: 71-85. 
Saussure,  H.  de  1858.  Memoire  sur  divers  Crustaces  nouveaux  des  Antilles  et  du  Mexique.  Memoires  de  la 

Societe  de  Physique  et  d'Histoire  Naturelle  de  Geneve  14: 41 7-496. 
Schlotterbeck,  R.  E.  1976.  The  larval  development  of  the  Lined  Shore  Crab,  Pachygrapsus  crassipes  Randall, 

1 840  (Decapoda,  Brachyurea,  Grapsidae)  reared  in  the  laboratory.  Crustaceana.  International  Journal  of 

Crustacean  Research.  Leiden.  30(2):  184-200. 
Soh  Chen  Lam  1969.  Abbreviated  development  of  a  non-marine  crab  Sesarma  (Geosesarma)  perracae 

(Brachyura:  Grapsidae)  from  Singapore.  Journal  of  Zoology.  Proceedings  of  the  Zoological  Society  of 

London  158:  357-370. 
Terada,  M.  1981.  Zoea  larvae  of  five  crabs  in  the  subfamily  Varuninae.  Researches  on  Crustacea.  The 

Carcinological  Society  of  Japan.  Tokyo.  11:  66-76. 


FIRST  ZOEA  41 

Wear,  R.  G.  1970.  Life-history  studies  on  New  Zealand  Brachyura  4.  Zoea  larvae  hatched  from  crabs  of  the 
family  Grapsidae.  New  Zealand  Journal  of  Marine  and  Freshwater  Research.  Wellington.  4(1):  3-35. 

Williamson,  H.  C.  1915.  VI.  Crustacea  Decapoda.  Larven.  Nordisches  Plankton.  Keil.  (6)  18:  315-588. 

Wilson,  K.  A.  1980.  Studies  on  Decapod  Crustacea  from  the  Indian  River  region  of  Florida.  XV.  The 
larval  development  under  laboratory  conditions  of  Euchirograpsus  americanus  A.  Milne  Edwards,  1880 
(Crustacea  Decapoda:  Grapsidae)  with  notes  on  grapsid  subfamilial  larval  characters.  Bulletin  of  Marine 
Science.  Coral  Gables.  30(4):  756-775. 

Villalobos,  C.  R.  1 97 1 .  First  zoeal  stage  of  Pachygrapsus  crassipes  Randall.  Revista  de  Biologica  Tropical.  San 
Jose,  Costa  Rica.  18(1  &  2):  107-1 13. 

Manuscript  accepted  for  publication  30  October  1985 


A  classification  of  the  phylum  Sipuncula 

Peter  E.  Gibbs 

Marine  Biological  Association  of  the  U.K.,  Plymouth,  Devon  PL1  2PB,  U.K. 

Edward  B.  Cutler 

Division  of  Science  and  Mathematics,  Utica  College  of  Syracuse  University,  Utica,  New  York 
13502,  U.S.A. 

Synopsis 

A  classification  of  the  phylum  Sipuncula  is  adopted  following  the  analysis  of  Cutler  &  Gibbs  (1985)  and 
comprises  two  classes,  four  orders  and  six  families.  This  replaces  the  earlier  classification  of  Stephen  & 
Edmonds  (1972)  which  was  based  on  four  families  only.  The  diagnostic  characters  are  reviewed.  Seventeen 
genera  are  redefined,  one  new  subgenus  is  described  and  twelve  other  subgenera  are  recognised. 

Introduction 

The  classification  of  the  phylum  Sipuncula  has  had  a  confused  history.  Early  attempts  to  define 
higher  taxa  by  grouping  genera  were,  to  a  large  extent,  thwarted  by  incomplete,  imprecise  or 
erroneous  descriptions  of  many  species.  Stephen  &  Edmonds  (1972)  classified  the  phylum  into 
four  families  in  providing  the  first  compilation  of  species  described  prior  to  about  1970.  How- 
ever, this  monograph  is  essentially  literature-based  and  consequently  many  errors  are  repeated; 
nevertheless,  it  provides  a  useful  base-line  to  the  present  revision. 

The  need  for  greater  precision  in  defining  genera  has  led  the  authors  to  re-examine  most  of 
the  available  type  specimens.  The  definitions  of  genera  presented  below  incorporate  both  novel 
observations  and  corrections  to  earlier  descriptions.  Where  possible,  nine  basic  characters  have 
been  checked  for  each  species  before  assigning  it  to  a  genus.  These  characters  are  summarised  for 
each  genus  in  Table  1 .  A  phylogenetic  interpretation  of  the  classification  used  here  will  be  found  in 
Cutler  &  Gibbs  (1985). 


Diagnostic  features  of  higher  taxa 

In  reviewing  the  diagnostic  characters  of  the  phylum,  particular  attention  has  been  paid  to  the 
structure  of  the  oral  disk  since  the  arrangement  of  the  tentacles  provides  a  useful  basis  for  dividing 
the  phylum  into  two  classes  -  Sipunculidea  and  Phascolosomatidea.  Certain  descriptions  of 
tentacle  arrangements  are  misleading  or  in  error  (see  for  example  Stephen  &  Edmonds  (1972)  p.  16 
and  Table  3).  No  doubt  these  errors  result  from  the  fact  that  some  species  are  not  amenable  to 
fixation  in  the  extended  state;  species  with  long  introverts  are  notoriously  difficult  to  preserve  with 
their  introvert  fully  extended.  Although  dissection  of  the  introvert  is  possible  the  details  of  the 
tentacular  arrangement  on  a  withdrawn  disk  are  often  difficult  to  interpret  and  have  yet  to  be 
satisfactorily  determined  in  some  small-sized  species  (e.g.  Apionsoma  trichocephala  Sluiter).  The 
following  summarises  the  distinctions  of  the  two  classes. 

The  sipunculan  tentacular  crown  exhibits  many  diverse  forms  but,  basically,  two  tentacular 
patterns  can  be  recognised.  In  one,  that  of  the  proposed  class  Sipunculidea,  the  tentacles  are 
arranged  peripherally  on  the  oral  disk  so  as  to  encircle  the  centrally-placed  mouth;  dorsally  this 
circle  is  inflected  to  form  an  arc  enclosing  the  nuchal  organ,  a  feature  well  developed  in  Thysano- 
cardia  spp  for  example  (see  Gibbs,  Cutler  &  Cutler,  1983,  Fig.  2).  In  the  other,  that  of  the  proposed 
class  Phascolosomatidea,  the  tentacles  are  restricted  to  a  dorsal  arc  enclosing  the  nuchal  organ  and 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  52(1):  43-58  Issued  29  January  1987 


44 


P.  E.  GIBBS  &  E.  B.  CUTLER 


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Fig.  1.  The  structure  of  the  tentacular  crown  of  Sipuncula:  some  examples  illustrating  the  form  and 
variation  within  the  classes  Sipunculidea  and  Phascolosomatidea.  Solid  lines  indicate  possible  evolu- 
tionary trends.  A.  Generalised  Sipunculidea  crown  such  as  might  have  been  possessed  by  ancestral 
stock  adults;  B.  Golfingia  margaritacea',  C.  Nephasoma  rimicola;  D.  Onchnesoma  squamatum;  E. 
Nephasoma  minutum;  F.  Sipunculus  norvegicus',  G.  Themis te  lageniformis;  H.  Phascolosoma 
granulatum;  J.  Antillesoma  antillarum;  K.  Aspidosiphon  johnstoni.  (B,  E,  F:  after  Theel,  1905.) 
Abbreviations:  cc,  cephalic  collar;  m,  mouth;  n,  nuchal  organ;  na,  dorsal  arc  of  tentacles  enclosing 
nuchal  organ;  1,2,3,  primary,  secondary  and  tertiary  tentacle  pairs  around  disk  periphery.  (Modified 
from  Cutler  &  Gibbs,  1985.) 


there  are  no  peripheral  tentacles  (Fig.  1).  Thus,  the  two  patterns  have  a  common,  perhaps  homolo- 
gous, feature  in  the  dorsal  arc  of  tentacles.  In  evolutionary  terms,  the  peripheral  tentacles  could  be 
interpreted  as  a  later  addition,  i.e.  the  Sipunculidea  have  evolved  from  a  Phascolosomatidea  stock. 
However,  around  the  margin  of  the  oral  disk  in  Phascolosomatidea  there  is  a  prominent  ridge,  the 
cephalic  collar,  and  it  is  thought  that  this  ridge  represents  a  vestige  of  the  peripheral  system.  If 
this  interpretation  is  correct  the  common  ancestor  must  have  possessed  a  Sipunculidea-type  of 
tentacular  crown,  probably  a  simple  form,  somewhat  similar  to  that  of  Thysanocardia  procera 
(Gibbs,  Cutler  &  Cutler,  1983,  Fig.  2B),  and  the  peripheral  tentacles  were  lost  during  an  early 
divergence  to  give  the  Phascolosomatidea  line.  The  Sipunculidea  tentacular  pattern,  peripheral 


46  P.  E.  GIBBS  &  E.  B.  CUTLER 

circle  plus  nuchal  arc,  can  perhaps  be  best  regarded  as  an  elaboration  of  simple  prostomial 
tentacles  possessed  by  the  early  protostomial  stock. 

The  development  of  the  tentacular  crown  in  the  Sipunculidea,  as  seen  for  example  in  Golfingia 
species,  commences  with  the  formation  of  four  primary  pairs  of  tentacles  in  the  dorsal,  ventral 
and  lateral  positions,  between  which  secondary  pairs  subsequently  develop  to  form  a  single  ring 
encircling  the  central  mouth  on  the  oral  disk  (Fig.  1  A).  Between  these  pairs  tertiary  pairs  usually 
develop:  in  the  adult  these  may  be  few  or  very  numerous;  in  the  latter  case  the  tentacles  are 
accommodated  in  loops  or  'festoons'  that  extend  aborally  on  to  the  anterior  introvert.  The  nuchal 
organ  situated  dorsally  between  the  two  primary  tentacles  thus  becomes  enclosed  by  an  arc  of 
tentacles.  As  a  general  rule,  the  number  of  tentacles  increases  with  increasing  size  and  age  of 
individuals  and  large-sized  species  have  more  tentacles  than  small-sized  species. 

Within  the  class  Sipunculidea  a  wide  range  of  tentacular  development  is  found.  The  most  highly 
evolved  crown  is  found  in  the  genus  Thysanocardia,  adult  specimens  of  which  often  possess 
well-developed  festoons  comprising  several  hundred  tentacles;  in  some  Thysanocardia  nigra 
(Ikeda)  the  number  exceeds  500  (see  Gibbs,  Cutler  &  Cutler,  1983).  Large  Golfingia  margaritacea 
(Sars)  have  100  or  more  tentacles  (Fig.  IB)  but  most  other  Sipunculidea  have  around  50  or  fewer 
with  only  a  limited  number  of  tertiary  tentacles  developing,  as  in  Golfingia  elongata  (Keferstein) 
with  20-34  and  Nephasoma  rimicola  (Gibbs)  with  12-20  (Fig.  1C;  see  Gibbs,  1973).  In  some  species 
only  the  primary  tentacles  appear,  as  in  Onchnesoma  squamatum  (Kor.  &  Dan.)  with  8  and 
Nephasoma  minutum  (Keferstein)  with  just  two  (Fig.  ID,  E).  Thus  the  evolution  of  the  tentacular 
crown  could  have  been  not  only  towards  greater  complexity  but  also  towards  simplification,  a 
trend,  possibly  neotenous,  seen  in  several  genera,  notably  Nephasoma  (e.g.  N.  minutum),  Onchne- 
soma (e.g.  O.  steenstrupi  Kor.  &  Dan.)  and  Phascolion  (e.g.  P.  pacificum  Murina).  Another  vari- 
ation is  seen  in  some  members  of  the  family  Sipunculidae  where  the  peripheral  tentacles  have 
become  flattened  and  fused  to  form  a  continuous  veil-like  structure,  as  for  example  in  Sipunculus 
norvegicus  Dan.  (Fig.  IF).  The  crown  of  Themiste  with  its  tentacles  arising  from  4-8  stems  appears 
anomalous  at  first  sight  but,  in  fact,  this  type  represents  yet  another  modification  of  the  basic 
Sipunculidea  pattern.  In  themistids  the  secondary  tentacles  develop  between  the  primary  pairs  but 
are  borne  on  outgrowths  of  the  oral  disk  so  that  with  subsequent  tertiary  tentacle  development, 
an  erect  dendritic  structure  results  rather  than  the  typical  festoon  which  is  contiguous  with  the 
introvert  wall.  In  the  themistid  type  the  dorsal  primary  tentacles  are  widely  spaced  and  do  not 
enclose  the  nuchal  organ  (Fig.  1G). 

All  six  genera  grouped  in  the  class  Phascolosomatidea  are  rather  similar  in  terms  of  the  tentacu- 
lar arrangement:  with  one  exception,  all  have  a  single  arc  of  up  to  30  tentacles  enclosing  the  nuchal 
organ  (Fig.  1H,K).  The  exception  is  Antillesoma  antillarum  (Grube  &  Oersted)  in  which  the 
tentacles  are  fairly  numerous  (Fig.  1 J)  presumably  as  a  result  of  secondary  proliferation. 

One  other  character  that  separates  the  Sipunculidea  and  Phascolosomidea  is  the  structure  of  the 
introvert  hooks  on  the  anterior  introvert,  when  present.  In  the  former  group  these  hooks  are 
somewhat  variable  but  generally  are  simple,  sharply-pointed  protrusions  of  the  epidermis  and 
scattered  in  their  distribution;  however,  in  the  latter  they  have  a  typical  recurved  shape,  usually  an 
internal  structure  is  apparent  and  they  are  closely-packed  in  distinct  rings  encircling  the  anterior 
introvert. 

Definitions  of  orders,  families  and  genera  are  given  below.  Four  orders  are  recognised.  In  the 
class  Sipunculidea,  members  of  the  order  Sipunculiformes  are  distinguished  by  the  presence  of 
banding  in  the  longitudinal  muscle  of  the  body  wall  found  in  five  genera,  all  of  which  are  placed  in 
the  family  Sipunculidae.  The  remaining  six  genera  within  this  class  all  have  a  uniform,  continuous 
layer  of  longitudinal  muscle  tissue  and  form  the  order  Golfingiiformes  comprising  three  families  — 
Golfingiidae,  Phascolionidae  and  Themistidae.  In  the  class  Phascolosomatidea  the  genera  are 
separated  into  two  orders,  each  with  a  single  family,  on  the  basis  of  the  presence  (Aspidosiphoni- 
formes:  Aspidosiphonidae)  or  absence  (Phascolosomatiformes:  Phascolosomatidae)  of  an  anal 
shield,  a  hardened  thickening  of  the  anterior  trunk  region.  It  should  be  noted  that  the  structure  of 
the  anal  shield  is  different  in  all  three  genera  within  the  Aspidosiphonidae  and  it  is  recognised  that 
this  character  may  have  evolved  several  times. 

Whilst  the  forms  of  the  tentacle  crown  and  of  the  introvert  hooks  are  useful  characters  for 


SIPUNCULA  47 

dividing  the  1 7  genera  into  two  natural  groups,  here  designated  as  classes,  few  other  major  charac- 
ters are  confined  to  one  or  other  of  these  two  classes  (Table  1):  coelomic  spaces  in  the  body  wall  is  a 
feature  exclusive  to  Sipunculidae  and  likewise  for  anal  shield  development  in  Aspidosiphonidae. 
Other  characters  are  found  in  both  classes,  notably  the  banding  of  the  longitudinal  muscle  layer  in 
the  body  wall,  the  attachment  of  the  spindle  muscle  to  the  posterior  trunk  and  an  increase  in  the 
volume  of  the  contractile  vessel  through  the  development  of  villi  in  conjunction  with  increased 
tentacular  volume  or  area.  Such  characters  would  appear  to  be  polyphyletic  in  origin.  There  is  little 
doubt  that  the  basic  number  of  introvert  retractor  muscles  is  four,  arranged  as  dorsal  and  ventral 
pairs.  Loss  of  the  dorsal  pair  appears  to  have  occurred  independently  in  a  number  of  generic  lines. 
Assessing  the  number  of  retractors  in  any  one  specimen  can  often  be  problematical  because  fusion 
frequently  occurs  but  may  not  be  evident.  In  some  species  only  one  retractor  is  apparent  in  the 
adult  form:  in  Phascolion  species  there  is  good  evidence  to  suggest  the  one  muscle  is  the  result  of  the 
fusion  of  all  four  muscles  (Gibbs,  1985)  whilst  in  Onchnesoma  the  single  muscle  is  thought  to 
comprise  only  the  fused  ventral  pair,  the  dorsal  pair  having  been  lost.  Use  of  the  number  of 
retractors  as  a  taxonomic  character  has  to  be  approached  with  some  caution  since  even  within  a 
single  population  the  number  is  liable  to  variation,  as  noted  for  Golfingia  elongata  (Gibbs,  1973). 
Morphological  variation  seems  to  be  one  of  the  hallmarks  of  the  phylum,  a  feature  that  may 
account  for  the  survival  of  this  small  group  but  one  that  does  not  facilitate  good  taxonomy. 

The  present  scheme  of  classification  (Table  2)  updates  that  given  in  Stephen  &  Edmonds  (1972) 
and  some  later  authors  by  incorporating  the  recent  revisions  of  several  major  genera,  notably, 
Siphonosoma,  Golfingia  and  Phascolosoma.  Synonymies  are  as  given  in  Stephen  &  Edmonds 
(1972):  any  more  recent  changes  are  noted  under  each  genus. 


Key  to  Families 

1  Tentacles  arranged  in  an  arc  encircling  dorsal  nuchal  organ;  peripheral  tentacles  absent;  hooks 

complex,  in  distinct  rings  [Class  PHASCOLOSOMA  TIDEA] 

Tentacles  arranged  peripherally  on  oral  disk  so  as  to  encircle  central  mouth;  may  be  borne 
on  stem-like  outgrowths  of  oral  disk  or  reduced  in  number  to  a  single  dorsal  pair;  hooks  simple, 
usually  scattered  [Class  SIPUNCULIDEA] 

2  Anal  shield  present Fam.  ASPIDOSIPHONIDAE  (p.  55) 

Anal  shield  absent Fam.  PHASCOLOSOMATIDAE(p.  54) 

3  Longitudinal  muscles  of  body  wall  gathered  into  separate  or  anastomosing  bands 

Fam.  SIPUNCULIDAE  (p.  48) 

Longitudinal  muscle  of  body  wall  in  a  uniform  continuous  layer 

4  Tentacles  carried  on  4-8  stem-like  outgrowths  of  oral  disk.        .    Fam.  THEMISTIDAE(p.  53) 
Tentacles  not  carried  on  disk  outgrowths 

5  A  single  nephridium  present Fam.  PHASCOLIONIDAE(p.  51) 

Two  nephridia  present Fam.  GOLFINGIIDAE(p.  50) 


Classification 

Phylum  SIPUNCULA 
Class  SIPUNCULIDEA 

Sipuncula  with  tentacles  encircling  a  central  mouth  on  the  oral  disk.  Introvert  hooks  (when 
present)  simple,  thorn-like  hollow  structures  that  are  usually  irregularly  distributed.  Spindle 
muscle  unattached  posteriorly  (except  in  Siphonosoma  and  Siphonomecus). 

Order  SIPUNCULIFORMES 

Sipunculidea  with  longitudinal  muscle  in  body  wall  gathered  into  bands  (likewise  for  circular 
muscle  in  two  genera — Sipunculus  and  Xenosiphon).  Coelomic  extensions  -  canals  or  sacs  -  in  body 
wall  (except  in  Phascolopsis). 


48  P.  E.  GIBBS  &  E.  B.  CUTLER 

Table  2.  Classification  of  the  phylum  Sipuncula 

Cl.  Sipunculidea 
Ord.  SIPUNCULIFORMES 

Fam.  SIPUNCULIDAE  Stephen  &  Edmonds,  1972 
Sipunculus  Linnaeus,  1 766 
S.  (Sipunculus) 

S.  ( Austrosiphon )  Fisher,  1954 
Xenosiphon  Fisher,  1947 
Siphonosoma  Spengel,  1912 
Siphonomecus  Fisher,  1 947 
Phascolopsis  Fisher,  1950 
Ord.  GOLFINGIIFORMES 

Fam.  GOLFINGIIDAE  Stephen  &  Edmonds,  1972 
Golfingia  Lankester,  1885 
Nephasoma  Pergament,  1946 
Thysanocardia  Fisher,  1950 
Fam.  PHASCOLIONIDAE  Cutler  &  Gibbs,  1985 
PhascolionTheel  1875 
P.  (Phascolion) 

P.  (Isomya)  Cutler  &  Cutler,  1985 
P.  (Montuga)  Gibbs,  1985 
P.  (Lesenka)  Gibbs,  1985 
P.  (Villiophora)  Cutler  &  Cutler,  1985 
Onchnesoma  Koren  &  Danielssen,  1875 
Fam.  THEMISTIDAE  Cutler  &  Gibbs,  1985 
ThemisteGray,  1828 
T.  (Themiste) 

T.  (Lagenopsis)  Edmonds,  1980 
Cl.  Phascolosomatidea 
Ord.  PHASCOLOSOMATIFORMES 

Fam.  PHASCOLOSOMATIDAE  Stephen  &  Edmonds,  1972 
Phascolosoma  Leuckart,  1 828 
P.  (Phascolosoma) 
P.  ( Edmondsius )  subgen.  nov. 
Apionsoma  Sluiter,  1902 
Antillesoma  Stephen  &  Edmonds,  1972 
Ord.  ASPIDOSIPHONIFORMES 
Fam.  ASPIDOSIPHONIDAE  Baird,  1 868 
Aspidosiphon  Diesing,  1851 
A.  (Aspidosiphon) 
A.  (Paraspidosiphon)  Stephen,  1964 
Cloeosiphon  Grube,  1 868 
Lithacrosiphon  Shipley,  1902 


Family  SIPUNCULIDAE  Baird,  1868 
Characters  are  those  of  the  order. 

Key  to  Genera 

1  Body  wall  circular  muscle  layer  continuous PHASCOLOPSIS 

Body  wall  circular  muscle  layer  gathered  into  bands 2 

2  Body  wall  circular  and  longitudinal  muscle  bands  anastomosing,  spindle  muscle  attached  to 

posterior  end  of  trunk 3 

Body  wall  circular  and  longitudinal  muscle  bands  not  anastomosing,  spindle  muscle  not  attached 
to  posterior  trunk 4 

3  Four  introvert  retractor  muscles SIPHONOSOMA 

Two  introvert  retractor  muscles  .  SIPHONOMECUS 


SIPUNCULA  49 

4      Gut  with  post-oesophageal  loop;  coelom  extends  into  body  wall  as  longitudinal  canals  running 
throughout  most  of  trunk  length   .........  SIPUNCULUS 

Gut  without  post-oesophageal  loop;  coelom  extends  into  body  wall  as  short  diagonal  canals 
running  across  the  width  of  one  circular  muscle  band      ....         XENOSIPHON 

Genus  SIPUNCULUS  Linnaeus,  1766 

DIAGNOSIS.  Introvert  much  shorter  than  trunk,  without  hooks,  covered  with  scattered  sub- 
triangular  papillae.  Trunk  cylindrical.  Body  wall  contains  coelomic  extensions  in  the  form  of 
parallel  longitudinal  canals  which  extend  most  of  trunk  length.  Circular  and  longitudinal  muscle 
layers  gathered  into  distinct  bands.  Oral  disk  carries  tentacles  arranged  around  the  mouth  some- 
times modified  with  the  development  of  an  inter-tentacular  membrane  (S.  nudus  L.,  S.  norvegicus 
Danielssen).  Four  introvert  retractor  muscles.  Two  protractor  muscles  may  be  developed  (S. 
mundanus  Sel.  &  Billow).  Two  contractile  vessels,  both  without  villi.  Gut  with  post-oesophageal 
loop,  caecum  on  rectum,  and  coil  attached  to  body  wall  along  its  entire  length  by  many  connective 
strands.  Spindle  muscle  not  attached  posteriorly.  Two  nephridia.  Species  usually  large-sized  (trunk 
greater  than  5  cm  long  in  adults). 

TYPE  SPECIES.  Sipunculus  nudus  Linnaeus,  1766,  subsequent  designation,  ?Fisher  1952. 


Subgenus  S/Pt/yVCt/LjyS  Linnaeus,  1766 
Sipunculus  (Sipunculus):  Cutler  &  Cutler,  1985a:  232. 

DIAGNOSIS.  Nephridia  anterior  to  anus.  Spindle  muscle  originates  on  body  wall  anterior  to  anus. 
TYPE  SPECIES.  Sipunculus  nudus  Linnaeus,  1766,  subsequent  designation,  ?Fisher,  1952. 

Subgenus  AUSTROSIPHON  Fisher,  1954,  emended 

Xenosiphon  (Austrosiphon)  Fisher,  1954:  314. 
Xenosiphon  (Xenopsis)  Johnson,  1969:  44. 
Sipunculus  (Contraporus)  Cutler  &  Cutler,  1985a:  241. 

DIAGNOSIS.  Nephridia  posterior  to  anus.  Spindle  muscle  originates  from  ventral  surface  of  rectum. 
TYPE  SPECIES.  Sipunculus  mundanus  Selenka  &  Biilow,  1883,  monotypy. 

Genus  XENOSIPHON  Fisher,  1947 

DIAGNOSIS.  Introvert  much  shorter  than  trunk,  and  without  hooks  but  covered  with  scattered 
subtriangular  papillae.  Body  wall  contains  coelomic  extensions  in  form  of  short,  diagonal  canals 
limited  in  length  to  width  of  one  circular  muscle  band.  Circular  and  longitudinal  muscle  layers 
divided  into  distinct  bands.  Oral  disk  carries  tentacles  arranged  around  mouth.  Four  introvert 
retractor  muscles  and  two  thin  protractor  muscles  present.  Contractile  vessel  without  villi,  gut 
without  post-oesophageal  loop,  caecum  present  on  rectum  and  coil  attached  to  body  wall  along 
entire  length  by  connective  strands.  Spindle  muscle  originates  on  ventral  wall  of  rectum  and  is  not 
attached  to  the  body  wall  posteriorly.  Anus  anterior  to  nephridiopores.  Two  nephridia.  Contains 
one  large-sized  species. 

TYPE  SPECIES.  Sipunculus  mundanus  var.  branchiatus  Fischer,  1895,  original  designation. 

REMARKS.  The  two  subgenera  previously  included  in  this  genus  are  now  assigned  to  Sipunculus  (see 
above). 

Genus  SIPHONOSOMA  Spengel,  1912 

Siphonosoma  (Siphonosoma):  Fisher,  1950b:  805. 
Siphonosoma  (Hesperosiphon)  Fisher,  1950b:  805. 
Siphonosoma  (Dasmosiphon)  Fisher,  1950b:  805. 

DIAGNOSIS.  Introvert  much  shorter  than  the  trunk  with  prominent  conical  papillae  (sometimes  also 
hooks)  arranged  in  rings.  Body  wall  with  coelomic  sac-like  extensions;  circular  and  longitudinal 


50  P.  E.  GIBBS  &  E.  B.  CUTLER 

muscle  layers  gathered  into  anastomosing  bands.  Oral  disk  carries  tentacles  arranged  around  the 
mouth.  Four  introvert  retractor  muscles.  Contractile  vessel  with  or  without  villi.  Spindle  muscle 
attached  posteriorly.  Two  nephridia.  Species  usually  large-sized  (trunk  greater  than  5  cm  long  in 
adults). 

TYPE  SPECIES.  Phascolosoma  australe  Keferstein,  1865,  subsequent  designation,  Gerould,  1913. 

REMARKS.  The  three  subgenera  recognised  by  Fisher  (1950b)  were  distinguished  by  the  presence  or 
absence  of  transverse  dissepiments  and  rectal  caeca.  These  characters  have  been  found  to  be 
subject  to  great  variation  and  of  limited  diagnostic  value:  consequently,  this  subgeneric  separation 
is  not  supportable  (Cutler  &  Cutler,  1982). 

Genus  SIPHONOMECUS  Fisher,  1947 

DIAGNOSIS.  Introvert  much  shorter  than  trunk  with  prominent  hooks  and  conical  papillae 
arranged  in  rings.  Body  wall  with  coelomic  extensions  (sacs);  circular  and  longitudinal  muscle 
layers  gathered  into  anastomosing  bands.  Oral  disk  carries  tentacles  arranged  around  the  mouth. 
Two  introvert  retractor  muscles.  Contractile  vessel  without  villi.  Spindle  muscle  attached 
posteriorly.  Two  nephridia.  Contains  one  large-sized  species. 

TYPE  SPECIES.  Siphonomecus  multicinctus  Fisher,  1947,  original  designation. 

Genus  PHASCOLOPSIS  Fisher,  1950 

DIAGNOSIS.  Introvert  shorter  than  trunk  with  deciduous  hooks  (present  in  juvenile  but  lost  in 
adult).  Body  wall  without  coelomic  extensions.  Circular  muscle  layer  continuous,  longitudinal 
muscle  layer  gathered  into  anastomosing  bands.  Oral  disk  carries  tentacles  arranged  around  the 
mouth.  Four  introvert  retractor  muscles.  Contractile  vessel  without  villi.  Spindle  muscle  not 
attached  posteriorly.  Two  nephridia.  Contains  one  large-sized  species. 

TYPE  SPECIES.  Sipunculus  gouldii  Portales,  1851,  monotypy. 

Order  GOLFINGIIFORMES 

Sipunculidea  with  body  wall  longitudinal  muscle  in  a  continuous  layer,  not  gathered  in  bands. 

Family  GOLFINGIIDAE  Stephen  &  Edmonds,  1972 
Golfingiiformes  with  two  nephridia.  Tentacles  not  borne  on  stem-like  extensions  of  oral  disk. 

Key  to  Genera 

1  Contractile  vessel  with  numerous  villi THYSANOCARD1A 

Contractile  vessel  without  villi 2 

2  Four  introvert  retractor  muscles GOLFINGIA 

Two  introvert  retractor  muscles NEPHASOMA 

Genus  GOLFINGIA  Lankester,  1885 

Golfingia  (Golfingia):  Fisher,  1950a;  549. 
Golfingia  (Dushana)  Murina,  1975:  1085. 
Themis te  (Stephensonum)  Edmonds,  1980:  33. 
Centrosiphon  Shipley,  1903:  173. 

DIAGNOSIS.  Introvert  about  equal  to  or  shorter  than  trunk;  hooks  when  present  are  usually  scat- 
tered (arranged  in  rings  in  G.  elongatd).  Body  wall  with  continuous  muscle  layers.  Oral  disk  carries 
tentacles  arranged  around  the  mouth.  Four  introvert  retractor  muscles.  Contractile  vessel  without 
villi.  Spindle  muscle  not  attached  posteriorly.  Two  nephridia.  Species  small-  to  large-sized. 

TYPE  SPECIES.  Golfingia  macintoshii  Lankester,  1885  [  =  Sipunculus  vulgaris  de  Blainville,  1827: 
Stephen,  1934],  monotypy. 


SIPUNCULA  5 1 

REMARKS.  This  genus  now  contains  only  those  species  previously  assigned  to  the  nominate  sub- 
genus  Golfingia  (Golfingia).  It  includes  Centrosiphon  Shipley,  1903:  Edmonds  (1980)  placed  the 
type  species  C.  herdmani  Shipley  within  the  genus  Golfingia;  the  Centrosiphon  specimens  recorded 
by  Cutler  &  Cutler  (1979)  are  now  considered  to  be  aberrant  Aspidosiphon. 

The  subgenus  G.  (Dushana)  Murina,  1975,  was  characterised  by  complete  or  partial  fusion  of 
the  dorsal  and  ventral  retractor  muscles  on  one  side  of  the  body.  However,  it  is  known  that  such 
fusion  of  the  retractors,  and  also  reduction  of  the  retractor  number  through  loss  of  one  or  both 
dorsal  retractors,  are  features  of  some  Golfingia  species,  for  example  G.  elongata  (see  Gibbs,  1973). 
The  holotype  of  G.  (Dushana)  adriatica  Murina  shows  a  similar  retractor  arrangement  (Murina, 
1975,  Fig.  1)  to  that  described  by  Watier  (1932)  for  aberrant  G.  vulgaris.  The  type  species  originally 
designated  for  G.  (Dushana),  G.  scutiger  (Roule),  does  not  differ  significantly  in  its  retractor 
arrangement  (Roule,  1906,  Fig.  95)  from  typical  Golfingia  species.  Thus  G.  (Dushana)  is  no  longer 
recognised. 

Genus  NEPHASOMA  Pergament,  1946 
Golfingia  ( Phascoloides )  Fisher,  1950a:  550. 

DIAGNOSIS.  Introvert  about  equal  to,  or  shorter  than,  trunk.  Hooks  when  present  usually  scattered 
(arranged  in  rings  in  N.  rimicola  (Gibbs),  in  spirals  in  TV.  abyssorum  (Kor.  &  Dan.)).  Body  wall  with 
continuous  muscle  layers.  Oral  disk  carries  tentacles  arranged  around  the  mouth  but  tentacles  may 
be  reduced  in  both  size  and  number  and  restricted  to  dorsal  region.  Two  introvert  retractor  muscles 
often  partially  fused.  Contractile  vessel  without  villi.  Spindle  muscle  not  attached  posteriorly.  Two 
nephridia.  Species  generally  small-  to  medium-sized  (trunk  less  than  5  cm  in  length). 

TYPE  SPECIES.  Nephasoma  marinki  Pergament,  1946  [  =  Onchnesoma  glaciale  Danielssen  &  Koren: 
Cutler  &  Murina,  1977;  =  Phascolosoma  HlljeborgiiDsimdssen  &  Koren:  Gibbs,  1982],  monotypy. 

REMARKS.  This  genus  now  contains  all  those  species  previously  assigned  to  the  Golfingia  subgenus 
Phascoloides  Fisher,  1950,  since  Nephasoma  Pergament  has  been  shown  to  have  priority  over 
Phascoloides  (Cutler  &  Murina,  1977). 

Genus  THYSANOCARDIA  Fisher,  1950 

DIAGNOSIS.  Introvert  longer  than  trunk,  without  hooks.  Body  wall  with  continuous  muscle  layers. 
Oral  disk  carries  tentacles  arranged  around  the  mouth;  those  enclosing  nuchal  organ  are  well 
developed.  Two  introvert  retractor  muscles.  Contractile  vessel  with  distinct  villi.  Spindle  muscle 
not  attached  posteriorly.  Two  nephridia.  Species  small-  to  medium-sized  (adults  generally  under 
5  cm  in  trunk  length). 

TYPE  SPECIES.  Phascolosoma  procerum  Mobius,  1875,  original  designation. 

REMARKS.  The  subgenus  Golfingia  (Thysanocardia)  was  recently  elevated  to  generic  rank  and  the 
number  of  species  reduced  to  three  by  Gibbs,  Cutler  &  Cutler  (1983). 

Family  PHASCOLIONIDAE  Cutler  &  Gibbs,  1985 

Golfingiiformes  with  one  nephridium  (usually  the  right).  Tentacles  not  borne  on  stem-like 
extensions  of  oral  disk.  Gut  coil  without  well-defined  axial  spindle  muscle. 

Key  to  Genera 

1       Anus  usually  situated  on  anterior  trunk;  epidermal  'holdfast'  or  'attachment'  papillae  often 
present.  Retractor  muscles  highly  fused  but  usually  2-4  roots  apparent  at  base  of  column 

PHASCOL1ON 

Anus  situated  on  distal  half  of  introvert;  epidermal  'attachment'  papillae  absent.  Retractor 
muscle(s)  appear  as  single  column  without  separate  roots        .        .        .      ONCHNESOMA 


52  P.  E.  GIBBS  &  E.  B.  CUTLER 

Genus  PHASCOLION  Thcel,  1875 

DIAGNOSIS.  Introvert  length  one-half  to  four  times  that  of  trunk  length,  with  or  without  hooks. 
Trunk  usually  with  modified  'holdfast'  papillae.  Body  wall  with  continuous  muscle  layers.  Oral 
disk  carries  tentacles  arranged  around  the  mouth.  Introvert  retractor  muscle  system  modified  by 
fusion  of  dorsal  and  ventral  pairs:  relative  size  and  degree  of  fusion  defines  subgenera  (see  below). 
Contractile  vessel  without  villi  (but  present  in  P.  cirratum).  Gut  coiling  generally  loose  and  without 
axial  spindle  muscle.  One  nephridium  (usually  right).  Species  small-  to  medium-sized  (less  than 
5  cm  in  length)  generally  inhabiting  mollusc  shells. 

TYPE  SPECIES.  Sipunculus  strombus  Montagu,  1804,  monotypy. 

Subgenus  PHASCOLION  Theel,  1875 
Phascolion  (Phascolion):  Gibbs,  1985:  314. 

DIAGNOSIS.  Retractor  column  divided  for  most  of  its  length:  oesophagus  detaches  from  retractor 
column  at  a  point  posterior  to  the  first  separation  of  the  retractor  muscles.  Dorsal  retractor(s) 
much  more  strongly  developed  than  ventral  retractor(s).  Contractile  vessel  without  villi. 

TYPE  SPECIES.  Sipunculus  strombus  Montagu,  1804,  monotypy. 

Subgenus  ISOMYA  Cutler  &  Cutler,  1985 
Phascolion  (Isomya)  Cutler  &  Cutler  1985A:  820 

DIAGNOSIS.  Characters  as  for  P.  (Phascolion)  except  that  dorsal  and  ventral  retractor  muscles  are 
about  equal  in  diameter. 

TYPE  SPECIES.  Phascolion  tuberculosum  Theel,  1875,  original  designation. 

Subgenus  MONTUGA  Gibbs,  1985 
Phascolion  (Montuga)  Gibbs,  1985:  315. 

DIAGNOSIS.  Retractor  column  divided  only  at  posterior  end:  oesophagus  detaches  from  retractor 
column  at  a  point  anterior  to  the  first  separation  of  the  retractor  muscles.  Contractile  vessel 
without  villi. 

TYPE  SPECIES.  Phascolion  Intense  Selenka,  1885,  original  designation. 

Subgenus  LESENKA  Gibbs,  1985 
Phascolion  (Lesenka)  Gibbs,  1985:  315. 

DIAGNOSIS.  Retractor  column  entire  with  retractor  muscles  fused  throughout  whole  length. 
Contractile  vessel  without  villi. 

TYPE  SPECIES.  Phascolion  cryptum  Hendrix,  1975,  original  designation. 

Subgenus  V1LLIOPHORA  Cutler  &  Cutler,  1985 
Phascolion  (Villiophora)  Cutler  &  Cutler,  19856:  821. 

DIAGNOSIS.  Retractor  column  entire  with  retractor  muscles  fused  throughout  whole  length. 
Contractile  vessel  with  numerous  villi. 

TYPE  SPECIES.  Phascolion  cirratum  Murina,  1968,  monotypy. 

Genus  ONCHNESOMA  Koren  &  Danielssen,  1875 

DIAGNOSIS.  Introvert  much  longer  than  trunk.  Body  wall  with  continuous  muscle  layers.  Oral  disk 
carries  tentacles  arranged  around  mouth  but  tentacles  may  be  highly  reduced  in  size.  Introvert 
retractor  muscle  system  modified  by  fusion  to  form  single  retractor  muscle.  Anus  situated  on 


SIPUNCULA  53 

introvert.  Contractile  vessel  rarely  apparent  and  without  villi.  Spindle  muscle  absent.  One 
nephridium  (right).  Species  small-sized  (trunk  less  than  1  cm  in  length). 

TYPE  SPECIES.  Onchnesoma  steenstrupii  Koren  &  Danielssen,  1875,  subsequent  designation, 
Stephen  &  Edmonds,  1972. 

Family  THEMISTIDAE  Cutler  &  Gibbs,  1985 
Golfingiiformes  with  two  nephridia.  Tentacles  borne  on  stem-like  extensions  of  oral  disk. 

Genus  THEMISTE  Gray,  1828 

DIAGNOSIS.  Introvert  less  than  trunk  length.  Body  wall  with  continuous  muscle  layers.  Oral  disk 
carries  tentacles  basically  surrounding  mouth  but  extending  with  growth  along  margins  of  stem- 
like  outgrowths  of  the  oral  disk.  With  or  without  hooks.  Two  introvert  retractor  muscles.  Contrac- 
tile vessel  with  villi.  Spindle  muscle  not  attached  posteriorly.  Two  nephridia.  Species  small-  to 
large-sized. 

TYPE  SPECIES.  Themiste  hennahi  Gray,  1 824,  monotypy . 

Subgenus  THEMISTE  Gray,  1828 
Themiste  (Themiste):  Edmonds,  1980:  33. 
DIAGNOSIS.  Contractile  vessel  with  long,  thread-like  villi. 
TYPE  SPECIES.  Themiste  hennahi  Gray,  1828,  monotypy. 

Subgenus  LAGENOPSIS  Edmonds,  1980 
Themiste  (Lagenopsis)  Edmonds,  1980:  33. 
DIAGNOSIS.  Contractile  vessel  with  short,  digitiform  villi. 
TYPE  SPECIES.  Themiste  lageniformis  Baird,  1868,  original  designation. 

REMARKS.  The  subgenus  T.  (Stephensonum)  Edmonds,  1980,  was  erected  to  include  two  species  of 
Themiste  having  four,  not  two,  retractor  muscles,  namely,  Themiste  stephensoni  (the  type  species, 
original  designation)  and  T.  pinnifolia.  The  type  material  of  Themiste  stephensoni  (Stephen) 
(described  under  the  name  Dendrostomum  Grube,  a  junior  synomym)  in  the  RSME  collections  has 
been  examined.  The  holotype  (1958.23.24)  has  a  golfingiid,  not  themistid,  tentacle  crown  (as  shown 
by  dissection  of  the  introvert)  and  the  'band  of  very  short  villi'  on  the  contractile  vessel  (Stephen, 
1 942,  p.  252)  do  not  appear  to  be  true  villi  but  rather  outpoutchings  of  a  relatively  voluminous 
vessel.  The  specimen  is  clearly  a  Golfingia  and  probably  G.  capensis  (Teuscher);  the  other  type 
specimens  comprise  further  Golfingia  but  also  include  some  Themiste  all  of  which  have  the  typical 
number  of  retractors  (two).  Thus  T.  (Stephensonum)  becomes  a  junior  synonym  of  Golfingia.  The 
species  Themiste  pinnifolia  (Keferstein)  is  based  on  a  single  specimen,  collected  more  than  100  years 
ago,  which  cannot  be  traced.  No  subsequent  record  appears  in  the  literature,  despite  extensive 
collecting  in  the  area  of  the  type  locality  (St  Thomas,  West  Indies).  Since  the  generic  identity  of  this 
specimen  is  in  doubt,  the  species  name  pinnifolia  is  regarded  as  a  nomen  dubium. 

Class  PHASCOLOSOMATIDEA 

Sipuncula  with  tentacles  confined  to  an  arc  enclosing  dorsal  nuchal  organ:  peripheral  tentacles 
absent.  Introvert  hooks  recurved,  usually  with  an  internal  structure  and  closely-packed  in 
regularly-spaced  rings  (absent  in  Antillesoma}.  Spindle  muscle  attached  posteriorly. 

Order  PHASCOLOSOMATIFORMES 

Phascolosomatidea  with  anterior  trunk  not  modified  to  form  anal  shield.  Four  introvert  retractor 
muscles. 


54  P.  E.  GIBBS  &  E.  B.  CUTLER 

Family  PHASCOLOSOMATIDAE  Stephen  &  Edmonds,  1972 
Characters  are  those  of  the  order. 

Key  to  Genera 

1  Introvert  hooks  absent.  Contractile  vessel  with  villi ANTILLESOMA 

Introvert  hooks  present.  Contractile  vessel  without  villi 2 

2  Longitudinal  muscle  in  body  wall  gathered  into  bands        ....  PHASCOLOSOMA 
Longitudinal  muscle  in  body  wall  a  uniform  continuous  layer    .        .        .        .  AP1ONSOMA 

Genus  PHASCOLOSOMA  Leuckart,  1828 

DIAGNOSIS.  Introvert  variable  in  length,  often  equal  to  trunk  with  numerous  rings  of  recurved 
hooks  (absent  in  P.  meteori  Herubel).  Body  wall  with  longitudinal  muscle  layer  gathered  into 
bands.  Oral  disk  carries  relatively  few  tentacles  (less  than  30)  enclosing  nuchal  organ.  Contractile 
vessel  without  true  villi  (may  have  bulbous  vesicles).  Four  introvert  retractor  muscles;  lateral  pairs 
sometimes  partially,  rarely  completely,  fused.  Spindle  muscle  attached  posteriorly  (except  in 
P.pectinatwri).  Two  nephridia. 

TYPE  SPECIES.  Phascolosoma  granulatum  Leuckart,  1828,  monotypy. 

Subgenus  PHASCOLOSOMA  Leuckart,  1828 

Phascolosoma  (Phascolosoma):  Stephen  &  Edmonds,  1972:  289. 
? Phascolosoma  (Rueppellisoma)  Stephen  &  Edmonds,  1972:  271. 
?Phascolosoma  (Satonus)  Stephen  &  Edmonds,  1972:  28  (in  part). 

DIAGNOSIS.  Spindle  muscle  attached  posteriorly.  Introvert  hook  without  accessory  spinelets. 
TYPE  SPECIES.  Phascolosoma  granulatum  Leuckart,  1828,  monotypy. 

Subgenus  EDMONDSIUS  subgen.  nov. 
Phascolosoma  (Satonus)  Stephen  &  Edmonds,  1972:  282  (in  part) 

DIAGNOSIS.  Spindle  muscle  not  attached  posteriorly.  Introvert  hook  with  accessory  spinelets  at 
base. 

TYPE  SPECIES.  Phascolosoma  pectinatum  Keferstein,  1867,  monotypy. 
The  subgenus  is  named  in  honour  of  Dr  Stanley  J.  Edmonds. 

REMARKS.  Stephen  &  Edmonds  (1972)  attempted  to  divide  this  large  genus  by  creating  four  sub- 
genera,  P.  (Phascolosoma),  P.  (Rueppellisoma),  P.  ( Antillesoma)  and  P.  (Satonus),  for  the  most 
part  using  published  descriptions  concerning  the  number  of  retractor  muscles  (four  or  two), 
presence  or  absence  of  contractile  vessel  villi  and  whether  or  not  the  spindle  muscle  is  attached 
posteriorly.  In  examining  all  of  the  available  type  material,  Cutler  &  Cutler  (1983)  found  that  the 
subgeneric  distinctions  were  highly  confused  because  many  of  the  original  descriptions  contained 
errors.  P.  (Rueppellisoma),  comprising  eight  putative  species  each  allegedly  with  two  retractor 
muscles,  is  now  considered  invalid  (all  Phascolosoma  are  now  interpreted  as  having  four  retrac- 
tors), the  type  species,  Phascolosoma  rueppellii  Grube,  1868,  by  original  designation,  being  placed 
as  incertae  sedis  since  the  type  is  lost.  P.  (Antillesoma),  formerly  containing  six  species,  now 
contains  only  the  type  species,  Phascolosoma  antillarum  Grube  &  Oersted,  1858,  original  desig- 
nation; this  subgenus  is  sufficiently  distinct  as  to  warrant  generic  rank  (see  below).  The  remaining 
subgenus,  P.  (Satonus),  is  distinguished  from  the  nominate  subgenus  by  the  absence  of  a  posterior 
attachment  of  the  spindle  muscle.  This  character  is  difficult  to  determine  with  any  degree  of 
certainty  in  any  specimen  that  has  been  damaged  internally,  become  macerated  or  has  dried,  as 
found  when  most  of  the  type  materials  of  the  eight  species  grouped  in  P.  (Satonus)  were  re- 
examined,  including  that  of  the  type  species,  Phymosoma  nigritorquatum  Sluiter,  1 882,  original 
designation.  Just  one  species,  Phascolosoma  pectinatum  Keferstein,  1867,  appears  to  fit  the  defi- 
nition off.  (Satonus).  Since  the  type  species  of  this  subgenus,  P.  nigritorquatum,  has  uncertain 


SIPUNCULA  55 

status  (it  may  be  a  junior  synonym  of  P.  ( Phascolosoma)  scolops  (Selenka  &  de  Man)),  it  has  been 
categorised  as  incertae  sedis  (Cutler  &  Cutler,  1983).  Thus  P.  (Satonus)  is  invalid  and  the  new 
subgenus  accommodates  P.  pectinatum. 

Genus  APIONSOMA  Sluiter,  1902 

Apionsoma  Sluiter,  1902:42. 

Golfingia  (Mitosiphon)  Fisher,  1950a:  550. 

Golfingia  (Phascolana)  Wesenberg-Lund,  1959:  183. 

Fisher  ana  Stephen,  1964:  460. 

Golfingia  (Siphonoides)  Murina,  1967:  1334. 

DIAGNOSIS.  Introvert  of  variable  length  in  relation  to  trunk  with  rings  of  recurved  hooks  (absent  in 
A.  trichocephald)  that  in  some  species  have  accessory  spinelets  at  base.  Body  wall  with  continuous 
muscle  layers.  Oral  disk  with  tentacles  enclosing  nuchal  organ  but  not  mouth.  Contractile  vessel 
without  villi.  Four  introvert  retractor  muscles.  Spindle  muscle  attached  posteriorly.  Two 
nephridia,  sometimes  bilobed.  Species  small-sized  (less  than  2  cm  in  length). 

TYPE  SPECIES.  Apionsoma  trichocephala  Sluiter,  1902,  monotypy. 

REMARKS.  Cutler  (1979)  reviewed  this  taxon  which  is  here  elevated  to  generic  status.  It  includes 
many  species  previously  assigned  to  various  Golfingia  subgenera  and  Fisher  ana  (see  above).  This 
genus  is  one  that  still  presents  problems,  in  particular,  the  precise  nature  of  the  oral  disk  in  A. 
trichocephala  remains  unknown.  The  variations  within  the  genus  may  justify  the  use  of  subgenera. 

Genus  ANT1LLESOMA  Stephen  &  Edmonds,  1972 
Phascolosoma  (Antillesoma)  Stephen  &  Edmonds,  1972:  277. 

DIAGNOSIS.  Introvert  variable  in  length,  often  about  equal  to  trunk,  without  hooks.  Body  wall  with 
longitudinal  muscle  layer  gathered  into  anastomosing  bands.  Oral  disk  carries  numerous  tentacles 
(more  than  30  in  adults)  enclosing  nuchal  organ.  Contractile  vessel  with  many  villi.  Four  introvert 
retractor  muscles,  lateral  pairs  often  extensively  fused.  Spindle  muscle  attached  posteriorly.  Two 
nephridia.  Contains  one  small-  to  medium-sized  species  (less  than  5  cm  in  length). 

TYPE  SPECIES.  Phascolosoma  antillarum  Grube  &  Oersted,  1858,  original  designation. 

REMARKS.  This  taxon  was  erected  as  a  subgenus  to  include  six  Phascolosoma  species  but  is  now 
considered  to  be  monospecific  (Cutler  &  Cutler,  1983)  and  of  generic  rank. 

Order  ASPIDOSIPHONIFORMES 

Phascolosomatidea  with  the  anterior  trunk  hardened  to  form  a  horny  or  calcareous  anal  shield. 
Two  retractor  muscles. 

Family  ASPIDOSIPHONIDAE  Baird,  1868 
Characters  are  those  of  the  order. 

Key  to  Genera 

1  Introvert  protrudes  from  centre  of  anal  shield.  Shield  calcareous  (white)  composed  of  numerous 

polygonal  plates .      CLOEOS1PHON 

Introvert  protrudes  from  ventral  margin  of  anal  shield       ...  .  .2 

2  Shield  composed  of  single  calcareous  cap  ....  .        LITHACROSIPHON 
Shield  composed  of  numerous  horny  (brown-black)  plates.        .        .        .     ASPIDOSIPHON 

Genus  ASPIDOSIPHON  Dieting,  1851 

DIAGNOSIS.  Introvert  usually  longer  than  trunk  with  recurved  hooks  in  numerous  rings.  Trunk 
with  anal  shield  composed  of  hardened  plates  (occasionally  inconspicuously  developed).  Introvert 


56  P.  E.  GIBBS  &  E.  B.  CUTLER 

protrudes  from  ventral  margin  of  shield.  Body  wall  either  with  continuous  longitudinal  muscle 
layer  or  with  longitudinal  muscle  layer  gathered  into  anastomosing,  sometimes  ill-defined, 
bands.  Oral  disk  with  tentacles  enclosing  nuchal  organ  but  not  mouth.  Contractile  vessel  without 
villi.  Two  introvert  retractor  muscles  often  almost  completely  fused.  Spindle  muscle  attached 
posteriorly.  Two  nephridia.  Species  small-  to  medium-sized. 

TYPE  SPECIES.  Aspidosiphon  muelleri  Diesing,  1851,  subsequent  designation,  Stephen  &  Edmonds, 
1972. 

Subgenus  ASPIDOSIPHON  Diesing,  1851 
Aspidosiphon  (Aspidosiphon):  Cutler,  1973:  174. 
DIAGNOSIS.  Longitudinal  muscle  layer  of  body  wall  continuous,  not  gathered  into  bands. 

TYPE  SPECIES.  Aspidosiphon  muelleri  Diesing,  1851,  subsequent  designation,  Stephen  &  Edmonds, 
1972. 

Subgenus  PARASPWOSIPHON  Stephen,  1964 

Paraspidosiphon  Stephen,  1964:  459. 
Aspidosiphon  (Paraspidosiphon):  Cutler,  1973:  168. 

DIAGNOSIS.  Longitudinal  muscle  layer  of  body  wall  gathered  into  bands. 
TYPE  SPECIES.  Aspidosiphon  steenstrupii  Diesing,  1859,  original  designation. 

REMARKS.  Earlier  diagnoses  of  this  genus  contain  serious  errors.  The  tentacular  arrangement  is 
phascolosomatid  (Gibbs,  1977;  Gibbs,  in  Edmonds,  1980)  and  there  are  always  two  retractor 
muscles.  Although  A.  semperi  ten  Broeke  and  A.  insular  is  Lanchester  are  described  as  having  four 
retractor  muscles,  the  type  of  the  former  (ZMUA  collection)  has,  in  fact,  two,  and  the  type  of  the 
latter  (BMNH:  Reg.  1 924.3. 1 .80)  is  not  an  Aspidosiphon  but  a  Phascolosoma  (possibly  P. perlucens 
Baird).  In  Aspidosiphon  species  the  spindle  muscle  is  always  attached  posteriorly. 

Genus  CLOEOSIPHON  Grube,  1868 

DIAGNOSIS.  Introvert  longer  than  trunk  with  numerous  rings  of  recurved  hooks.  Trunk  with 
conspicuous  anal  shield  composed  of  small  rectangular  calcareous  plates.  Introvert  protrudes 
through  centre  of  shield.  Body  wall  with  continuous  muscle  layers.  Oral  disk  carries  tentacles 
enclosing  nuchal  organ,  but  not  mouth.  Contractile  vessel  without  villi.  Two  introvert  retractor 
muscles  often  almost  completely  fused.  Spindle  muscle  attached  posteriorly.  Two  nephridia. 
Contains  one  medium-sized  species. 

TYPE  SPECIES.  Loxosiphon  aspergillus  Quatrefages,  1865,  monotypy. 

Genus  L1THACROSIPHON  Shipley,  1902 

DIAGNOSIS.  Introvert  about  equal  to  trunk  with  numerous  rings  of  recurved  hooks.  Trunk  with 
anal  shield  formed  by  internal  calcareous  conical  structure.  Body  wall  with  longitudinal  muscle 
layer  gathered  into  bands.  Oral  disk  with  tentacles  enclosing  nuchal  organ  but  not  mouth. 
Contractile  vessel  without  villi.  Two  introvert  retractor  muscles,  often  almost  completely  fused. 
Spindle  muscle  attached  posteriorly.  Two  nephridia.  Species  small-  to  medium-sized  (less  than 
4  cm  in  length). 

TYPE  SPECIES.  Lithacrosiphon  maldiviense  Shipley,  1902,  monotypy. 
REMARKS.  This  genus  now  contains  two  species  (see  Cutler  &  Cutler,  1981). 

Acknowledgements 

We  are  indebted  to  numerous  colleagues  for  generously  providing  specimens  and  for  kindly  arranging  loans 


SIPUNCULA  57 

of  type  materials,  in  particular,  Mr  R.  W.  Sims -British  Museum  (Natural  History),  Dr  S.  van  der  Spoel- 
Zoologisch  Museum,  Universiteit  van  Amsterdam,  Dr  R.  Olerod  -  Naturhistoriska  Riksmuseet,  Stockholm, 
Dr  J.  B.  Kirkegaard  -  Zoologisk  Museum,  Copenhagen,  Dr  C.  B.  Goodhart  -  University  Museum  of 
Zoology,  Cambridge,  and  Dr  S.  Chambers  -  Royal  Scottish  Museum,  Edinburgh.  We  are  grateful  to  Mr 
R.  W.  Sims  for  his  help  with  the  preparation  of  this  paper. 

References 

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58  P.  E.  GIBBS  &  E.  B.  CUTLER 

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Manuscript  accepted  for  publication  6  December  1985 


Two  new  species  of  Garra  (Teleostei-Cyprinidae) 
from  the  Arabian  peninsula 

K.  £.  Banister 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7  5BD 


Introduction 

Continued  interest  in  speleology  in  Oman  has  led  to  the  discovery  of  yet  another  hypogean,  but  in 
this  case  microphthalmic,  species  of  the  cyprinid  genus  Garra  from  an  extremely  isolated  sink  hole 
in  the  Jabal  Qara  mountains  of  Oman.  In  the  course  of  comparing  this  new  species  with  its 
congeners,  using  the  recent  revision  of  Krupp  (1983),  the  discovery  of  more  specimens  of  Krupp's 
' Garra:  incertae  sedis  (1)'  enabled  a  second  new  species  to  be  described.  In  all,  eight  species  of  Garra 
are  now  known  from  the  Arabian  peninsula. 


Garra  dun  sire  i  sp.  nov. 
(Fig.  1) 

The  first  four  specimens  received  (BMNH  1984.3.6:  577-580)  were  collected  by  Mr  A.  Dunsireand 
Mr  D.  Green  on  16  May  1 980,  but  were  in  too  poor  a  condition  to  be  used  for  description.  A  further 
consignment  of  19  live  specimens  (13  of  which  were  still  alive  at  the  time  of  writing,  July  1985)  was 
collected  by  Mr  D.  Maclelland  on  26  February  1983. 

HOLOTYPE.  BMNH  1984.3.6:  571,  68  mm  SL;  Tawi  Atair  (17°06'N,  54°34'E)  in  the  Jabal  Qara 
(variously  Jabal  Samhan)  mountains,  Dhofar,  Oman. 

PARATYPES.  BMNH  1984.3.6:  572-576,  34-^9  mm,  same  data  as  holotype. 

LOCALITY.  The  fishes  were  caught  in  a  pool  in  a  side  passage  200  m  down  the  sink  hole  shown  in 
Plates  1  and  2.  The  surface  drainage  is  southwards  to  the  Arabian  Sea.  The  significance  of  the 
drainage  direction  and  the  isolated  of  the  locality  will  be  discussed  below. 

DESCRIPTION.  The  description  is  based  on  the  holotype  and  five  paratypes  (34-68  mm  SL),  all  of 
which  were  radiographed.  Additionally  two  of  the  first  four  specimens  (1984.3.6:  577-580)  were 
cleared  and  stained  with  alizarin.  All  measurements  are  expressed  as  a  percentage  of  the  standard 
length.  Because  the  sample  is  so  small,  the  measurements  of  the  holotype  are  included  in  the  range 
as  well  as  being  given  in  parentheses. 

MORPHOMETRIC  DATA.  Body  depth  x  =  22-9,  range  =  21-4-23-8  (23-8);  head  length,  x  =  27-3,  range  =  25-0-30-3 
(25-0);  eye  diameter  x  =  3-3,  range  =  2-8-3-7  (3-7);  interorbital  width  x  =  9-4,  range  =  8-2-10-0  (8-8);  pectoral 
fin  length  x  =  20-2,  range=  18-8-21-4  (19-3);  caudal  peduncle  length  x  =  13-7,  range=  12-0-16-0  (14-7);  caudal 
peduncle  depth  x  =  10-1,  range  =  9-2-1 1-4  (10-3);  anterior  barbel  length  x  =  4-2,  range  =  3-7-4-5  (3-7);  poster- 
ior barbel  length  x  =  3-5,  range  =  2-2^-0  (2-6);  dorsal  fin  height  x  =  23- 1 ,  range  =  22-2-23-8  (22-8);  mental  disc 
maximum  length  x  =  7-4,  range  =  6-5-8-8  (6-5);  mental  disc  maximum  width  x  =  7-3,  range  =  7- 1-8-1  (7-2); 
distance  between  snout  and  anus  x  =  76-6,  range  =  73-5-78-5  (74-4);  distance  between  snout  and  anal  fin  origin 
x  =  80-9,  range  =  77-9-83-6  (78-7);  distance  between  snout  and  pelvic  fin  origin  x  =  60-5,  range  =  58 -8-63 -3 
(59-1);  distance  between  snout  and  dorsal  fin  origin  x  =  53-0,  range  =  45-9-55-9  (51-5). 

The  body  shape  and  details  of  the  mental  disc  are  shown  in  Figs  1  and  2a.  The  abdomen  of  the 
holotype  has  collapsed,  creating  an  uncharacteristic  flat-bellied  appearance.  The  eyes  are  very 
small  and  not  visible  in  ventral  view.  The  mental  disc  is  approximately  circular.  In  the  smallest 
specimen  (34  mm  SL),  only  the  posterior  margin  of  the  disc  is  free,  but  as  the  fish  grows,  the  rest  of 


Bull.  Br.  Mm.  nat.  Hist.  (Zool.)  52(1):  59-70  Issued  29  January  1987 


60 


K.  E.  BANISTER 


Plate  1 .    The  sink  hole  at  Tawi  Atair . 


SPECIES  OF  GARRA 


61 


Plate  2.    A  close-up  of  the  arrowed  area  in  Plate  1  to  indicate  the  size  of  the  sink  hole. 


Fig.  1.     Garra  dunsirei  Holotype.  Scale  =  1 0  mm. 


62 


K.  E.  BANISTER 


Fig.  2.     Ventral  views  of  the  heads  of  a.  Garra  dunsirei  and  b.  Garra  lautior.  Scale  =  5  mm. 


the  rim  becomes  free.  The  papillae  are  scarcely  developed,  the  papillar  bed  being  only  just  differen- 
tiated in  a  specimen  of  50  mm  SL  (Fig.  2a).  Only  on  the  holotype  are  the  papillae  well  formed.  The 
size  and  extent  of  the  papillae  are  less  than  in  all  other  Garra  in  the  region. 

None  of  the  specimens,  alive  or  preserved,  has  tubercles  on  the  snout,  although  a  39-5  mm  SL 
specimen  is  a  gravid  female. 

SQUAMATION.  The  scales  are  less  well  developed  than  those  of  the  hypogean  population  of  Garra 
barreimiae  (Banister,  1 984)  which,  although  thinner  than  the  scales  of  epigean  populations,  are 
scarcely  less  extensive.  In  Garra  dunsirei  the  scales  are  as  deep  as,  or  deeper,  than  long  and  although 
they  fill  the  scale  pocket  vertically,  they  often  fail  to  reach  the  posterior  edge  of  the  pocket.  A  scale 
and  its  pattern  of  striations  is  shown  in  Fig.  3.  Scales  of  the  holotype  have  eleven  'growth'  rings.  It  is 
not  known  if  these  represent  annual  or  seasonal  spawning  marks  or  are  the  result  of  growth 
changes  caused  by  food  availability.  Food  only  comes  into  the  cave  .during  the  annual  rains.  The 
scales  of  the  ventral  region  of  the  body  are  severely  reduced  or  absent.  In  the  lateral  line  series  there 
are  34(f2),  35(f3)  or  36(fl)  scales.  There  are  3^(f6)  scale  rows  from  the  dorsal  fin  base  to  the  lateral 
line  and  2^(f3)  or  3|(f3)  scale  rows  from  there  to  the  pelvic  fin  base.  On  the  two  specimens  on  which 
a  count  was  possible  there  are  6|  scale  rows  from  the  lateral  line  to  the  ventral  mid-line.  Twelve  (f6) 
scale  rows  encircle  the  least  circumference  of  the  caudal  peduncle. 

VERTEBRAE.  Radiographs  revealed  the  presence  of  32(f3)  or  33(f3)  vertebrae,  excluding  those 
forming  the  Weberian  mechanism.  It  was  difficult  to  identify  with  certainty  the  first  caudal  vertebra 
but  there  appear  to  be  18(f2)or  1 9(f4)  abdominal  vertebrae.  There  are  14(f2)or  15(f4)  pairs  of  ribs. 
Characteristic  of  this  species  is  the  most  unusual  change  of  shape  of  the  neural  arches  and  spines 
below  the  dorsal  fin.  In  all  the  other  Arabian  peninsular  Garra  species  and  in  all  other  Garra  species 
investigated  the  neural  spines  are  shorter  in  this  region,  but  of  the  same  general  configuration  and 
angle  as  the  other  neural  spines  (Fig.  9).  In  Garra  dunsirei  (Fig.  4)  the  neural  spines  are  bent  sharply 
back  and  come  to  lie  almost  in  parallel  with  the  axis  of  the  centrum. 


SPECIES  OF  GARRA 


63 


Fig.  3.     A  scale  from  the  row  above  the  lateral  line  of  the  holotype  of  Garra  dunsirei.  Scale  =  0-5  mm. 


Fig.  4.     Garra  dunsirei.  The  vertebral  column  below  the  dorsal  fin  to  show  the  unusual  profile  of  the 
neural  spines.  For  clarity  the  ribs  are  omitted.  Scale  =  1  mm. 


FINS.  The  dorsal  fin  has  only  3(f6)  unbranched  and  7(f5)  or  8(fl)  branched  rays.  The  foremost 
unbranched  ray  is  minute  and  only  visible  in  a  radiograph  or  an  alizarin  preparation.  The  anal  fin 
has  3  unbranched  and  5  branched  rays  (f6).  The  first  unbranched  ray  is  again  minute  and  not  visible 
externally. 

GILL  RAKERS.  The  gill  rakers  are  small  in  size  and  few  in  number.  There  are  6(f2)  or  7(f2)  on  the 
lower  limb  of  the  first  gill  arch.  They  could  not  be  counted  in  the  two  smallest  specimens. 

PHARYNGEAL  BONES  AND  TEETH.  The  pharyngeal  teeth  number  2.4.5-5.4.2  (Fig.  5).  The  five 
posterior  teeth  of  the  innermost  row  are  thin  and  have  hooked  crowns,  quite  unlike  those  typical  of 
other  Garra  species  from  the  Arabian  peninsula  (see  Krupp,  1983:  figs  23,  27, 40). 


64  K.  E.  BANISTER 


Fig.  5.     Left  pharyngeal  bone  of  Garra  dunsirei.  Scale  =  1  mm. 


COLORATION.  Alcohol  preserved  specimens  are  uniform  pale,  yellowish  grey.  The  ventral  surface  is 
only  slightly  paler  than  the  rest  of  the  body.  There  are  traces  of  dark  pigment  near  the  anterior  edge 
of  the  pectoral  fins,  on  the  dorsal  edge  of  the  dorsal  fin  membrane  as  well  as  near  the  base  of  the 
dorsal  fin  (the  last  being  the  typical  Garra  markings).  Other  fins  are  colourless. 

Living  fishes  are  a  dirty  white,  slightly  more  heavily  dusted  with  dark  pigment  dorsally.  The 
cheeks  and  operculum  reflect  greenish-gold.  The  post-opercular  spot  is  inconspicuous  but  the  red 
of  the  blood  in  the  gills  is  clearly  visible  through  the  adjacent  notch. 

ETYMOLOGY.  This  species  is  named  in  honour  of  the  collector,  Mr  Andy  Dunsire,  who  has 
encouraged  so  many  people  to  search  for  subterranean  fishes,  as  well  as  collecting  such  fish  himself 
in  isolated  and  hazardous  regions. 

NOTES  ON  LIVING  SPECIMENS.  The  fishes  swim  slowly,  but  continually,  usually  in  a  slightly  head 
down  position.  When  resting,  they  are  indifferent  to  their  orientation  provided  that  their  ventral 
surface  is  in  contact  with  a  solid  object.  They  stay  near  the  bottom  of  the  aquarium,  very  rarely 
approaching  the  surface  even  to  take  food. 

In  these  aspects  they  contrast  markedly  with  the  blind,  hypogean  population  of  Garra  barrei- 
miae  which  swims  rapidly  and  swarms  at  the  surface  on  the  introduction  of  food  (Banister,  1984). 

Garra  dunsirei  shows  no  preference  for  either  light  or  dark  conditions,  although  a  strong  light 
beam  shone  on  an  individual  will  cause  it  to  jerk  away  after  one  or  two  seconds.  After  that, 
however,  the  fishes  will  come  and  investigate  a  localised  illuminated  patch  on  the  substrate. 

DISTRIBUTION  AND  RELATIONSHIPS.  Of  particular  interest  is  the  geographical  isolation  of  Garra 
dunsirei  horn  its  congeners.  In  fact,  no  primary  freshwater  fishes  have  been  recorded  from  this  area 
of  Dhofar,  the  closest  natural  populations  being  nearly  400  miles  away  at  Tarim  in  the  Wadi 
Hadramut  to  the  west  and  also  some  450  miles  to  the  northeast  in  the  Omani  Jabal  Akhdar.  The 
drainage  of  this  part  of  the  peninsular  coast  is  a  series  of  isolated  wadis  carrying  the  run-off  due 
south  from  the  Jabal  Qara  range  to  the  Arabian  sea.  On  the  north  side  of  the  Jabal  Qara  is  an 
interlinked  series  of  wadis  that  in  wetter  times  would  have  carried  water  northwards  into  the 
extensive  lake  or  lake  system  that  occupied  the  site  of  the  Rub  al  Khali.  Krupp  (1983)  points  out 
that  the  Jabal  Qara  range  is  part  of  an  ancient  feature  whose  presence  caused  the  formation  of  the 
internal  drainages.  Although  the  internal  basin  would  have  permitted  fish  dispersal  northward 
from  Jabal  Qara,  whether  or  not  it  allows  fishes  from  the  north  access  to  the  streams  south  of  Jabal 
Qara  is  unknown.  It  might  be  possible  to  determine  the  hydrological  affinities  of  this  isolated  water 
source  were  the  phylogenetic  relationships  of  Garra  dunsirei  known.  For  the  moment,  therefore, 
neither  the  hydrological  affinities  of  the  sink  hole  nor  the  phylogenetic  relationships  of  Garra 
dunsirei  can  be  determined. 

DIAGNOSIS.  This  species  can  be  characterised  by  the  late  development  of  the  papillar  beds  on 
the  mental  disc,  the  papillar  bed  being  only  just  differentiated  at  50  mm  SL;  also  the  small  eye 


SPECIES  OF  GARRA 


65 


diameter  (x  =  3-3)  and  the  highly  unusual  shape  of  the  neural  spines  below  the  dorsal  fin  (see  Fig.  4 
and  p.  62). 


Garra  lautior  sp.  nov. 

The  recognition  of  this  species  stems  from  Krupp  (1983: 615)  who  described  six  specimens  from  the 
Wadi  Hadramut  as  'Garra:  incertae  sedis  (1)'  but  was  reluctant  to  base  a  species  on  such  a  small 
sample.  A  search  through  the  collections  of  the  British  Museum  (Natural  History)  revealed  13 
more  specimens. 

HOLOTYPE.  BMNH  1976.4.7:  398,  74  mm  SL  from  the  Qasam  area,  Wadi  Hadramut,  Yemen,  coll. 
King-Webster. 

PARATYPES.  BMNH  1976.4.7:  399-404,  64-74  mm  SL  (other  details  as  above);  1976.4.7:  647-648, 
71  &  75  mm  SL  (other  detals  as  above);  1976.4.7: 377-378, 61  &  74  mm  SL:  (other  details  as  above); 
1976.4.7:  645,  80  mm  SL  from  Al-Ghurf,  Wadi  Hadramut,  coll.  King- Webster;  BMNH  1976.4.7: 
366  79  mm  SL,  from  Gheil  Umar,  Wadi  Hadramut,  coll.  King- Webster. 

DESCRIPTION.  The  description  is  based  on  the  holotype  as  12  paratypes  (61-80  mm  SL).  The 
measurements  are  expressed  as  a  percentage  of  the  standard  length  those  of  the  holotype  are 
included  in  the  range  also  also  given  separately  in  parentheses. 

MORPHOMETRIC  DATA.  Body  depth  x  =  22-3,  range  =  20-2-25-0  (22-3)  (n  =  8);  head  length  x  =  22-4, 
range  =  21 -6-23-4  (21-6);  eye  diameter  x  =  4-8,  range  =  4-0-5-4  (5-4);  mouth  width  x  =  5-7,  range  =  4-4-6-6 
(6- 1);  pectoral  fin  length  x=  19-4,  range=  18-7-21 -5  (19-3);  caudal  peduncle  length  x=  17-1,  range  =15-2-20-0 
(15-9);  caudal  peduncle  depth  x  =  8-6,  range  =  7-7-9-4  (9-3);  anterior  barbel  length  x  =  2-6,  range  =  1-3-3-2 
(3-0);  posterior  barbel  length  x  =  2-0,  range  =  1  -5-3-7  (1-9);  dorsal  fin  height  x  =  24-8,  range  =  22-9-28- 1  (25-5); 
mental  disc  maximum  length  x  =  5-4,  range  =  4-7-6-3  (5-5);  mental  disc  maximum  width  x  =  7-0, 
range  =  5-7-7-7  (7-3);  distance  between  snout  and  anus  x  =  71-4,  range  =  66- 6-75-0  (70-2);  distance  between 
snout  and  anal  fin  origin  x  =  74-3,  range  =  69-3-77-0  (72-2);  distance  between  snout  and  pelvic  fin  origin 
x  =  50-6,  range  =  47-9-52-6  (50-4);  distance  between  snout  and  dorsal  fin  origin  x  =  45-2,  range  =  43-2-46-8 
(43-5). 

The  body  has  a  characteristic,  streamlined  shape  (Fig.  6  and  Krupp,  1983:  fig.  30).  From  the 
pointed  snout,  the  dorsal  profile  rises  smoothly  to  the  insertion  of  the  dorsal  fin.  Behind  the  dorsal 
fin,  the  trunk  diminishes  in  depth,  terminating  in  a  slender  caudal  peduncle  almost  exactly  half  as 
deep  as  long.  In  five  specimens  the  abdomen  had  collapsed,  so  a  reliable  body  depth  measurement 
could  not  be  taken.  In  ventral  view,  the  upper  lip  is  thick  and  has  many  small  papillae.  The  shape  of 


Fig.  6.     Garra  lautior.  Holotype.  Scale  =  10  mm. 


66 


K.  E.  BANISTER 


Fig.  7.  Details  of  the  distribution  of  papillae  on  the  frenum,  lips  and  disc  of  (left)  a  paratype  of  Garra 
mamshuqua  (74  mm  SL  ex  1976.4.7:  381-387),  and  (right)  a  paratype  of  Garra  lautior  (74  mm  SL  ex 
1976. 4.7:  399^04)  Scale  =  0-1  mm. 


Fig.  8.    A  scale  from  the  row  above  the  lateral  line  of  Garra  lautior.  Scale  =  0-5  mm. 


the  disc  and  the  disposition  of  the  papillar  beds  are  shown  in  Figs  2  &  7B.  None  of  the  specimens 
has  any  tubercles  on  the  head  although  both  mature  males  and  females  are  present  in  the  sample 
(see  below). 

The  small  size  range  of  the  specimens  available  was  insufficient  to  establish  any  marked  instances 
of  allometric  growth. 

SQUAMATION.  The  scales  are  well  developed  and  slightly  lobate.  A  scale  and  its  striations  are  shown 
in  Fig.  8.  Only  two  poorly  defined  growth  rings  were  discernible.  In  the  lateral  line  series  there  are 
32(f3),  33(f4),  34(f4)  or  35(f2)  scales.  From  the  dorsal  mid-line  to  the  lateral  line  there  are  3^(f9)  or 
4|(f3)  scales  and  from  the  lateral  line  to  the  pelvic  fin  base  3^(fl  3)  scales.  In  front  of  the  anal  fin  the 


SPECIES  OF  GARRA 


67 


ventral  surface  is  scaleless.  There  are  12(fl3)  scale  rows  around  the  least  circumference  of  the 
caudal  peduncle. 

VERTEBRAE.  In  the  nine  specimens  radiographed  there  are  27(f2),  28(f3),  29(f3)  or  30(fl )  vertebrae, 
excluding  those  comprising  the  Weberian  mechanism.  The  abdominal  vertebrae  number  12(f3)  or 
1 3(f6)  (allowing  for  the  difficulty  in  identifying  the  first  caudal  vertebra).  The  neural  spines  below 
the  dorsal  fin  pterygiophores  display  the  normal  alignment  and  reduction  in  size  of  most  Garra  spp 
(Fig.  9)  in  contrast  to  the  unique  condition  in  Garra  dunsirei  (Fig.  4). 

In  all  the  other  species  of  Garra  radiographed:  viz  G.  mamshuqua  Krupp,  1983,  G.  barreimiae 
Fowler  &  Steinitz,  1956,  G.  sahilia  Krupp,  1983,  G.  tibanica  Trewavas,  1941,  and  G.  dunsirei,  there 
are  4  interhaemal  spine  spaces  corresponding  to  the  anal  fin  pterygiophores  (Fig.  10),  but  in  G. 
lautior  only  three  interhaemal  spine  spaces  do  so  correspond  (Fig.  1 1).  No  particular  significance  is 
attached  to  variations  in  the  shape  of  the  last  anal  fin  pterygiophore.  There  are  1 4(f7)  or  1 5(f2)  pairs 
of  ribs. 

GILL  RAKERS.  The  gill  rakers  are  small,  hooked  and  number  ll(fl),  12(fl),  13(f3),  14(fl),  15(f3), 
16(fl)  and  17(fl)  on  the  lower  limb  of  the  first  gill  arch. 

PHARYNGEAL  BONES  AND  TEETH.  The  pharyngeal  teeth  number  2.4.5-5.4.2  (Fig.  12).  The  crowns 
have  shallow  spoon-edged  depressions,  the  depression  being  most  sharply  edged  in  newly  replaced 
teeth. 


Fig.  9.     The  vertebral  column  below  the  dorsal  fin  of  Garra  lautior  to  show  the  shape  of  the  neural  spines. 

Scale  =  1  mm. 


Fig.  10.     The  anal  fin  pterygiophores  of  an  unregistered  BMNH  specimen  of  Garra  mamshuqua,  54  mm 
SL,  to  show  their  opposition  to  four  interhaemal  spaces.  Scale  =  1  mm. 


68 


K.  E.  BANISTER 


Fig.  11.     The  anal  fin  pterygiophores  of  a  specimen  of  Garra  lautior  (unregistered)  to  show  their 
opposition  to  only  three  interhaemal  spaces.  Scale  =  1  mm. 


Fig.  12.     Left  pharyngeal  bone  of  the  holotype  of  Garra  lautior.  Scale  =  0-5  mm. 


COLORATION.  Alcohol  preserved  specimens  are  a  uniform  sandy  brown,  darker  dorsally.  The 
post-opercular  spot  is  a  deeper  brown,  but  the  'Garra '  marks  at  the  base  of  the  dorsal  fin  membrane 
are  not  especially  conspicuous.  The  fin  membranes  are  clear. 

DISTRIBUTION.  This  species  is  known  only  from  localities  within  the  Wadi  Hadramut  drainage, 
Yemen. 

ETYMOLOGY.  The  trivial  name  is  the  comparative  oflautus,  the  Latin  for  smart  or  neat  and  alludes 
to  the  neat,  streamlined  appearance  of  the  fish. 

DIAGNOSIS.  Garra  lautior  is  sympatric  only  with  Garra  mamshuqua  (see  below).  Although  the  two 
species  are  somewhat  similar  in  body  shape,  Garra  mamshuqua  can  be  distinguished  by  the  pres- 
ence of  tubercles  on  the  snout,  the  very  conspicuous  'Garra '  marks  on  the  dorsal  fin  and  behind  the 
operculum  and  the  different  disc  shape  (Fig.  2b  &  7b).  Very  small  specimens  can  be  most  easily 
separated  on  the  greater  intensity  of  the  post-opercular  spot  in  Garra  mamshuqua. 


SPECIES  OF  GARRA  69 

TUBERCLES.  Tubercles  on  the  snout  are  often  called  nuptial  or  breeding  tubercles  (e.g.  Wiley  & 
Collette,  1970)  or  multicellular  horny  tubercles  (Roberts,  1982).  The  latter  author  points  out  that 
they  may  occur  in  both  sexes  as  well  as  being  present  before  the  onset  of  sexual  maturity  in  Labeo 
species  and  the  homalopterids.  In  Garra  mamshuqua,  the  tubercles  are  present  in  both  males  and 
females  at  all  stages  of  sexual  maturity  and  first  appear  in  specimens  of  27  mm  SL  (e.g.  in  BMNH 
1967.4.7: 407-418).  It  seems  probable  therefore  that  in  Garra  mamshuqua  the  tubercles  do  not  have 
a  solely  sexual  or  reproductive  function. 

A  hydrodynamic  function  was  suggested  by  Reid  (1978)  for  their  occurrence  in  Labeo,  since 
fishes  from  faster  flowing  waters  had  more  and  larger  tubercles  than  those  from  quiet  waters.  The 
localities  where  the  smooth  Garra  lautior  and  the  tuberculate  Garra  mamshuqua  are  sympatric  were 
described  by  the  collector  (original  letter  in  the  BMNH  Fish  Section  archives)  as  'a  clear  stream 
with  stony  shallows  and  deep  holes'  (Gheil  Umar),  and  also  an  'isolated  muddy  pot-hole  below  a 
dam'  (Al-Ghurf).  Such  scanty  and  seemingly  inconsistent  information  adds  nothing  to  Reid's 
hydrodynamic  hypothesis. 

Although  the  function  of  the  tubercles  is  not  known,  it  does  seem  in  this  case  that  their  presence 
can  be  used  as  a  sound  diagnostic  character  to  distinguish  these  two  sympatric  species.  However,  it 
is  not  suggested  that  the  presence  or  absence  of  tubercles  is  diagnostic  for  other  species. 


Discussion 

Krupp  ( 1 983)  also  recorded  both  Garra  tibanica  and  Garra  sahilia  from  the  Wadi  Hadramut.  Garra 
tibanica  was  included  as  a  member  of  the  Hadramut  fauna  solely  on  specimens  collected  by 
Scortecci  at  Bir  el  Manzil  (14°32'N,  4&°5\'Ejide  Krupp).  Scortecci's  Bir  el  Manzil  is  shown  on 
the  map  in  Balletto  &  Spano  (1977),  which  is  concerned  with  the  Scortecci  expedition  and  is 
approximately  14°30'N,  44°30'E  or  well  to  the  west  of  the  Hadramut. 

The  Wadi  Hadramut  record  of  Garra  sahilia  was  based  on  four  fishes,  two  from  Sayun  (BMNH 
1980.4.24:  8,9)  and  two  from  nearby  Shibam  (BMNH  1980.4.24:  6,7).  The  latter  specimens 
were  identified  by  Krupp  but  not  listed  in  his  1983  paper.  The  four  fishes  do  not  correspond  to 
the  description,  especially  in  having  much  longer  barbels  and  the  anus  closer  to  the  anal  fin  than 
in  Garra  sahilia.  However,  their  poor  condition  precludes  confirmation  of  Krupp's  specific 
determination. 

There  are  some  difficulties  in  establishing  which  specimens  of  Garra  sahilia  are  types.  Krupp 
(1983:  601)  lists  63  specimen  (BMNH  1976.4.7:419,420-425;  1951.5.9:  12-65  and  1944.4.3:  1-10) 
as  paratypes  but  used  only  25  specimens  in  his  description.  Presumably,  the  63  listed  paratypes 
included  the  24  actually  described  (although  the  largest  specimen  in  his  sample,  100- 5  mm  SL 
BMNH  1940.2.15:  12-18  was  used  in  the  description  but  not  designated  a  paratype). 

Although  twice  as  many  Garra  lautior  specimens  were  available  to  me  than  to  Krupp,  in 
most  respects  our  descriptions  are  similar.  However,  there  is  substantial  discrepancy  in  our  scale 
counts  around  the  caudal  peduncle.  In  the  13  specimens  used  here  (6  of  which  were  those  used  by 
Krupp)  I  could  count  only  12  scales,  whereas  Krupp  gives  14(fl)  or  16(f5).  A  similar  discrepancy 
occurs  with  the  same  count  in  Garra  buettikeri  Krupp,  1983,  Krupp  giving  18(f2)  or  20(fl8)  as  the 
diagnostically  high  circumpeduncular  scale  count,  whereas  in  the  six  BMNH  specimens  he  used  in 
his  description  I  count  only  16(f2)  or  18(f4)  scales.  In  Krupp's  fig.  21  twenty  scales  would  be  too 
many,  unless  the  squamation  in  the  specimen  illustrated  was  unusually  asymmetrical.  Although 
Krupp  did  not  indicate  how  he  made  his  counts  I  can  imagine  only  one  way  of  counting  the  number 
of  scale  rows  around  the  least  circumference  of  the  caudal  peduncle.  It  seems  unwise,  therefore,  to 
attribute  diagnostic  significance  to  this  particular  meristic  feature.  No  attempt  is  made  in  this  paper 
to  produce  a  key  to  the  Garra  species  of  the  Arabian  peninsula.  Even  a  cursory  glance  at  the  means 
and  ranges  of  any  particular  morphometric  or  meristic  feature  used  here,  in  Krupp  (1983),  in 
Banister  &  Clarke  (1977)  and  in  many  other  papers  shows  that  the  similarity  of  means  and  the 
extensive  overlaps  in  range  usually  precludes  the  use  of  such  characters  in  a  key.  Even  if  the  eight 
peninsular  species  were  initially  subdivided  by  drainage  regions  (giving  groups  of  3,  2,  2  and  1)  a 
key  based  on  morphometric  and  meristic  characters  would  not  infallibly  separate  the  sympatric 


70  K.  E.  BANISTER 

species.  The  major  diagnostic  features  are  regrettably  very  difficult  to  quantify.  At  the  moment  the 
most  useful  characters  are  the  overall  body  shape,  the  shape  of  the  mental  disc  and  the  distribution 
of  papillae  thereon,  and  the  colour  pattern.  Although  details  of  the  mental  disc  serve  to  distinguish 
species,  based  on  the  samples  available,  the  variation  in  at  least  one  species,  Garra  tibanica  (Balletto 
&  Spano,  1977:  fig.  6)  makes  one  wonder  whether  it  will  remain  useful  when  more  populations  are 
discovered. 

Krupp  1983:  603-615  provided  a  useful  list  of  all  the  specimens  he  examined.  There  are, 
however,  some  confusions  in  the  BMNH  register  numbers  in  his  list  and  these  and  some  other 
errors  are  corrected  below. 

Garra  sahilia  sahilia  (p.  603) 

Sample  No.  10  for  Wadi  Abd  read  Wadi  Anad  44°50'E,  13°17'N 

No.  14  for  1976.4.7:  443^60  read  1976.4.7:  460 

No.  15  for  1910.1.28:  1-3  read  1870.1.28:  1-3 

Garra  sahilia  gharbia  (p.  604) 

Sample  No.  5  for  1976.4.7:  646-354  read  1976.4.7:  346-349 

Garra  tibanica  tibanica  (p.  608) 

Sample  No.    5  for  1976.4.7:  443^60  read  1976.4.7:  443^59 

No.  1 1  for  1952.5.7:  13-18     read  1952.5.7:  13-17 

No.  17  for  1976.4.7:  346-356  read  1976.4.7:  350-354 

Garra  incertae  sedis  (p.  615) 

Sample  No.  1  for  1976.4.7:  374-377  read  1976.4.7:  377 

No.  3  for  1976.4.7:  380-406  read  1976.4.7:  380-396 


Acknowledgements 

I  particularly  wish  to  thank  the  collectors  for  their  enthusiasm  in  seeking  for  fish  life  in  such  an  inhospitable 
environment.  Without  the  efforts  of  such  people,  our  knowledge  of  fish  distribution  and  habitats  would 
increase  more  slowly.  My  colleagues,  Dr  P.  J.  P.  Whitehead  and  Mr  A.  C.  Wheeler  offered  constructive 
comments  on  the  manuscript.  My  thanks  go  to  them  and  to  Gordon  Howes  for  illustrating  the  new  species 
and  to  Joan  Ellis  for  typing  the  paper. 


Bibliography 

Balletto,  E.  &  Spano,  S.  1977.  Ciprinidi  del  genere  Garra  Hamilton  1822,  raccolti  nello  Yemen  dal  Prof. 

Guiseppe  Scortecci.  Annali  del  Museo  Civico  di  Storia  Naturale  (di  Genova)  Giacomo  Doria,  Genova  81: 

246-287. 
Banister,  K.  E.  1984.  A  subterranean  population  of  Garra  barreimiae  (Teleostei:  Cyprinidae)  from  Oman, 

with  comments  on  the  concept  of  regressive  evolution.  Journal  of  Natural  History  18: 927-938. 
Banister,  K.  E.  &  Clarke,  M.  A.  1977.  The  freshwater  fishes  of  the  Arabian  peninsula.  The  scientific  results  of 

the  Oman  Flora  and  Fauna  survey  1975.  Journal  of  Oman  Studies  1977:  1 1 1-154. 
Krupp,  F.  1983.  Fishes  of  Saudi  Arabia  and  adjacent  regions  of  the  Arabian  Peninsula.  Fauna  of  Saudi  Arabia 

5:  568-636. 
Reid,  G.  McG.  1978.  A  systematic  study  oflabeine  cyprinid fishes  with  particular  reference  to  the  comparative 

morphology  and  morphometrics  of  African  Labeo  species.  Ph.D.  thesis,  University  of  London,  770  pp. 
Roberts,  T.  R.  1982.  Unculi  (horny  projections  arising  from  single  cells),  an  adaptive  feature  of  the  epidermis 

of  ostariophysan  fishes.  Zoologica  Scripta  11:  55-76. 
Wiley,  M.  L.  &  Collette,  B.  B.  1970.  Breeding  tubercles  and  contact  organs  in  fishes:  their  occurrence, 

structure  and  significance.  Bulletin  of  the  American  Museum  of  Natural  History  143:  143-216. 

Manuscript  accepted  for  publication  6  November  1985 


British  Museum  (Natural  History) 

The  birds  of  Mount  Nimba,  Liberia 


Peter  R.  Colston  &  Kai  Curry-Lindahl,  with  a  section  on  the  biogeographic 
context  by  Malcolm  Coe. 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  on  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Tring,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1986,  129pp.  Hardback.  0  565  00982  6  £17.50. 


Titles  to  be  published  in  Volume  52 

Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophom  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barbm  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata).  By  P.  J. 

Chimonides 


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


A  revision  of  the  Suctoria  (Ciliophora, 
Kinetofragminophora)  5.  The  Paracineta 
and  Corynophrya  problem 

Colin  R.  Curds 


Zoology  series    Vol52    No  2    27  February  1987 


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Vol52  No.  2  pp  71-106 
British  Museum  (Natural  History) 
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London  SW7  5BD  Issued  27  February  1987 


PRESENTED 


II    f 
A  revision  of  the  Suctoria  (Ciliophor a,  f L  1     2  7  F  E  B 1987 

Kinetofragminophora)  5.  The  Paradneta  and 
Corynophrya  problem 


Colin  R.  Curds 

Zoology  Department,  British  Museum  (Natural  History),  Cromwell  Road,  London 


Contents 


Synopsis 
Introduction   . 
Genus  Actinocyathula 
Genus  Corynophrya 
Genus  Pelagacineta 
Genus  Paradneta    . 
Genus  Loricophrya  . 
Genus  Anthacineta  . 
Genus  Flectacineta  . 
References 
Index  to  species 


71 

71 

72 

76 

82 

86 

91 

98 

101 

104 

106 


Synopsis 


The  continual  drift  in  the  diagnosis  of  the  unrelated  genera  Paradneta  and  Corynophrya  causes  considerable 
taxonomic  problems  and  confusion  at  several  levels  in  classification.  The  transfer  of  Paradneta  crenata  and 
Paradneta  homari  into  the  genus  Actinocyathula  has  allowed  the  present  review  to  be  based,  as  far  as  possible, 
on  the  original  diagnoses  of  the  genera.  In  addition  to  those  mentioned  above,  the  species  of  four  other  genera, 
Pelagacineta,  Loricophrya,  Anthacineta  and  Flectacineta  are  reviewed  since  some  have  been  previously 
associated  in  some  way  with  the  Paradneta-Corynophrya  problem  in  the  past. 

A  new  diagnosis  for  each  genus  is  given  with  a  key  to  its  constituent  species  and  where  appropriate  a 
genotype  has  been  designated  to  encourage  taxonomic  stability.  All  species  are  described  and  figured. 


Introduction 

There  is  still  considerable  confusion  and  disagreement  on  the  generic  diagnoses  of  Paradneta 
Collin,  1911  and  Corynophrya  Kahl,  1934.  The  purpose  of  this  publication  is  to  review  the  species 
involved,  to  amend  previous  diagnoses  and  to  assign  type  species  to  the  genera  in  an  attempt  to 
establish  taxonomic  stability.  The  genus  Paradneta  was  erected  in  order  to  take  account  of  those 
loricate  suctoria  with  an  apical  group  of  tentacles  that  reproduced  by  external  budding  and  that 
were  longitudinally  symmetrical.  In  his  original  generic  description,  Collin  (1911)  included  the 
three  species  Paradneta  crenata  (Fraipont,  1878),  P.  homari  (Sand,  1899)  and  P.patula  (Claparede 
&  Lachmann,  1861)  but  failed  to  designate  the  type  species.  In  his  later  taxonomic  revision, 
Collin  (1912)  transferred  several  more  species  into  the  genus  including  Paradneta  limbata 
(Maupas,  1881),  P.  vorticelloides  (Fraipont,  1878),  P.  jorisi  (Sand,  1895),  P.parva  (Sand,  1899), 
P.  multitentaculata  (Sand,  1895),  P.  livadiana  (Mereschkowsky,  1881),  P.  elegans  (Imhoff,  1883) 
and  P.  bifaria  (Stokes,  1887).  Collin  (1911,  1912)  stressed  that  although  external  budding  was  a 
prime  feature  of  the  genus  both  Paradneta  crenata  and  P.  homari  in  fact  reproduced  by  semi- 
external  budding  (the  semi-invaginative  budding  of  Batisse,  1975).  At  the  time  this  method  was 
thought  to  be  only  a  slight  variation  on  the  external  budding  theme  and  of  little  significance. 


Bull.  Br.  Mas.  not.  Hist.  (Zool.)  52(2):  71-106 


Issued  27  February  1987 


71 


72  C.  R.  CURDS 

Modern  workers  however  consider  the  different  modes  of  budding  to  be  of  great  taxonomic 
importance  and  that  there  is  a  distinct  difference  between  semi-invaginative  and  external  budding. 
Nevertheless,  the  two  species  remained  in  their  original  genus  until  Batisse  (1975)  suggested  their 
transfer  into  the  genus  Corynophrya  Kahl,  1934  which  had  been  originally  erected  for  a  hetero- 
genous  assemblage  of  aloricate  suctoria  reproducing  by  internal  budding.  Although  the  suggestion 
by  Batisse  (1975)  may  appear  strange,  since  the  two  species  in  question  are  loricate  and  reproduce 
differently,  it  should  be  pointed  out  that  the  generic  diagnosis  of  Corynophrya  has  drifted  consider- 
ably since  that  originally  outlined  by  Kahl  (1934).  However,  Batisse  (1975)  had  not  taken  into 
account  that  Paracineta  crenata  can  be  regarded  to  be  congeneric  with  Actinocyathus  cidaris  Kent, 
1882  and  would  be  more  neatly  transferred  into  the  latter  older  genus.  Jankowski  (1981)  is  also 
apparently  of  a  similar  opinion  since  he  suggested  that  the  name  Actinocyathus  might  replace  that 
of  Paracineta.  The  name  Actinocyathus  was  shown  by  Corliss  (1960)  to  be  preoccupied  and  he 
suggested  the  replacement  name  Actinocyathula  Corliss,  1960. 

Kahl  (1934)  erected  the  genus  Corynophrya  to  include  the  mostly  marine  assemblage  of  suctoria 
which  Collin  (1912)  had  gathered  together  in  his  third  group  within  the  genus  Discophrya.  The 
major  diagnostic  features  were  that  they  reproduced  by  internal  budding,  did  not  possess  a  lorica, 
were  rounded  in  cross-section,  had  one  type  of  tentacle  that  was  restricted  to  the  apical  surface  and 
had  a  rounded,  compact  nucleus.  According  to  Kahl  (1934)  the  following  species  held  these 
features  in  common,  Corynophrya  marina  (Andrusov,  1886),  C.  conipes  (Mereschkowsky,  1879), 
C.  macropus  (Meunier,  1910),  C.  lyngbyi  (Ehrenberg,  1833),  C.  francottei  (Sand,  1895), 
C.  campanula  (Schroder,  1907),  C.  interrupta  (Shroder,  1907)  and  C.  stueri  (Schroder,  1911). 
Kahl  agreed  with  Collin  (1912)  and  placed  the  genus  in  the  family  Discophryidae  where  it  remained 
until  Batisse  (1975)  transferred  it  into  the  Thecacinetidae.  More  recently  Jankowski  (1978)  has 
transferred  three  of  the  species,  which  clearly  have  elongate  to  branched  macronuclei  and  multiple 
endogenous  buds,  into  the  new  genus  Pelagacineta  Jankowski,  1978. 

Genus  ACTINOCYATHULA  Corliss,  1960 

Actinocyathus  Kent,  1882 
Corynophrya  sensu  Batisse,  1975 
Paracineta  sensu  Jankowski,  1978 
Faltacineta  Jankowski,  1982 

The  genus  Actinocyathus  was  erected  by  Kent  (1882)  for  those  resembling  Ephelota  in  general 
form  but  borne  upon  a  stalked  lorica.  Kent's  (1882)  diagnosis  also  stated  that  the  tentacles  were 
retractile  but  not  capitate.  However,  Kent  further  stated  in  his  description  of  the  type  species 
Actinocyathus  cidaris  Kent,  1882  that  he  only  saw  the  tentacles  in  the  contracted  state  which 
leaves  the  absence  of  capitate  tentacles  open  to  considerable  doubt.  There  seems  to  be  little  doubt 
that  the  organism  depicted  by  Dons  (1922)  which  he  calls  Paracineta  crenata  (Fraipont)  forma 
pachyteca  Collin  (Dons  mispelling  ofpachytheca)  is  congeneric  with  Actinocyathus  and  conspecific 
with  Acineta  crenata  Fraipont,  1878.  In  view  of  this  the  two  species  Paracineta  crenata  (Fraipont, 
1878)  and  P.  homari  (Sand,  1899)  which  both  reproduce  by  semi-invaginative  budding  are  trans- 
ferred to  Actinocyathula  Corliss,  1960.  Jankowski  (1982)  erected  the  genus  Faltacineta  Jankowski, 
1982  for  the  two  marine  epizoic  species  Paracineta  pleuromammae  Steuer,  1928  and  Paracineta 
gaetani  Sewell,  1951.  However,  the  former  species  P.  pleuromammae  is  clearly  depicted  showing 
semi-invaginative  budding  and  for  this  reason  the  two  are  transferred  to  Actinocyathula  for  the 
first  time. 

Diagnosis  of  Actinocyathula 

Marine  suctorians  whose  ovoid-shaped  body  is  restricted  to  the  anterior  half  of  the  lorica. 
Lorica  cup-shaped,  never  laterally  compressed,  borne  upon  a  stalk  and  attached  to  marine 
invertebrates  such  as  Crustacea,  hydroid  colonies  and  calcareous  sponges.  Tentacles  in  a  single 
group  that  is  restricted  to  the  apical  region  of  the  body.  Actinophores  absent.  Reproduction  by 
semi-invaginative  budding. 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 

Key  to  the  species  of  Actinocyathula 

1  Stalk  equal  to  or  less  than  lorica  length,  epizoic  on  Crustacea 

Stalk  greater  than  lorica  length,  epizoic  on  invertebrates  other  than  Crustacea 

2  Lorica  smooth 

Lorica  striated  transversely        .        .        .        .        .        ... 

3  Posterior  region  of  lorica  broadly  rounded 

Posterior  of  lorica  distinctly  narrow 

4  Lorica  elongate,  stalk  usually  less  than  half  lorica  length      .... 
Lorica  width  and  stalk  length  approximately  equal  to  lorica  length 


73 


3 

..         .         .2 

.  A.  cidaris 

.  A.  crenata 

.  A.  ho  mar  i 

.       4 

A .  pic  lit- o  mammae 
.  A.gaetani 


Species  descriptions 

Actinocyathula  cidaris  Corliss,  1960 
Actinocyathus  cidaris  Kent,  1882 

DESCRIPTION  (Fig.  1).  This  the  type  species  is  a  small  (40  ^m  long),  marine,  loricate  suctorian.  The 
ovoid  body  has  a  flattened  base  and  protrudes  from  the  apical  region  of  the  lorica.  Tentacles 
retractile,  radiating  from  the  anterior  surface  of  body .  Lorica  surface  smooth,  triangular  in  outline, 
rounded  in  cross-section.  Apical  edge  of  lorica  bends  inwards  to  form  a  thin  cup-like  platform  in 
which  the  zooid  is  located.  Lorica  mounted  on  slender  but  rigid  stalk  that  is  3-4  times  the  lorica 
length.  Epizooic  on  the  calcareous  sponge  Grantia  compressa.  Contractile  vacuole  may  be  single  or 
double.  Nuclear  and  reproductive  features  not  described. 


Fig.  1     Actinocyathula  cidaris  after  Kent,  1 882  (called  Actinocyathus  cidaris). 

Actinocyathula  crenata  n.  comb. 

Acineta  crenata  Fraipont,  1878 

Acineta  saifulae  Mereschkowsky,  1877 

Paracineta  crenata  Collin,  1911 

Paracineta  crenata  var.  pachytheca  Collin,  1912 

Paracineta  crenata  forma  pachyteca  Dons,  1922 

Corynophrya  crenata  Batisse,  1975 

Miracineta  saifulae  Jankowski,  1981 


74 


C.  R.  CURDS 


Fig.  2  Actinocyathula  crenata:  (a-c)  after  Collin,  1912  (called  Paracineta  crenata);  (d,e)  after 
Mereschkowsky,  1877  (called  Acineta  saifulae);  (f)  after  Collin,  1912  (called  Paracineta  crenata); 
(g)  after  Fraipont,  1878  (called  Acineta  crenata);  (h)  after  Dons,  1922  (called  Paracineta  crenata  var. 
pachytheca);  (i)  after  Wailes,  1928  (called  Paracineta  crenata  var.  pachytheca). 


DESCRIPTION  (Fig.  2).  Medium  (75  |im  long),  marine,  loricate  suctorian.  The  ovoid  body  protrudes 
from  the  apical  region  of  the  lorica.  Capitate  tentacles  sometimes  retractile,  radiating  from  the 
anterior  surface  of  body.  Lorica  surface  crenulated  with  three  to  many  transverse  striations, 
triangular  to  elongate  in  outline,  rounded  in  cross-section.  There  is  a  thin  cup-like  platform  in 
which  the  zooid  is  located.  Lorica  mounted  on  slender  stalk  that  is  3-4  times  the  lorica  length. 
Epizooic  on  a  variety  of  marine  invertebrates  including  the  hydroids  Clytia  volubilis,  Leptoscyphus 
grigoriewi  and  Perigonimus  repens  and  the  polychaete  Aphrodite  aculeata.  Single  contractile 
vacuole  located  laterally.  Spherical  macronucleus  centrally  positioned.  Reproduction  by  semi- 
invaginative  budding.  Swarmer  not  described. 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


75 


Fig.  3    Actinocyathula  gataeni:  (a-d)  various  growth  stages;  (e-g)  adults;  all  after  Sewell,  1951  (called 

Paracineta  gataeni). 


A ctinocyathula  gataeni  (Sewell,  1951)  n.  comb. 

Paracineta  gataeni  Sewell,  195 1 
Faltacineta  gataeni  Jankowski,  1 982 

DESCRIPTION  (Fig.  3).  Small  (30-55  um  diameter),  marine,  loricate  suctorian.  The  ovoid  body 
protrudes  from  the  apical  region  of  the  lorica.  Tentacles  radiate  out  from  the  anterior  body  surface. 
Lorica  surface  usually  smooth  but  sometimes  with  transverse  wrinkles,  triangular  in  outline, 
rounded  in  cross-section.  Lorica  mounted  on  a  robust  rigid  stalk  that  is  usually  less  than  the  lorica 
length.  Lorica  sometimes  mounted  eccentrically  on  stalk.  Epizooic  on  the  copepods  Gaetanus 
antarcticus  Wolfendon  and  G.  curvicornis  Sars.  Macronucleus  spherical.  Reproduction  and 
swarmer  not  described. 

Actinocyathula  homari  n.  comb. 

Acineta  homari  Sand,  1899 
Paracineta  homari  Collin,  1911 
Corynophrya  homari  Batisse,  1975 

DESCRIPTION  (Fig.  4).  Small  (25-40  urn  long),  marine,  loricate  suctorian.  The  ovoid  body 
protrudes  from  the  apical  region  of  the  lorica.  Tentacles  retractile,  radiating  out  from  the  anterior 
body  surface.  Lorica  surface  smooth,  triangular  to  bell-shaped  in  outline,  rounded  in  cross- 
section.  Lorica  mounted  on  a  robust  rigid  stalk  that  rarely  exceeds  the  lorica  length.  Lorica 
sometimes  mounted  eccentrically  on  stalk.  Epizooic  on  a  variety  of  decapod  Crustacea.  Single 
contractile  vacuole  located  centrally  or  laterally.  Macronucleus  spherical,  located  at  posterior  of 
body.  Reproduction  by  semi-invaginative  budding.  Swarmer  not  described. 


76 


C.  R.  CURDS 


Fig.  4    Actinocyathula  homari:  (a-d)  after  Collin,  1912  (called  Paracineta  homari);  (e,f)  after  Sand,  1 899 

(called  Acineta  homari}. 


Actinocyathula pleuromammae  (Steuer,  1928)  n.  comb. 

Paracineta  pleuromammae  Steuer,  1928 
Faltacineta  pleuromammae  Jankowski,  1982 

DESCRIPTION  (Fig.  5).  Medium  (60-1 15  jam  long),  marine,  loricate  suctorian.  The  ovoid  body 
protrudes  from  the  apical  region  of  the  lorica.  Tentacles  radiate  out  from  the  anterior  body  surface. 
Lorica  surface  with  irregular  transverse  striations,  elongated  cone,  rounded  in  cross-section. 
Lorica  mounted  on  a  robust  rigid  stalk  that  is  less  than  half  the  lorica  length.  Epizoic  on  the 
copepods  Pleuromamma  abdominalis  and  P.  xiphias.  Single  contractile  vacuole  located  laterally. 
Macronucleus  spherical,  located  centrally.  Reproduction  by  semi-invaginative  budding.  Swarmer 
ovoid  with  many  transverse  ciliary  rows. 

Genus  CORYNOPHRYA  Kahl,  1934 
Pelagacineta  Jankowski,  1 978  pro  pane 

The  genus  was  orginally  erected  by  Kahl  (1934)  to  include  a  heterogenous  collection  of  mainly 
marine  species.  He  stated  that  the  major  features  distinguishing  it  from  other  genera  included 
internal  budding,  a  single  apical  group  of  tentacles  and  a  rounded,  compact  macronucleus.  Kahl 
(1934)  included  eight  species  in  his  genus  but  three  have  recently  been  transferred  to  the  new  genus 
Pelagacineta  by  Jankowski  (1978).  Kahl  (1934)  followed  the  original  higher  classification  system 
of  Collin  (1912)  and  placed  the  genus  in  the  family  Discophryidae  where  it  remained  until  Batisse 
(1975)  transferred  it  into  the  Thecacinetidae  which  demands  reproduction  by  semi-invaginative 
budding.  The  latter  step  was  taken  because  Batisse  (1975)  had  included  Actinocyathula 
(Paracineta)  crenata  and  A.  homari  in  the  genus.  In  fact  the  mode  of  budding  has  only  been 
described  for  one  of  the  five  remaining  species,  where  in  Corynophrya  lyngbyi  it  is  endogenous. 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


77 


10 


Fig.  5    Actinocyathula pleuromammae:  (a-c)  after  Steuer,  1928  (called  Paracineta pleuromammae). 


However,  Jankowski  (1981)  was  recently  of  the  opinion  that  genera  in  his  family  Corynophryidae 
reproduce  exogenously  although  he  gave  no  practical  evidence  for  that  conclusion.  Of  those 
which  Kahl  (1934)  originally  included  in  the  genus  only  four,  Corynophrya  macropus,  C.  conipes, 
C.  lyngbyi  and  C.  francottei  remain  in  the  present  review.  The  anterior  notch  in  the  body  of 
Corynophrya  marina  has  been  interpreted  to  indicate  invaginative  budding  and  will  be  transferred 
to  an  appropriate  genus  in  a  later  publication.  One  other  species,  Ephelota  columbiae  Wailes,  1943 
is  included  in  the  genus  for  the  first  time  since  it  bears  only  one  type  of  tentacle  whereas  there  are 
two  types  in  Ephelota.  The  five  species  that  are  included  have  several  features  in  common,  they  all 
have  a  compact  rounded  macronucleus,  a  single  apical  group  of  tentacles  that  are  both  retractile, 
prehensile  and  suctorial  and  in  most  there  is  a  conical  stalk  that  clearly  narrows  towards  its  base. 
The  species  most  completely  described  is  Corynophrya  lyngbyi  and  this  is  designated  to  be  the  type 
species  in  an  attempt  to  establish  taxonomic  stability. 

Diagnosis  of  Corynophrya 

Mainly  marine,  aloricate  suctorians  whose  body  shape  is  spherical  to  ovoid,  rounded  in  cross 
section.  Borne  upon  a  stalk  which  is  commonly  stout  near  to  the  zooid  narrowing  markedly 
towards  its  base.  Usually  epizooic  on  hydroids,  Crustacea  and  polychaetes  but  also  noted  on 
marine  algae.  Tentacles  prehensile  and  retractile  in  a  single  group  that  is  restricted  to  the 
apical  region  on  the  body.  Actinophores  absent.  Macronucleus  usually  spherical.  Reproduction  by 
endogenous  budding. 


Key  to  the  species  of  Corynophrya 

1       Stalk  long,  at  least  3  times  length  of  body 
Stalk  short,  up  to  twice  length  of  body 


78  C.  R.  CURDS 

2  Freshwater,  tentacles  wide  at  base,  narrowing  towards  capitate  ends    . 
Marine,  sides  of  tentacles  parallel,  do  not  narrow  towards  capitate  ends 

3  Body  spherical  and  regular         ........ 

Body  ovoid,  uneven  with  folds 

4  Stalk  striated  transversely 

Stalk  striated  longitudinally  or  without  striations        .... 

5  Macronucleus  spherical 

Macronucleus  in  shape  of  horseshoe 

6  Stalk  markedly  wider  near  zooid,  narrowing  towards  base  . 
Sides  of  stalk  parallel,  stalk  does  not  narrow  towards  base  . 


.  C.  tumida 
3 

C.  columbine 

C.  symbiotica 

.  C.  conipes 

5 

.       6 

.  C.  lyngbyi 

C.  macropus 

C.francottei 


Species  descriptions 

Corynophrya  lyngbyi  (Ehrenberg,  1833)Kahl,  1934 

Acineta  lyngbyi  Ehrenberg,  1833 

Podophrya  lyngbyei  Claparede  &  Lachmann,  1859  non  Robin,  1879 

Tokophrya  lyngbyei  Biitschli,  1889 

Discophrya  lyngbyei  Collin,  1912 

DESCRIPTION  (Fig.  6).  This  the  type  species  is  a  small  to  medium  (40-80  urn),  marine,  aloricate 
suctorian.  The  ovoid  body  is  oval  in  section  and  slightly  wider  anteriorly.  The  retractile,  capitate 
tentacles  located  on  the  anterior  body  surface.  Stalk  long  (1 20-400  um),  at  least  four  times 


Fig.  6     Corynophrya  lyngbyi  after  Fraipont,  1 878  (called  Podophrya  lyngbyi). 


PARACINETA  AND  COR  YNOPHR  YA  PROBLEM  79 


10 


Fig.  7    Corynophrya  columbiae  after  Wailes,  1943  (called  Ephelota  columbiae}. 


the  body  length.  Stalk  wider  near  zooid  than  at  its  base.  Attached  to  hydroid  colonies  such  as 
Sertularia  and  Clytia  as  well  as  marine  algae.  There  are  one  or  two  contractile  vacuoles. 
Macronucleus  spherical  in  the  young  adult  but  this  elongates  into  a  horse-shoe  shape  at  maturity. 
Reproduction  by  endogenous  budding  which  may  be  multiple.  Swarmer  not  described. 

NOTE.  The  specific  epithet  has  been  consistently  mispelt  by  several  authors  over  many  years. 
Ehrenberg's  (1833)  original  spelling  was  lyngbyi  but  later  (1838)  in  his  atlas  the  name  appears  as 
lyngbyei  and  it  was  this  spelling  that  was  used  by  several  later  authorities. 

Corynophrya  columbiae  n.  comb. 
Ephelota  columbiae  Wailes,  1943 

DESCRIPTION  (Fig.  7).  This  is  a  small  (30-60  um),  marine,  aloricate  suctorian.  The  spherical  to 
ovoid  body  is  round  in  section.  The  retractile,  capitate  tentacles  located  on  the  anterior  half  of 
body  surface.  Stalk  usually  short  (50-200  um),  and  usually  less  than  three  times  the  body  length. 
Stalk  wide  near  zooid  narrowing  towards  the  base.  Attached  to  Crustacea  in  large  numbers. 
Macronucleus  spherical,  centrally  located.  Reproduction  not  described. 

Corynophrya  conipes  (Mereschkowsky,  1877)Kahl,  1934 

Acineta  conipes  Mereschkowsky,  1877 
Podophrya  conipes  Mereschkowsky,  1879 
Tokophrya  conipes  Biitschli,  1889 

DESCRIPTION  (Fig.  8).  This  is  a  large  (100-190  um),  marine,  aloricate  "suctorian.  The  ovoid  to 
pyriform  body  is  oval  in  section  and  widens  anteriorly.  The  retractile,  capitate  tentacles  located 
mainly  on  the  anterior  body  surface.  Stalk  long  (800-1 500  urn),  usually  8-10  times  the  body  length. 
Stalk  distinctly  wider  near  zooid  than  at  its  base,  finely  striated  transversely  and  usually  with  two 
distinct  annuli  situated  about  a  third  of  the  way  down  the  stalk.  Attached  to  marine  algae  such  as 
Ptilota  and  Ceramium.  Single  anterior  contractile  vacuole.  Macronucleus  spherical,  located 
centrally  or  subcentrally.  Reproduction  and  swarmer  not  described. 

Corynophryafrancottei(Sand,  1895)Kahl,  1934 

Tokophrya  francottei  Sand,  1895 
Discophryafrancottei  Collin,  1912 

DESCRIPTION  (Fig.  9).  This  is  a  small  (50-60  um),  marine,  aloricate  suctorian.  The  retractile, 
capitate  tentacles  are  located  on  the  anterior  surface  of  the  spheroidal  body.  Stalk  long 
(1 00-230  um),  at  least  three  times  the  body  length,  retaining  a  constant  diameter  along  its 
entire  length.  Attached  to  hydroid  colonies  such  as  Sertularia  and  Ceramium.  There  is  a  single 


80 


C.  R.  CURDS 


Fig.  8    Corynophrya  conipes:  (a)  after  Mereschkowsky,  1879  (called  Podophrya  conipes);  (b)  after 

Meunier,  1910  (called  Podophrya  conipes). 


10 


Fig.  9     Corynophrya  francottei  after  Sand,  1895  (called  Tokophryafrancottei). 


PAR  A  CINETA  AND  COR  YNOPHR  YA  PROBLEM  8 1 


Fig.  10     Corynophrya  macropus  after  Meunier,  1910  (called  Podophyra  macropus). 

marginal  contractile  vacuole.  Macronucleus  oval  to  spherical,  located  centrally  or  subcentrally. 
Reproduction  and  swarmers  not  described. 

Corynophrya  macropus  (Meunier,  1910)Kahl,  1934 
Podophrya  macropus  Meunier,  1910 

DESCRIPTION  (Fig.  10).  This  is  an  incompletely  defined  species  whose  size  has  not  been  recorded, 
marine,  aloricate.  The  body  is  spherical  in  shape  and  carries  retractile,  capitate  tentacles  on  its 
anterior  surface.  Stalk  long,  at  least  three  times  the  body  length.  Stalk,  which  is  wider  near  the 
zooid  than  at  its  base,  is  distinctly  striated,  longitudinally  along  its  entire  length.  Macronucleus 
spherical,  located  centrally.  Reproduction  and  swarmer  not  described. 

Corynophrya  symbiotica  Jankowski,  1981 

DESCRIPTION  (Fig.  1 1).  This  is  a  medium  (80-105  urn),  marine,  aloricate  suctorian.  The  ovoid  body 
has  rather  bumpy  irregular  appearance  with  some  longitudinal  folds.  The  retractile  tentacles 
occupy  the  entire  domed  anterior  body  surface.  Stalk  comparatively  short  (up  to  90  jam),  about 
same  as  the  body  length.  Stalk  slightly  wider  near  zooid  than  at  its  base.  Attached  to  arctic 
polychaete  worms  belonging  to  the  family  Aphroditidae.  There  is  a  single  anterior  contractile 
vacuole.  Macronucleus  spherical,  located  centrally.  Reproduction  and  swarmer  not  described. 

Corynophrya  tumida  (Gajewskaja,  1933)  Matthes,  1954 
Discophrya  tumida  Gajewskaja,  1933 

DESCRIPTION  (Fig.  12).  This  is  a  small  (50  um),  freshwater,  aloricate  suctorian.  The  ovoid  body  is 
round  in  section  and  slightly  wider  posteriorly.  The  retractile,  capitate  tentacles  are  rather  wider 
at  the  base  and  occupy  the  anterior  half  of  the  body  surface.  Stalk  short  (60-70  urn),  only  just 
longer  than  the  body.  Stalk  wider  near  zooid  than  at  its  base  and  distinctly  striated  transversely 
at  infrequent  intervals  along  its  length.  The  stalk  is  also  irregularly  striated  longitudinally. 


82 


C.  R.  CURDS 


10 


Fig.  1 1     Corynophrya  symbiotica  after  Jankowski,  1 98 1 , 


Fig.  12     Corynophrya  tumida  after  Gajewskaja,  1933  (called  Discophrya  tumidd). 


Attached  to  gammarid  Crustacea  in  Lake  Baikal.  There  is  a  single  anterior  contractile  vacuole. 
Macronucleus  spherical,  located  centrally.  Reproduction  and  swarmer  not  described. 

Genus  PELAGACINETA  Jankowski,  1978 

Schroder  (1907)  first  described  the  two  marine  species  Tokophrya  interrupta  and  T.  campanula 
which  resembled  Ephelota  in  some  respects  and  Podocyathus  in  others.  They  resembled  Ephelota  in 
their  multiple  endogenous  method  of  budding  but  Ephelota  is  without  a  thecostyle  and  has  two 
different  types  of  tentacles.  Similarly  they  resembled  Podocyathus  in  their  overall  structure  but 
reproduced  differently  from  that  genus.  Schroder  (191 1)  later  added  a  further  species  T.  steueri  to 
the  group  but  still  placed  it  in  Tokophrya  a  genus  typified  by  the  absence  of  a  lorica.  Collin  (1912) 
was  the  first  to  transfer  the  three  species  out  of  Tokophrya  and  he  grouped  them  with  several  other 
misfits  into  his  third  section  of  the  genus  Discophrya.  Kahl  (1934)  later  erected  the  new  genus 
Corynophrya  for  Collin's  third  section  where  they  remained  until  the  genus  Pelagacineta  was 


PAR  A  CINETA  AND  COR  YNOPHR  YA  PROBLEM  8  3 

defined  by  Jankowski  (1978)  for  those  species  'like  Podocyathus  but  with  multiple  endogenous 
budding'.  Jankowski  (1978)  designated  P.  interrupta  (Schroder,  1907)  to  be  the  type  species  and 
included  P.  campanula  (Schroder,  1907)  in  the  new  genus.  In  the  current  revision  the  diagnosis  is 
elaborated  for  the  sake  of  clarity  and  some  other  species  are  transferred  to  the  genus  for  the  first 
time. 

Diagnosis  of  Pelagadneta 

Marine  suctoria  with  lorica-like  thecostyle.  Body  shape  ovoid,  discoidal  or  pyriform,  rounded  in 
cross  section,  actinophores  absent.  Stalk  widens  anteriorly  to  form  lorica-like  thecostyle.  Single 
type  of  retractile  tentacle  present,  arranged  in  one  or  two  anterior  groups.  Attached  to  copepods  or 
marine  algae.  Macronucleus  typically  elongate  and  often  branched.  Reproduction  by  multiple 
endogenous  budding.  Swarmers  ovoid  partially  ciliated  with  several  longitudinal  kinetics. 

Key  to  the  species  of  Pelagadneta 

1  Tentacles  in  single  anterior  group P.  campanula 

Tentacles  in  two  anterior  groups 2 

2  Only  2  tentacles  present,  attached  to  algae P.  dibdalteria 

Many  tentacles  present,  attached  to  copepods 3 

3  Macronucleus  elongate  but  not  branched,  body  ovoid  but  not  discoidal       .         .         .       P.  euchaetae 
Macronucleus  elongate  and  branched,  body  sometimes  discoidal         ....       P.  interrupta 


Species  descriptions 
Pelagadneta  interrupta  (Schroder,  1907)  Jankowski,  1978 

Tokophrya  interrupta  Schroder,  1907 
Discophrya  interrupta  Collin,  1912 
Corynophrya  interrupta  Kahl,  1934 

DESCRIPTION  (Fig.  1 3).  This  the  type  species  is  a  medium  (100-140  urn  long),  marine  suctorian  with 
thecostyle.  The  ovoid  body  may  be  dorso-ventrally  compressed  and  discoidal  in  shape  lying  at  the 
top  of  a  thecostyle  that  widens  considerably  to  form  a  lorica-like  anterior  region.  Stalk  region 
hollow,  2-3  times  the  length  of  the  lorica  part  of  the  thecostyle,  terminating  in  a  longitudinally 
striated  basal  disc.  Many  retractile,  capitate  tentacles  located  anteriorly  arranged  in  two  fascicles. 
Attached  to  marine  copepods  such  as  Euchaeta  and  Metridia  reported  from  antarctic  waters. 
Shape  of  macronucleus  variable,  always  elongate  and  frequently  branched.  Reproduction  by 
multiple  endogenous  budding  producing  oval  swarmers  partially  ciliated  with  many  kinetics  on 
part  of  the  ventral  body  surface. 

Pelagadneta  campanula  (Schroder,  1907)  Jankowski,  1978 

Tokophrya  campanula  Schroder,  1907 
Tokophrya  steueri  Schroder,  1911 
Discophrya  campanula  Collin,  1912 
Discophrya  steueri  Collin,  1912 
Corynophrya  campanula  Kahl,  1934 
Corynophrya  steueri  Kahl,  1934 

DESCRIPTION  (Fig.  14).  This  is  a  medium  (100-1 50  um  long),  marine  suctorian  with  thecostyle.  The 
ovoid  body  may  be  dorso-ventrally  compressed  and  discoidal  in  shape  lying  at  the  top  of  a 
thecostyle  that  widens  considerably  to  form  a  cupped  lorica-like  anterior  region.  Stalk  region 
hollow,  1-3  times  the  length  of  the  lorica  part  of  the  thecostyle,  terminating  in  a  longitudinally 
striated  basal  disc.  Many  retractile,  capitate  tentacles  located  anteriorly  arranged  in  a  single 
fascicle  sometimes  surrounded  by  an  outer  ring  of  short  tentacles.  Attached  to  marine  copepods 
such  as  Euchaeta  and  Metridia  reported  from  antarctic  waters.  Shape  of  macronucleus  variable  but 


84 


C.  R.  CURDS 


Fig.  13     Pelagacineta  interrupta:  (a,b)  after  Schroder,  1907  (called  Tokophrya  interruptd). 


always  elongate  and  highly  branched.  Reproduction  by  multiple  endogenous  budding  producing 
oval  swarmers  partially  ciliated  with  many  kineties  on  part  of  the  ventral  body  surface. 

Pelagacineta  dibdalteria  (Parona,  1881),  n.  comb. 
Acineta  dibdalteria  Parona,  1881 

DESCRIPTION  (Fig.  1 5).  This  is  a  small  (50-60  um  long),  marine  suctorian  with  thecostyle.  The  body 
is  pyriform  in  outline,  rounded  in  cross  section  and  lies  at  the  top  of  a  thecostyle  that  widens 
considerably  to  form  a  cupped  lorica-like  anterior  region.  Stalk  region  hollow,  equal  to  or  slightly 
less  than  the  length  of  the  lorica  part  of  the  thecostyle.  There  are  only  two  capitate,  prehensile 
mobile  tentacles,  one  located  anteriorly  on  either  side  of  the  body.  Attached  to  marine  algae. 
Contractile  vacuole  positioned  centrally.  Macronucleus  elongate  sausage-shaped.  Reproduction 
and  swarmers  not  described. 


Pelagacineta  euchaetae  (Sewell,  1951)  n.  comb. 
Acineta  euchaetae  Sewell,  1951 

DESCRIPTION  (Fig.  16).  This  is  a  medium  (80-90  um  diameter),  marine  suctorian  with  thecostyle. 
The  ovoid  body  lies  at  the  top  of  a  thecostyle  that  widens  considerably  to  form  a  lorica-like  anterior 
region.  Young  forms  without  lorica  portion  of  the  thecostyle.  Stalk  region  hollow,  usually  shorter 
than  length  of  the  lorica  part  of  the  thecostyle,  terminating  in  a  longitudinally  striated  basal  disc. 
Many  retractile,  capitate  tentacles  located  anteriorly  arranged  in  two  fascicles.  Attached  to  the 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


85 


Fig.  14     Pelagacineta  campanula:  (a-c)  after  Schroder,  1907  (called  Tokophrya  campanula);  (d,e)  adult 
and  swarmer,  after  Schroder,  1911  (called  Tokophrya  steueri). 


10 


Fig.  15     Pelagacineta  dibdalteria  after  Parona,  1881  (called  Acineta  dibdalteria). 


86 


C.  R.  CURDS 


Fig.  16     Pelagacineta  euchaetae:  various  growth  stages,  after  Sewell,  1951  (called  Acineta  euchaetae). 


marine  copepod  Euchaeta  reported  from  antarctic  waters.  Shape  of  macronucleus  variable,  always 
elongate  and  curved.  Reproduction  by  endogenous  budding  producing  oval  swarmers. 

Genus  PARACINEIA  Collin,  191 1 

Luxophrya  Jankowski,  1978 
Proluxophrya  Jankowski,  1978 
Stemacineta  Jankowski,  1978 

The  genus  Paracineta  Collin,  1911  was  erected  in  order  to  provide  for  those  loricate  suctoria 
with  an  apical  group  of  tentacles  that  reproduced  by  external  budding  and  were  longitudinally 
symmetrical.  The  inclusion  of  Paracineta  crenata,  and  P.  hotnari  which  reproduce  by  semi- 
invaginative  budding  has  already  been  dealt  with  above,  but  even  after  their  removal,  the  species 
included  by  Collin  (1912)  in  the  genus  Paracineta  form  a  heterogenous  group.  Several  other 
transfers  have  been  suggested  and  are  dealt  with  in  other  parts  of  this  paper.  After  the  removal 
of  these  from  the  genus  the  following  four  species  remain  from  Collin's  (1912)  list,  Paracineta 
jorisi  (Sand,  1895),  P.  limbata  (Maupas,  1881),  P.  patula  (Claparede  &  Lachmann,  1861)  and 
P.  vorticelloides  (Fraipont,  1878).  Since  that  time,  one  other  valid  species  has  been  added.  One  of 
the  remaining  major  problems  is  the  lack  of  a  type  species  that  will  give  some  stability  to  the  genus 
and  enable  a  modern  diagnosis  to  be  proposed.  This  omission  is  rectified  here  by  designating 
Paracineta  patula  (Claparede  &  Lachmann,  1861)  Collin,  1911  as  type  species  for  the  genus. 
This  species  is  well  described  and  includes  good  illustrated  accounts  of  the  budding  and  general 


87 

morphology.  Furthermore  it  is  the  only  surviving  species  of  the  three  originally  placed  in  the  genus 
by  Coffin  (1911). 

Diagnosis  of  Paracineta 

Marine  suctorians  whose  body  shape  is  spherical  to  ovoid,  rounded  in  transverse  section.  Long 
thecostyle  with  a  semi-lorica  that  is  variable  in  size.  Semi-lorica  may  be  sufficient  to  enclose  half  the 
zooid's  volume  or  be  reduced  sufficiently  for  the  body  to  be  perched  on  top  of  a  small  cone-like 
widening  at  the  top  of  the  stem.  Capitate  tentacles  usually  restricted  to  apical  body  face  but  may 
radiate  out  from  other  areas  when  the  semi-lorica  is  very  small.  Reproduction  by  exogenous 
budding,  swarmers  covered  in  many  transverse  ciliary  rows. 

Key  to  the  species  of  Paracineta 

1  Zooid  perched  on  top  of  very  small  semi-lorica 2 

Approximately  half  of  zooid  enclosed  within  semi-lorica 4 

2  Tentacles  emerge  from  all  over  zooid 3 

Tentacles  restricted  to  apical  surface P.jorisi 

3  Zooid  with  thick  gelatinous  outer  covering P.  limbata 

Zooid  without  gelatinous  outer  covering P.  vorticelloides 

4  Stem  of  thecostyle  with  narrow  flexible  portion  near  junction  with  zooid      ....    P.patula 
Stem  of  thecostyle  not  narrowed,  not  flexible 5 

5  Thecostyle  striated  transversely  regularly  along  entire  length P.  moebiusi 

Thecostyle  smooth,  unstriated 6 

6  Semi-lorica  with  border-like  rim P.jorisi 

Semi-lorica  without  border-like  rim 7 

7  Small,  (semi-lorica  1 5-25  urn  long),  epizoic  on  polychaetes P.  irregularis 

Medium,  (semi-lorica  30-80  urn  long),  epizoic  on  hydroids  and  marine  algae        .        .        .    P.patula 


Species  descriptions 

Paracineta  pat ula  (Claparede  &  Lachmann,  1861)  Collin,  1911 

Acineta  patula  Claparede  &  Lachmann,  1861 
Acineta  divisa  Fraipont,  1878 
Paracineta  divisa  Kahl,  1934 
Stemacineta  patula  Jankowski,  1978 

DESCRIPTION  (Fig.  17).  This  the  type  species  is  a  small  (50-60  um  long),  marine  suctorian  with  a 
thecostyle.  The  ovoid  to  elongate  body  protrudes  to  a  greater  or  lesser  extent  beyond  the  apical  rim 
of  thecostyle  although  the  latter  is  sufficiently  large  to  enclose  at  least  half  of  the  zooid.  Capitate 
tentacles  not  in  fascicles,  usually  covering  the  apical  surface  of  the  exposed  part  of  the  zooid.  Apical 
part  of  thecostyle  is  triangular,  tapering  posteriorly  to  form  a  hollow  tube-like  stem  that  is  at  least 
three  times  the  length  of  the  lorica-like  part.  The  junction  between  the  two  parts  of  the  thecostyle 
often,  secondarily,  narrowed  and  flexible.  Attached  to  hydroid  colonies  and  marine  algae. 
Single  contractile  vacuole  usually  positioned  laterally.  Spherical  macronucleus  located  centrally. 
Reproduction  by  exogenous  budding  resulting  in  an  ovoid  swarmer  covered  in  transverse  ciliary 
rows  with  some  anterior  short  residual  tentacles. 

NOTE.  The  observation  by  Collin  (1912)  that  the  formation  of  a  narrow  flexible  junction  between 
stem  and  lorica  is  a  secondary  event  allows  the  inclusion  of  Acineta  divisa  Fraipont,  1 878  as  a  junior 
synonym. 

Paracineta  irregularis  Dons,  1928 

DESCRIPTION  (Fig.  18).  This  is  a  small  (15-25  um  long),  marine  suctorian  with  a  thecostyle.  The 
ovoid  to  irregularly  shaped  body  protrudes  to  a  greater  or  lesser  extent  beyond  the  apical  rim  of 
thecostyle  although  the  latter  half  of  the  zooid  is  always  enclosed.  Tentacles  cover  the  apical 


C.  R.  CURDS 


Fig.  17  Paracineta  patula:  (a-c)  after  Collin,  1912;  (d-e)  after  Claparede  &  Lachmann,  1861  (called 
Acineta  patula);  (f)  after  Fraipont,  1877  (called  Acineta  divisa);  (g)  after  Calkins,  1902  (called  Acineta 
divisd). 


surface  of  the  exposed  part  of  the  zooid.  Apical  part  of  thecostyle  irregularly  triangular,  tapering 
posteriorly  to  form  a  rigid  hollow  tube-like  stem  that  is  at  least  half  the  length  of  the  lorica-like 
part.  Epizoic  on  chaetae  of  the  polychaete  worm  Pherusa  plumosa.  Spherical  macronucleus  located 
centrally.  Reproduction  not  described. 

Paracineta  jorisi  (Sand,  1895)  Collin,  1912 
AcinetajorisiSand,  1895 

DESCRIPTION  (Fig.  19).  This  is  a  small  to  medium  (30-80  urn  long),  marine  suctorian  with  a 
thecostyle.  The  ovoid  to  pyriform  body  protrudes  to  a  great  extent  beyond  the  apical  rim  of 
the  semi-lorica  part  of  the  thecostyle  which  is  not  normally  large  enough  to  enclose  the  zooid. 
Tentacles  not  in  fascicles,  usually  covering  the  apical  surface  of  the  exposed  part  of  the  body. 
Apical  part  of  thecostyle  is  triangular  or  cup-like.  The  rim  is  prominently  flared  and  folds  back  on 
itself  to  form  an  internal  layer  upon  which  the  zooid  is  mounted.  Thecostyle  tapers  posteriorly  to 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


89 


Fig.  18    Paracineta  irregularis:  (a-e)  various  forms  after  Dons,  1928. 


Fig.  19     Paracineta  jorisi  after  Sand,  1 895  (called  Acinetajorisi). 


form  a  rigid  hollow  tube-like  stem  that  is  at  least  three  times  the  length  of  the  lorica-like  part. 
Attached  to  hydroid  colonies  such  as  Vesicularia  and  Sertularia.  Single  contractile  vacuole. 
Spherical  macronucleus  located  centrally.  Reproduction  by  exogenous  budding. 


90 


C.  R.  CURDS 


Fig.  20 


Paracineta  limbata:  (a)  adult  with  swarmer,  after  Collin,  1912;  (b)  after  Wailes,  1928;  (c)  after 
Dons,  1922;  (d,e)  after  Moebius,  1888  (called  Podophrya  limbata). 


Paracineta  limbata  (Maupas,  1881)  Collin,  1912 

Podophrya  limbata  Maupas,  1881 
Tokophrya  limbata  Biitschli,  1889 
Paracineta  limbata  forma  convexa  Dons,  1922 
Luxophrya  limbata  Jankowski,  1978 

DESCRIPTION  (Fig.  20).  This  is  a  small  (20-45  um  diameter),  marine  suctorian  with  a  thecostyle. 
The  spherical  body  is  mounted  on  the  rim  of  a  greatly  reduced  lorica-like  part  of  the  thecostyle. 
Zooid  often  covered  by  a  thick  gelatinous  outer  coat.  Capitate  tentacles  not  in  fascicles,  radiate  out 
from  the  entire  surface  of  the  exposed  zooid.  Reduced  apical  part  of  thecostyle  is  cone-like, 
tapering  posteriorly  to  join  a  rigid  hollow  tube-like  stem  that  is  at  least  four  times  the  diameter  of 
the  zooid  in  length.  Attached  to  hydroid  colonies.  Two  contractile  vacuoles  usually  positioned 
laterally.  Spherical  macronucleus  located  centrally.  Reproduction  by  exogenous  budding  resulting 
in  an  ovoid  swarmer  covered  in  transverse  ciliary  rows  with  some  residual  tentacles. 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


91 


Fig.  21     Paracineta  moebiusi  after  Moebius,  1 888  (called  Acineta  crenatd). 


Paracineta  moebiusi  (Moebius,  1888)Kahl,  1934 
Acineta  crenata  Moebius,  1888 

DESCRIPTION  (Fig.  21).  This  is  a  medium  (76  um  long),  marine  suctorian  with  a  thecostyle. 
Approximately  half  the  elongate  body  protrudes  beyond  the  apical  rim  of  thecostyle.  Tentacles  not 
in  fascicles,  covering  only  the  apical  surface  of  the  exposed  part  of  the  zooid.  The  thecostyle  is 
prominently  and  totally  ribbed  transversely.  The  apical  part  is  cup-shaped,  and  tapers  posteriorly 
to  form  a  rigid  hollow  tube-like  stem  that  is  about  one  and  a  half  times  the  length  of  the  lorica- 
like  part.  Epizoic  on  the  crustacean  Holocarus.  Single  anterior  contractile  vacuole.  Spherical 
macronucleus  located  posteriorly.  Reproduction  not  described. 

Paracineta  vorticelloides  (Fraipont,  1877)  Collin,  1912 

Acineta  vorticelloides  Fraipont,  1877 
Proluxophrya  vorticelloides  Jankowski,  1978 

DESCRIPTION  (Fig.  22).  This  is  a  small  (30-40  um  diameter),  marine  suctorian  with  a  thecostyle. 
The  spherical  body  is  mounted  on  the  greatly  reduced  anterior  part  of  the  thecostyle.  Capitate 
tentacles  not  in  fascicles,  radiating  out  from  the  entire  surface  of  the  exposed  zooid.  Reduced  apical 
part  of  thecostyle  is  cup-like,  tapering  posteriorly  to  join  a  rigid  hollow  tube-like  stem  that  is  at 
least  four  times  the  diameter  of  the  body  in  length.  Epizoic  on  hydroid  colonies,  Crustacea  and 
marine  algae.  Single  central  contractile  vacuole.  Spherical  macronucleus  located  posteriorly. 
Reproduction  by  exogenous  budding. 


Genus  LORICOPHRYA  Matthes,  1956 


Acineta  Ehrenberg,  \S33proparte 
Thecacineta  Collin,  \9Q9proparte 
Paracineta  Collin,  191 1  pro  par  te 


92 


C.  R.  CURDS 


Fig.  22    Paracineta  vorticelloides:  (a,b)  after  Fraipont,  1878  (called  Acineta  vorticelloides). 


Corynacineta  Jankowski,  1978 
Heliotheca  Jankowski,  1978 
Paraloricophrya  Jankowski,  1978 
Spongiophrya  Jankowski,  1978 

The  genus  was  originally  erected  by  Matthes  (1956)  for  loricate  suctoria  with  a  single  apical 
group  of  tentacles  but  with  an  unknown  method  of  budding.  He  designated  Loricophrya  parva 
(Schulz,  1932)  as  the  type  species  and  listed  the  following  species  to  constitute  the  genus: 
Loricophrya  cattanei  (Parona,  1883),  L.  simplex  (Maskell,  1886),  L.  lasanicola  (Maskell,  1887), 
L.  tulipa  (Maskell,  1887),  L.  solenophryaformis  (Sand,  1899),  L.  cypridinae  (Collin,  1912), 
L.  caepula  (Penard,  1920),  L.  edmondsoni(Kmg,  1932),  L.  sivertseni  (Allgen,  1951),  L.  trichophora 
(Allgen,  1951)  and  L.  longe-petiolatus  (Allgen,  1951).  The  present  author  does  not  consider  all  of 
these  species  to  be  congeneric  although  the  majority  are  retained  in  this  revision.  The  three  species 
described  by  Maskell  (1886,  1887)  have  already  been  transferred  back  (Curds,  1985)  into  the 
genus  Acineta  but  the  generic  position  of  L.  cattanei  (Parona,  1883)  is  still  uncertain.  Similarly, 
L.  cypridinae  (Collin,  1912)  will  be  returned  back  to  its  original  genus  Thecacineta.  All  the  others 
in  Matthes  (1956)  original  list  have  been  retained  within  the  genus  although  the  specific  epithet 
may  be  different  to  that  used  by  him  and  several  additions  have  been  made. 

Diagnosis  of  Loricophrya 

Freshwater  or  marine  sectoria  with  a  thecostyle.  When  clearly  differentiated  the  stem  is  shorter 
than  the  lorica  part  of  the  thecostyle.  Body  ovoid  to  elongate,  rounded  in  cross-section.  Capitate 
tentacles  restricted  to  a  single  group  on  the  apical  surface  of  the  zooid.  Mode  of  reproduction  not 
yet  recorded. 

Key  to  the  species  of  Loricophrya 

1  Thecostyle  continually  narrows  posteriorly  without  a  stalk  region  being  clearly  differentiated       .       2 
A  narrow  stalk  region  is  clearly  differentiated  from  the  rest  of  the  thecostyle         ....       4 

2  Most  of  zooid  projects  out  of  short  thecostyle L.ovifornus 

Most  of  zooid  enclosed  within  long  thecostyle 3 

3  Zooid  small,  pyriform,  lying  in  apical  quarter  of  thecostyle L.tuba 

Zooid  large,  elongate,  filling  most  of  thecostyle  cavity L.  sivertseni 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


93 


10 


Fig.  23     Loricophrya  parva:  (a,b)  after  Schulz,  1932  (called  Thecacinetaparvd). 


10 


Zooid  longer  than  wide,  never  dorso-ventrally  flattened  nor  discoidal. 
Zooid  wider  than  length,  flattened  dorso-ventrally  or  discoidal  in  shape 

Lorica  part  of  thecostyle  striated  transversely 

Lorica  part  of  thecostyle  without  striations  or  ribs 

Thecostyle  wider  than  height,  covered  in  tubercles 

Thecostyle  longer  than  wide,  smooth 

Stalk  part  of  thecostyle  is  half  length  of  lorica  part,  and  may  be  striated 

Stalk  part  of  thecostyle  very  short,  about  1  /8  of  lorica  part,  not  striated 

Stalk  part  of  thecostyle  striated,  lorica  part  triangular  in  outline  . 

Stalk  part  of  thecostyle  not  striated,  lorica  part  oval  in  outline     . 

Stalk  region  very  short,  about  1/8  length  of  lorica  region.  Rim  without  collar, 

Stalk  region  short,  about  1/2  length  of  lorica  region.  Rim  of  thecostyle 

surrounding  wide  aperture 

Stalk  part  of  thecostyle  conical  in  shape 

Stalk  part  of  thecostyle  tubular 


5 
9 

.     L.  lauterborni 
6 

.  L.  bifaria 
1 
8 

.     L.  multitentaculata 

.    L.  stresemanni 

.    L.  trichophora 

small  aperture  L.  caepula 

with  collar  region 

.      10 

L.  parva 

.  L.  solenophryaformis 


Species  descriptions 

Loricophrya  parva  (Schulz,  1932)  Matthes,  1956 
Thecacineta  parva  Schulz,  1932 

DESCRIPTION  (Fig.  23).  This  the  type  species  is  a  small  (36-41  um  long),  brackish- water  suctorian 
with  a  thecostyle.  The  discoidal  body  is  rounded  in  cross-section  and  lies  within  an  urn-like 
thecostyle.  There  is  a  single  apical  group  of  capitate  tentacles  on  the  apical  surface.  The 
thecostyle  narrows  somewhat  posteriorly  to  form  a  cone-like  stalk  region.  Attached  to  inanimate 
objects.  Single  lateral  contractile  vacuole.  Macronucleus  oval,  centrally  located.  Reproduction 
not  described. 

Loricophrya  bifaria  (Stokes,  1887)  n.  comb. 

Acineta  bifaria  Stokes,  1887 
Paracineta  bifaria  Collin,  1912 
Paraloricophrya  bifaria  Jankowski,  1978 


94 


C.  R.  CURDS 


Fig.  24 


Loricophrya  bifaria:  (a)  adult;  (b)  budding;  (c)  swarmer;  all  after  Stokes,  1887  (called  Acineta 

bifarid). 


Fig.  25    Loricophrya  caepula  after  Penard,  1920  (called  Thecacineta  caepuld). 


DESCRIPTION  (Fig.  24).  This  is  a  small  (45  urn  diameter),  freshwater  suctorian  with  a  thecostyle.  The 
elongate  body  is  rounded  in  cross-section  and  projects  out  well  beyond  the  rim  of  the  thecostyle. 
There  is  a  single  group  of  apical  capitate  tentacles.  Stalk  region  a  short,  button-like  projection. 
Lorica  region  ovoid,  covered  in  tubercles,  width  greater  than  height.  Single  lateral  contractile 
vacuble.  Ovoid  macronucleus  centrally  located.  Reproduction  by  exogenous  budding  resulting  in 
an  elongate  swarmer  with  longitudinal  rows  of  cilia  and  some  residual  tentacles. 

Loricophrya  caepula  (Penard,  1920)  Matthes,  1956 

Thecacineta  caepula  Penard,  1920 
Heliotheca  caepula  Jankowski,  1978 

DESCRIPTION  (Fig.  25).  This  is  a  small  (33  \im  diameter),  freshwater  suctorian  with  a  thecostyle. 
The  ovoid  body  is  rounded  in  cross-section  and  just  projects  out  beyond  the  rim  of  the  thecostyle. 
There  is  a  single  group  of  apical  capitate  tentacles.  Stalk  region  a  short,  button-like  projection. 
Lorica  region  ovoid,  width  greater  than  height.  Single  anterio-lateral  contractile  vacuole.  Ovoid 
macronucleus  centrally  located.  Reproduction  not  described. 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


95 


Fig. '26    Loricophrya  lauterborni:  (a)  after  Sondheim,  1929  (called  Paracineta  lauterborni);  (b)  after 

King,  1932  (called  Thecacineta  edmondsi). 


Loricophrya  lauterborni  (Sondheim,  1929)  n.  comb. 

Paracineta  lauterborni  Sondheim,  1929 
Thecacineta  edmondsi  King,  1932 
Paraloricophrya  lauterborni  Jankowski,  1978 

DESCRIPTION  (Fig.  26).  This  is  a  small  (40-55  um  diameter),  freshwater  suctorian  with  a  thecostyle. 
The  ovoid  body  is  rounded  in  cross-section  and  projects  out  beyond  the  rim  of  the  thecostyle. 
Capitate  tentacles  radiate  out  from  the  surface  of  the  exposed  part  of  the  zooid.  Stalk  region  a 
short,  button-like  projection  or  up  to  half  the  lorica  length.  Lorica  region  cup-like  with  about 
four  transverse  rings.  Attached  to  inanimate  objects.  Two  or  three  contractile  vacuoles.  Ovoid 
macronucleus  centrally  located.  Reproduction  possibly  by  exogenous  budding. 

Loricophrya  multitentaculata  (Sand,  1895)  n.  comb. 

Hallezia  multitentaculata  Sand,  1895 
Acineta  multitentaculata  Sand,  1899 
Paracineta  multitentaculata  Collin,  1912 
Spongiophry a- multitentaculata  Jankowski,  1978 

DESCRIPTION  (Fig.  27).  This  is  a  large  (304  um  long),  marine  suctorian  with  a  thecostyle.  The 
cylindrical  body  is  rounded  in  cross-section  and  only  the  small  posterior  part  is  housed  in  the 
cup-like  thecostyle.  There  is  a  single  apical  group  of  capitate  tentacles  on  the  apical  surface.  The 
thecostyle  follows  the  outline  of  the  body  and  there  is  a  short  button-like  stalk  region.  Epizoic  on 
sponges  such  as  Leucosolenia.  Contractile  vacuole  not  observed.  Macronucleus  large,  elongate, 
centrally  located.  Reproduction  not  described. 


96 


C.  R.  CURDS 


Fig.  27 


Loricophrya  multitentaculata  after  Sand,  1895  (called  Hallezia  multitentaculatd).  Note  that  the 
theca  was  described  but  not  illustrated  in  the  original  description. 


NOTE.  The  presence  of  a  lorica  was  not  shown  in  the  diagram  of  this  species  but  was  mentioned 
clearly  in  the  description.  Here  the  presence  of  a  lorica  is  indicated  means  of  dotted  lines. 

Loricophrya  oviformis  (Dons,  1918)  n.  comb. 
Par  acineta  oviformis  Dons,  1918 

DESCRIPTION  (Fig.  28).  This  is  a  medium  (85  urn  long),  marine  suctorian  with  a  thecostyle.  The 
ovoid  body  is  only  partially  enclosed  within  the  thecostyle  whose  rim  is  smooth.  There  is  a  single 
group  of  tentacles  which  are  scattered  over  much  of  the  exposed  body  surface.  The  thecostyle 
follows  the  outline  of  the  body  posterior  and  there  is  a  short  button-like  stalk-region.  Epizoic  on 
the  worm  Spirorbis.  Nuclear  and  reproductive  features  not  described. 

Loricophrya  sivertseni  (Allgen,  1951)  Matthes,  1956 
Thecacineta  sivertseni  Allgen,  1951 

DESCRIPTION  (Fig.  29).  This  is  a  large  (108um  long),  marine  suctorian  with  a  thecostyle.  The 
elongate  body  is  totally  enclosed  within  the  cone-shaped  thecostyle  whose  rim  is  scalloped. 


Fig.  28     Loricophrya  oviformis  after  Dons,  1918  (called  Paracineta  oviformis). 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


97 


10 


Fig.  29     Loricophrya  sivertseni  after  Allgen,  1951  (called  Thecacineta  sivertseni). 


Fig.  30    Loricophrya  solenophryaformis  after  Sand,  1 899  (called  Acineta  solenophryaformis). 


Capitate  tentacles  in  a  single  apical  group.  There  is  no  distinct  stalk  region,  the  lorica  gradually 
and  continually  narrows  posteriorly  to  join  the  attachment  plate.  Epizoic  on  the  nematode  worm 
Spirina  parasitifera.  Ovoid  macronucleus  centrally  located.  Mode  of  reproduction  not  described. 

Loricophrya  solenophryaformis  (Sand,  1899)  Matthes,  1956 

Acineta  solenophryaformis  Sand,  1899 
Thecacineta  solenophryaformis  Collin,  1909 

DESCRIPTION  (Fig.  30).  This  is  a  small  (30-35  um  long),  freshwater  suctorian  with  a  thecostyle.  The 
discoid  body  is  totally  enclosed  within  an  urn-like  thecostyle  whose  rim  is  surrounded  by  a  collar- 
like  region.  Capitate  tentacles  located  in  a  single,  tightly-packed,  apical  group  which  are  enclosed 
within  the  thecostyle.  There  is  a  short  but  distinct,  tubular  stalk  region.  Attached  to  freshwater 
algae.  Ovoid  macronucleus  located  posteriorly.  Mode  of  reproduction  not  described. 

Loricophrya  stresemanni  (Allgen,  1951)  Matthes,  1956 
Paracineta  stresemanni  Allgen,  1951 
DESCRIPTION  (Fig.  31).  This  is  a  small  (40  urn  long),  marine  suctorian  with  a  thecostyle.  The 


98 


C.  R.  CURDS 


Fig.  31     Loricophrya  stresemanni:  (a,b)  after  Allgen,  1951  (called  Paracineta  stresemanni). 


elongate  body  is  mostly  enclosed  within  a  cone-shaped  thecostyle  whose  rim  is  smooth.  Capitate 
tentacles  in  a  single  apical  group.  There  is  a  distinct  stalk  region  which  is  about  half  the  lorica 
length  and  is  striated  transversely.  Epizoic  on  the  nematode  worm  Spirina  parasitifera.  Ovoid 
macronucleus  centrally  located.  Mode  of  reproduction  not  described. 

Loricophrya  trichophora  (Allgen,  1951)  Matthes,  1956 

Thecacineta  trichophora  Allgen,  1951 
Thecacineta  longe-petiolatus  Allgen,  1951 

DESCRIPTION  (Fig.  32).  This  is  a  medium  (80  um  long),  marine  suctorian  with  a  thecostyle. 
The  elongate  body  is  totally  enclosed  within  an  ovoid  thecostyle  whose  rim  is  smooth.  Capitate 
tentacles  in  a  single  apical  group.  There  is  a  distinct  stalk  region  which  is  about  half  the  lorica 
length,  not  striated.  Epizoic  on  the  nematode  worm  Spirina  parasitifera.  Ovoid  macronucleus 
centrally  located.  Mode  of  reproduction  not  described. 

Loricophrya  tuba  (Zelinka,  1914)  n.  comb. 

Acineta  tuba  Zelinka,  1914 
Paracineta  tuba  Kahl,  1934 
Corynacineta  tuba  Jankowski,  1978 

DESCRIPTION  (Fig.  33).  This  is  a  small  (25-32  um  long),  marine  suctorian  with  a  thecostyle.  The 
pyriform  body  is  enclosed  within  the  apical  quarter  of  the  elongated  cone-like  thecostyle.  Tentacles 
emerge  from  the  apical  surface,  not  in  fascicles.  There  is  no  distinct  stalk  region,  the  lorica 
gradually  and  continually  narrows  posteriorly  to  join  the  substratum.  Epizoic  on  the  shells  of 
echinoderms.  Ovoid  macronucleus  centrally  located.  Mode  of  reproduction  not  described. 

Genus  ANTH ACINETA  Jankowski,  1978 

Acineta  Ehrenberg,  1833  pro  pane 
Noracine  ta  Jankowski,  1978 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


99 


10 


Fig.  32     Loricophrya  trichophora  after  Allgen,  1951  (called  Paracineta  trichophord). 


10 


Fig.  33    Loricophrya  tuba  after  Zelinka,  1914  (called  Acineta  tuba). 


The  genus  Anthacineta  was  erected  by  Jankowski  (1978)  for  Acineta  craterellus  Collin,  1909  giving 
the  following  brief  diagnosis  'semi-lorica  -  stylotheca'.  According  to  that  brief  definition  the  genus 
could  be  transferred  to  Paracineta  and  several  other  similar  genera  as  a  junior  synonym.  It  can  only 
be  classified  as  a  distinct  genus  if  the  two  fascicles  of  tentacles  and  rounded  transverse  section  to  the 
body  are  taken  into  account.  Here  the  diagnosis  has  been  expanded  and  one  other  species,  Acineta 
infundibuliformis  Wang  &  Nie,  1933,  has  been  transferred  to  it  for  the  first  time. 

Diagnosis  of  Anthacineta 

Marine  suctorians  with  thecostyle.  Zooid  only  partly  enclosed  in  the  semi-lorica  part  of  the 
thecostyle  which  has  a  long  stem.  Body  ovoid,  rounded  in  cross-section.  Two  fascicles  of  capitate 
tentacles  present,  one  either  side  of  the  zooid.  Mode  of  reproduction  not  recorded. 


100 


C.  R.  CURDS 


JO. 


Fig.  34    Anthacineta  craterellus  after  Collin,  1912  (called  Acineta  craterellus). 


Key  to  the  species  of  Anthacineta 

1       Zooid  wider  than  long,  contractile  vacuole  in  posterior  body  half.  Thecostyle  narrows  abruptly  to  form 

stalk-like  region     . A.  infundibuliformis 

Zooid  longer  than  wide,  contractile  vacuole  in  anterior  body  half.  Thecostyle  narrows  consistently  to 
form  the  stalk-like  region 5.  craterellus 


Species  descriptions 

Anthacineta  craterellus  (Collin,  1909)  Jankowski,  1978 

Acineta  tuber osa  Sand,  1901  propane 
Acineta  craterellus  Collin,  1909 

DESCRIPTION  (Fig.  34).  This  the  type  species  is  a  small  (50  ^m  long),  marine  suctorian  with  a 
thecostyle.  The  ovoid  to  pyriform  body  is  rounded  in  cross-section  and  about  half  of  it  protrudes 
beyond  the  rim  of  the  semi-lorica  part  of  the  thecostyle.  There  are  two  anterio-lateral  fascicles 
of  capitate  tentacles.  The  lorica  part  of  the  thecostyle  is  short  and  cone-like,  narrowing  gently 
posteriorly  to  form  the  hollow  stem  region  that  is  at  least  twice  the  length  of  the  zooid.  Epizoic  on 
bryozoa.  Single  contractile  vacuole  situated  apically  between  fascicles.  Spherical  macronucleus 
centrally  located.  Reproduction  not  described. 

Anthacineta  infundibuliformis  (Wang  &  Nie,  1933)  n.  comb. 

Acineta  infundibuliformis  Wang  &  Nie,  1933 
Noracineta  infundibuliformis  Jankowski,  1978 

DESCRIPTION  (Fig.  35).  This  is  a  small  (50  um  long),  marine  suctorian  with  a  thecostyle.  The 
wedge-shaped  body  is  rounded  in  cross-section  and  about  half  of  it  protrudes  beyond  the  rim  of  the 
semi-lorica  part  of  the  thecostyle.  There  are  two  lateral  fascicles  of  capitate  tentacles.  The  lorica 
part  of  the  thecostyle  is  short  and  cone-like,  narrowing  abruptly  posteriorly  to  form  the  hollow 
stem  region  that  is  about  the  length  of  the  lorica.  Attached  to  marine  algae.  Single  contractile 
vacuole  situated  posteriorly.  Ovoid  macronucleus  centrally  located.  Reproduction  not  described. 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


101 


10 


Fig.  35     Anthacineta  infundibuliformis  after  Wang  &  Nie,  1933  (called  Acineta  infundibuliformis). 


Genus  FLECTACINETA  Jankowski,  1978 

Acineta  Ehrenberg,  \S33proparte 
Podophrya  Ehrenberg,  1833  pro  par  te 
Alderia  Alder,  1851 
Paracineta  Collin,  191 1  pro  par  te 

The  genus  was  erected  by  Jankowski  (1978)  for  Acineta  livadiana  Mereschowsky,  1881  who  gave 
the  following  brief  diagnosis,  'with  stylotheca  and  apical  tentacles'.  The  stalk  is  normally  shown  as 
being  hollow  but  not  as  an  extension  of  the  lorica  as  the  term  stylotheca  implies.  Thus  the  diagnosis 
has  been  emended  slightly  and  expanded  for  the  sake  of  clarity.  Two  species  Paracineta  dadyi 
(Daday,  1886)  Kahl,  1934  and  Acineta  elegans  Imhoff,  1883  have  been  transferred  to  the  genus  for 
the  first  time. 

Diagnosis  of  Flectacineta 

Marine  loricate  suctorians.  Ovoid  body,  rounded  in  cross-section  lying  within  lorica.  Capitate 
tentacles  restricted  to  single  apical  group.  Thecostyle  lorica  rim  characteristically  inverted  at  apex, 
mounted  upon  a  hollow  stalk.  Reproduction  by  exogenous  budding. 


Key  to  the  species  of  Flectacineta 

1  Rim  of  lorica  smooth,  junction  between  stalk  and  lorica  simple    . 
Rim  of  lorica  scalloped,  junction  between  stalk  and  lorica  complex 

2  Wall  or  lorica  divided  into  an  inner  and  outer  wall  near  aperture . 
Wall  of  lorica  not  divided  . 


.  F.  elegans 

F.  dadyi 

F.  livadiana 


Species  descriptions 
Flectacineta  livadiana  (Mereschkowsky,  1881)  Jankowski,  1978 


Cothurnia  havniensis  Ehrenberg,  1838 
Alderia  pyriformis  Alder,  1851 
Podophrya  pyriformis  Pritchard,  1861 
Acineta  livadiana  Mereschkowsky,  1881 
Acineta  neapolitana  Daday,  1886 
Acineta  sp.  Robin,  1879 
Paracineta  neapolitana  Kahl,  1934 


102 


C.  R.  CURDS 


Fig.  36  Flectacineta  livadiana:  (a)  after  Sand,  1895  (called  Acineta  livadiana);  after  Mereschkowsky, 
1881  (called  Acineta  livadiana);  (c)  after  Wang  &  Nie,  1933  (called  Acineta  livadiana);  (d)  after  Daday, 
1886  (called  Acineta  neapolitana). 


DESCRIPTION  (Fig.  36).  This  the  type  species  is  a  small  to  medium  (30-80  um  long),  marine,  loricate 
suctorian.  The  small  ovoid  body  is  rounded  in  cross-section  and  is  completely  enclosed  within  the 
lorica.  There  is  a  single  apical  group  of  capitate  tentacles.  The  lorica  is  ovoid  with  an  inverted  rim 
that  forms  a  small  aperture.  The  stem  region  is  distinct  and  most  diagrams  show  that  there  is 
usually  at  least  a  narrow  channel  through  the  centre.  Length  of  stem  variable.  Epizoic  on  hydroids 
and  marine  algae.  Single  contractile  vacuole  situated  laterally.  Ovoid  macronucleus  centrally 
located.  Reproduction  by  exogenous  budding. 

Flectacineta  dadayi  (Daday,  1886)  n.  comb. 

Acineta  livadiana  Daday,  1886 
Paracineta  livadiana  Collin,  1912  pro  pane 
ParacinetadadayiKahl,  1934 

DESCRIPTION  (Fig.  37).  This  is  a  small  (45  urn  long),  marine,  loricate  suctorian.  The  small  ovoid 
body  is  rounded  in  cross-section  and  is  completely  enclosed  within  the  lorica.  There  is  a  single 
apical  group  of  capitate  tentacles.  The  lorica  is  ovoid  to  cone-shaped  with  an  inverted  rim  that 
forms  a  small  aperture.  The  lorica  surrounding  the  aperture  is  divided  into  an  inner  and  an 
outer  wall.  The  hollow  stem  region  is  distinct  and  some  diagrams  show  that  there  is  a  narrow 


PARACINETA  AND  CORYNOPHRYA  PROBLEM 


103 


Fig.  37    Flectadneta  dadayi:  (a)  after  Daday,  1886  (called  Acineta  livadiana);  (b)  after  Collin,  1912 

(called  Paracineta  livadiana}. 


Fig.  38    Flectadneta  elegans  after  Imhoff,  1 884  (called  Acineta  elegans). 

channel  through  the  centre.  Length  of  stem  region  about  that  of  lorica.  Epizoic  on  hydroids  and 
marine  algae.  Single  contractile  vacuole  situated  laterally.  Ovoid  macronucleus  centrally  located. 
Reproduction  not  described. 


104  C.  R.  CURDS 

Flectacineta  elegans  (Imhoff,  1883)  n.  comb. 

Acineta  elegans  Imhoff,  1883  non  Maskell,  1886 
Paracineta  elegans  Collin,  1912 

DESCRIPTION  (Fig.  38).  This  is  a  medium  (70  jim  long),  marine,  loricate  suctorian.  The  rectangular 
body  is  rounded  in  cross-section  and  is  completely  enclosed  within  the  lorica.  There  is  a  single 
apical  group  of  capitate  tentacles.  The  lorica  is  pyriform  with  an  inverted  scalloped  rim  that 
forms  a  small  aperture.  The  hollow  stem  region  is  distinct  and  joins  the  lorica  via  an  intervening 
ball-like  joint.  Length  of  stem  at  least  twice  that  of  the  lorica.  Epizoic  on  the  cladoceran 
Bythotrephes  longimanus.  Single  apical  contractile  vacuole.  Ovoid  macronucleus  centrally  located. 
Reproduction  not  described. 


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106 


C.  R.  CURDS 


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Gesellschaft  Deutscher  Naturforscher.  Leipzig  85: 680-683. 

Manuscript  accepted  for  publication  7  April  1986 


Index  to  Species 

Names  given  in  roman  refer  to  synonyms 


Acineta  bifaria  93 

conipes  79 

craterellus  100 

crenata  Fraipont  73 

crenata  Moebius  90 

dibdalteria  84 

divisa  87 

elegans  104 

euchaetae  84 

homari  75 

infundibuliformis  100 

jorisi  88 

livadiana  Daday  102 

livadiana  Mereschkowsky  101 

lyngbyi  78 

multitentaculata  95 

neopolitana  101 

patula  87 

sai  fulae  73 

solenophryaformis  97 

sp.  Robin  101 

tuba  98 

tuberosa  100 

vorticelloides91 
Actinocyathula  cidaris  73 

crenata  73 

gaetani  75 

homari  75 

pleuromammae  76 
Actinocyathus  cidaris  73 
Alderia  pyriformis  101 
Anthacineta  craterellus  100 

infundibuliformis  100 

Corynophrya  campanula  83 
columbiae  79 
conipes  79 
crenata  73 
francottei  79 
homari  75 
interrupta  83 
lyngbyi  78 
macropus  8 1 
steueri  83 
symbiotica%\ 
tuba  98 


tumida  8 1 
Cothurnia  havniensis  101 

Discophrya  campanula  83 
francottei  79 
interrupta  83 
lyngbyei  78 
steueri  83 
tumida  8 1 

Ephelota  columbiae  79 

Faltacineta  gataeni  75 

pleuromammae  76 
Flectacineta  dadayi  102 

elegans  104 

livadiana  101 

Hallezia  multitentaculata  95 
Heliotheca  caepula  94 

Loricophrya  bifaria  93 

caepula  94 

lauterborni  95 

multitentaculata  95 

oviformis  96 

parva  93 

sivertseni  96 

solenophryaformis  97 

stresemanni  97 

trichophora  98 

tuba  98 
Luxophrya  limbata  90 

Miracineta  saifulae  73 
Noracineta  infundibuliformis  100 

Paracineta  bifaria  93 
crenata  73 

crenata  forma  pachytheca  73 
crenata  var.  pachytheca  73 
dadayi  102 
divisa  87 
elegans  104 
gataeni  75 


homari  75 

irregularis  87 

jorisi  88 

lauterborni  95 

limbata  90 

limbata  forma  convexa  90 

livadiana  102 

moebiusi9\ 

multitentaculata  95 

neapolitana  101 

oviformis  96 

patula  87 

pleuromammae  76 

stresemanni  97 

tuba  98 

vorticelloides  91 
Paraloricophrya  bifaria  93 

lauterborni  95 
Pelagacineta  campanula  83 

dibdalteria  84 

euchaetae  84 

interrupta  83 
Podophrya  conipes  79 

limbata  88 

lyngbyei  78 

macropus  81 

pyriformis  101 
Proluxophrya  vorticelloides  91 

Spongiophrya  multitentaculata  95 
Stemacineta  patula  87 

Thecacineta  caepula  94 

edmondsi  95 

longe-petiolatus  98 

parva  93 

sivertseni  96 

solenophryaformis  97 

trichophora  98 
Tokophrya  campanula  83 

conipes  79 

francottei  79 

interrupta  83 

limbata  88 

lyngbyei  78 

steueri  83 


C.  G.  Ogden  &  R.  H.  Hedley 


1980,  Hardcovers,  222pp,  £17.50  (£18.00  by  post)  Co-published  by  British  Museum  (Natural  History)  and 
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Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitm  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
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systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

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Notes  on  spiders  of  the  family  Salticidae. 
1 .  The  genera  Spartaeus,  Mintonia  and 
Taraxella 


F.  R.  Wanless 


Zoology  series    Vol52    No  3    26  March  1987 


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ISSN  0007-1 498  Zoology  series 

Vol52  No.  3  pp  107-1 37 
British  Museum  (Natural  History) 
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Notes  on  spiders  of  the  family  Salticida^. 

Ir^.1  o  » J.  .         *jj  ^  2  6nAS.i7o7 

.  1  he  genera  bpart aeus,  Mintonia  and;  Taraxi 


F.  R.  Wanless 

Department  of  Zoology,  British  Museum  (Natural  History), 

"X£^   ^ffAl     Hl^>  ' 

&^ 

I 

Synopsis 

The  genus  Taraxella  is  redefined  to  include  those  spartaeines  in  which  the  embolus  of  the  male  palp  is  largely 
obscured  by  tegular  apophyses.  The  male  of  Spartaeus  thailandica  Wanless  and  the  female  of  Mintonia 
melinauensis  Wanless  are  described  for  the  first  time.  One  new  species  of  Spartaeus,  two  new  species  of 
Mintonia  and  four  new  species  of  Taraxella  are  described  from  the  Oriental  Region.  Diagnoses  and  figures  are 
provided.  The  presence  of  mytiliform  organs  on  the  legs  and  filamentous  metatarsal  leg  fringes  are  described 
for  the  first  time.  They  are  illustrated,  together  with  femoral  organs,  muscle  attachment  sites  and  apophyses 
by  scanning  electron  micrographs. 


Introduction 

The  purpose  of  the  present  paper  is  to  describe  seven  new  species  and  the  previously  unknown  male 
and  female  of  two  described  species  belonging  in  the  subfamily  Spartaeinae.  The  subfamily  is  of 
particular  interest  because  some  species  spin  large  webs  that  are  used  to  capture  prey.  Typical 
salticids  are  cursorial  hunters  with  good  vision  that  do  not  spin  webs  to  capture  prey,  although  they 
will  spin  silk  nests  in  which  to  lay  eggs,  moult  and  sometimes  mate,  and  generally  rest  at  night  or 
during  other  periods  of  inactivity  (Jackson,  1979). 

The  first  reports  of  web-spinning  in  jumping  spiders  (Coleman,  1978, 1980;  Murphy,  in  Wanless, 
19786)  were  followed  by  several  important  studies  (Jackson  &  Blest,  1982;  Jackson,  1982;  Jackson 
&  Hallas,  in  press  a)  that  confirmed  the  phenomenon  and  provided  a  rare  insight  into  the  biology  of 
a  small  group  of  tropical  salticids.  These  spiders  all  belonged  in  the  old  world  genus,  Portia  Karsch, 
that  is  presently  classified  along  with  12  other  genera  in  the  subfamily  Spartaeinae.  In  addition  to 
building  large  prey-capture  webs,  Portia  species  will  leave  their  web  and  stalk  prey  as  cursorial 
hunters  i.e.  in  the  same  manner  as  other  salticids.  Furthermore,  they  may  invade  the  webs  of  other 
spiders  and  feed  on  trapped  insects  (kleptoparasitism),  the  resident  spider,  or  even  its  eggs 
(oophagy).  In  life  they  resemble  tatty  mouldy  leaves  or  detritus,  their  ornate  hair  tufts  and  fringes 
(Fig.  3)  providing  a  form  of  camouflage  that  enables  them  to  stalk  prey  without  being  noticed,  an 
important  guise  since  they  show  a  marked  preference  for  other  spiders,  including  salticids.  Portia 
species  are  also  'aggressive  vibratory  mimics'  for  when  they  invade  other  spider  webs  they  pluck 
the  threads  and  deceive  the  owner  into  accepting  Portia  as  potential  prey  only  to  be  attacked 
themselves  on  approaching  within  jumping  distance. 

Occurrence  of  the  unusual  behaviour  patterns  of  Portia  species  correlate  to  some  degree  in  this 
and  related  spartaeines  by  the  presence  of  morphological  structures  that  are  not  known  to  occur  in 
other  spiders.  These  include  femoral  organs  (Figs  5E;  14A)  and  pore-bearing  apophyses  (Figs  9E 
arrowed;  20 A,  B)  both  of  which  are  especially  evident  in  some  of  the  species  described  below.  Also 
present  on  the  legs  of  one  species  (Spartaeus  wildtrackii  sp.  n.)  are  mytiliform  organs  (Fig.  15A-C; 
16A,  B),  structures  that  have  hitherto  only  been  found  grouped  together  as  a  discrete  patch  on  the 
dorsal  surface  of  the  abdomen  (Fig.  20C)  of  species  ofCyrba  Simon,  Portia,  Gelotia  Thorell,  and 
Mintonia  Wanless.  The  function  of  these  structures  is  unknown  but  previous  studies  (Wanless, 
1984a,  b;  1985)  have  suggested  that  they  may  be  associated  with  pheromone  dispersal.  Jackson  & 
Hallas  (in  press  b}  have  demonstrated  that  sex  pheromones  are  involved  in  mate  recognition  in 
some  Portia,  Brettus  Thorell  and  Cyrba,  thus  supporting  earlier  work  by  Legendre  and  Llinares, 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  52(3):  107-137  Issued  26  March  1987 

107 


108 


F.  R.  WANLESS 


Fig.  1.    (above)  Spartaeus  wildtrackii  sp.  n.  Subadult  <$,  under  web  on  surface  of  tree  trunk.  Fig.  2 

(below)  web  of  Spartaeus  wildtrackii  sp.  n. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


109 


Fig.  3.     Portia  labiata  (Thorell),  9  from  Malaysia. 

1970  who  noted  that  in  Cyrba  algerina  (Lucas)  pheromones  left  by  the  female  stimulate  the  male 
and  elicit  searching  behaviour. 

Another  feature  clearly  demonstrated  by  some  of  the  species  described  below  is  the  disparity  in 
size  of  the  posterior  median  eyes.  In  most  spartaeines  and  also  in  a  few  other  subfamilies  the 
posterior  median  eyes  are  relatively  large  (Fig.  5A,  B)  and  fully  functional,  whereas  in  most 
jumping  spiders  they  are  relatively  small  (Fig.  10A,  B)  with  no  demonstrable  function.  It  is 
therefore  of  interest  to  note  that  in  the  species  of  Taraxella  Wanless,  described  below  there  is  a  clear 
trend  towards  the  development  of  small  posterior  median  eyes.  In  practice  the  distinction  between 
relatively  large  or  relatively  small  becomes  somewhat  blurred  in  Taraxella,  although  there  has 
hitherto  been  no  difficulty  in  assigning  one  state  or  the  other. 

The  presence  of  large  posterior  median  eyes  and  web  spinning  behaviour  in  Portia  species  gave 
rise  to  the  hypothesis  (Jackson  &  Blest,  1982)  that  the  ancestors  of  modern  day  salticids  evolved 
from  web  building  spiders  with  poorly  developed  vision  and  that  acute  vision,  evolved  originally  in 
a  spider  like  Portia  that  became  an  araneophagic  predator,  proficient  at  invading  diverse  types  of 
webs.  Subsequent  papers  on  morphology  (Wanless,  19840,  b)  the  evolution  of  salticid  eyes  (Blest, 
1984;  Blest  &  Sigmund,  1985)  and  behaviour  (Jackson,  19856;  Jackson  &  Hallas,  in  press  a,  b) 
have  given  support  to  the  hypothesis  and  suggest  that  although  spartaeine  salticids  are  highly 
specialized  they  may  nevertheless  represent  one  of  the  most  primitive  branches  of  the  family. 

Studies,  however,  are  at  an  early  stage  for  the  biology  of  tropical  salticids  is  very  poorly  known, 
in  fact  the  majority  of  species  cannot  even  be  identified  with  confidence.  Also,  it  is  important  to 
appreciate  that  the  subfamily  Spartaeinae,  with  more  than  60  described  species,  represents  less 
than  1-5%  of  the  worlds  salticid  fauna.  Despite  this  there  are  indications  that  jumping  spiders 
make  far  more  use  of  silk  than  has  hitherto  been  supposed,  for  recent  studies  on  other  salticid 
groups  have  revealed  species  that  build  large  prey-capture  webs  (Jackson,  19850)  and  even  species 
that  live  in  groups  forming  nest  complexes  within  the  webs  of  other  spiders  (Jackson,  in  press). 


110  F.  R.  WANLESS 

The  standard  abbreviations  and  measurements  are  those  made  by  Wanless  (1978)  but  for  the  leg 
spination  the  system  adopted  is  that  used  by  Platnick  &  Shadab  ( 1 975).  Note  also,  that  the  covering 
hairs  on  the  male  palps  are  not  shown  in  any  of  the  figures,  because  they  are  usually  rather  dense 
and  obscure  details. 

Genus  SPA  R  TA  BUS  Thorell 

Spartaeus:  Wanless  1984a:  147  [synonymy,  definition  and  species  descriptions].  Blest  &  Sigmund,  1985:  129. 
Blest,  1985:96. 

Spartaeus  is  a  small  oriental  genus  comprising  three  species,  S.  spinimanus  Thorell,  from  Indonesia, 
Malaysia  and  Sri  Lanka,  S.  thailandicus  Wanless  from  Thailand  and  S.  wildtrackii  sp.  n.,  from 
Malaysia.  They  are  easily  distinguished  by  the  structure  of  the  genitalia. 

All  three  species  possess  relatively  large  posterior  median  eyes  (Fig.  4A),  unusually  long  slender 
legs  bearing  numerous  spines,  and  well  developed  femoral  organs  (Fig.  4E;  5E;  14A-C)  on  the  first 
pair  of  legs  of  adult  males.  Also  present  on  the  legs  of  both  males  and  females  are  disc-like 
mytiliform  organs  (Figs  15C;  16B).  Those  on  the  femora  are  more  or  less  rounded  and  sparsely 
distributed  (Fig.  16A),  whereas  those  on  the  tibiae  are  ovoid  and  located  distally  on  the  dorsal 
surface  of  the  segment  (Fig.  15A,  B).  As  mentioned  above,  mytiliform  organs  have  hitherto  only 
been  found  in  the  form  of  a  patch  on  the  dorsal  surface  of  the  abdomen.  Their  occurrence  on  legs  is 
therefore  of  particular  interest,  especially  as  some  are  grouped  on  the  distal  end  of  the  tibiae,  an 
arrangement  that  may  be  unique  to  Spartaeus.  However,  at  present  these  structures  cannot  be 
used  to  determine  relationships  as  their  distribution  on  the  legs  of  other  salticids  is  unknown. 
Furthermore,  they  are  almost  certainly  homologous  with  pustuliform  organs  (see  Hill,  1977  and 
Wanless,  1984)  that  occur  as  scattered  pore-bearing  pustules  on  the  abdomen,  legs  and  pedipalps 
of  Icius  Simon,  Metaphidippus  F.O.P-C  and  Phidippus  Koch,  and  as  a  group  on  the  abdomen  of 
Holcolaetis  Simon  and  Sonoita  Peckham  and  Peckham. 

The  dorsal  surface  of  the  tibiae  are  characterised  by  hinge  lines  or  rows  of  muscle  attachment 
sites  (Fig.  15A,  B)  that  are  electron  dense  and  smooth  in  contrast  to  the  surrounding  cuticle.  They 
resemble  mytiliform  organs,  but  lack  pores  and  raised  rims.  It  is  also  worth  noting  that  they  differ 
from  the  rows  of  triangular  muscle  attachment  sites  found  on  the  tibiae  and  other  leg  segments  of 
Holcolaetis  species  (see  Wanless,  1985). 

The  biology  of  Spartaeus  is  unknown  except  for  observations  (see  below)  made  by  Mr  P.  D. 
Hillyard  (BMNH)  who  also  provided  the  photographs  (Fig.  1 ;  2)  showing  S.  wildtrackii  beneath  its 
sheet  web  on  a  tree  trunk. 

Spartaeus  thailandicus  Wanless 
(Fig.  4A-I) 

Spartaeus  thailandica  Wanless,  1984a:  151.  Holotype  ?,  Thailand,  BMNH,  [examined]. 

DIAGNOSIS.  5".  thailandicus  seems  to  be  most  closely  related  to  S.  wildtrackii  sp.  n.,  but  may  be 
distinguished  by  the  presence  of  a  tegular  apophysis  (Fig.  4G,  arrow)  and  the  form  of  the  retro- 
lateral  tibial  apophysis  (Fig.  4F)  in  males;  females  are  separated  by  the  absence  of  a  postepigynal 
furrow  (see  Wanless  \9S4a,  Fig.  5D). 

MALE,  in  good  condition,  from  Khas  Yai  National  Park,  Thailand.  Carapace  (Fig.  4A-D):  weakly 
iridescent  under  some  angles  of  illumination;  orange-brown  lightly  tinged  and  mottled  black  with  a 
broad  tapering  pale  yellow  band  on  thoracic  part;  rubbed  except  for  some  black  and  pale  amber 
hairs  on  sides.  Eyes:  laterals  with  black  surrounds;  fringed  by  white  hairs.  Clypeus:  yellow-brown 
with  sooty  markings;  sparsely  clothed  in  black  and  whitish  hairs.  Chelicerae:  long  robust  and 
diverging;  posterior  surface  with  series  of  transverse  furrows;  orange-brown  lightly  tinged  black; 
sparsely  clothed  in  black  hairs  with  dense  promarginal  scopula;  fang  robust  and  curved;  fang 
groove  with  eight  promarginal  teeth  and  12  retromarginal  denticles.  Maxillae  and  labium:  pale 
brownish  yellow  faintly  tinged  with  some  black.  Sternum  (Fig.  4B):  pale  yellow-brown  with  darker 
margins.  Coxae:  pale  yellow-brown  except  for  blackish  sides  of  I  and  II.  Abdomen:  pale  yellow- 
brown  with  black  markings,  ventrally  a  broad  sooty  band  from  epigastric  furrow  to  spinnerets; 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


111 


Fig.  4.  Spartaeus  thailandicus  Wanless  £,  A,  dorsal  view;  B,  sternum;  C,  chelicerae,  maxillae  and 
labium;  D,  carapace,  lateral  view;  E,  leg  I;  F,  palp,  retrolateral  view;  G,  palp,  ventral  view;  H,  cheliceral 
teeth;  I,  palpal  tibia,  dorsal  view.  Abbreviations:  e,  embolus;  fo,  femoral  organ;  Ml,  fan-shaped 
element  of  distal  haematodocha. 


112  F.  R.  WANLESS 

rubbed;  spinnerets  long  and  robust.  Legs:  very  long  and  slender,  femoral  organ  pronounced;  legs  I 
yellow-brown  except  for  blackish  streaks  on  sides  of  femora,  a  blackish  tinge  towards  apices  of 
metatarsi  and  whitish  yellow  tarsi;  other  legs  whitish  yellow  to  yellow-brown  with  apices  of 
metatarsi  and  tarsi  tinged  black;  clothed  in  scattered  simple  and  feathery  hairs — mostly  rubbed; 
tarsi  I-II  and  apices  of  metatarsi  I  with  pro  ventral  row  of  specialised  prey-capture  setae;  metatarsi 
II-III  with  basal  fringe  of  fine  curved  filamentous  setae.  Spines  strong  and  numerous;  spination  of 
leg  I:  metatarsus  v  2^M),  tibia  v  4-6-7,  femur  p  1-1-1,  d  0-2-2,  r  0-1-1.  Palp  (Fig.  4F,  G,  I): 
element  M 1  of  the  distal  haematodocha  (see  Wanless,  19840)  fan-shaped  (Fig.  4G);  element  M2  is 
obscure  and  appears  to  have  fused  with  the  base  of  the  embolus. 

Dimensions  (mm):  total  length  8-5;  carapace  length  3-84,  breadth  3-14,  height  2-75;  abdomen 
length  4-6;  eyes,  anterior  row  2-7,  middle  row  2-04,  posterior  row  2-32;  quadrangle  length  1-9 
(49%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

6-40 

3-84 

3-56 

4-56 

1-60 

Patella 

2-60 

1-72 

1-44 

1-60 

0-88 

Tibia 

7-04 

3-32 

3-16 

4-28 

0-56 

Metatarsus 

5-36 

3-16 

3-64 

5-28 

Tarsus 

1-96 

1-14 

1-16 

1-36 

1-68 

Total  23-36        13-18        12-96        17-08         4-72 


Ratios:  AM  :  AL  :  PM  :  PL  ::  22  :  13  :  8-5  :  12;  AL— PM— PL  :  12-14;  AM  :  CL  ::  22  :  5. 
DISTRIBUTION.  Thailand. 

MATERIAL  EXAMINED.  Thailand:  Khas  Yai  National  Park,  tropical  evergreen  forest,  1^  under  bark  of 
decomposing  log,  17.iii.1984,  P.  D.  Hillyard,  BMNH.  1985.8.16.1. 

NATURAL  HISTORY.  The  male  described  above  was  found  together  with  several  harvestmen 
(Opiliones)  under  the  bark  of  a  fallen  decomposing  log.  There  was  no  evidence  of  a  web,  but  it  was 
noticeable  that  the  legs  of  this  species  were  comparatively  much  longer  than  those  of  5".  wildtrackii, 
a  character  that  may  enable  future  collectors  to  distinguish  the  species  in  the  field. 


Spartaeus  wildtrackii  sp.  n. 
(Figs  1;  2;  5A-E;  6A-D;  14A-C;  15A&C;  16A-D;  17A-G;  18A-D) 

DIAGNOSIS.  S.  wildtrackii  seems  to  be  most  closely  related  to  S.  thailandica,  but  may  be  dis- 
tinguished by  the  absence  of  a  tegular  apophysis  and  the  form  of  the  retrolateral  tibial  apophysis  in 
males  (Fig.  6A,  B,  D),  and  the  presence  of  a  postepigynal  furrow  in  females  (Fig.  5D). 

FEMALE  HOLOTYPE,  in  fair  condition.  Carapace  (Fig.  5A,  B):  weakly  iridescent  under  some  angles  of 
illumination;  orange-brown  lightly  tinged  and  mottled  black  with  a  broad  yellow-brown  tapering 
band  on  thoracic  part  and  vague  patches  on  sides;  irregularly  clothed  in  whitish  hairs  (mostly 
rubbed).  Eyes:  laterals  with  black  surrounds;  fringed  by  whitish  and  pale  yellow  hairs.  Clypeus: 
sparsely  clothed  in  black  hairs.  Chelicerae:  robust,  moderately  long  and  more  or  less  parallel; 
brownish  orange  lightly  tinged  black;  thinly  clothed  in  black  hairs  with  dense  promarginal  scopula; 
promargin  with  six  teeth,  retromargin  with  nine  or  12  denticles.  Maxillae  andlabium:  pale  orange- 
brown  lightly  tinged  with  some  grey.  Sternum:  pale  yellow  with  darker  margins.  Coxae:  pale  yellow 
except  first  pair  with  blackish  promarginal  sides.  Abdomen:  pale  yellow  with  blackish  markings  and 
scattered  clumps  of  whitish  guanin;  ventrally  a  broad  black  band  from  epigyne  to  spinnerets; 
spinnerets  long  and  robust.  Legs:  long  and  slender;  legs  I  pale  yellow  to  light  orange-brown  with 
vague  sooty  markings  and  a  proventral  black  stripe  on  femora  that  appears  iridescent  green  under 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


pm 


Fig.  5.  Spartaeus  wildtrackii  sp.  n.,  holotype  <$:  A,  dorsal  view;  B,  carapace,  lateral  view;  D,  epigyne. 
Paratype  <$:  C,  sternum;  E,  leg,  I.  Paratype  $:  F,  vulva,  ventral  view;  G,  vulva,  dorsal  view. 
Abbreviations:  pe,  postepigynal  furrow;  pm,  posterior  median  eye;  fo,  femoral  organ. 


some  angles  of  illumination;  other  legs  similar  except  femoral  stripe  lacking;  also,  dorsal  spines 
arise  from  black  spots;  sparsely  clothed  in  simple  and  feathery  hairs,  (Fig.  16D),  mostly  rubbed, 
with  pro  ventral  row  of  specialized  prey-capture  setae  on  tarsi  I-II  (Fig.  17B,  G)  and  apices  of 
metatarsi  I;  proximal  half  of  metatarsi  II-III  with  scanty  ventral  fringe  of  fine  curved  filamentous 
setae  (Fig.  1 7  A,  C-F).  Spines  strong  and  numerous;  spination  of  leg  I:  metatarsus  v  3-2-0,  p  0-1-0, 
tibia  v  5-8-7,  femur  d  0-2-1.  Palp:  with  terminal  claw;  pale  yellow  with  greyish  patch  on  tarsus; 
clothed  in  pale  yellow  and  light  greyish  hairs. 


114  F.  R.  WANLESS 

Dimensions  (mm):  total  length  6-4;  carapace  length  2-72,  breadth  2-28,  height  1-52;  abdomen 
length  3-68;  eyes,  anterior  row  2-08,  middle  row  1-48,  posterior  row  1-75;  quadrangle  length  1-58 
(58%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

3-24 

2-56 

2-56 

3-16 

1-06 

Patella 

1-50 

1-16 

1-00 

1-06 

0-60 

Tibia 

3-16 

2-20 

2-20 

3-04 

0-64 

Metatarsus 

1-92 

1-74 

2-36 

3-20 

Tarsus 

0-84 

0-78 

0-92 

1-08 

1-16 

Total 

10-66 

8-44 

9-04 

11-54 

3-46 

:  PM  :  PL  :: 

16-5:11:7: 

11;  AL- 

-PM- 

-PL  ::  10-10;  AM 

MALE  PARATYPE,  in  good  condition.  Carapace:  dark  brown  with  dull  orange-brown  eye  region  and 
yellow-brown  markings  on  thoracic  part;  clothed  in  fine  recumbent  light  greyish  and  pale  amber 
hairs.  Eyes:  laterals  with  black  surrounds;  fringed  by  pale  yellow,  whitish  and  amber  hairs;  also,  a 
dense  matt  of  short  hairs  behind  anterior  medians.  Clypeus:  tinged  black,  sparsely  clothed  in  black 
hairs.  Chelicerae:  long,  robust  and  slightly  diverging;  dark  brown  heavily  tinged  black;  shiny; 
thinly  clothed  in  black  hairs  with  dense  promarginal  scopula;  fang  robust  and  curved  with  basal 
protuberance;  promargin  with  eight  teeth,  retromargin  with  12  (Fig.  6C).  Maxillae  and  labium: 
orange-brown  to  yellow-brown  tinged  grey.  Sternum  (Fig.  5C):  pale  greenish  yellow  with  darker 
margins;  thinly  clothed  in  greyish  simple  and  feathery  hairs.  Coxae:  pale  greenish  yellow  with  black 
promarginal  stripe.  Abdomen:  yellow-brown  suffused  and  mottled  black;  clothed  in  light  and  dark 
amber  hairs  with  two  spots  comprised  of  whitish  guanin;  venter  yellow-brown  with  grey-black 
band  clothed  in  black  feathery  hairs  from  epigastric  furrow  to  spinnerets;  spinnerets  long  and 
robust.  Legs:  long  and  slender;  femoral  organ  well  developed  (Fig.  5E;  14A-C);  legs  I  pale  greenish 
yellow  to  light  orange  brown  with  sooty  markings  and  blackish  longitudinal  stripes  on  femora  that 
shine  iridescent  green  under  some  angles  of  illumination;  thinly  clothed  in  simple  and  black 
feathery  hairs  with  some  whitish  ones  on  tarsi;  specialized  prey  capture  and  filamentous  hairs  as 
in  female.  Spination  of  leg  I:  metatarsus  v  3-1-1,  p  0-0-1,  r  1-0-0;  tibia  v  5-7-6;  femur  d  0-2-1, 
p  0-1-1.  Palp  (Fig.  6A,  B,  D):  yellow-brown  to  light  orange-brown  with  black  iridescent  stripe  on 
underside  of  femora. 

Dimensions  (mm):  total  length  6-96;  carapace  length  3-36,  breadth  2-68,  height  1-92;  abdomen 
length  3-8;  eyes,  anterior  row  2-32,  middle  row  1-62,  posterior  row  1-96;  quadrangle  length  1-8 
(54%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

5-40 

3-44 

3-36 

4-28 

2-20 

Patella 

2-20 

1-48 

1-28 

1-36 

1-38 

Tibia 

5-68 

3-16 

3-08 

4-20 

1-08 

Metatarsus 

3-48 

2-56 

3-24 

4-58 

Tarsus 

1-22 

0-92 

1-04 

1-20 

1-88 

Total  17-98        11-56        12-00        15-62         6-54 

Ratios:  AM  :  AL: :  PM  :  PL  ::  19  :  1 1  :  8  :  1 1;  AL— PM— PL  ::  10  :  13;  AM  :  CL  ::  19  :  5. 

VARIATION.  Male  total  length  varies  from  6-24  to  7-44  mm,  carapace  length  2-72-3-36  mm  (five 
specimens);  female  total  length  6-0-6-96  mm,  carapace  length  2-56-3-36  mm  (10  specimens). 

In  two  females  the  arthrodial  membrane  between  the  chelicerae  and  the  clypeus  is  evident  as  a 
narrow  white  band,  whereas  it  is  broad  and  conspicuous  in  other  specimens.  However,  to  judge 
from  the  set  of  the  chelicerae  it  is  apparent  that  the  phenomenon  is  the  result  of  postmorten 
changes. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


115 


Fig.  6.  Spartaeus  wildtrackii  sp.  n.,  paratype  cJ,  A,  palp,  ventral  view;  B,  palpal  tibia,  dorsal  view;  C, 
cheliceral  teeth;  D,  palp,  retrolateral  view;  E,  chelicerae,  maxillae  and  labium.  Abbreviations:  rta, 
retrolateral  tibial  apophysis,  tf,  tegular  furrow;  va,  ventral  apophysis. 


NATURAL  HISTORY.  S.  wildtrackii  has  so  far  only  been  found  in  lowland  rainforest  on  the  trunks  of 
large  trees  covered  in  lichen  or  moss,  against  which  the  spiders  are  well  camouflaged.  A  number  of 
specimens  were  seen  both  at  night  and  during  the  day  resting  on  bark  beneath  large  silken  webs  ca. 
5-6  cm  constructed  of  glossy  translucent  silk  that  was  often  torn  (Fig.  1,2).  Their  prey  is  unknown 
except  for  that  of  one  specimen  which  was  seen  feeding  on  newly  emerged  moths,  that  were  resting 
and  evidently  drying  their  wings.  The  spiders  were  not  seen  to  jump,  but  it  was  noted  that  they  were 
fast  runners  (P.  D.  Hillyard,  pers.  comm.). 

DISTRIBUTION.  West  Malaysia. 

MATERIAL  EXAMINED.  West  Malaysia,  P.  D.  Hillyard:  Pahang  State,  Taman  Negara,  nr.  Kuala  Tahan, 
lowland  primary  rain  forest:  on  tree  trunks  under  sheet  webs,  9.iii.l984,  holotype  §,  BMNH.  1985.8.16.2, 
paratypes  3  ??,BMNH.  1985.8. 16.3-5;  on  tree  trunks,  iii.  1985,  paratypes  4??,  4<JcJ,  BMNH  1985.8.16.6-13. 
Negeri  Sembilan  State,  Pasoh  Forest  Reserve,  on  tree  trunks  in  lowland  primary  rain  forest,  iii.1985, 
paratypes  3  <J<J,  9  ??,  BMNH,  1985.8.16.14-25. 

ETYMOLOGY.  This  species  is  named  for  the  BBC  television  programme  'Wildtrack'  which  has  done 
much  to  encourage  children  to  care  for  the  environment  and  take  an  interest  in  natural  history. 


116  F.  R.  WANLESS 

REMARKS.  1 .  Postepigynal  furrows  (Fig.  5D)  are  an  unusual  feature  of  salticid  epigynes  and  to  date 
have  only  been  found  in  one  other  spartaeine  i.e.  Gelotia  bimaculata  Thorell.  Their  function  is 
uncertain,  but  they  may  form  part  of  the  supporting  mechanism  that  holds  the  male  palp  in 
position  during  copulation. 

2.  Loerbroks  ( 1 984)  has  recently  drawn  attention  to  conspicuous  similarities  in  palpal  structure 
between  Misumena  vatia  (Clerck),  a  crab  spider  (Family  Thomisidae)  and  Phaeacius  Koch,  a  genus 
of  flattened  salticid  that  has  also  been  classified  in  the  Spartaeinae.  Futhermore,  he  has  shown  that 
in  M .  vatia,  and  probably  all  other  thomisids,  the  ventral  apophysis  locks  into  the  regular  ridge  as 
the  palpal  elements  expand  and  rotate  during  copulation.  The  ventral  apophysis  (Figs  6A;  13F) 
and  tegular  furrow  (Fig.  6A)  ( =  tegular  ridge  of  Loerbroks)  characteristic  of  all  spartaeines  are 
evidently  homologous  with  those  of  thomisids  and  probably  function  in  a  similar  manner.  In  M. 
vatia  the  inner  surface  of  the  ventral  apophysis  is  covered  in  papillae  (see  Loerbroks  1984,  Fig.  6) 
that  evidently  reduce  friction  between  the  apophysis  and  the  rotating  tegulum.  Similar  papillae 
might  therefore  be  expected  on  the  ventral  apophysis  of  5.  wildtrackii,  but  are  absent  (Fig.  18A). 
However,  spicule-like  papillae  do  occur  on  the  inner  surface  of  the  retrolateral  tibial  apophysis 
(Fig.  18B-D)  and  presumbly  they  too  could  reduce  surface  friction,  although  on  the  otherhand 
they  may  serve  to  prevent  the  apophysis  from  sliding  out  of  position  during  copulaton. 

Although  genital  structures  are  not  generally  used  in  assessing  relationships  at  the  familial  level, 
they  are  considered  here  because  ventral  apophyses  and  tegular  furrows  are  not  known  to  occur  in 
other  spider  families.  They  may  have  arisen  independently,  but  as  Loerbroks  (1984)  has  already 
stated  they  may  provide  evidence  of  a  phylogenetic  link  between  salticids  and  thomisids. 
Additional  evidence  is  provided  by  Homann  (1971)  who  has  shown  that  the  anterior  median  eyes 
(principal  eyes)  of  most  spiders  are  small  and  have  few  visual  cells,  whereas  those  of  salticids  and 
thomisids  are  unique  in  possessing  many  visual  cells  which  provide  for  'sharp  vision'.  These  optical 
similarities  may  be  convergent  as  Homann  regards  thomisid  eyes  as  being  structurally  closer  to 
those  of  wolf  spiders  (Family  Lycosidae).  Future  studies  will  have  to  take  account  of  the  genitalial 
similarities  noted  by  Loerbroks  since  these  can  be  interpreted  as  supportive  of  a  sister  group 
relationship  between  salticids  and  thomisids. 

Genus  MINTONIA  Wanless 
Mintonia  Wanless,  1984a:  157.  [definition,  diagnosis  and  key  to  species]. 

This  small  oriental  genus  comprises  nine  species  including  two  new  taxa  described  below.  The 
majority  of  species  have  been  collected  from  Borneo,  but  the  genus  is  also  known  from  Java, 
Sumatra  and  Peninsular  Malaya.  Males  are  of  particular  interest  because  they  possess  femoral 
organs  (Fig.  9D;  19A-C),  a  presumptive  sex  pheromone  dispersal  site,  and  retrolateral  tibial 
apophyses,  some  of  which  bear  openings  (Fig.  9E  arrowed;  20A,  B).  Unfortunately  nothing  is 
known  of  their  natural  history. 

Mintonia  melinauensis  Wanless 
(Fig.  7A-E) 

Mintonia  melinauensis  Wanless,  1984a:  165,  $  holotype,  Sarawak  (BMNH)  [examined]. 

DIAGNOSIS.  Males  can  be  recognized  by  the  heavy  inward  curving  embolus  and  by  the  form  of  the 
retrolateral  tibial  apophysis  (see  Wanless,  1984a,  Fig.  13);  females  by  the  structure  of  the  epigyne 
(Fig.  7E)  which  is  clearly  different  from  that  of  other  females  of  the  genus. 

FEMALE,  formerly  undescribed,  in  fair  condition.  Carapace  (Fig.  7A,  D):  weakly  iridescent  under 
some  angles  of  illumination;  orange-brown  lightly  mottled  black  with  paler  lateral  markings  and 
band  from  fovea  to  posterior  margin;  rubbed,  but  otherwise  clothed  in  whitish  pubescent  hairs. 
Eyes:  laterals  with  black  surrounds;  fringed  by  whitish  hairs.  Clypeus:  lightly  tinged  with  some 
black;  sparsely  clothed  in  fine  whitish  hairs  with  several  long  bristles.  Chelicerae:  orange-brown 
suffused  with  some  black  proximally;  shiny,  thinly  clothed  in  long  pale  orange  hairs  with  modera- 
tely dense  promarginal  scopulae;  promargin  with  three  teeth,  retromargin  with  six  (Fig.  7B). 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


117 


B 


Fig.  7.    Mintonia  melinauensis  Wanless,  9:  A,  dorsal  view;  B,  cheliceral  teeth;  C,  sternum;  D,  carapace, 

lateral  view;  E,  epigyne. 

Maxillae:  light  orange-brown  with  whitish  yellow  inner  distal  margins.  Labium:  brownish  orange 
tipped  whitish  yellow.  Sternum  (Fig.  7C):  pale  yellow  with  darker  margins;  thinly  clothed  in  fine 
hairs.  Coxae:  pale  greyish  yellow.  Abdomen:  rubbed;  whitish  yellow  with  vague  greyish  markings; 
spinnerets  whitish  yellow  with  outer  sides  of  anteriors  tinged  black.  Legs:  moderately  long  and 
robust;  whitish  yellow  to  orange  brown;  spines  strong  and  numerous.  Spination  of  leg  I;  metatar- 
sus v  2-2-1,  p  1-0-1,  d  0-0-2,  r  1-0-0;  tibia  v  2-2-2,  p  1-0-1,  r  0-0-1;  patella  p  0-1-0,  r  0-1-0; 
femur  d  0-2-3.  Palp:  whitish  yellow  with  sooty  markings  except  for  pale  orange  brown  tarsi. 
Epigyne  (Fig.  7E):  clothed  in  fine  pale  yellowish  hairs. 

Dimensions  (mm):  total  length  5-2;  carapace  length  2-24,  breadth  1-76,  height  1-44;  abdomen 
length  3-04;  eyes,  anterior  row  1-64,  middle  row  1-48,  posterior  row  1-64;  quadrangle  length  1-2 
(53%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-48 

1-46 

1-48 

1-80 

0-80 

Patella 

0-86 

0-81 

0-71 

0-76 

0-48 

Tibia 

1-12 

1-04 

1-12 

1-44 

0-52 

Metatarsus 

0-96 

0-94 

1-16 

1-60 

Tarsus 

0-58 

0-57 

0-60 

0-68 

0-72 

Total 


5-00 


4-82 


5-07        6-28 


2-52 


Ratios:  AM  :  AL  :  PM  :  PL::  13:8:5:8;  AL— PM— PL  ::  7-5-8-5;  AM  :  CL  ::  13  :  2-5. 
DISTRIBUTION.  Sarawak. 


118 


F.  R.  WANLESS 


rta 


Fig.  8.  Mintonia  caliginosa  sp.  n.,  holotype  $:  A,  dorsal  view;  B,  carapace,  lateral  view;  C,  palp,  ventral 
view;  D,  palp,  retrolateral  view;  E,  palp,  dorsal  view.  Abbreviations:  e,  embolus;  rta,  retrolateral  tibial 
apophysis;  v,  vacuole;  va,  ventral  apophysis. 

MATERIAL  EXAMINED.  Type  data  given  in  synonymy.  Sarawak:  Gunung  Mulu  National  Park,  Environs  of 
camp  3,  moss  forest,  1$,  from  moss  covered  tree  trunk,  27.5.78,  F.  R.  Wanless,  Royal  Geographic  Society/ 
Sarawak  Government  Expedition.  (BMNH). 

Mintonia  caliginosa  sp.  n. 

(Fig.  8A-E;19A-C) 

DIAGNOSIS.  A  distinctive  species  easily  recognized  by  the  long  retrolateral  tibial  apophysis 
(Fig.  8D). 

FEMALE.  Unknown. 

MALE  HOLOTYPE,  rubbed  otherwise  in  fair  condition.  Carapace  (Fig.  8A,  B):  orange-brown  lightly 
tinged  and  mottled  black;  shiny  and  weakly  iridescent  under  some  angles  of  illumination.  Eyes: 
laterals  with  black  surrounds;  fringed  by  whitish  and  pale  amber  hairs,  mostly  rubbed.  Clypeus 
edged  black  below  anterior  median  eyes;  rubbed — a  few  whitish  hairs  remaining.  Chelicerae:  light 
brown  with  black  markings,  shiny,  thinly  clothed  in  scattered  fine  hairs;  promargin  with  three 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE  1 1 9 

teeth,  retromargin  with  four  or  five.  Maxillae:  light  yellowish  brown.  Labium:  blackish  edged  pale 
yellow-brown.  Sternum:  yellow-brown,  shiny,  thinly  clothed  in  fine  greyish  hairs.  Abdomen: 
yellow-brown  lightly  tinged  and  mottled  black  with  a  poorly  defined  orange-brown  scutum  and 
chevrons  dorsally,  and  three  rather  vague  longitudinal  bands  ventrally;  spinnerets  yellow-brown 
lightly  tinged  black.  Legs:  moderately  long  and  slender;  femoral  organ  (Fig.  19A-C)  a  low 
tubercle;  yellow-brown  tinged  with  some  black  except  metatarsi  and  tarsi  which  are  darker — 
orange-brown  tinged  black,  also  on  underside  of  femora  I-II  a  transverse  blackish  patch;  spines 
strong  and  numerous.  Spination  of  leg  I:  metatarsus  v  2-0-0,  p  1-1-1,  d  0-1-2,  r  1-1-1;  tibia  v 
2-2-2,  p  1-1-0,  d  1-1-0,  r  1-1-0;  patella  p  0-1-0,  r  0-1-0;  femur  d  0-2^.  Palp  (Fig.  8C-E):  the 
retrolateral  tibial  apophysis  is  broken  at  point  arrowed  in  Fig.  8E;  however,  note  that  in  Fig.  8C  i.e. 
the  same  palp  but  drawn  from  a  different  angle,  the  appearance  of  the  retrolateral  tibial  apophysis 
has  been  reconstructed  from  the  apophysis  of  the  other  palp. 

Dimensions  (mm):  total  length  3-8;  carapace  length  1-76,  breadth  1-36,  height  1-1;  abdomen 
length  1-92;  eyes,  anterior  row  1-27,  middle  row  1-2,  posterior  row  1-35;  quadrangle  length  0-94 
(53%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-16 

1-12 

1-19 

1-46 

0-60 

Patella 

0-60 

0-60 

0-52 

0-56 

0-32 

Tibia 

0-84 

0-80 

0-88 

1-16 

0-24 

\^of  Q  t  Q  ¥*C1  1  C 

0-80 

0-78 

0-88 

1  -78 

IVItldltll  oUS 

Tarsus 

\J  OU 

0-52 

u  /  o 

0-48 

u  oo 

0-56 

i  _  <> 

0-60 

0-72 

Total  3-90         3-78         4-03         5-06         1-88 

Ratios:  AM  :  AL  :  PM  :  PL  ::  9-5  :  5-5  :  3-5  :  5-5;  AL— PM— PL  ::  6-7;  AM  :  CL  ::  9-5  :  3-3. 
DISTRIBUTION.  Borneo,  Sabah. 

MATERIAL  EXAMINED.  Borneo:  Sabah,  Tuaran  Division,  Mt.  Kinabalu  National  Park,  Power  Station — 
Layang  Layang,  cloud  forest,  holotype  <$,  2000-2800  m,  7.H.1976  P.  T.  Lehtinen,  (TU,  Turku). 

REMARK.  The  presence  of  a  vacuole  in  the  base  of  the  retrolateral  tibial  apophysis  suggests  that 
there  is  probably  a  distal  opening.  There  is  insufficient  material  for  this  to  be  confirmed  by  SEM. 

ETYMOLOGY.  The  specific  name  is  from  the  Latin  meaning  misty,  cloudy  places. 


Mintonia  silvicola  sp.  n. 
(Fig.  9A-G) 

DIAGNOSIS.  M.  silvicola  seems  to  be  most  closely  related  to  M.  tauricornis  Wanless,  but  may  be 
readily  distinguished  by  the  syringe-shaped  retrolateral  tibial  apophysis  (Fig.  9E). 

FEMALE.  Unknown. 

MALE  HOLOTYPE,  rubbed,  also  right  leg  I  missing,  otherwise  in  fair  condition.  Carapace  (Fig.  9A, 
B):  weakly  iridescent  under  some  angles  of  illumination;  orange-brown  with  faint  blackish 
mottling  on  sides.  Eyes:  laterals  with  black  surrounds;  anteriors  fringed  by  whitish  hairs.  Clypeus: 
orange-brown  with  blackish  margin  below  anterior  median  eyes  and  vague  yellow-brown  mark- 
ings clothed  in  whitish  hairs  below  anterior  laterals.  Chelicerae:  yellow-brown,  shiny,  clothed  in 
white  hairs  proximally  and  scattered  brown  hairs  distally  with  dense  promarginal  scopulae; 
promargin  with  three  teeth,  retromargin  with  eight  (Fig.  9G).  Maxillae:  yellow-brown  with  inner 
distal  margins  paler.  Labium:  yellow-brown  faintly  tinged  grey.  Sternum:  pale  yellow  with  vague 
darker  margins;  thinly  clothed  in  fine  pale  yellow  hairs.  Coxaei  pale  yellow.  Abdomen:  pale  yellow 


120 


F.  R.  WANLESS 


B 


Fig.  9.  Mintonia  silvicola  sp.  n.,  holotype  <$:  A,  dorsal  view;  B,  carapace,  lateral  view;  C,  palp,  ventro- 
lateral  view;  D,  leg  I;  E,  palp,  retrolateral  view;  F,  palp,  ventral  view;  G,  cheliceral  teeth.  Abbreviation: 
fo,  femoral  organ. 


with  vague  sooty  markings  and  two  pairs  of  sigilla;  rubbed;  spinnerets  moderately  long,  yellow- 
brown.  Legs:  moderately  long  and  slender;  specialized  prey  capture  and  metatarsal  setae  lacking; 
femoral  organ  appearing  as  a  low  dark  amber  mound;  legs  I  pale  yellow  to  yellow-brown  with 
underside  of  femora  tinged  black;  other  legs  pale  yellow  grading  to  yellow-brown  distally  with 
ventral  longitudinal  grey  stripe  on  tibiae  II-IH;  spines  numerous  and  moderately  strong.  Spination 
of  leg  I:  metatarsus  v  2-0-0,  r  1-1-1,  d  0-2-2,  p  1-1-1;  tibia  v  2-2-2,  p  1-1-0,  d  1-1-0,  r  1-1-0; 
patella  p  0-1-0,  r  0-1-0;  femur  d  0-2^.  Palp  (Fig.  9C,  E,  F):  element  M2  lies  above  the  embolus 
and  the  tegular  ledge  is  poorly  developed;  the  opening  of  the  retrolateral  tibial  apophysis  (arrow, 
Fig.  9E)  is  distinct. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE  1 2 1 

Dimensions  (mm):  total  length  5-1;  carapace  length  2-24,  breadth  1-88,  height  1-44;  abdomen 
length  2-6;  eyes,  anterior  row  1-71,  middle  row  1-48,  posterior  row  1-64;  quadrangle  length  1-32 
(58%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-72 

1-72 

1-72 

2-04 

0-84 

Patella 

0-88 

0-84 

0-76 

0-80 

0-40 

Tibia 

1-36 

1-28 

1-34 

1-64 

0-34 

\^f*t  d  t  d  rcnc 

1  -78 

1  -96 

1  -44 

1  -80 

IVlCLd.  Ldl  S  US 

Tarsus 

i  —'i 

0-64 

1    —O 

0-64 

1  HH 

0-72 

I    OU 

0-76 

1-12 

Total  5-88         5-74         5-98         7-04         2-70 

Ratios:  AM  :  AL  :  PM  :  PL  ::  14  :  8  :  5-4  :  8;  AL— PM— PL  ::  8-9-5;  AM  :  CL  ::  14  :  3. 
DISTRIBUTION.  West  Malaysia. 

MATERIAL  EXAMINED.  West  Malaysia:  Pahang  State,  Taman  Negara,  holotype  $,  from  buttress  of  large  tree, 
lowland  rain  forest  nr.  Kuala  Tahan,  3-10.iii.1984.  P.  D.  Hillyard,  BMNH.  1985.8.21.1. 

ETYMOLOGY.  The  specific  name  is  from  the  Latin  meaning  inhabiting  woods. 

Genus  TARAXELLA  Wanless 
Taraxella  Wanless,  1984a:  155.  [definition  and  diagnosis]. 

This  genus  was  originally  erected  on  the  basis  of  a  single  male  of  Taraxella  solitaria  Wanless, 
from  Sarawak.  Subsequent  collections  have  produced  four  new  species,  described  below,  that 
necessitate  modifications  to  the  original  generic  definition. 

DEFINITION.  Spiders  small  to  medium  in  size,  i.e.  between  2-0  and  8-0  mm  in  length;  males  some- 
times with  conspicuous  encircling  band  on  the  carapace;  sexual  dimorphism  sometimes  evident  in 
colour  patterns. 

Carapace,  high,  longer  than  broad,  widest  at  about  level  of  coxae  II-III;  fovea  long  and  sulci- 
form,  apex  at  level  of  centre  of  posterior  lateral  eyes.  Eyes:  anterior  medians  more  or  less  level  or 
weakly  procurved  in  frontal  view;  posterior  medians  small  to  relatively  large;  posterior  laterals 
with  outer  margins  of  lenses  set  inside  or  at  level  of,  lateral  margins  of  carapace  when  viewed  from 
above;  entire  quadrangle  length  between  57-65%  of  carapace  length.  Clypeus:  low  to  moderately 
high.  Chelicerae:  promargin  with  five  or  seven  teeth,  retromargin  with  seven  or  nine  denticles. 
Legs:  moderately  long  and  slender;  femoral  organs  lacking;  specialized  prey  capture  tarsal  setae 
and  filamentous  metatarsal  setae  also  lacking.  Female  palps:  moderately  long  and  slender  with 
apical  claw.  Epigynes:  interspecifically  distinct,  see  descriptions;  vulvae  not  examined,  insufficient 
material.  Male  palps:  complex  and  interspecifically  distinct;  retrolateral  tibial  apophyses  complex, 
sometimes  bifid  with  sharp  slender  prongs,  or  evidently  reduced  with  associated  stout  setae; 
apophyses  X  and  Y  variable  in  development;  embolus  short  slender  and  gently  curved,  and  for  the 
most  part  obscured,  in  ventral  view,  by  tegular  apophyses  X  and  occasionally  Y;  tegular  furrow 
and  ventral  apophysis  usually  conspicuous;  Ml,  see  Wanless  19840,  a  delicate  fan-shaped  lamella 
that  protrudes  beyond  the  distal  edge  of  the  tegulum.  Expanded  palps  not  examined. 

DIAGNOSIS.  Distinguished  from  other  spartaeines  by  the  conformation  of  the  embolus  of  the 
male  palp  which  is  almost  completely  obscured,  in  ventral  view,  by  tegular  apophyses  'X'  and 
occasionally  'Y'. 

An  identification  key  is  not  provided  as  the  five  known  species  are  easily  separated  from  one 
another  by  the  structure  of  the  palpal  organs  and  epigynes. 

INTERSPECIFIC  RELATIONSHIPS.  To  judge  from  the  structure  of  the  tibia  of  the  male  palpal  organs  T. 
solitaria,  T.  petrensis  sp.  n.,  and  T.  hilly ardi  sp.  n.,  form  a  closely  related  group  since  they  all  possess 
retrolateral  tibial  apophyses  with  a  sharp  dorsal  prong.  T.  sumatrana  sp.  n.,  and  T.  reinholdae 


122 


F.  R.  WANLESS 


am 


al 


B 


Fig.  10.  Taraxella  hillyardi  sp.  n.,  holotype  c?:  A,  dorsal  view;  B,  carapace,  lateral  view;  C,  leg  I;  D, 
palpal  tibia,  dorsal  view;  E,  palp,  retrolateral  view;  F,  palp,  ventral  view.  Abbreviations:  al,  anterior 
lateral  eye;  am,  anterior  median  eye;  dh,  distal  haematodocha;  e,  embolus;  pi,  posterior  lateral  eye;  pm, 
posterior  median  eye;  tf,  tegular  furrow;  va,  ventral  apophysis. 

would  also  appear  to  form  a  natural  group  as  they  possess  conspicuous  fringes  of  unusually  stout 
setae,  and  lack  retrolateral  tibial  apophyses  with  sharp  dorsal  prongs. 


Taraxella  hillyardi  sp.  n. 
(Fig.  10 A,  F) 

DIAGNOSIS.  T.  hillyardi  seems  to  be  most  closely  related  to  T.  petrensis  and  T.  solitaria,  but  can  be 
easily  distinguished  by  the  bifid  retrolateral  tibial  apophysis  (Fig.  10E). 

FEMALE.  Unknown. 

MALE  HOLOTYPE,  in  fair  condition.  Carapace  (Fig.  10A,  B):  weakly  iridescent  under  some  angles  of 
illumination;  light  orange-brown  tinged  black  with  a  broad  yellowish  encircling  band  on  sides  and 
also  a  black  marginal  band  extending  posteriorly  from  level  of  coxae  I;  rubbed.  Eyes:  laterals  with 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE  1 23 

black  surrounds;  sparsely  fringed  in  pale  amber  and  whitish  hairs.  Clypeus:  greyish  with  black 
markings  below  anterior  median  eyes  and  whitish  yellow  stripes  below  anterior  laterals;  sparsely 
fringed  by  whitish  hairs.  Chelicerae:  yellow-brown  with  extensive  patches  on  facies;  shiny;  thinly 
clothed  in  fine  clear  hairs  and  some  blackish  ones  with  dense  promarginal  scopulae;  promargin 
with  five  teeth;  retromargin  with  eight  denticles.  Maxillae  andlabium:  pale  yellow  with  vague  sooty 
markings.  Coxae:  yellow-brown.  Abdomen:  dorsum  and  sides  pale  yellow-brown  suffused  and 
mottled  black,  venter  pale  yellow  brown  suffused  black  in  region  of  tracheal  spiracle;  spinnerets 
moderately  long;  anteriors  and  posteriors  suffused  black,  medians  pale  yellow.  Legs  Fig.  IOC): 
moderately  long  and  slender;  generally  yellow-brown  tinged  with  some  black,  with  incomplete 
annuli  on  femora  and  blackish  tibiae  particularly  of  legs  I  and  IV;  spines  moderately  strong  and 
numerous.  Spination  of  leg  I:  metatarsus  v  2-0-0,  p.  1-1-1 ,  d  0-2-2,  r  1-1-1 ;  tibia  v  2-2-2,  p  0-1-1 , 
d  i_i_o,  r  1-1-0;  patella  p  0-1-1,  r  0-1-0;  femur  d  0-2-3,  p  0-0-1.  Palp  (Fig.  10D-F):  yellow- 
brown  to  orange-brown  mottled  black  with  patches  of  white  hairs  on  patella  and  apices  of  femur, 
otherwise  clothed  in  black  hairs  and  scattered  white  ones  with  greyish  scopula  on  cymbium. 

Dimensions  (mm):  total  length  3-9;  carapace  length  1-76,  breadth  1-53,  height  1.2;  abdomen 
length  1-84;  eyes,  anterior  row  1-56,  middle  row  1-24,  posterior  row  1-52;  quadrangle  length  1-07 
(60%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-36 

1-28 

1-28 

1-64 

0-68 

Patella 

0-76 

0-63 

0-56 

0-64 

0-33 

Tibia 

1-12 

0-96 

1-00 

1-39 

0-36 

Metatarsus 

1-02 

0-96 

1-12 

1-60 

Tarsus 

0-53 

0-52 

0-56 

0-66 

0-72 

Total  4-79         4-35         4-52         5-93         2-09 

Ratios:  AM  :  AL  :  PM  :  PL  ::  13  :  7  :  3  :  7;  AL— PM— PL  ::  7-8;  AM  :  CL  ::  13  :  3. 
DISTRIBUTION.  West  Malaysia. 

MATERIAL  EXAMINED.  West  Malaysia:  Gunong  Jerai,  (Kedeh),  ca.  700  m,  holotype  <$,  shrub  layer,  along  forest 
edge  during  middle  of  dry  season,  15.ii.1983,  P.  D.  Hillyard(BMNH.  1985.9.5.2). 

ETYMOLOGY.  This  species  is  named  after  my  colleague  Mr  P.  D.  Hillyard,  BMNH,  who  collected 
many  of  the  new  species  described  in  this  paper. 

Taraxella  petrensis  sp.  n. 
(Fig.  11A-J) 

DIAGNOSIS.  T.  petrensis  seems  to  be  most  closely  related  to  T.  solitarius  and  T.  hillyardi  sp.  n.,  but 
may  be  easily  separated  by  the  broad  flange  of  the  retrolateral  tibial  apophysis  (Fig.  1 U)  in  males 
and  the  structure  of  the  epigyne  (Fig.  1  ID)  in  females. 

MALE  HOLOTYPE,  rubbed  otherwise  in  good  condition.  Carapace  (Fig.  1 1  A,  F):  shiny  and  weakly 
iridescent  in  eye  region;  dark  orange-brown  suffused  black  with  broad  encircling  creamy  white 
band.  Eyes:  laterals  with  black  surrounds;  anteriors  sparsely  fringed  in  greyish  hairs.  Clypeus: 
creamy  white  with  black  spots  near  lower  rims  of  anterior  median  eyes.  Chelicerae:  shiny  black 
except  for  orange-brown  inner  margins;  sparsely  clothed  in  greyish  hairs  with  dense  promarginal 
scopulae;  promargin  with  five  teeth,  retromargin  with  nine  denticles  (Fig.  11C).  Maxillae  and 
labium:  greyish  yellow  faintly  tinged  black.  Sternum:  pale  yellow-brown  with  darker  margins; 
thinly  clothed  in  light  brownish  hairs.  Coxae:  yellow-brown  tinged  grey.  Abdomen:  yellow-brown 
with  dorsum  and  sides  mottled  black,  also  a  vague  black  patch  in  area  of  tracheal  slit;  rubbed; 
spinnerets  moderately  long,  black  except  for  light  greyish  medians.  Legs:  moderately  long  and 
slender;  yellow-brown  heavily  suffused  black,  especially  on  femora,  patellae  and  tibiae;  shiny  and 


124 


F.  R.  WANLESS 


Fig.  11.  Taraxella  petrensis  sp.  n.,  holotype  (J:  A,  dorsal  view;  C,  cheliceral  teeth;  F,  carapace,  lateral 
view;  G,  palpal  tibia,  dorsal  view;  H,  palp,  ventral  view;  J,  palp,  retrolateral  view.  Paratype  $: 
B,  cheliceral  teeth;  E,  carapace,  dorsal  view;  I,  leg  I.  Abbreviation:  e,  embolus;  'x'  and  'y'  tegular 
apophyses. 

iridescent  under  some  angles  or  illumination;  spines  strong  and  numerous.  Spination  of  leg  I: 
metatarsus  v  2-2-2,  p  1-0-0,  d  0-1-2,  r  1-0-0;  tibia  v  1-3-2,  p  0-1-1,  d  1-1-0,  r  0-0-1;  patella  p 
0-1-0,  r  0-1-0;  femur  d  0-2-3.  Palp  (Fig.  1 1G,  H,  J):  yellow  to  orange-brown  suffused  with  some 
black  especially  on  femur  and  cymbium;  clothed  in  pale  grey  and  black  hairs. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE  1 25 

Dimensions  (mm):  total  length  3-72;  carapace  length  1-76,  breadth  1-56,  height  1-24;  abdomen 
length  2-16;  eyes,  anterior  row  1-6;  middle  row  1-21,  posterior  row  1-52;  quadrangle  length  1-08 
(61  %  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-52 

1-34 

1-36 

1-72 

0-76 

Patella 

0-76 

0-60 

0-60 

0-66 

0-32 

Tibia 

1-24 

1-00 

1-04 

1-42 

0-36 

\ji  t 

1-19 

1  -09 

1  •  1  X 

1  -fis 

ivieiaiarsus 
Tarsus 

1    1Z 

0-60 

1   \JZ. 

0-56 

1     1  0 

0-56 

1   U  J 

0-70 

0-84 

Total  5-24         4-52         4-74         6-15         2-28 

Ratios:  AM  :  AL  :  PM  :  PL  ::  13-5  :  7-5  :  2-5  :  7-5;  AL— PM— PL  ::  7-5  :  7;  AM  :  CL  ::  13-5  :  5. 

FEMALE  PARATYPE,  in  fair  condition.  General  habitus  as  in  male  except  encircling  cephalic  band 
lacking.  Carapace  (Fig.  1  IE):  light  orange-brown  lightly  and  finely  reticulated  black  in  eye  region 
with  blackish  mottling  on  sides,  also  weakly  iridescent  under  some  angles  of  illumination;  rubbed. 
Eyes:  generally  as  in  male,  but  sparsely  fringed  by  pale  amber  hairs.  Clypeus:  light  orange-brown 
faintly  reticulated  black;  bald  except  for  scattered  fine  marginal  hairs  and  several  long  stiff  hairs 
including  usual  triad  in  lower  space  between  anterior  median  eyes.  Chelicerae:  yellow-brown 
lightly  tinged  with  some  black;  shiny;  sparsely  clothed  in  brown  hairs  with  dense  promarginal 
scopulae;  promargin  with  five  teeth,  retromargin  with  nine  denticles  (Fig.  11B).  Maxillae  and 
labium:  yellow-brown.  Sternum:  pale  yellow-brown  with  darker  margins;  sparsely  clothed  in  fine 
hairs  centrally  and  darker,  longer  ones  towards  margins.  Abdomen:  generally  as  in  male  except 
venter  pale  yellow-brown  with  scattered  dark  brown  simple  hairs  and  vague  light  greyish  feathery 
hairs,  otherwise  rubbed;  spinnerets  similar  to  male,  yellow-brown  tinged  black.  Legs  (Fig.  Ill): 
moderately  long  and  slender;  light  orange-brown  faintly  tinged  with  some  black;  sparsely  clothed 
in  brownish  hairs;  spines  strong  and  numerous.  Spination  of  leg  I:  metatarsus  v  2-2-0,  p  1-0-1,  d 
0-1-2,  r  0-0-1;  tibia  v  2-3-1,  p  0-1-1;  patella  p  0-1-0;  femur  d  0-2-2.  Epigyne  (Fig.  1  ID). 

Dimensions  (mm):  total  length  4-32;  carapace  length  1-96,  breadth  1/72,  height  1-32;  abdomen 
length  2-28;  eyes,  anterior  row  1-76,  middle  row  1-36,  posterior  row  1-72;  quadrangle  length  1-2 
(61%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-58 

1-44 

1-44 

1-76 

0-76 

Patella 

0-84 

0-74 

0-64 

0-70 

0-43 

Tibia 

1-18 

1-00 

1-08 

1-46 

0-48 

Metatarsus 

1-04 

1-00 

1-20 

1-68 

Tarsus 

0-60 

0-56 

0-63 

0-64 

0-67 

Total  5-24         4-74         4-99         6-24         2-34 

Ratios:  AM  :  AL  :  PM  :  PL  ::  15  :  8  :  2  :  8;  AL— PM— PL  ::  8-9;  AM  :  CL  ::  15  :  2. 
DISTRIBUTION.  West  Sumatra. 

MATERIAL  EXAMINED.  West  Sumatra:  Harau  Nature  Reserve,  near  Payakumbu,  holotype  c?,  on  rock  walls  on 
edge  of  forest,  ca.  600  m,  ii.  1985,  P.  D.  Hillyard,  (BMNH.  1985.9.5. 1);  Taram,  near  Payakumbu,  in  secondary 
forest  litter,  paratype  ?,  ii.1985,  P.  D.  Hillyard,  (BMNH.  1985.9.5.2). 

ETYMOLOGY.  The  specific  name  is  from  the  Latin  meaning  rocky  places. 

Taraxella  sumatrana  sp.  n. 

(Fig.  12A-J) 

DIAGNOSIS.  Easily  separated  from  other  species  of  Taraxella  by  the  dorsal  fringe  of  stout  setae  on 
the  palpal  tibia  (Fig.  12G,  J)  in  males  and  by  the  structure  of  the  epigyne  in  females  (Fig.  12D). 


126 


F.  R.  WANLESS 


Fig.  12  Taraxella  sumatrana  sp.  n.,  holotype  ?:  A,  dorsal  view.  B,  cheliceral  teeth;  D,  epigyne;  F, 
sternum;  I,  maxillae  and  labium.  Paratype  <$:  C,  cheliceral  teeth;  E,  carapace,  lateral  view;  G,  palpal 
tibia,  dorsal  view;  H,  palp,  ventral  view;  J,  palp,  retrolateral  view.  Abbreviations:  e,  embolus;  Ml, 
fan-shaped  element  of  distal  haematodocha. 

FEMALE  HOLOTYPE,  in  fair  condition.  Carapace  (Fig.  12A):  yellow-brown  faintly  tinged  and 
mottled  black  with  a  metallic  sheen  under  some  angles  of  illumination;  rubbed  except  for  some 
scattered  brown  hairs.  Eyes:  laterals  with  black  surrounds;  fringed  by  light  brownish  hairs  and 
some  whitish  ones  below  anterior  median  eyes.  Clypeus  yellow-brown  tinged  black  with  a  few 
black  bristles.  Chelicerae:  pale  yellow-brown,  shiny,  sparsely  clothed  in  brown-black  hairs  and 
some  bristles  with  dense  promarginal  scopulae;  promargin  with  six  teeth,  retromargin  with  eight 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE  1 27 

denticles  (Fig.  12B).  Maxillae  and  labium:  pale  yellow-brown.  Sternum  (Fig.  12F):  pale  yellow- 
brown  with  darker  margins;  thinly  clothed  in  stiff  brown  hairs.  Coxae:  pale  yellow-brown. 
Abdomen:  pale  yellow-brown  tinged  and  mottled  black  with  vague  chevrons  dorsally;  venter  pale 
greyish  yellow;  mostly  rubbed  otherwise  sparsely  clothed  in  patches  of  recumbent  dull  amber 
lanceolate  hairs,  thinly  interspersed  with  erect  black  hairs;  spinnerets  moderately  long,  pale  yellow 
brown  except  anteriors  tinged  with  some  black.  Legs:  moderately  long  and  slender;  generally  pale 
yellow-brown  faintly  tinged  with  some  black;  thinly  clothed  in  pale  greyish  and  black  hairs;  spines 
moderately  strong  and  numerous.  Spination  of  leg  I:  metatarsus  v  2-0-0,  r  0-1-1 ,  d  0-1-2,  p  1-1-1 ; 
tibia  v  1-3-2,  p  0-1-1;  patella  p  0-1-0;  femur  d  0-2-3.  Epigyne  (Fig.  12D):  clothed  in  dark  grey 
hairs. 

Dimensions  (mm):  total  length  4-44;  carapace  length  2-0,  breadth  1-68,  height  1-32;  abdomen 
length  2-32;  eyes,  anterior  row  1-72,  middle  row  1-38,  posterior  row  1-61;  quadrangle  length  1-12 
(56%  of  carapace  length). 


Leg 

1 

2 

3 

4 

Palp 

Femur 

1-52 

1-36 

1-36 

1-76 

0-76 

Patella 

0-78 

0-68 

0-62 

0-68 

0-40 

Tibia 

1-12 

0-96 

1-00 

1-40 

0-52 

IMpf  H  tfl  TQ11Q 

1-02 

0-96 

1.1  1 

1  •% 

1  VI  I/  Id  Id  1    T  LI  .> 

Tarsus 

1    \  t  ~ 

0-56 

\J   .s\J 

0-56 

1    11 

0-58 

1    J\J 

0-70 

0-66 

Total  5-00         4-52         4-67         6-10         2-34 

Ratios:  AM  :  AL  :  PM  :  PL  ::  14  :  8  :  4  :  8;  AL— PM— PL  ::  7-7-5;  AM  :  CL  ::  14  :  3. 

MALE  PARATYPE,  abdomen  and  legs  IV  missing,  otherwise  in  fair  condition.  Similar  to  female 
except  for  the  following.  Carapace  (Fig.  12E):  heavily  suffused  and  mottled  black  especially  on 
sides;  rubbed  except  for  shining  violet  hairs  behind  anterior  eyes.  Clypeus:  with  scattered  fine 
whitish  hairs.  Chelicerae:  yellow-brown  with  sooty  markings,  shiny,  clothed  in  scattered  black 
hairs  with  dense  promarginal  scopulae;  promargin  with  five  teeth  retromargin  with  seven  denticles 
(Fig.  12C).  Spination  of  leg  I:  metatarsus  v  2-2-2,  r  1-0-0,  d  0-2-2,  p  1-0-0;  tibia  v  1-3-2,  p  0-1-1, 
r  1-0-1,  d  1-0-0;  patella  p  0-1-0,  r  0-1-0;  femur  d  0-2-3.  Palp  (Fig.  12G,  H,  J):  femur  and  patella 
pale  yellowish  lightly  suffused  black;  tibia  and  cymbium  light  to  dark  amber  mottled  with  some 
black;  clothed  in  brownish  hairs  with  patch  of  white  hairs  on  patella  and  apices  of  femora. 

Dimensions  (mm):  total  length  ?;  carapace  length  1  -92,  breadth  1  -22,  height  1  -32;  abdomen  length 
?;  eyes,  anterior  row  1-64,  middle  row  1-29,  posterior  row  1-52;  quadrangle  length  1-12  (58%  of 
carapace  length). 


Leg 
Femur 
Patella 
Tibia 
Metatarsus 
Tarsus 

1 
1-58 
0-76 
1-26 
1-16 
0-62 

2 
1-44 
0-68 
1-04 
1-08 
0-60 

3 
1-40 
0-60 
1-08 
1-22 
0-64 

4           Palp 
fl-RO 

0-^fi 

0-d.fi 

0-92 

Total  5-38         4-84         4-94  2-54 

Ratios:  AM  :  AL  :  PM  :  PL  ::  14  :  8  :  4  :  8;  AL— PM— PL  ::  6-7;  AM  :  CL ::  14  :  5. 
DISTRIBUTION.  Sumatra. 

MATERIAL  EXAMINED.  Sumatra,  Bohorok,  Gunung  Leuser  Reserve:  holotype  ?,  in  litter,  14.vi.  1983,  P.  R. 
Deeleman  and  C.  L.  Deeleman- Reinhold;  paratype  £,  in  bamboo  litter  on  plateau  with  large  bamboo  trees, 
50  m,  lO.ii.  1983,  P.  R.  Deeleman  and  S.  Djojosudharma  (Rijksmuseum  van  Natuurlijke  Histoire,  Leiden). 

ETYMOLOGY.  The  specific  name  refers  to  the  country  in  which  the  holotype  was  collected. 


128 


F.  R.  WANLESS 


B 


Fig.  13.  Taraxella  reinholdae  sp.  n.,  holotype  $:  A,  dorsal  view;  B,  carapace,  lateral  view;  C,  leg  I;  D, 
palpal  tibia,  dorsal  view;  E,  cheliceral  teeth;  F,  palp,  ventral  view;  G,  palp,  retrolateral  view.  Abbrevi- 
ations: M 1 ,  fan-shaped  element  of  distal  haematodocha;  tf,  tegular  furrow;  va,  ventral  apophysis;  'x' 
and  'y',  tegular  apophyses. 


Taraxella  reinholdae  sp.  n. 
(Fig.  13A-G) 

DIAGNOSIS.  Easily  separated  from  other  species  of  Taraxella  by  the  conspicuous  tuft  of  stout  setae 
arising  from  the  retrolateral  surface  of  the  palpal  tibial  apophysis  (Fig.  13D,  F,  G). 

FEMALE.  Unknown. 

MALE  HOLOTYPE  in  fair  condition.  Carapace  (Fig.  13 A,  B):  mottled  black  with  metallic  sheen  and 
pale  yellowish  green  markings  on  thoracic  part;  irregularly  clothed,  in  scattered  black  and  white 
hairs  with  a  central  white  haired  stripe  behind  anterior  median  eyes.  Eyes:  laterals  with  black 
surrounds;  fringed  by  white  hairs  with  scattered  dark  grey  hairs  around  laterals  and  upper  rims  of 
anterior  medians.  Clypeus:  clothed  in  scattered  white  hairs.  Chelicerae:  black  with  metallic  sheen 
except  for  light  greyish  amber  inner  margins;  sparsely  clothed  in  light  greyish  hairs  with  moderately 
dense  promarginal  scopulae;  promargin  with  five  teeth,  retromargin  with  seven  denticles  (Fig. 
13E).  Maxillae  and  labium:  pale  yellow-brown  heavily  mottled  black  with  whitish  inner  distal 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE  1 29 

margins  on  maxillae  and  labial  tip.  Sternum:  yellow-brown  suffused  black;  shiny,  with  scattered 
fine  blackish  hairs.  Coxae:  pale  greyish  yellow  suffused  with  some  black.  Abdomen:  dorsum  and 
sides  yellowish  green  heavily  tinged  black;  clothed  in  brown-black  hairs  with  vague  patches  of 
whitish  hairs  laterally;  venter  pale  yellow  with  black  surrounds,  clothed  in  grey  black  hairs; 
spinnerets  black  with  blackish  hairs.  Legs  (Fig.  1 3C):  moderately  long  and  slender;  generally  black 
except  for  pale  yellowish  brown  tarsi  and  yellow-brown  annuli  on  metatarsi,  tibiae  and  patellae; 
spines  strong  and  numerous.  Spination  of  leg  I:  metatarsus  v  2-0-0,  p  1-0-2,  d  1-1-2;  tibia  v  2-2-2, 
p  1-0-1 ,  d  1-1-0;  patella  p  0-1-0,  r  0-1-0;  femur  d  0-2-3.  Palp  (Fig.  1 3D,  F,  G):  patella  pale  yellow 
tinged  with  some  black,  other  segments  heavily  mottled  black;  clothed  in  black  hairs  with  patch  of 
whitish  hairs  on  patella  and  apices  of  femora. 

Dimensions  (mm):  total  length  ca.  3-2  (bent);  carapace  length  1-52,  breadth  1-44,  height  1-16; 
abdomen  length  1-68;  eyes,  anterior  row  1-48,  middle  row  1-16,  posterior  row  1-40;  quadrangle 
length  1-0  (65%  of  carapace  length). 


Leg 

Femur 
Patella 
Tibia 
Metatarsus 
Tarsus 

1 
1-28 
0-66 
1-02 
0-96 
0-52 

2 
1-18 
0-58 
0-84 
0-88 
0-48 

3 
1-20 
0-54 
0-88 
0-96 
0-52 

4 
1-56 
0-60 
1-20 
1-42 
0-61 

Palp 
0-64 
0-32 
0-32 

0-72 

Total  4-44         3-96         4-10         5-39         2-00 

Ratios:  AM  :  AL  :  PM  :  PL  ::  7  :  12-5  :  2  :  7;  AL— PM— PL  ::  8-7;  AM  :  CL  ::  12-5  :  3-5. 
DISTRIBUTION.  Borneo,  Sarawak. 

MATERIAL  EXAMINED.  Sarawak:  Bako  National  Park,  holotype  $,  in  litter,  in  swampy  forest,  29-30.iii.  1 985.  C. 
L.  Deeleman-Reinhold  and  P.  R.  Deeleman,  (RNH,  Leiden). 

ETYMOLOGY.  This  species  is  named  for  one  of  the  collectors  Dr  C.  L.  Deeleman-Reinhold. 


Acknowledgements 

I  would  like  to  thank  Dr  C.  L.  Deeleman-Reinhold,  Ossendrecht,  Holland  and  Dr  P.  Lehtinen,  University  of 
Turku,  Turku,  Finland  for  allowing  me  to  study  their  collections  of  oriental  salticids. 

I  am  also  indebted  to  my  colleague  Mr  P.  D.  Hillyard  for  collecting  spiders  in  Indonesia  and  Malaysia 
and  for  providing  photographs  for  Figs  1-2.  Mrs  F.  Murphy,  London  kindly  supplied  the  photograph  for 
Figure  3. 

Finally  I  would  like  to  thank  Dr  R.  R.  Jackson,  University  of  Canterbury,  Christchurch,  New  Zealand  and 
Mr  D.  Macfarlane,  Commonwealth  Institute  for  Entomology,  London  for  reading  the  manuscript. 


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19856.  Web-building,  predatory  versatility  and  the  evolution  of  the  Salticidae.  In:  W.  A.  Shear,  Ed.,  The 

evolution  of  spiders  webs.  Stanford  University  Press. 
(in  press).  Communal  jumping  spiders  (Araneae:  Salticidae)  from  Kenya:  interspecific  nest  complexes, 

co-habitation  with  web-building  spiders,  and  intraspecific  interactions.  N.Z.  J.  Zool. 
&  Blest,  A.  D.  1982.  The  biology  of  Portia  fimbriata,  a  web-building  jumping  spider  (Araneae,  Salticidae) 

from  Queensland:  utilization  of  webs  and  predatory  versatility.  J.  Zool.  Lond.  196: 255-293. 
&  H alias,  S.  E.  A.  (in  press  a).  Comparative  biology  of  Portia  africana,  P.  albimana,  P.  fimbriata,  P. 

labiata  and  P.  schultzi,  araneophagic  web-building  jumping  spiders  (Araneae,  Salticidae):  utilization  of 

webs,  predatory  versatility,  and  intraspecific  interactions.  N.Z.  Jl.  Zool. 

& (in  press  b).  Predatory  versatility  and  intraspecific  interactions  of  spartaeine  jumping  spiders 


(Araneae:  Salticidae):  Brettus  adonis,  B.  cingulatus,  Cyrba  algerina,  and  Phaeacius  sp.  n.  N.Z.  Jl.  Zool. 
Legendre,  R.  &  Llinares,  D.  1970.  L'accouplement  de  1'araignee  salticide  Cyrba  algerina  (Lucas,  1846).  Annls 

Soc.  Hon.  Hist.  nat.  Herault  110(4):  169-174. 
Loerbroks,  A.  L.  1984.  Mechanik  der  Kopulationsorgane  von  Misumena  vatia  (Clerck,  1757)  (Arachnida: 

araneae:  Thomisidae).  Verh.  naturwisc.  Ver.  Hamburg  (NF)  27:  383-403. 
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. 
Wanless,  F.  R.  1978a.  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. 
19786.  A  revision  of  the  spider  genus  Portia  (Araneae:  Salticidae).  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  34(3): 

83-124. 
1984a.  A  review  of  the  spider  subfamily  Spartaeinae  nom.  n.  (Araneae:  Salticidae)  with  descriptions  of 

six  new  genera.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  46(2):  135-205. 

19846.  A  revision  of  the  spider  genus  Cyrba  (Araneae:  Salticidae)  with  the  description  of  a  new  presump- 
tive pheromone  dispersing  organ.  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  47(7):  445-481. 
1985.  A  revision  of  the  spider  genera  Holcolaetis  and  Sonoita  (Araneae:  Salticidae).  Bull.  Br.  Mus.  nat. 

Hist.  (Zool.)  48(4):  249-278. 

Manuscript  accepted  for  publication  21  February  1986 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


131 


Fig.  14.     Spartaeus  wildtrackii  sp.  n.,  paratype  (J:  femoral  organ:  A,  x  316;  B,  x  820;  C,  x  1040.  Scale 

bars  urn.  Arrow  indicates  direction  of  coxa. 


132 


F.  R.  WANLESS 


Fig.  15.  Spartaeus  wildtrackii  sp.  n.,  cJ,  tibia  II:  A,  dorsal  view  of  distal  region  showing  distribution  of 
mytiliform  organs  and  muscle  attachment  sites,  x  570;  B,  mytiliform  organs  and  muscle  attachment 
sites,  x  1600;  C,  detail  of  mytiliform  organ,  x  4200.  Abbreviations:  h,  hair  sockets;  hi,  hinge  line;  m, 
muscle  attachment  site;  my,  mytiliform  organ.  Scale  bars  urn. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


133 


Fig.  16.  Spartaeus  wildtrackii  sp.  n.,  J:  A,  femora  I,  mytiliform  organs  and  seta,  x  1000;  B,  femora  II, 
detail  of  mytiliform  organ,  x  6200;  C,  metatarsus  II,  base  of  filimentous  seta  showing  dendritic  pore, 
x  10300;  D,  femora  I,  feathery  setae,  x  620.  Scale  bars  ^m. 


134 


F.  R.  WANLESS 


Fig.  17.  Spartaeus  wildtrackii,  sp.  n.,  $:  A,  basal  half  of  metatarsi  II,  lateral  view  showing  fringe  of 
filamentous  setae;  C,  two  filamentous  setae,  x  1300;  D,  E,  F,  mid  region,  distal  region  and  tip  of 
filamentous  setae,  x  7000,  x  8000,  x  8000;  B,  basal  half  of  tarsi  II,  ventral  view  showing  row  of  prey 
capture  setae,  arrowed;  x  210;  G,  detail  of  prey  capture  setae,  x  2000.  Scale  bars  urn. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


135 


Fig.  18.  Spartaeus  wildtrackii  sp.  n.,  tibia  of  male  palpal  organ:  A,  ventral  apophysis  inner  view,  x  1000; 
B,  retrolateral  apophysis,  outer  view,  x  100;  C,  retrolateral  apophysis,  inner  view,  x  100;  D,  detail  of 
dorsal  prong  of  retrolateral  apophysis,  x  350.  Scale  bars  ^im. 


136 


F.  R.  WANLESS 


Fig.  19.  Mintonia  caliginosa  sp.  n.,  $:  A,  underside  of  femora  I  showing  position  of  femoral  organ, 
arrowed,  x  75;  B,  femoral  organ,  x  550;  C,  femoral  organ  showing  pores,  x  2800.  Scale  bars  A,  mm; 
B,  C,  urn. 


NOTES  ON  SPIDERS  OF  THE  FAMILY  SALTICIDAE 


137 


Fig.  20.  (A,  B)  Mintonia  tauricornis  Wanless,  <$,  palpal  organ:  A,  tibia  showing  ventral  apophysis 
and  pore  bearing  retrolateral  apophysis,  x  150;  B,  tip  of  retrolateral  apophysis  showing  pore,  x  850. 
C,  Portia  labiata  Thorell,  <$,  showing  mytiliform  field  on  dorsal  surface  of  abdomen,  x  730. 
Abbreviations:  m,  mytiliform  organs;  va,  ventral  apophysis;  rta,  retrolateral  tibial  apophysis.  Scale 
bars  |im. 


British  Museum  (Natural  History) 

The  birds  of  Mount  Nimba,  Liberia 


Peter  R.  Colston  &  Kai  Curry-Lindahl 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  on  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Tring,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1986,  129pp.  Hardback.  0  565  00982  6  £17.50. 


Titles  to  be  published  in  Volume  52 


Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophora  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicm,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata).  By  P.  J. 

Chimonides 


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


Bulletin  of  the 

British  Museum  (Natural  History) 


Mites  of  the  genus  Holoparasitus  Oudemans 
1936  (Mesostigmata:  Parasitidae)  in  the 
British  Isles 

Keith  H.  Hyatt 


Zoology  series    Vol52    No  4    30  April  1987 


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ISSN  0007-1498  Zoology  series 

Vol52  No.  4  pp  139-1 64 
British  Museum  (Natural  History) 
Cromwell  Road 
London  SW7  5BD  Issued  30  April  1 987 


Mites  of  the  genus  Holoparasitus  Oudemans,  1936 
(Mesostigmata:  Parasitidae)  in  the  British  Isles 

Keith  H.  Hyatt 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7  5BD 


Contents 


Synopsis 

Introduction    .... 

Material 

External  morphology 
Summary  of  classification 
Genus  Holoparasitus  Oudemans 

Key  to  species 

Descriptions  of  species . 
Acknowledgements . 
References 


139 
139 
141 
141 
141 
142 
142 
143 
163 
163 


Synopsis 

The  mites  of  the  genus  Holoparasitus  Oudemans,  1936  occurring  in  the  British  Isles  and  the  Channel  Islands 
are  revised.  Five  species  are  recorded  of  which  two,  H.  lawrencei  and  H.  maritimus,  are  new  to  science,  and  H. 
calcaratus  (C.  L.  Koch,  1839)  is  recorded  for  the  first  time.  A  neotype  is  designated  for  H.  calcaratus.  Habitat 
and  distribution  data  are  given  and  keys  to  species  for  males  and  females  are  provided. 


Introduction 

The  family  Parasitidae  comprises  two  subfamilies,  the  Parasitinae  Oudemans,  1901  and  the 
Pergamasinae  Juvara-Bals,  1972.  Hyatt  (1980)  has  revised  the  British  species  of  Parasitinae. 
Following  Evans  and  Till  (1979)  the  British  Pergamasinae  comprises  the  genera  Pergamasus 
Berlese,  1904,  Amblygamasus  Berlese,  1904,  Paragamasus  Hull,  1918,  Holoparasitus  Oudemans, 
1936  and  Pergamasellus  Evans,  1957.  Of  these,  the  first  three  were  included  by  Bhattacharyya 
(1963)  in  his  revision  of  the  British  species  of  Pergamasus  sensu  lato,  whilst  the  monotypic  genus 
Pergamasellus  (of  which  specimens  have  so  far  been  found  only  in  two  localities  in  southern 
England)  is  figured  by  Evans  and  Till  (1979).  The  remaining  genus,  Holoparasitus,  is  revised  in  the 
present  paper. 

The  genus  Holoparasitus  has  been  mentioned  in  the  British  literature  scarcely  a  dozen  times  and 
only  four  authors,  Halbert  (1915),  Hull  (1918),  Turk  and  Turk  (1952)  and  Turk  (1953),  discussed 
their  determinations.  Halbert  (1915)  recorded  calcaratus  Koch,  pollicipatus  Berlese  and  inornatus 
Berlese  from  the  west  coast  of  Ireland,  but  he  attached  reservations  to  his  identifications.  Hull 
(1918)  recorded  the  same  three  taxa  from  north-east  England.  Turk  and  Turk  (1952)  recorded 
berlesei  Oudemans  and  commented  on  the  confused  synonymy  of  this  species,  echoing  the  remarks 
of  Oudemans  (1936),  whilst  Turk  (1953),  in  his  'Synonymic  Catalogue  of  British  Acari',  listed  all 
the  above  names  plus  four  of  Berlese's  'varieties'  based  on  specimens  in  his  own  collection.  The 
remaining  authors  have  given  species  determinations  without  comment.  Where  these  specimens 
are  available  for  study  they  are  referred  to  under  the  taxa  they  are  considered  to  represent. 


Bull.  Br.  Mus.  not.  Hist.  (Zool.)  52(4):  139-164 


Issued  30  April  1987 


139 


140 


K.  H.  HYATT 


Fig.  1     The  National  and  Irish  Grids,  showing  to  the  nearest  10-kilometre  square  the  extent  of  material 
examined  during  the  preparation  of  the  present  work. 


Mead-Briggs  and  Hughes  (1966)  recorded  Holoparasitus  pollicipatus  from  Cambridgeshire  and 
Davis  (1963)  recorded  Holoparasitus  sp.  from  Northamptonshire.  Unfortunately  none  of  these 
specimens  is  available  for  study. 

The  most  comprehensive  revision  of  the  European  species  of  the  genus  is  by  Karg  (1971)  who 
keys  and  illustrates  nine  species  of  which  three,  calcaratus,  inornatus  and  stramenti  Karg,  are 
represented  in  the  British  fauna. 


MITE  GENUS  HOLOPARASITUS  1 4 1 

Material 

This  revision  is  based  on  the  examination  of  over  six  hundred  specimens.  The  majority 
were  removed  from  Berlese-funnel  extractions  during  the  course  of  my  revision  of  the  British 
Parasitinae  (Hyatt,  1980).  A  few  were  removed  from  subsequent  samples,  mainly  from  areas  of 
northern  England  and  south-west  Scotland  not  previously  sampled.  With  the  exception,  however, 
of  the  monotypic  genus  Pergamasellus,  Holoparasitus  is  not  found  so  abundantly  as  the  other 
genera  of  Pergamasinae,  nor  is  it  apparently  so  widely  distributed. 

Few  named  specimens  were  already  in  the  Museum  collections  and  of  these  the  majority  were  in 
the  J.  H.  Murgatroyd  and  Harry  Britten  collections  from  the  New  Forest  area  and  the  north  of 
England  respectively. 

The  map,  figure  1,  shows  on  the  10-kilometre  squares  of  the  National  and  Irish  Grids  the  extent 
of  the  material  from  which  I  have  examined  Holoparasitus  in  the  British  Isles  and  the  Channel 
Islands.  Localities  for  Great  Britain  are  given  under  the  current  English  and  Welsh  counties  and 
Scottish  regions  or  island  areas,  but  if  a  record  is  from  only  a  small  part  of  the  area  the  former 
county  name  is  inserted  in  parentheses.  Additionally,  large  urban  areas,  e.g.  Manchester  and 
London,  and  islands  or  individual  but  prominent  localities,  e.g.  Isle  of  Wight,  Island  of  Mull,  or 
Dungeness,  are  given  as  such. 


External  morphology 

The  external  morphology  of  Holoparasitus  is  essentially  typical  of  the  Parasitidae  and  the  terms 
used  in  the  descriptions  follow  Evans  and  Till  (1979)  and  Hyatt  (1980).  As  in  the  majority  of  the 
Pergamasinae  the  dorsal  setae  of  the  idiosoma  are  usually  of  uniform  length  and  thickness,  where- 
as in  the  majority  of  species  of  Parasitinae  these  setae  are  heterogeneous.  Holoparasitus  species  are 
usually  readily  separated  from  the  other  pergamasines  by  being  conspicuously  more  spherical  in 
body  outline,  and  even  under  low  magnifications  the  ventral  fusion  of  the  holodorsal  and  opistho- 
gastric  shields  in  the  British  species  of  Holoparasitus  sensu  stricto  is  easily  seen.  The  exception  to 
this  last  character  is  in  the  subgenera  Ologamasiphis  Holzmann,  1969  and  Heteroparasitus  Juvara- 
Bals,  1976  (both  as  yet  not  recorded  from  the  British  Isles)  in  which  the  two  shields  are  not  fused, 
although  the  holodorsal  shield  is  continued  ventrally,  unlike  Pergamasus  sensu  lato  which  has  the 
holodorsal  shield  entirely  dorsal. 


Summary  of  classification 

Oudemans  (1936)  proposed  Holoparasitus  as  a  new  name  for  Ologamasus  Berlese,  1906 
( =  Hologamasus  [lapsus]  Berlese,  1 892)  with  the  type  species  Gamasus  calcaratus  C.  L.  Koch,  1 839. 
The  type  species  of  Ologamasus  Berlese,  1888  is,  by  monotypy,  Gamasus  aberrans  Berlese,  1888,  a 
member  of  the  family  Rhodacaridae  (Ryke,  1962,  Lee,  1970). 

The  first  comprehensive  review  of  the  genus  Holoparasitus  was  contained  in  Berlese's  'Mono- 
grafia  del  genere  Gamasus  Latr.'  published  in  1906.  In  this  work  the  following  ten  taxa  are 
recognised:  Gamasus  (Ologamasus)  calcaratus  Koch,  1839  and  its  two  varieties,  excisus  Berlese, 
1906  and  siculus  Berlese,  1906;  Gamasus  (Ologamasus)  inornatus  Berlese,  1906;  Gamasus 
(Hologamasus)  pollicipatus  Berlese,  1904  and  its  five  varieties  appeninorum,  cultriger,  excipuliger, 
peraltus,  andpseudoperforatus,  all  Berlese,  1906. 

Oudemans  (1936)  considered  that  Berlese's  pollicipatus  and  Koch's  calcaratus  were  both 
synonyms  of  the  earlier  Acarus  lichenis  Schrank,  1781.  However,  Micherdzinski  (1969)  considered 
that  Oudemans  had  no  real  evidence  for  this,  bearing  in  mind  that  Schrank's  figures  were  so  lacking 
in  detail. 

Of  the  recent  authors  Holzmann  (1969)  uses  the  name  Ologamasus  Berlese,  1892,  with  which 
she  erroneously  considers  Holoparasitus  to  be  synonymous.  She  recognises  two  subgenera, 
Ologamasus  s.  str.  and  Ologamasiphis  nov.,  separated  by  several  characters  of  the  deutonymph 


142  K.  H.  HYATT 

and  the  female.  In  the  subgenus  Ologamasus  s.  str.  she  includes  calcaratus  Koch,  1839, 
inornatus  Berlese,  1906,  hemisphaericus  Vitzthum,  1923,  absoloni  Willmann,  1940  and  intermedius 
Holzmann,  1969,  whilst  in  Ologamasiphis  she  includes  rotulifer  Willmann,  1940  and  a  new  species, 
minimus  Holzmann,  1969. 

Micherdzinski  (1969)  follows  Oudemans  in  his  preference  for  Holoparasitus  and  divides  the 
genus  into  four  species-groups:  1 .  calcaratus-group  and  2.  pollicipatus-group,  both  based  on  the 
form  of  the  femoral  spurs  on  leg  II  of  the  male,  3.  Ologamasiphis-group  with  H.  rotulifer  and  H. 
minimus  following  Holzmann,  and  4.  a  group  to  include  females  which  do  not  belong  to  previous 
groups,  H.  excisus  (Berlese)  and  H.  hemisphaericus  (Vitzthum). 

Karg  (1971),  at  couplet  twelve  in  his  key  to  females,  recognises  the  subgenus  Ologamasiphis 
for  three  species,  minimus  Holzmann,  coronarius  Karg  (nom.  nov.  pro  rotulifer  Holzmann,  1969, 
non  Willmann,  1941)  and  tirolensis  Sellnick,  1968,  implying  that  the  preceding  species  belong  to 
Holoparasitus  s.  str. 

Genus  HOLOPARASITUS  Oudemans 

Hologamasus  Berlese,  1904:  235.  Non  Berlese,  1892:  62. 
Ologamasus  Berlese,  1906:  242.  Non  Berlese,  1888:  194. 
Holoparasitus  Oudemans,  1936:  164. 

TYPE  SPECIES:  Gamasus  calcaratus  C.  L.  Koch,  1839. 

Dorsal  and  ventral  shields  of  the  adults  well  sclerotised;  males  with  holodorsal  and  opisthogastric 
shields  always  fused  posteriorly;  females  with  holodorsal  and  opisthogastric  shields  fused 
posteriorly  in  Holoparasitus  s.  str.,  but  free  in  the  subgenus  Ologamasiphis  Holzmann. 
Deutonymphs  with  separate  podonotal  and  opisthonotal  shields,  not  strongly  sclerotised.  In  all 
stages  setae  of  dorsal  hexagon,  i.e./5,  z5  and 76,  similar  to  each  other  and  not  differing  markedly 
from  the  remaining  dorsal  stage  which  are  generally  short  (not  exceeding  50  um)  and  often 
inconspicuous.  Tritosternum  of  male  biramous  and  sometimes  modified,  base  closely  associated 
with  genital  orifice.  Tristosternum  of  deutonymph  and  female  normal,  biramous.  Junction 
between  sternal  and  metasternal  shields  of  female  clique.  Genital  shield  of  female  broadly 
pentagonal.  Opisthogaster  with  usually  not  more  than  15  pairs  of  setae.  Setaea/ofpalptrochanter 
bifid  and  with  one  or  more  distinct  slender  processes;  setae  alv  and  al2  of  palp  femur  entire, 
spatulate  or  setiform.  Corniculi  strong,  entire  or  notched  internally.  Legs  of  deutonymph  and 
female  without  spurs;  only  leg  II  of  male  spurred.  Lobes  of  pulvilli  normal,  rounded. 

Key  to  species 
Males 

1  Apophysis  on  femur  II  thumb-shaped,  about  twice  as  long  as  axillary  process  (Fig.  2G)       .        .         2 
Apophysis  on  femur  II  short,  hemispherical  and  not  extending  beyond  the  blunt  axillary  process 

(Fig.  80) 3 

2  Sternogenital  shield  with  a  conspicuous  'excipulum'  medially  between  coxae  II  and  HI  (Fig.  2B); 

majority  of  dorsal  setae  extremely  short  (c.  10-20  um)  (Fig.  2A);  idiosoma  590-635  um 

...        .        .         .        .        .     Holoparasitus  calcaratus  (C.  L.  Koch)  (p.  143) 

Sternogenital  region  without  such  a  median  'excipular'  mark;  dorsal  setae  generally  exceeding 
25  um  in  length;  idiosoma  520-570  um    .         .         .         .          Holoparasitus  stramenti  Karg  (p.  146) 

3  Anterior  margin  of  Sternogenital  shield  strongly  concave  medially  to  behind  sternal  setae  I;  a 

conspicuous  and  shallow  transverse  structure  line  present  at  sternal  setae  II  (Fig.  8B);  idiosoma 

530-590  um   .         . Holoparasitus  inornatus  ( Berlese)  (p.  151) 

Anterior  margin  of  Sternogenital  shield  not  strongly  concave  medially;  sternal  ornamentation 
otherwise;  idiosoma  exceeding  680  um 4 

4  Large  species  -  idiosoma  780-840  um;  sternal  region  with  a  light  but  strongly  procurved  line 

reaching  forward  from  sternal  setae  II  (Fig.  1  OB);  tectum  trispinate,  centre  spine  long  (Fig.  IOC); 

corniculi  smooth Holoparasitus  lawrencei  sp.  n.  (p.  1 55) 

Smaller  species  -  idiosoma  680-750  um;  sternal  region  with  reticulations  only;  tectum  broadly 
triangular,  granular  (Fig.  13C);  corniculi  cleft  on  inner  margins  (Fig.  13G) 

i Holoparasitus  maritimussv.  n.  (p.  158) 


MITE  GENUS  HOLOPARASITUS  143 

Females 

1  Genital  shield  produced  anteriorly  to  a  tongue-shaped  apex  (Fig.  6B);  endogynium  as  in  figure  6C; 

idiosoma  590-670  urn     .        .  ......          Holoparasitus  stramenti  Karg  (p.  146) 

Genital  shield  with  anterior  margin  forming  an  obtuse  to  right  angle,  with  or  without  a  small 
protruding  tip  (Fig.  9B)         .....         ........         2 

2  Sternal  shield  divided  longitudinally  into  two  (Fig.  9B);  endogynium  appears  as  a  circular  granular 

structure  (Fig.  9C);  idiosoma  6  10-690  um        .         .         .    Holoparasitus  inornatus  (Berlese)  (p.  151) 
Sternal  shield  entire  (Fig.  11  B)  .............         3 

3  Sternal  shield  with  a  pair  of  indentations  on  the  anterior  margin  internally  to  sternal  setae  I  (Fig. 

3B);  endogynium  as  in  figure  3C;  majority  of  dorsal  setae  extremely  short  (c.  10-25  um)  (Fig. 
3A);  idiosoma  670-720  jam      .....     Holoparasitus  calcaratus  (C.  L.  Koch)  (p.  143) 
Sternal  shield  with  anterior  margin  smooth,  without  indentations       ......         4 

4  Sternal  shield  granular,  with  a  distinct  median  pattern,  pre-sternal  shields  entire  or  divided, 

strongly  denticulate  (Fig.  14B);  endogynium  as  in  figure  14C;  idiosoma  770-810  urn 

...........         Holoparasitus  maritimus  sp.  n.  (p.  158) 

Sternal  shield  smooth,  no  distinct  median  pattern;  pre-sternal  shields  entire  or  divided,  granular 

(Fig.  1  IB);  endogynium  as  in  figure  1  1C;  idiosoma  850-920  um 

.        ..........         Holoparasituslatvrenceisp.n.(p.\55) 

Descriptions  of  species 

Holoparasitus  calcaratus  (C.  L.  Koch) 
(Figs  2A-G, 


Gamasus  calcaratus  C.  L.  Koch,  1839:  Heft  26,  Taf.  6. 
Gamasus  timidulus  C.  L.  Koch,  1839:  Heft  26,  Taf.  7. 
Gamasus  (Ologamasus)  calcaratus:  Berlese,  1906:  245. 
Holoparasitus  calcaratus:  Micherdzinski,  1969:  354. 
Ologamasus  (Ologamasus)  calcaratus:  Holzmann,  1969:  47. 
Holoparasitus  excipuliger:  Karg,  1971:  361.  Non  Berlese,  1906. 
Ologamasus  pollicipatus  Berlese,  1904;  1913:  203  (in  part). 

Designation  of  a  neotype 

Oudemans  (1906,  1929,  1936)  opined  that  Gamasus  timidulus  C.  L.  Koch  (?)  and  Gamasus 
calcaratus  C.  L.  Koch  (^)  were  junior  synonyms  of  Acarus  lichenis  Schrank,  1781.  However, 
Schrank's  and  Koch's  specimens  are  no  longer  in  existence  and  although  Micherdzinski  (  1  969)  and 
Karg  (1971)  have  accepted  that  timidulus  is  a  synonym  of  calcaratus,  they  have  not  accepted  the 
validity  of  lichenis.  Since  Oudemans  (1936)  it  has  been  universally  accepted  that  calcaratus  is 
the  type  of  Holoparasitus,  but,  unfortunately,  authors'  concepts  of  calcaratus  have  not  been 
consistent.  Specimens  labelled  Ologamasus  pollicipatus  in  the  set  of  slides  accompanying  the 
Acarotheca  Italica  (Berlese,  1913)  in  the  BM(NH)  collection  are  referable  to  the  present  species, 
whereas  the  specimens  similarly  labelled  in  the  set  in  the  Oudemans  collection  are  rotulifer 
Willmann,  1940. 

Through  the  courtesy  of  Dr  L.  van  der  Hammen  I  have  been  able  to  examine  material  from 
several  places  very  close  to  Regensburg,  the  type  locality.  Of  the  15  samples  of  Gamasina 
examined,  seven  contained  Holoparasitus  and  of  these  six  contained  specimens  of  H  .  calcaratus. 
One  sample  contained  a  single  male  and  female  of  another  species. 

A  female  from  Dechbetten,  1-5  miles  (2km)  west  of  Regensburg,  Bavaria,  19  July  1959,  in 
rotting  dry  litter,  L.  van  der  Hammen  coll.,  is  designated  as  the  neotype  of  Gamasus  calcaratus  C.  L. 
Koch.  The  specimen  is  deposited  in  the  Rijksmuseum  van  Natuurlijke  Historic,  Leiden,  together 
with  deutonymphs,  males  and  other  females  from  the  same  sample.  Three  males,  three  females  and 
one  deutonymph  from  this  sample  are  retained  in  the  British  Museum  (Natural  History)  through 
the  courtesy  of  Dr  van  der  Hammen. 

MALE.  The  holodorsal  and  opisthogastric  shields,  which  are  fused  ventrally  posterior  to  coxae  IV, 
are  strongly  sclerotised  and  completely  reticulated  (Figs  2A,  B).  The  dorsum  (Fig.  2A)  measures 
590-635  um  long  x  430-480  um  wide  and  bears  over  50  pairs  of  simple  setae  ranging  in  length  from 


144 


K.  H.  HYATT 


Fig.  2    Holoparasitus  calcaratus  (C.  L.  Koch),  male  -  A  dorsum,  B  venter,  C  tectum,  D  chelicera,  E  palp 
trochanter,  femur  and  genu,  F  venter  of  gnathosoma,  G  leg  II. 

35  urn  (the  vertical  setae, y'l)  to  10  um  or  less  for  the  majority  in  the  opisthonotal  region.  The  setae 
are  not  arranged  entirely  symmetrically. 

The  tritosternum  comprises  two  slender  pilose  laciniae  that  arise  from  below  the  anterior  margin 
of  the  genital  lamina  (Fig.  2B).  The  sternogenital  region  is  strongly  reticulated  and  bears  medially 
between  coxae  II  and  III  a  conspicuous  oval  mark  (the  'excipulum'  of  Berlese).  The  sternogenital 


MITE  GENUS  HOLOPARASITUS 


145 


Fig.  3    Holoparasitus  calcaratus  (C.  L.  Koch),  female  -  A  dorsum,  B  venter,  C  endogynium,  D  tectum,  E 
chelicera,  F  palp  trochanter,  femur  and  genu,  G  venter  of  gnathosoma. 

and  median  opisthogastric  setae  are  up  to  40  nm  in  length,  whilst  the  three  anal  setae  and  those 
situated  posteriorly  in  the  opisthogastric  region  are  as  little  as  10  um.  The  stigma  is  situated 
opposite  the  posterior  margin  of  coxa  III  and  the  peritreme  extends  anteriorly  to  coxa  I. 

The  tectum  is  shown  in  figure  2C.  The  median  portion  is  broad  and  blunt  in  all  specimens 
examined  and  the  lateral  teeth  vary.  The  chelicera  is  shown  in  detail  in  figure  2D.  The  movable  digit 


146  K.H.HYATT 

is  75  um  long  and  bears  one  large  tooth  and  5-6  small,  sometimes  irregular,  teeth.  The  fixed  digit 
bears  one  sometimes  blunt  tooth  towards  the  tip  and  up  to  seven  very  small  teeth  in  the  region  of 
the  pilus  dentilis.  The  palp  trochanter,  femur  and  genu  are  shown  in  figure  2E.  The  anterolateral 
setae  of  the  femur  and  genu  are  spatulate,  that  on  the  genu  is  more  slender  and  is  bifurcate  near  to 
its  tip.  The  corniculi  (Fig.  2F)  are  strong  and  stalked  and  all  the  gnathosomal  setae  are  simple.  Leg 
II  is  shown  in  detail  in  figure  2G.  The  femoral  apophysis  is  thumb-shaped  and  is  about  twice  as  long 
as  the  axillary  process.  The  ventral  process  on  the  genu  is  short  and  rounded  whilst  that  on  the  tibia 
is  elongate  and  bean-shaped.  All  leg  setae  are  slender  whilst  a  number  on  the  tarsi  of  legs  II-IV  are 
stronger  and  pilose  on  one  margin.  The  ambulacra  are  normal. 

FEMALE.  The  holodorsal  and  opisthonotal  shields,  which  are  strongly  sclerotised  and  completely 
reticulated,  are  fused  ventrally  anterior  to  the  anus  (Figs  3A,  B).  The  dorsum  (Fig.  3A)  measures 
670-720  (im  long  x  500-560  |im  wide  and  bears  up  to  57  or  more  pairs  of  simple  setae,  about  20 
pairs  being  in  the  podonotal  region  and  from  34-37  pairs  in  the  opisthonotal  region.  The  vertical 
setae,y7,  are  the  longest  and  measure  c.  40  urn,  whilst  the  shortest  setae,  the  majority  of  those  in  the 
opisthonotal  region,  measure  as  little  as  10  urn  and  are  extremely  fine. 

The  tritosternum  (Fig.  3B)  has  a  narrow  base  and  the  paired  laciniae  are  simple.  The  presternal 
shields  are  fused  to  form  a  single  narrow  bar.  The  ventral  shields  are  strongly  reticulated  and  all 
setae  are  simple.  The  sternal  shield  has  a  pair  of  indentations  on  the  anterior  margin  inwards  from 
sternal  setae  I.  The  reticulations  show  a  bold  procurved  transverse  line  extending  from  posterior  to 
coxae  II  through  sternal  pores  II.  The  genital  shield  measures  1 30-1 60  um  long  x  210-220  urn 
wide.  Its  anterior  margin  is  formed  into  a  slight  obtuse  angle  or  right  angle  and  bears  a  slender 
pointed  tip.  The  endogynium  is  shown  in  figure  3C  and  in  the  specimens  examined  shows  con- 
stancy in  form  despite  clearing  for  examination.  The  opisthogastric  region  bears  8-9  pairs  of  setae. 
The  longest  of  the  ventral  setae  are  sternal  setae  II  and  III  which  measure  up  to  75  |im.  The  shortest 
ventral  setae  are  around  the  posterior  margin  and  measure  10  uin  or  less.  In  most  of  the  specimens 
examined  the  outline  of  the  deutonymphal  anal  shield  is  retained.  The  stigma  is  situated  opposite 
the  posterior  margin  of  coxa  III  and  the  peritreme  extends  anteriorly  to  coxa  I. 

The  tectum  (Fig.  3D)  has  the  centre  portion  tapered  and  sinuous.  The  chelicera  is  shown  in  figure 
3E.  The  movable  digit  measures  87  urn  and  bears  three  prominent  teeth  and  two  small  teeth,  whilst 
the  fixed  digit  bears  five  teeth  decreasing  in  size  towards  its  tip.  The  palp  trochanter,  femur  and 
genu  are  shown  in  figure  3F.  The  anterolateral  seta  of  the  genu  is  bifurcate  whilst  those  of  the  tibia 
are  spatulate.  The  venter  of  the  gnathosoma  is  shown  in  figure  3G.  The  internal  posterior  hypo- 
stomatic  setae  are  the  longest  (70  um  or  more)  and  with  the  palpcoxal  setae  are  pilose,  at  least  on 
one  margin.  The  hypognathal  groove  shows  only  about  seven  rows  of  denticles.  The  leg  setae  are 
slender  and  a  few  on  the  tarsi  are  pilose  on  one  margin.  The  ambulacra  are  normal. 

MATERIAL  EXAMINED.  14  samples -7^^,  169$. 

ENGLAND:  Hampshire,  Surrey,  Sussex/Kent  border  near  the  coast,  Cambridgeshire  (including 
Huntingdonshire). 
This  species  is  recorded  mainly  from  grasses  and  leaf-litter  in  damp  habitats. 

As  far  as  I  can  establish,  there  are  no  previous  authenticated  records  from  the  British  Isles. 

Holoparasitus  stramenti  Karg 
(Figs  4A-F,  5A-G,  6A-G) 

Holoparasitus  stramenti  Karg,  1971:  356. 

Gamasus  (Ologamasus)  calcaratus  var.  excisus  Berlese,  sensu  Halbert,  1915:  54  (in  part). 

Gamasus  (Ologamasus)  pollicipatus  Berlese,  sensu  Halbert,  1915:  55. 

Holoparasitus  pollicipatus  (Berlese)  sensu  Browning,  1956:  386. 

DEUTONYMPH.  The  lightly  reticulated  dorsal  shields  are  weakly  sclerotised  and  yellowish-brown  in 
colour.  The  podonotal  shield  averages  235  um  long  x  325  um  wide.  It  bears  17  pairs  of  simple 
slender  setae,  none  longer  than  30  um  (Fig.  4A).  The  opisthonotal  shield  averages  180um 
long  x  225  um  wide  and  bears  12  pairs  of  simple  slender  setae  from  20-25  um  in  length.  The 
surrounding  membrane  bears  dorsally  on  each  side  about  20  short,  simple  setae. 


MITE  GENUS  HOLOPARASITUS 


147 


Fig.  4    Holoparasitus  stramenti  Karg,  deutonymph  -  A  dorsum,  B  venter,  C  tectum,  D  chelicera,  E  palp 
trochanter,  femur  and  genu,  F  venter  of  gnathosoma. 


The  tritosternum  has  a  narrow  base  and  pilose  laciniae.  The  sternal  shield  (Fig.  4B)  averages 
1 80  um  long  and  is  lightly  reticulated  and  weakly  sclerotised.  The  sternal  setae  average  25  urn  long 
and  are  simple.  Pre-sternal  shields  absent.  The  oval  anal  shield  bears  the  usual  three  setae,  each 
simple  and  about  12  ^im  in  length.  The  stigma  is  situated  opposite  the  anterior  margin  of  coxa  IV, 
and  the  granular  peritreme  and  peritrematal  shield  extend  anteriorly  to  the  level  of  coxa  I.  The  1 1  to 
15  or  more  pairs  of  simple  opisthogastric  setae  are  slender. 

The  triangular  tectum  is  serrated  anteriorly  and  bears  a  slender  tip  (Fig.  4C).  The  chelicerae  are 
as  in  figure  4D,  the  movable  digit  measures  58  um  in  the  figured  specimen.  The  palp  trochanter, 
femur  and  genu  are  shown  in  figure  4E.  The  anterolateral  setae  of  the  femur  and  genu  are  spatulate. 
The  corniculi  and  the  venter  of  the  gnathosoma  are  as  in  figure  4F,  the  gnathosomal  setae  being 
simple  with  the  internal  hypostomatics  the  longest.  The  hypognathal  denticles  are  not  discernible. 
All  leg  setae  are  slender,  the  majority  are  simple  but  some,  especially  on  tarsus  II,  are  finely  pilose 
on  one  margin.  The  pulvilli  are  normal,  rounded  and  with  two  claws. 


148 


K.  H.  HYATT 


r    tT^  ^  . 


7.  v    \i.  -VM 


Fig.  5    Holoparasitus  stramenti  Karg,  male -A  dorsum,  B  venter,  C  tectum,  D  chelicera,  E  palp 
trochanter,  femur  and  genu,  F  venter  of  gnathosoma,  G  leg  II. 

MALE.  The  holodorsal  and  opisthogastric  shields,  which  are  strongly  sclerotised  and  completely 
reticulated,  are  fused  ventrally  anterior  to  the  anus  (Figs  5A,  B).  The  dorsum  (Fig.  5A)  measures 
520-570  um  long  x  330-400  um  wide,  is  finely  granular  and  bears  45  pairs  of  simple  setae,  20  pairs 
in  the  podonotal  region  and  25  pairs  in  the  opisthonotal  region.  The  vertical  setae,  jl,  are  the 
longest,  measuring  up  to  45  um.  The  remaining  setae  are  20-25  um  long. 

The  tritosternum  comprises  two  slender  laciniae  which  arise  from  below  the  anterior  margin  of 
the  genital  lamina  (Fig.  5B).  The  sternogenital  region  is  reticulated  and  the  pattern  is  constant  for 


MITE  GENUS  HOLOPARASITUS  1 49 

the  species.  The  sternogenital  setae  and  the  median  opisthogastric  setae  are  the  longest  -  up  to 
45  um  -  whilst  the  remainder  are  considerably  shorter.  The  three  anal  setae  are  simple.  The  stigma 
is  situated  opposite  the  posterior  margin  of  coxa  IV  and  the  peritreme  extends  to  coxa  I. 

The  tectum  (Fig.  5C)  is  normally  symmetrically  formed  and  is  trispinate.  The  chelicera  is  shown 
in  figure  5D;  the  movable  digit  is  50  um  long  in  the  figured  specimen  and  bears  a  single  tooth  near 
the  tip,  whilst  the  fixed  digit  bears  two  prominent  teeth  and  about  four  smaller  ones.  The  palp 
trochanter,  femur  and  genu  are  shown  in  figure  5E.  The  anterolateral  setae  of  the  femur  and  genu 
are  spatulate.  The  venter  of  the  gnathosoma  is  as  in  figure  5F.  The  corniculi  are  stalked  and  strong. 
The  gnathosomal  setae  are  simple  and  the  hypognathal  groove  bears  about  nine  rows  of  denticles. 
Leg  II  is  shown  in  detail  in  figure  5G.  The  femoral  apophysis  is  strong  and  thumb-like  whilst  the 
axillary  process  is  short,  but  elongate.  The  ventral  process  on  the  genu  is  swollen,  whilst  that  on  the 
tibia  is  not  prominent.  All  the  leg  setae  are  slender,  some  on  the  tarsi  being  pilose  on  one  margin. 
The  ambulacra  are  normal,  with  rounded  pulvilli  and  two  claws. 

FEMALE.  The  holodorsal  and  opisthogastric  shields,  which  are  strongly  sclerotised  and  completely 
reticulated,  are  fused  ventrally  anterior  to  the  anus  (Figs  6A,  B).  The  dorsum  (590-670  um 
long  x  410-475  um  wide)  is  finely  granular  and  bears  48-49  pairs  of  simple  setae  (Fig.  6A),  20  pairs 
in  the  podonotal  region  and  28-29  pairs  in  the  opisthonotal  region.  The  vertical  setae,  jl,  are  the 
longest,  measuring  up  to  40  um.  The  remaining  setae  are  about  20-25  um  in  length. 

The  tritosternum  (Fig.  6B)  has  a  narrow  base  and  pilose  laciniae.  The  presternal  shields  are 
fused  to  form  a  strong  transverse  bar,  occasionally  with  a  small  section  at  each  end  being  almost 
or  entirely  detached.  The  ventral  shields  are  completely  reticulated  and  are  granular.  The 
reticulations  on  the  sternal  shield  follow  a  definite,  but  simple,  pattern  with  a  pair  of  oblique  lines 
originating  from  the  angle  between  coxae  II  and  III  and  passing  through  sternal  pores  II,  but 
petering  out  before  reaching  the  centre  of  the  shield.  The  genital  shield  measures  130-183  um 
long  x  190-225  um  wide,  its  size  appearing  to  be  related  to  the  actual  size  of  the  specimen.  The 
genital  shield  is  unique  among  the  British  species  of  the  genus  in  that  it  is  produced  anteriorly  into  a 
strong  tongue-shaped  apex.  The  endogynium  is  shown  in  figure  6C;  its  general  content  appears 
constant,  but  the  position  of  the  'teeth'  varies  considerably  due  to  distortion  in  life  or  during 
preparation  for  examination.  The  opisthogastric  region  bears  8-9  pairs  of  setae.  The  sternal  setae 
are  the  longest  -  c.  45  um  -  and  the  metasternal,  genital  and  opisthogastric  setae  decrease  slightly 
in  length  in  that  order.  The  three  anal  setae  are  short  (c.  12  um)  and  simple.  The  stigma  is  situated 
opposite  the  posterior  margin  of  coxa  III  and  the  peritreme  extends  anteriorly  to  the  level  of  coxa  I. 

The  tectum  (Fig.  6D)  is  similar  to  that  of  the  male,  but  in  some  specimens  irregularity  is  present. 
The  chelicera  is  as  in  figure  6E.  The  movable  digit  measures  82  um  in  the  figured  specimen  and 
bears  three  teeth,  whilst  the  fixed  digit  bears  up  to  six  less  prominent  teeth.  The  palp  trochanter, 
femur  and  genu  are  shown  in  figure  6F.  The  anterolateral  setae  of  the  femur  and  genu  are  spatulate. 
The  venter  of  the  gnathosoma  is  shown  in  figure  6G.  The  palpcoxal  setae  are  pilose  on  one  margin 
whilst  the  other  gnathosomal  setae  are  simple.  The  hypognathal  groove  bears  about  eight  rows  of 
denticles.  Some  of  the  leg  setae  are  finely  pilose  on  one  margin.  The  ambulacra  are  normal. 

MATERIAL  EXAMINED.  93  samples  -  3  DNN,  80  £&  178  ??. 

ENGLAND:  Isles  of  Scilly,  Devon,  Somerset,  Berkshire,  Hampshire,  Surrey,  Sussex,  Kent, 
Middlesex,  Hertfordshire,  Buckinghamshire,  Gloucestershire,  Bedfordshire,  Norfolk,  Suffolk, 
Cambridgeshire  (including  Huntingdonshire),  Hereford  and  Worcester,  Leicestershire, 
Cheshire,  North  Yorkshire,  Cumbria  (Cumberland,  Westmorland),  Northumberland. 
SCOTLAND:  Lothian/Borders,  Strathclyde  (Argyllshire),  Tayside  (Perthshire),  Inner  Hebrides 
(Mull,  lona,  Eigg). 

WALES:  Gwynedd  (Caernarvonshire),  Dyfed  (Cardiganshire),  Clwyd,  Gwent,  Glamorgan. 
IRELAND:  Galway,  Mayo,  Clare,  Leitrim. 
CHANNEL  ISLANDS:  Jersey. 

Although  this  species  has  been  collected  from  leaf-litter,  compost,  moss  and  grassland,  it  has  a 
marked  preference  for  wet,  marshy  habitats.  One  female  from  Co.  Mayo,  Ireland,  was  taken  on  the 
seashore. 


150 


K.  H.  HYATT 


Fig.  6    Holoparasitus  stramenti  Karg,  female -A  dorsum,  B  venter,  C  endogynium,  D  tectum, 
E  chelicera,  F  palp  trochanter,  femur  and  genu,  G  venter  of  gnathosoma. 


DISTRIBUTION.  Karg  (1971)  stated  that  the  holotype  of  this  species  came  from  the  Baltic  coast  and 
gave  its  distribution  as  central  Europe.  The  name  has  not  appeared  subsequently  in  the  literature. 
H.  stramenti  is  one  of  the  two  most  abundant  British  members  of  the  genus.  It  was  recorded  from 
Co.  Mayo,  Ireland,  by  Halbert  (1915)  as  Gamasus  (Ologamasus)  calcaratus  var.  excisus  Berlese 
from  Achill  Island  and  Westport,  and  as  Gamasus  (Ologamasus)  pollicipatus  Berlese  from  Clare 
Island.  Two  females  of  this  species  were  recorded  from  Jersey  by  Browning  (1956)  as  Holoparasitus 
pollicipatus  (Berlese).  A  single  undetermined  female  was  recorded  from  Moor  House  National 
Nature  Reserve,  Westmorland,  by  Block  (1965),  whilst  Davis  (1970)  recorded  a  single  female  from 
Monks  Wood  National  Nature  Reserve  as  Holoparasitus  ?pollicipatus. 


MITE  GENUS  HOLOPARASITUS  1 5 1 

Holoparasitus  inornatus  (Berlese) 
(Figs  7A-F,  8A-G,  9A-G) 

Gamasus  (Ologamasus)  inornatus  Berlese,  1906:  257. 

Holoparasitus  inornatus:  Schweizer,  1961:  34  ($  only),  Micherdzinski,  1969:  366. 

Ologamasus  (Ologamasus)  inornatus'.  Holzmann,  1969:  47. 

Holoparasitus  calcaratus  (Koch,  1839)  sensu  Schweizer,  1961:  36  (J  only).  Karg,  1971:  361. 

Gamasus  (Ologamasus)  calcaratus  Koch,  1839  sensu  Halbert,  1915:  54  (part). 

DEUTONYMPH.  The  dorsal  shields  are  light  yellowish-brown  in  colour,  weakly  sclerotised  and 
faintly  reticulated  (Fig.  7A).  The  podonotal  shield  averages  460  um  long  x  300  um  wide  when 


Fig.  7    Holoparasitus  inornatus  (Berlese),  deutonymph  -  A  dorsum,  B  venter,  C  tectum,  D  chelicera, 
E  palp  trochanter,  femur  and  genu,  F  venter  of  gnathosoma. 


1 52  K.  H.  HYATT 

flattened  and  depending  on  one  or  more  of  the  marginal  (r-series)  setae  being  on  or  off  the  shield, 
bears  16-17  pairs  of  fine  simple  setae,  none  measuring  more  than  about  35  um.  The  opisthonotal 
shield  averages  180  um  long  x  210  um  wide  and  bears  12  pairs  of  simple  setae  up  to  c.  25  um  in 
length.  All  the  dorsal  setae  taper  extremely  finely.  The  surrounding  membrane  bears  dorsally  on 
each  side  up  to  about  20  short  simple  setae. 

The  tritosternum  has  a  narrow  base  and  pilose  laciniae.  The  sternal  shield  (Fig.  7B)  measures 
160-170  um  long  and  is  lightly  reticulated  and  weakly  sclerotised.  The  setae  are  simple.  Presternal 
shields  absent.  The  oval  anal  shield  bears  the  usual  three  setae,  each  simple  and  about  12-15  um  in 
length.  The  stigma  is  situated  opposite  the  anterior  margin  of  coxa  IV  and  the  granular  peritreme 
and  peritrematal  shield  extend  anteriorly  to  the  level  of  coxa  I.  The  opisthogastric  setae,  which 
number  about  1 7  pairs,  are  fine  and  simple. 

The  tectum  (Fig.  7C)  is  triangular  and  is  flanked  by  small  tooth-like  projections  each  side.  The 
chelicera  is  shown  in  figure  7D;  the  movable  digit  measures  72  um  in  the  figured  specimen  and  bears 
three  teeth,  whilst  the  fixed  digit  bears  about  five  teeth.  The  palp  trochanter,  femur  and  genu  are 
shown  in  figure  7E.  The  anterolateral  seta  of  the  femur  is  broad  and  pectinate  on  one  margin  and 
the  two  anterolateral  setae  of  the  genu  are  spatulate.  The  corniculi  and  venter  of  the  gnathosoma 
are  shown  in  figure  7F.  The  four  pairs  of  gnathosomal  setae  are  simple  whilst  the  hypognathal 
denticles  are  indistinct.  All  leg  setae  are  slender,  the  majority  are  simple,  but  a  few  on  tarsus  II  are 
finely  pilose  on  one  margin.  The  ambulacra  are  normal,  with  rounded  pulvilli  and  two  claws. 

MALE.  The  holodorsal  and  opisthogastric  shields  are  fused  ventrally  posterior  to  coxae  IV  and  are 
heavily  sclerotised  (Figs  8A,  B).  The  dorsum,  which  measures  530-600  um  long  x  390-460  um 
wide,  is  strongly  granular  with  almost  no  trace  of  reticulations  (Fig.  8A).  It  bears,  apparently,  a 
fairly  constant  and  symmetrically  arranged  number  of  setae,  20  pairs  in  the  podonotal  region  and 
30-31  pairs  in  the  opisthonotal  region.  The  longest  setae,  the  verticals  (/7),  measure  c.  30  um  and 
the  tendency  is  for  the  setae  to  decrease  in  length  towards  the  posterior  of  the  dorsum  where  some 
are  as  short  as  10  um. 

The  tristosternum  comprises  a  short  base  and  two  pilose  laciniae  (Fig.  8B).  The  sternogenital 
region  is  strongly  reticulated  and  characteristically  shaped.  The  anterior  margin  of  the  sterno- 
genital shield  is  recessed  deeply  to  accommodate  the  genital  lamina  and  immediately  posterior  to 
sternal  setae  II  there  is  a  strong  procurved  ridge  right  across  the  shield.  These  two  formations  give 
this  anterior  region  a  very  characteristic  appearance.  The  posterior  two-thirds  of  the  sternogenital 
region  are  characteristically  ornamented  and  are  clearly  separated  from  the  opisthogastric  region 
level  with  the  posterior  third  of  coxae  IV,  only  the  endopodal  shields  retaining  their  fusion.  The 
sternogenital  and  opisthogastric  setae  are  up  to  50  um  in  length,  whilst  the  three  anal  setae  and 
those  around  the  posterior  margin  of  the  ventral  surface  are  about  18  um  long.  The  stigma  is 
situated  opposite  the  posterior  margin  of  coxa  III  and  the  peritreme  extends  anteriorly  to  coxa  I. 

The  tectum  is  shown  in  figure  8C.  The  broad  central  part  is  acutely  tapered  and  is  flanked  on  each 
side  by  a  short  prong.  The  chelicera  is  shown  in  figure  8D.  The  movable  digit  is  83  um  long  and 
bears  four  or  five  small  teeth.  The  fixed  digit  bears  six  to  eight  small  teeth.  The  palp  trochanter, 
femur  and  genu  are  shown  in  figure  8E.  The  anterolateral  setae  of  the  genu  and  femur  are  spatulate. 
The  corniculi  (Fig.  8F)  are  strong  and  stalked  and  are  cleft  to  a  varying  degree  on  their  inner 
margins.  The  palpcoxal  setae  are  finely  plumose,  the  remaining  three  pairs  of  gnathosomal  setae 
are  simple.  There  are  about  10  rows  of  hypognathal  denticles.  Leg  II  is  shown  in  detail  in  figure  8G. 
The  apophysis  on  femur  II  is  short,  hemispherical,  and  does  not  extend  beyond  the  tip  of  the 
similarly  shaped  axillary  process.  The  ventral  processes  on  the  genu  and  tibia  are  smooth  and 
elongate.  All  leg  setae  are  slender,  some  on  tarsi  II-IV  are  finely  pilose  on  one  margin.  The 
ambulacra  are  normal. 

FEMALE.  The  holodorsal  and  opisthonotal  shields  are  fused  ventrally  posterior  to  coxae  IV  and 
are  heavily  sclerotised  (Figs  9 A,  B).  The  dorsum  (Fig.  9A),  which  measures  61 0-690  um 
long  x  420-570  urn  wide,  is  strongly  granular  and,  like  the  male,  has  almost  no  trace  of  reticulation. 
It  bears  about  50  pairs  of  simple  setae,  up  to  23  pairs  in  the  podonotal  region  and  up  to  27  pairs  in 
the  opisthonotal  region.  Setae//,  the  verticals,  are  the  longest,  measuring  about  35  um  and  the 
remainder  are  slightly  shorter  with  a  minimum  length  of  c.  18  um  posteriorly. 


MITE  GENUS  HOLOPARASITUS 


153 


Fig.  8    Holoparasitus  inornatus  (Berlese),  male  -  A  dorsum,  B  venter,  C  tectum,  D  chelicera,  E  palp 
trochanter,  femur  and  genu,  F  venter  of  gnathosoma,  G  leg  II. 


The  tritosternum  has  a  narrow  base  and  pilose  laciniae  (Fig.  9B).  The  presternal  shields  are 
fused  into  a  single  narrow  bar.  The  ventral  shields  are  strongly  reticulated  and  all  setae  are  simple. 
The  sternal  shield  is  finely  granular  and  is  divided  longitudinally  at  the  centre.  Additionally,  a 
conspicuous  procurved  line  passes  through  sternal  pores  II  and  spans  the  entire  shield.  The 
genital  shield  measures  135-160  urn  long  x  210-260  um  wide.  Its  apex  is  formed  almost  as  a  right 


154 


K.  H.  HYATT 


Fig.  9    Holoparasitus  inornatus  (Berlese),  female -A  dorsum,  B  venter,  C  endogynium,  D  tectum, 
E  chelicera,  F  palp  trochanter,  femur  and  genu,  G  venter  of  gnathosoma. 

angle  and  it  does  not  bear  an  elongate  tip.  The  endogynium  appears  to  comprise  simply  an  oval 
punctate  area  with  a  central  detail  (Fig.  9C).  The  opisthogastric  region  bears  8-9  pairs  of  simple 
setae.  The  longest  of  the  ventral  setae  are  the  sternals  which  measure  c.  65  um  and  the  shortest  are 
around  the  posterior  margin  and  measure  c.  12  um.  The  stigma  is  situated  opposite  the  posterior 
margin  of  coxa  III  and  the  finely  granular  peritreme  extends  anteriorly  to  coxa  I. 

The  tectum  (Fig.  9D)  has  the  centre  portion  finely  tapered  and  it  is  flanked  on  each  side  by  a 
single  prong.  The  chelicera  is  shown  in  figure  9E.  The  movable  digit  measures  95  um  in  the  figured 
specimen  and  bears  three  teeth,  one  large  and  two  smaller  but  of  equal  size.  The  fixed  digit  bears 


MITE  GENUS  HOLOPARASITUS  1 5  5 

four  teeth,  the  two  distals  being  small,  the  two  proximals  larger.  The  palp  trochanter,  femur  and 
genu  are  shown  in  figure  9F.  The  anterolateral  seta  on  the  femur  is  pilose  distally  whilst  the  two  on 
the  genu  are  spatulate.  The  venter  of  the  gnathosoma  is  shown  in  figure  9G.  The  corniculi  are 
strongly  formed  and  the  four  pairs  of  gnathosomal  setae  are  simple.  Ten  rows  of  hypognathal 
denticles  are  visible.  The  leg  setae  are  slender,  some  on  tarsi  II-IV  are  pilose  on  one  margin.  The 
ambulacra  are  normal. 

MATERIAL  EXAMINED.  74  samples  -  4  DNN,  85  <$<$,  225  ??. 

ENGLAND:  Cornwall,  Devon,  Dorset,  Hampshire,  Sussex,  Kent,  Hertfordshire,  Gloucestershire, 
Suffolk,  Cambridgeshire  (including  Huntingdonshire),  Norfolk,  Herefordshire,  Warwick- 
shire, Nottinghamshire,  Lincolnshire,  Cumbria  (Westmorland),  N.  Yorkshire,  Durham, 
Northumberland. 

SCOTLAND:  Strathclyde  (Argyllshire,  Mull,  Ulva),  Tayside  (Perthshire),  Highland  (Inverness- 
shire,  Ross  and  Cromarty),  Sutherland,  Shetland  (Fair  Isle). 

WALES:  West  Glamorgan,  Dyfed  (Cardiganshire),  Gwynedd  (Caernarvonshire,  Anglesey), 
Clwyd  (Denbighshire). 
IRELAND:  Leitrim,  Mayo,  Sligo,  Clare,  Kerry. 

This  species  is  recorded  mainly  from  mosses,  litter  and  soil  in  damp  habitats  and  is  one  of  the  two 
most  abundant  representatives  of  the  genus  in  the  British  Isles. 

DISTRIBUTION.  The  only  previous  records  from  the  British  Isles  that  I  have  been  able  to  trace  and 
authenticate  are  of  specimens  recorded  by  Halbert  (191 5)  as  Gamasus  ( Ologamasus )  calcaratus  (in 
part)  from  Co.  Mayo  and  by  Davis  (1970)  as  Holoparasitus  ?inornatus  from  Huntingdonshire. 
Davis'  (loc.  cit.)  single  female  of  ?pollicipatus  is  Holoparasitus  stramenti  Karg. 

It  is  recorded  from  France  (Berlese,  1916),  Germany  (Berlese,  1 906,  Karg,  1971)  and  Switzerland 
(Schweizer,  1961). 

X  .  1^ 

Holoparasitus  lawrencei  sp.  nov. 
(FigslOA-G,  11A-G) 

MALE.  The  holodorsal  and  opisthogastric  shields,  which  are  fused  ventrally  posterior  to  coxae  IV, 
are  strongly  sclerotised  and  completely  reticulated  (Figs  10A,  B).  The  dorsum  (Fig.  10A)  measures 
780-840  urn  long  x  550-610  um  wide  and  bears  around  50  pairs  of  simple  setae  that  range  in  length 
from  63  um  (/7)  to  18um  in  the  opisthonotal  region.  The  figured  specimen  measures  820  um 
long  x  610  um  wide  and  bears  apparently  46  setae  on  the  left  side  of  the  dorsum  and  51  on  the  right 
side.  As  can  be  seen  from  the  figure,  the  setae  are  not  entirely  arranged  symmetrically. 

The  tritosternum  comprises  two  slender  pilose  laciniae  that  arise  from  below  the  anterior  margin 
of  the  genital  lamina  (Fig.  10B).  The  sternogenital  region  is  strongly  reticulated  and  bears  a 
strongly  procurved  line  between  coxae  II  and  a  similar,  but  less  conspicuous,  line  between  coxae  III 
and  IV.  The  sternogenital  and  median  opisthogastric  setae  are  up  to  65  um  in  length,  whilst  the 
three  simple  anal  setae  and  the  posterior  ventral  setae  are  about  20  um  long.  The  stigma  is  situated 
opposite  the  posterior  margin  of  coxa  III  and  the  strongly  granular  peritreme  extends  to  coxa  I. 

The  tectum  (Fig.  IOC)  is  trispinate  and  the  centre  prong  is  long  and  sinuous  and  may  be  broken 
off.  The  chelicera  is  shown  in  figure  10D.  The  movable  digit  is  92  um  long  in  the  figured  specimen 
and  bears  no  distinct  teeth.  The  fixed  digit  bears  apparently  only  one  rudimentary  tooth  adjacent  to 
the  pilus  dentilis.  The  palp  trochanter,  femur  and  genu  are  shown  in  figure  10E.  The  anterolateral 
setae  of  the  femur  and  genu  are  spatulate,  that  of  the  genu  is  pilose  on  one  margin.  The  venter  of  the 
gnathosoma  is  shown  in  figure  10F.  The  corniculi  are  strong  and  stalked,  the  palpcoxal  setae  are 
pilose  whilst  the  remaining  three  pairs  are  simple,  and  the  hypognathal  groove  bears  about  eleven 
rows  of  denticles.  Leg  II  is  shown  in  detail  in  figure  10G.  The  femoral  apophysis  is  short  and  does 
not  extend  beyond  the  blunt  axillary  process.  The  ventral  processes  on  the  genu  and  tibia  are 
shallow  and  directed  anteriorly.  All  leg  setae  are  slender,  a  number  on  the  tarsi  and  tibiae  of  legs 
II-IV  are  stronger  and  pilose  on  one  margin.  The  ambulacra  are  normal. 

FEMALE.  The  holodorsal  and  opisthonotal  shields,  which  are  strongly  scelerotised  and  completely 
reticulated,  are  fused  ventrally  anterior  to  the  anus  (Figs  1 1  A,  B).  The  dorsum  (Fig.  1 1  A)  measures 


156 


K.  H.  HYATT 


Fig.  10    Holoparasitus  lawrencei  sp.  nov.,  male- A  dorsum,  B  venter,  C  tectum,  D  chelicera,  E  palp 
trochanter,  femur  and  genu,  F  venter  of  gnathosoma,  G  leg  II. 

850-920  urn  long  x  650-710  um  wide  and  bears  up  to  49  or  more  pairs  of  simple  setae,  about  20 
pairs  in  the  podonotal  region  and  from  29-31  pairs  in  the  opisthonotal  region.  The  figured 
specimen -the  holotype  -  measures  870  urn  long  x  660  urn  wide  and  bears  35  setae  on  the 
podonotum,  1 7  on  the  left  and  1 8  on  the  right,  and  60  setae  on  the  opisthonotum,  3 1  on  the  left  and 
29  on  the  right.  The  vertical  setae,y7,  and  setaey'2  are  the  longest,  measuring  50  urn  or  more,  whilst 


MITE  GENUS  HOLOPARASITUS 


157 


Fig.  11     Holoparasitus  lawrencei  sp.  nov.,  female -A  dorsum,  B  venter,  C  endogynium,  D  tectum, 
E  chelicera,  F  palp  trochanter,  femur  and  genu,  G  venter  of  gnathosoma. 

the  shortest  setae,  the  majority  of  those  in  the  opisthonotal  region,  measure  as  little  as  12  \am  and 
are  extremely  fine. 

The  tritosternum  (Fig.  1  IB)  has  a  narrow  base  and  pilose  laciniae.  The  presternal  shields  are 
coarsely  granular  and  may  be  entire  or  divided  medially.  The  ventral  shields  are  reticulated  and  all 
setae  are  simple.  The  sternal  shield  bears  a  pair  of  lines  originating  from  the  angle  between  coxae  II 
and  III  and  passing  through  sternal  pores  II  and  almost  meeting  at  the  centre  of  the  shield.  The 
genital  shield  measures  170-180  urn  long  x  265-280  um  wide.  In  the  figured  specimen-  the  holo- 
type  -  it  measures  170  um  x  280  um.  Its  anterior  margin  is  formed  into  slightly  more  than  a  right 


1 58  K.  H.  HYATT 

angle  and  does  not  have  an  extended  tip.  The  endogynium  is  shown  in  figure  1 1C.  It  appears  to 
distort  easily.  The  opisthogastric  region  bears  8-10  pairs  of  setae.  The  longest  of  the  ventral  setae 
are  probably  sternal  setae  II,  being  up  to  75  urn,  whilst  the  shortest,  in  the  opisthogastric  region, 
measure  only  c.  12  um.  The  three  simple  anal  setae  are  also  short.  In  the  type  the  post-anal  seta  is 
bifid.  The  stigma  is  situated  opposite  the  posterior  margin  of  coxa  III  and  the  peritreme  extends  to 
the  level  of  coxa  I. 

The  tectum  (Fig.  1  ID)  is  very  similar  to  that  of  the  male.  The  chelicera  is  as  in  figure  1  IE.  The 
movable  digit  measures  125  um  in  the  figured  specimen  and  bears  three  blunt  teeth,  whilst  the  fixed 
digit  bears  five  blunt  teeth.  The  palp  trochanter,  femur  and  genu  are  shown  in  figure  11F.  The 
anterolateral  seta  of  the  femur  is  broad  with  one  edge  pectinate,  whilst  those  of  the  genu  are 
spatulate.  The  venter  of  the  gnathosoma  is  shown  in  figure  1 1G.  The  setae  are  simple,  and  the 
hypognathal  groove  bears  ten  rows  of  denticles.  The  majority  of  the  leg  setae  are  fine  and  simple, 
but  some  of  the  distal  setae  on  tarsi  II-IV  are  pilose  on  one  margin.  The  ambulacra  are  normal. 

MATERIAL  EXAMINED.  16  samples  -  20  <3<$,  16  $?. 

ENGLAND:  Cornwall  -  Hayle,  the  holotype  $  (1984.12.4.  1)  collected  by  Mr  P.  N.  Lawrence  from 
dry,  light,  leaf-litter,  24.5.1975:  Lelant,  St  Ives,  1  3  in  a  carrion  trap  on  salt  marsh  and  1  $  in  a 
garden  trap,  1943  (Dr  F.  A.  Turk)  (these  specimens  not  included  in  the  type  series);  Isles  of 
Scilly-St  Agnes,  1  $  paratype  (1984.12.4.  9)  from  litter  under  Pittosporum,  5.11.1959  (K.  H. 
Hyatt);  Somerset  -  Bath,  Kennet  and  Avon  Canal,  \<S  paratype  (1984.12.4.  16)  from  moss, 
humus,  etc.,  10.3.1962  (P.  N.  Lawrence);  Hampshire  -  Milford-on-Sea,  1  9  paratype  (1984.12.4: 
30)  with  no  data  (A.  S.  Hirst):  Isle  of  Wight,  1  £  paratype  (1984.12.4.  3)  with  no  habitat  data, 
April  1948  (T.  A.  Lloyd);  Oxfordshire  -  Oxford,  1  cJ,  1  $  paratypes  (1984.12.4. 17-18)  from  the 
nest  of  blackbird  Turdus  merula,  August  1979  (Miss  A.  Warburton);  Norfolk  -  Blackborough,  1 
cJ  paratype  (1984.12.4.  2)  with  no  habitat  data,  25.2.1969  (Miss  A.  Reeve);  Suffolk  -  Westleton 
Heath,  3  33  paratypes  (1984. 12.4. 13-1 5)  from  algae  on  rotten  wood,  7.3. 1964  (P.  N.  and  Mrs  K. 
Lawrence);  Lincolnshire  -  no  locality,  1  ?  paratype  (1925.6.24.  584)  with  no  habitat  data,  1900 
(C.  F.  George);  Cumbria  (Lancashire)  -  Grange-over-Sands,  3  33,  2  $?  paratypes  (1984.12.4. 
4-8)  from  tree-holes,  27.1.1954  (D.  Macfarlane);  Cumbria  (Cumberland)  -  Newton  Arlosh, 
Carlisle,  1  3,  2  $$  paratypes  (1973.28)  with  no  data  (J.  E.  Hull). 

SCOTLAND:  Tayside  (Perthshire)  -  Glen  Farg,  2  33  paratypes  ( 1 984. 12.4.1 9-20)  from  mosses  on 
deciduous  trees,  24.9.1982  (K.  H.  Hyatt);  Dumfries  and  Galloway  (Wigtownshire)  -  Moss  of 
Cree,  5  33,  4  ??  paratypes  (1984.12.4.  21-29)  from  moss  in  birch  tree-holes,  16.9.1982  (K.  H. 
Hyatt). 

WALES:  Dyfed  (Cardiganshire)  -  Dol-y-Bont,  1  ?  paratype  (1984.12.4.  12)  from  damp  moss, 
1 5.8. 1 957  (Dr  G.O.Evans). 

IRELAND:  Clare-  Lough  Inchiquin,  1  (J,  1  $  paratypes  (1984.12.4. 10-1 1)  from  litter  near  a  weir, 
June/July  1971  (P.  N.  Lawrence). 

This  species  is  named  after  Mr  P.  N.  Lawrence  whose  diligent  collecting  of  soil  arthropods  has 
done  much  to  increase  our  knowledge  of  the  British  and  Irish  faunas. 

Holoparasitus  maritimus  sp.  nov. 
(Figs  12A-F,  13A-H,  14A-G) 

Holoparasitus  calcaratus:  Browning,  1956:  386,  Non  Koch,  1839. 

DEUTONYMPH.  The  dorsal  shields  are  pale  yellowish  brown  in  colour,  lightly  sclerotised  and 
reticulated  (Fig.  12 A).  The  podonotal  shield  measures  260-280  um  long  x  up  to  480  um  wide, 
depending  on  the  degree  of  lateral  displacement  of  the  posterior  region  as  shown  in  the  figure.  The 
figured  specimen  bears  essentially  18  pairs  of  setae  although  the  lefty'2  is  missing.  Setae  jl,  the 
verticals,  measure  c.  35  um  whilst  the  remainder  reduce  in  length  to  c.  18  um  on  the  margins.  The 
opisthonotal  shield  measures  1 90-240  um  long  x  260-290  um  wide  and  bears  13  pairs  of  setae, 
although  in  the  figured  specimen  the  right  Z3  is  missing.  The  setae  range  in  length  from  1 8 — 22  um. 
The  surrounding  membrane  bears  dorsally  on  each  side  from  30-40  fine  setae  not  exceeding  1 8  um 
in  length. 


MITE  GENUS  HOLOPARASITUS 


159 


Fig.  12    Holoparasitus  maritimus  sp.  nov.,  deutonymph  -  A  dorsum,  B  venter,  C  tectum,  D  chelicera, 
E  palp  trochanter,  femur  and  genu,  F  venter  of  gnathosoma. 

The  tritosternum  has  a  narrow  base  and  simple  laciniae.  The  sternal  shield  (Fig.  12B)  measures 
160-170  |im  long  and  is  lightly  sclerotised  and  entirely  reticulated.  The  setae  are  simple.  Presternal 
shields  absent.  The  oval  anal  shield  bears  the  usual  three  setae  and  is  reticulated.  The  stigma  is 
situated  opposite  the  anterior  margin  of  coxa  IV  and  the  granular  peritreme  and  irregularly 
outlined  peritrematal  shield  extend  to  coxa  I.  The  opisthogastric  setae  number  upwards  of  16  pairs 
depending  on  the  actual  position  of  those  towards  the  posterior  margin. 


160  K.  H.HYATT 

The  tectum  (Fig.  12C)  is  essentially  triangular  and  bears  strong  lateral  teeth,  but  is  irregularly 
outlined.  The  chelicera  is  shown  in  figure  12D.  The  movable  digit  measures  78  um  in  the  figured 
specimen  and  bears  three  widely  spaced  teeth.  The  fixed  digit  bears  five  or  six  smaller  teeth.  The 
palp  trochanter,  femur  and  genu  are  shown  in  figure  12E.  The  anterolateral  setae  of  the  femur  and 
genu  are  spatulate.  The  corniculi  and  the  venter  of  the  gnathosoma  are  shown  in  figure  12F.  The 
anterior  and  the  internal  posterior  hypostomatic  setae  are  simple  whilst  the  external  hypostomatic 
and  the  palpcoxal  setae  are  lightly  pilose  on  one  margin.  About  seven  rows  of  hypognathal 
denticles  are  present.  All  the  leg  setae  are  slender  and  the  majority  are  simple,  but  a  few  on  tarsus  II 
are  finely  pilose  on  one  margin.  The  ambulacra  are  normal,  with  rounded  pulvilli  and  two  claws. 

MALE.  The  holodorsal  and  opisthogastric  shields  are  fused  ventrally  posterior  to  coxae  IV.  They 
are  heavily  sclerotised  and  entirely  reticulated  (Figs  13 A,  B).  The  dorsum  (Fig.  13 A)  measures 
680-750  jim  long  x  425-500  um  wide  and  bears  about  60  pairs  of  simple  setae  that  range  in  length 
from  c.  55  um  (setaey'7)  to  1 2  um  in  the  opisthonotal  region.  The  figured  specimen  measures  730  um 
long  x  450  um  wide  and  bears  apparently  60  pairs  of  setae  on  the  left  side  and  58  pairs  on  the  right 
side.  The  podonotal  region  bears  19  pairs  of  setae  arranged  symmetrically,  whilst  the  opisthonotal 
region  bears  41  setae  on  the  left  side  and  39  on  the  right  side. 

The  tritosternum  comprises  two  slender  pilose  laciniae  that  arise  from  below  the  anterior  margin 
of  the  genital  lamina  (Fig.  13B).  The  anterior  margin  of  the  sternogenital  shield  is  moderately 
recessed  medially.  The  ornamentation  of  the  sternogenital  region  is  without  a  characteristic 
pattern.  The  sternogenital  setae  are  about  60  um  in  length  whilst  the  opisthogastric  setae  are 
shorter.  The  three  anal  setae  are  simple  and  like  those  in  the  posterior  region  of  the  opisthogastric 
shield  measure  approximately  18  um.  The  stigma  is  situated  opposite  the  posterior  margin  of  coxa 
III  and  the  peritreme  extends  anteriorly  to  coxa  I. 

The  tectum  is  strongly  granular  and  produced  normally  into  a  triangular  process  (Fig.  13C). 
However,  in  the  figured  specimen  it  is  irregularly  formed  as  shown  in  figure  13D.  The  chelicera  is 
shown  in  figure  13E.  The  movable  digit  is  97  um  long  and  bears  one  large  tooth  and  four  to  five 
small  teeth.  The  fixed  digit  bears  up  to  seven  very  small  teeth.  The  palp  trochanter,  femur  and  genu 
are  shown  in  figure  1 3F.  The  anterolateral  seta  of  the  femur  is  broad  and  slightly  pectinate  on  one 
margin  and  the  two  anterolateral  setae  of  the  genu  are  spatulate.  The  corniculi  (Fig.  13G)  are 
stalked  and  are  deeply  cleft  on  their  inner  margins.  The  gnathosomal  setae  are  all  simple  and  there 
are  up  to  13  rows  of  hypognathal  denticles.  Leg  II  is  shown  in  figure  13H.  The  apophysis  on  femur 
II  is  short,  hemispherical,  and  does  not  extend  beyond  the  tip  of  the  swollen  axillary  process.  The 
ventral  processes  on  the  genu  and  tibia  are  smooth  and  elongate.  All  leg  setae  are  slender,  some  on 
tarsi  II-IV  are  finely  pilose  on  one  margin.  The  ambulacra  are  normal. 

FEMALE.  The  holodorsal  and  opisthonotal  shields  are  strongly  sclerotised  and  fused  ventrally 
anterior  to  the  anus  (Figs  14A,  B).  The  dorsum  (Fig.  14A)  is  reticulated  except  in  the  median 
podonotal  region  where  it  is  strongly  granular.  It  measures  770-810  um  long  x  520-560  um  wide 
and  bears  up  to  about  60  pairs  of  simple  setae,  about  20  pairs  in  the  podonotal  region  and  up  to  40 
pairs  in  the  opisthonotal  region.  The  figured  specimen -the  holotype  -  measures  800  um 
long  x  530  um  wide  and  bears  20  pairs  of  setae  in  the  podonotal  region,  whilst  in  the  opisthonotal 
region  there  are  39  setae  on  the  left  side  and  34  on  the  right.  The  vertical  setae,y'7,  are  the  longest, 
measuring  c.  55  um,  whilst  the  remainder  reduce  gradually  in  length  towards  the  posterior  of  the 
dorsum  where  the  shortest  are  c.  \  5  um. 

The  tritosternum  has  a  narrow  base  and  pilose  laciniae  (Fig.  14B).  The  presternal  shields  are 
fused  medially  and  are  strongly  denticulate  over  most  of  their  surface.  The  sternal  shield  bears  a 
characteristic  ornamentation  which  shows  up  clearly  in  alchohol  under  low  magnification.  There 
is  a  longitudinal  median  design  and  two  pairs  of  liniae  which  form  part  of  the  reticulation. 
The  anterior-most  lines  run  almost  diagonally  from  the  centre  of  the  shield  towards  the 
anterior  corners,  whilst  the  second  pair  runs  from  the  centre  through  sternal  pores  II.  The  genital 
shield  measures  160-165  um  long  x  240-260  um  wide.  In  the  figured  specimen  -  the  holotype -it 
measures  165  um  x  250  um.  Its  anterior  margin  is  almost  right-angled  medially  and  forms  a  short 
broad  tip.  The  endogynium  is  shown  in  figure  14C  and  appears  to  distort  easily.  The  opisthogastric 


MITE  GENUS  HOLOPARASITUS 


161 


Fig.  13    Holoparasitus  maritimus  sp.  nov.,  male  -  A  dorsum,  B  venter,  C,  D  tectum,  E  chelicera,  F  palp 
trochanter,  femur  and  genu,  G  venter  of  gnathosoma,  H  leg  II.    . 

region  bears  8-9  pairs  of  setae.  The  three  anal  setae  are  short  (c.  18  um)  and  similar  in  length  to  the 
posterior-most  ventral  setae.  The  stigma  is  situated  opposite  the  posterior  margin  of  coxa  III  and 
the  peritreme  extends  anteriorly  to  coxa  I. 

The  tectum  (Fig.  14D)  is  granular  and  produced  into  a  strong  median  spine  and  small  lateral 
spines.  The  chelicera  is  shown  in  figure  14E.  The  movable  digit  measures  c.  108  um  long  and  bears 


162 


K.  H.  HYATT 


r  •>        t 
/    f              V        1 

\ 

j  „ 

£  ''  .'T 

V  '  t  > 

f 

\T'   1 

\ 

A',  ', 

r 

rt^ 


Fig.  14     Holoparasitus  maritimus  sp.  nov.,  female -A  dorsum,  B  venter,  C  endogynium,  D  tectum, 
E  chelicera,  F  palp  trochanter,  femur  and  genu,  G  venter  of  gnathosoma. 


three  strong  teeth.  The  fixed  digit  bears  at  least  six  smaller  teeth.  The  palp  trochanter,  femur  and 
genu  are  shown  in  figure  14F.  The  anterolateral  setae  on  the  femur  and  genu  are  spatulate,  that  on 
the  femur  being  pilose  on  one  margin.  The  venter  of  the  gnathosoma  is  shown  in  figure  14G.  The 
external  posterior  hypostomatic  setae  are  simple,  the  remaining  three  pairs  are  lightly  pilose.  There 
are  up  to  1 1  rows  of  hypognathal  denticles.  The  majority  of  the  leg  setae  are  fine  and  simple,  but 
some  on  tarsi  II-IV  are  pilose  on  one  margin.  The  ambulacra  are  normal. 


MITE  GENUS  HOLOPARASITUS  1 63 

MATERIAL  EXAMINED.  8  samples  -  6  DNN,  16  <$<$,  20  $9- 

ENGLAND:  Cornwall  -  Kelsey  Head,  the  holotype  ?  (1984.12.4.  31)  and  4  DNN,  8  &J,  16  ?? 
paratypes  (1984.12.4.  32^3)  from  thrift  Armeria  maritima,  etc.,  on  cliffs,  20.5.1975  (P.  N. 
Lawrence  coll.):  Land's  End,  1  ?  paratype  (1984.12.4.  46)  from  mossy  turf,  26.5.1975  (P.  N. 
Lawrence):  Phillack,  Hayle,  2  ??  paratypes  (1984.12.4. 47^8)  from  dried  seaweed,  24.5. 1975  (P. 
N.  Lawrence):  Porthleven,  2  DNN,  3  ^  paratypes  (1984.12.4.  49-53)  from  mossy  cliff  turf, 
29.5.1975  (P.  N.  Lawrence);  Isles  of  Scilly-St  Agnes,  2  $$  paratypes  (1984.12.4.  44-45)  from 
thrift  and  grasses  on  rocks  by  seashore,  7.4.1957  (K.  H.  Hyatt). 

SCOTLAND:  Inner  Hebrides  -  lona,  1  $  paratype  (1984.12.4.  55)  from  sandy  beach  grass  with 
Fucus,  3.6.1970  (P.  N.  Lawrence). 

CHANNEL  ISLANDS:  Jersey  -  Elizabeth  Castle,  1  $  paratype  (1954.3.19.  49)  from  vegetation  on 
cliff-face,  30.8.1951  (Dr  G.  O.  Evans).  This  specimen  was  recorded  by  Browning  (1956)  as 
Holoparasitus  calcaratus  (C.  L.  Koch). 


Acknowledgements 

Of  the  many  collectors  who  have  donated  specimens  to  the  British  Museum  (Natural  History)  my  special 
thanks  go  to  Mr  P.  N.  Lawrence,  formerly  in  the  Department  of  Entomology. 

Colleagues  in  other  institutions  have  kindly  loaned  material:  Dr  L.  van  der  Hammen,  Rijksmuseum  van 
Natuurlijke  Historic,  Leiden  (Oudemans),  Dr  M.  V.  Hounsome,  Manchester  Museum  (Britten)  and  Dr  J.  P. 
O'Connor,  National  Museum  of  Ireland,  Dublin  (Halbert).  Dr  F.  A.  Turk,  Camborne,  Cornwall,  kindly 
loaned  specimens  from  his  own  collection.  Miss  A.  S.  Baker  compared  my  drawings  with  material  in  the 
Berlese  collection  in  Florence  and  Mr  K.  P.  Martyn  prepared  the  distribution  map. 


References 

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(Pahstwowe  Wydawnictwo  Naukowe),  690  pp.  [German.  Polish  and  Russian  summaries]. 

Oudemans,  A.  C.  1901.  Notes  on  Acari,  Ser.  3.  Tijdschr.  ned.  dierk.  Vereen.  7:  50-88. 

-  1929.  Kritisch  Historisch  Overzicht  der  Acarologie.  II,  1759-1804.  Tijdschr.  Ent.  72,  Suppl.:  1-1097. 

—  1936.  Kritisch  Historisch  Overzicht  der  Acarologie  (Critico-historical  survey  of  Acarology).  Leiden 
(Brill),  IIIA,  1805-1 850, 430  pp. 

Ryke,  P.  A.  J.  1962.  The  subfamily  Rhodacarinae  with  notes  on  a  new  subfamily  Ologamasinae  (Acarina: 

Rhodacaridae).  Ent.  Ber.,  Amst.  22: 155-162. 

Schrank,  F.  von  P.  von.  1781.  Enumeratio  Insectorum  Austriae  Indigenorum.  Augustae  Vindelicorum,  548  pp. 
Schweizer,  J.  1961.  Die  Landmilben  der  Schweiz  (Mittelland,  Jura  und  Alpen).  Parasitiformes  Reuter. 

Denkschr.  schweiz.  naturf.  Ges.  84: 1-207. 
Sellnick,  M.  1968.  Zwei  neue  Pergamasus-Arten  aus  Osterreich.  Ber.  naturw.-med.  Ver.  Innsbruck  56: 

463^72. 
Turk,  F.  A.  1953.  A  synonymic  catalogue  of  British  Acari.  Ann.  Mag.  nat.  Hist.  (12)  6: 1-26,  81-99. 

—  &  Turk,  S.  M.  1952.  Studies  on  Acari.  7th  series:  Records  and  descriptions  of  mites  new  to  the  British 
fauna,  together  with  short  notes  on  the  biology  of  sundry  species.  Ann.  Mag.  nat.  Hist.  (12)  5: 475-506. 

Vitzthum,  H.  v.  1923.  Acarologische  Beobachtungen.  7.  Reihe.  Arch.  Naturgesch.  89A,  2:  97-181. 
\\illmann,  C.  1940.  Neue  Milben  aus  Hohlen  der  Balkanhalbinsel,  gesammelt  von  Prof.  Dr  K.  Absolon, 
Brunn  (2.  Mitteilung).  Zoo/.  Anz.  130: 209-218. 

—  1941.  Die  Acari  de  Hohlen  der  Balkanhalbinsel.  (Nach  dem  Material  der  "Biospeologica  balcanica".) 
Studie  Oboru  vseol.  kras.  Nauky  B.  Biol.  Ser.  8  (14),  80  pp. 


Peter  R.  Colston  &  Kai  Curry-Lindahl 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  on  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Tring,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1986,  129pp.  Hardback.  0  565  00982  6  £17.50. 


Titles  to  be  published  in  Volume  52 

Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophora  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holopamsitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata).  By  P.  J. 

Chimonides 


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


Bulletin  of  the 

British  Museum  (Natural  History) 


The  phylogenetic  position  of  the 
Yugoslavian  cyprinid  fish  genus  Aulopyge 
Heckel,  1841,  with  an  appraisal  of  the  genus 
Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae 


Gordon  J.  Howes 


Zoology  series    Vol52    No  5    28  May  1987 


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ISBN  0  565  05029  X 

ISSN  0007 -1 498  Zoology  series 

Vol52  No.  5  pp  165-1 96 
British  Museum  (Natural  History) 
Cromwell  Road 
London  SW75BD  Issued  28  May  1987 


The  phylogenetic  position  of  the  Yugoslavian  cyprinid 
fish  genus  Aulopyge  Heckel,  1841,  with  an  appraisal  of 
the  genus  Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae 

Gordon  J.  Howes 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7  5BD 

Contents 

Introduction 165 

Methods  and  materials 166 

Nomenclature 167 

Abbreviations  used  in  the  figures 168 

Anatomical  characters  in  Aulopyge  and  their  phylogenetic 

significance ^r**~-Lr~"J~"    ...        169 

169 
174 
177 

Other  osteological  features   .        .        .    ,/.     /.''       .        '.•"'.        .       \       .VN    182 
Sexual  dimorphism  and  genitalia  .         .   £ ;    .     /  .         .         .         ,,,.,-,         •         •  V  j    186 

Discussion !  £  I   .      38  {1^2937        .|   #.    J    186 

Aulopyge  relationships  and  barbin  classification!    .        i "~     .     '"."j    .        .j      .  H    186 
Immediate  relationships  of  Aulopyge     .  V\  •     \-         |        •       _•_,,;.    •         •'        7/y     190 

Acknowledgements V\.      ^^*s^   <•>,».  _.. —,]    *«:**' t-      /J      192 

References  ' ;-^:^  1Q7 

IVC1C1  C11CC&  ......  ^^S.     *"*^     / "  f'  *^  •         »•»••  fi    •   *    •  Sy^  *  ^ 

Appendix  1 .  The  genus  Barbus  sensu  stricto       .  ^^^^ :-  "»AL  ^&_**\l>^&r        •  193 

Definition  and  included  species     ....     <i!^rr:=™irrTF' ^=^'^-- *^     .        .  193 
The  systematic  positions  of  Barbus  brachycephalus  Kessler,  1872  and  B.  mursa 

(Giildenstadt,  1773) 195 

The  systematic  positions  of  Barbus  andrewi  Barnard,  1937  and  B.  serra  Peters,  1864  195 

Appendix  2.  Characteristics  of  the  subfamilies  Cyprininae  and  Leuciscinae         .        .  196 

Introduction 

A ulopyge  is  a  monotypic  genus  represented  by  the  species  A.huegeliiHeckel,  1841  (Fig.  1),  endemic 
to  rivers  and  lakes  of  the  Yugoslavian  karst  regions  of  Dalmatia.  Regrettably,  there  appears  to  be 
no  published  information  on  the  ecology  of  Aulopyge.  Populational  data  are  lacking  and  the 
species  is  classified  as  rare  (Lelek,  1980:  122). 

Aulopyge  possesses  several  unique  characters  (detailed  below)  which  have  led  to  its  being  placed 
in  a  separate  taxonomic  category,  the  Aulopygini  (Bleeker,  1863;  Karaman,  1971).  In  Karaman's 
(1971)  view  Aulopyge  represents  a  relic  of  an  earlier  Eurasian  barbine  assemblage  having  a  close 
relationship  with  the  schizothoracine  cyprinids — a  group  now  confined  to  high-Asia.  Lelek  (1980: 
122)  simply  comments '. . .  it  is  difficult  to  compare  it  with  other  taxa'.  Arai  (1982:  146)  concluded 
from  his  study  of  karyotypes  that  Aulopyge,  which  is  polyploid  (2n  =  100),  possesses  a  'mosaic  of 
barbine  and  gobionine  characters'. 

The  recent  acquisition  by  the  British  Museum  (Natural  History)  of  well-preserved  specimens  of 
Aulopyge  huegelii  makes  possible,  for  the  first  time,  a  detailed  anatomical  study  of  the  species.  The 
information  gained  from  this  study  has  provided  a  basis  not  only  for  a  discussion  of  the  phylo- 
genetic status  of  Aulopyge  but  also  of  the  classification  and  relationships  of  the  genus  Barbus  and 
other  cyprinines. 


Bull.  Br.  A/MS.  not.  Hist.  (Zool.)  52(5):  165-196  Issued  28  May  1987 

165 


166 


G.  J.  HOWES 


Fig.  1     Aulopyge  huegelii,  female  (above)  127  mm  SL  and  male  (below)  106  mm  SL. 


In  an  earlier  paper  (Howes,  1981:  47-49)  the  classification  of  barbelled  and  non-barbelled 
Cyprinidae  was  discussed.  It  was  reasoned  that  one  lineage,  the  barbelled  cyprinids,  could  be 
defined  on  the  synapomorphic  presence  of  paired  maxillary  barbels  each  associated  with  a 
maxillary  foramen  (or  its  suggested  past  presence)  and  a  rostrally  extended  supraethmoid. 
Following  the  original  division  of  the  European  Cyprinidae  by  Bonaparte  (1846),  the  barbelled 
carps  were  recognised  as  the  subfamily  Cyprininae,  while  the  non-barbelled  taxa  were  ranked  as 
the  subfamily  Leuciscinae.  The  latter  group  was  defined  simply  by  absence  of  maxillary  barbels 
and  associated  nerve  foramen,  no  synapomorphy  having  been  discovered  that  would  define  it  as  a 
natural  group. 

It  can  be  assumed  from  the  outset  therefore  that  Aulopyge  belongs  to  the  Cyprininae,  since  it 
possesses  a  pair  of  maxillary  barbels  supplied  by  a  branch  of  the  VII  facial  nerve.  From  this 
standpoint  a  more  refined  hypothesis  of  the  relationship  between  Aulopyge  and  other  cyprinids 
may  be  attempted. 


Methods  and  materials 

The  osteology  of  Aulopyge  huegelii  was  studied  from  an  alcian-alizarin  stained  and  a  dry  skeletal 
preparation  (BMNH)  1903.12.4:  41-5),  and  from  X-Radiographs  of  specimens  106,  112  and 
127  mm  SL  (BMNH  1985.8.20:  1-3;  Busko  Lake,  south  Bosnia).  Genital  anatomy  was  studied  in 
dissected  specimens  of  this  latter  series. 

Comparative  osteology  of  a  wide  range  of  cyprinoids  was  studied  both  from  alizarin  stained  and 
dry  skeletal  preparations.  A  principal  data  source  has  been  the  large  collection  of  X-Radiographs 
of  cyprinoid  specimens  in  the  BMNH.  The  following  list  is  ofBarbus  specimens  used  in  this  study. 

A  =  alizarin  stained  preparation;  D  =  dissected  specimen;  S  =  dry  skeleton.  All  catalogue 
numbers  are  BMNH. 

Barbus  ablabes  1983.3.30:  7-14(D),  B.  albanicus  1970.9.24:  265-67(D),  B.  altianalis  eduardianus  Uncat.,  (S), 
B.  a.  radcliffi  1981.4.9:  42-66(D),  B.  altus  1898.4.2:  196-205(D),  B.  amphigramma  1980.7.18:  319-332(D),  B. 
andrewi  1900.11.6:  58(D),  1903.4.27:  94^95(S),  B.  antinorii  (type)  1908.10.14:  7,  B.  arabicus  1976.4.7: 


CYPRINID  FISH  GENUS  A  ULOPYGE  1 67 

201-272(D),  B.  argenteus  1907.6.29: 217(S),  B.(  =  Puntius)  arulus  1978.8.31 : 234-259(D),  B.  aspilus  1909.4.29: 
74(S),  B.  barbulus  1974.2.22:  1275-77(A),  B.  barbus  1864.4.11:  41-42(D),  1908.12.28:  123(S),  1985.10.16: 
62-ll(A),\9S5.\Q.l6:5l,B.(^Puntius)bimaculatusl9Sl.\.2l:209-2\l(D),B.binotatus\9S43.3:39-60(D), 
1970.9.3:  56-85(A),  B.  biscarensis  1970.3.1:  100-125(D,A),  B.  bocagei  1980.8.20:  1-6(D),  B.  brachycephalus 
1899.7.25:  25-27,  B.  burmanicus  1894.5.21:  46-55(D),  B.  bynni  1861.9.9:  65(S),  B.  callemis  1951.4.10: 
1-20(A,D),  1 869. 1 .29: 4(S),  B.  camptacanthus  Uncat.  (S),  B.  canis  1 974.2.22: 1292-94(D),  Uncat.,  (S),  B.  chola 
1935.10.18:  32^6(D),  B.  (  =  Puntius)  collingwoodi  1892.9.2:  52-56(A),  1982.4.21:  37-38(D),  B.  comiza 
(syntype)  1909.7.29:  1,  B.  conchonius  1978.8.31:  21-35(D),  B.  cummingi  1978.8.31:  186-222(A),  B. 
dorsolineatus  1965.3.15:  406-435(D).  B.  esocinus  1920.3.3:  80-82(D),  B.  eutaenia  1965.3.15:  93-122(D),  B. 
(  =  Puntius)filamentosus  1981.1.21: 242-260(D),  B.fritschi  1904.1 1.28:  59(S),  B.graellsii  1908.2.12:  21^9(D), 
B.  grahami  1907.5.4:  52-57(D),  B.  grypus  1920.3.3:  1-18(D),  B.  guirali  1902.11.12:  119(S),  B.  haasianus 
1976.3.18:  892-93(A),  B.  harterti  1902.7.28:  35(S),  B.  holotaenia  1984.7.5:  22-27(D),  B.  holubi  1937.10.4: 
12-14(D),  B.  hospes  1980.7.18:  434-438,  B.  hypsolepis  1971.11.26:  28^1(A),  B.  intermedius  intermedius 
1974.1.16:  128-162(A),  166-179(D),  1902.12.13:  338(S),  intermedius  australis  1893.12.2:  36(S),  B.johnstoni 
1975.8.3:  576-80,  B.  kersteni  1978.8.3:  632-84(D),  B.  ksibi  1934.10.25:  1-14(D),  B.  leonensis  1974.9.18: 
77-177(A),  B.  lineomaculatus  1974.1.16:  396-41 1(D),  B.  litamba  1974.1.11:  88-93(D),  B.  lithopides  1889.2.1: 
559-61(D),  B.  longiceps  1936.4.6:  5-11,  1949.9.16:  90-92,  1864.8.20:  21(S),  B.  luteus  1874.4.28:  23(S), 
1968.12.13:  201-212(D),  B.  macrolepis  1972.11.28:  9-1 2(D),  B.  macrops  1960.6.7:  111-160(D),  B.  mattozi 
1962.8.22:  2-6(D),  B.  meridionalis  1935.10.28:  14-17(D,S),  B.  minimus  1974.1.16:  276-292(D),  B.  mursa 
1872.5.30:  67-68,  B.  nasus  1902.1.4:  22(S),  B.  natalensis  1862.8.28:  8(S),  B.  neglectus  1980.7.10:  1-26(D),  B. 
neumayeri  1969.3.6:  31-50(D),  B.  (  =  Puntius)  orphoides  1974.10.10:  865-872(D),  B,  oxyrhynchus  1893.12.2: 
31(D),  1906.8.25:  17(S),  B.  paludinosus  1979.3.1:  1-53(D),  1908.1.20:  84(S),  cf.  paludinosus  Uncat.,  (A),  B. 
paytoni  1976.2.2:  29-31(D),  B.  (  =  Puntius)  pentazona  1954.11.23  7-82(A),  B.perince  1907.12.2:  1268-77(D), 
B.  plebejus  plebejus  1887.4.5:  15-16,  1982.2.24:  149-1 55(D),  plebejus  peloponnesius  1964.6.12:  20-26(D),  B. 
poechi  1962.7.5: 4-15(D),  B.progenys  1903.7.28: 155(S),  B.profundus  1970.5.14: 19-30(D),  B.(=Tor)putitora 
1884.2.1:  52(S),  B.  radiatus  1982.4.13:  4597^605(D),  B.  reinii  1903.10.29:  10(S),  B.  rocadasi  191 1.6.1:  26(S), 
B.  rothschildi  1902.7.28:  22-26(D),  B.  (  =  Puntius)  sarana  1933.8.19:  7-14(D),  B.  schejch  1931.12.21:  4(D),  B. 
sclateri  (syntypes)  1861.11.20:  9-13,  B.  serra  1937.10.4:  6-ll(D,S),  B.  setivemensis  1905.11.28:  59(S),  B. 
sharpeyi  1920.3.3:  71-75(D),  B.  (  =  Puntius)  sophore  1889.2.1:  777-782(D),  B.  subquincunciatus  1934.10.29: 
1(D),  B.  (  =  Tor)  tambroides  1982.4.21:  39(D),  B.  tenuis  1975.12.29:  250-265(D),  B.  thalamakanensis 
1976.3.18:  363-550(D),  B.  (  =  Puntius)  titteya  1974.6.11:  8-12(A),  B.  (=Tor)tor  1893.6.30:  31-38(D),  B. 
trimaculatus  1907.4.9:  98(S),  B.  tropidolepis  1936.6.15:  599-629(A),  B.  xanthopterus  1973.5.21:  198(D). 

Species  without  a  suffix  and  those  cited  in  the  text  but  not  listed  above  have  been  examined  by 
X-Radiography  only. 

Nomenclature 

Because  the  concept  of  cyprinid  subfamilies  and  other  higher  categories  used  here  differs  from  that 
of  previous  authors  (see  Discussion)  I  have  adopted  the  following  nomenclature. 

Subfamily  Cyprininae  (cyprinines):  a  monophyletic  assemblage  (see  text)  which  includes  the 
following  subgroups: 

*barbins:  a  possibly  monophyletic  group,  the  members  of  which  possess  a  foraminate  dilatator 
fossa  (see  text  and  Table  3  for  included  taxa).  This  group  embraces,  in  part,  the  Barbinae  and 
Barbini  of  previous  authors. 

*labeins:  a  monophyletic  group  sensu  Reid,  1982  and  1985;  includes  Labeinae,  Labeini, 
Labeoinae  and  Garrini  of  previous  authors. 

*squaliobarbins:  a  monophyletic  group  sensu  Howes,  1981. 

*schizothoracins:  a  supposed  monophyletic  group  (see  text);  the  Schizothoracinae  and 
Schizothoracini  of  previous  authors. 

*other  cyprinines:  an  unresolved  assemblage  of  taxa  not  included  in  any  of  the  above  categories 
and  lacking  a  foraminate  dilatator  fossa  (see  text  and  Table  3). 

Subfamily  Leuciscinae  (leuciscines):  a  possibly  non-monophyletic  assemblage  including 
Abraminae,  Cultrinae  etc.  of  previous  authors. 

After  this  paper  had  been  submitted  for  refereeing,  my  attention  was  drawn  to  a  publication  by 
Chen  et  al.  (1984).  These  authors  have  proposed  an  hypothesis  of  cyprinoid  relationships  whereby 
they  recognise  the  Cyprinidae  as  comprising  two  'series',  the  Barbini  and  Leuciscini.  They  further 
recognise  two  monophyletic  groups  (tribes)  within  the  Barbini,  viz.  Barbines  and  Tincanes,  of 


168 


G.  J.  HOWES 


which  the  Tincanae,  Cyprininae,  Barbinae  and  Labeoninae  (sic)  are  the  constituent  lineages.  My 
concept  of  Cyprininae  corresponds  to  Chen  et  al.  'Barbini',  whilst  my  subgroups  embrace  their 
subfamilies. 

The  appellations  'small'  and  'large'  are  often  given  to  African  Barbus  species.  As  used  here, 
'small'  refers  to  those  species  in  which  the  striae  on  the  exposed  part  of  the  scale  are  radiate,  the  fish 
usually  less  than  1 50  mm  SL  adult  size,  and  the  body  often  marked  with  spots  or  lateral  stripes; 
'large'  refers  to  those  species  in  which  the  scale  striae  are  more  or  less  parallel,  the  fish  more  than 
1 50  mm  SL  adult  size,  and  the  body  lacking  any  noticeable  markings. 


Abbreviations  used  in 


aa  anguloarticular 

abr  1  1  st  branched  anal  fin  ray 

afsl-3  anal  fin  rays  (unbranched) 

ah  anterohyal 

asn  anterior  supraneural 

at  anal  tube 

bb  basibranchials 

bh  basihyal 

bo  basioccipital 

bp  basioccipital  process 

bsr  branchiostegal  ray 

cb  ceratobranchials 

ccf  coracoid-cleithral  foramen 

cl  cleithrum 

cor  coracoid 

csi  cavum  sinus  imparis 

ct  connective  tissue 

de  dentary 

df  dilatator  fossa 

dfo  dilatator  foramen 

dfs  dorsal  fin  rays 

dh  dorsohyal 

eb  epibranchials 

ect  ectopterygoid 

enf  ectopterygoid  facet 

ent  entopterygoid 

epf  entopterygoid-palatine  facet 

ep  epural 

epo  epioccipital 

fc  frontal  canal 

fm  foramen  magnum 

fr  frontal 

frl  frontal  lamina 

hb  hypobranchial 

hmf  hymandibular  fossa 

hs  haemal  spine 

hyo  hyomandibula 

hyp  hypurals 

hys  hypurapophysis 

ic  intercalar 

ih  interhyal 

int  intestine 

io  infraorbitals 

iop  interoperculum 

ip  infrapharyngobranchials 

lac  lachrymal 

let  lachrymal  canal  tube 

le  lateral  ethmoid 


lef 

lien 

loc 

me 

met 

mp 

nc 

nca 

ns4 

nspu2 

op 

OS 

ov 

pa 

pc 

pel 

pe 

ph 

phy 

po 

poc 

pro 

ps 

pte 

pts 

ptt 

qf 

ra 

rp 

sb 

sec 

scp 

se 

so 

sop 

sor 

sp 

spr 

srp 

sy 

vh 

vo 

I 
II 

V 

VII 
IX 
X 


the  figures 

lateral  ethmoid  facet 

lateral  ethmoid-entopterygoid  ligament 

lateral  occipital  fenestra 

mesethmoid 

metapterygoid 

masticatory  plate  of  basioccipital 

neural  complex 

neural  canal 

neural  spine  of  4th  centrum 

neural  spines  of  2nd  preural  centrum 

operculum 

orbitosphenoid 

oviduct 

parietal 

parietal  canal 

postcleithrum 

preethmoid 

posterohyal 

parhypural 

preoperculum 

preopercular  canal  (bone  enclosed) 

prootic 

parasphenoid 

pterotic 

pterosphenoid 

posttemporal 

quadrate  facet 

retroarticular 

proximal  radials 

splenial  bone 

subcutaneous  canal 

scapula 

supraethmoid 

supraoccipital 

suboperculum 

supraorbital 

sphenotic 

sphenotic  process 

supraethmoid  rostral  process 

symplectic 

ventrohyal 

vomer 

olfactory  nerve  foramen 

optic  fenestra 

trigeminal  nerve  foramen 

facial  nerve  foramen 

glossopharyngeal  nerve  foramen 

vagus  nerve  foramen 


CYPRINID  FISH  GENUS  AULOPYGE 

-srp 
se 


169 


sor 


exo 


bo 


Fig.  2    Aulopyge  huegelii,  neurocranium  in  dorsal  (left)  and  ventral  (right)  views.  Scale  bar  in  mm. 


Anatomical  characters  in  Aulopyge  and  their  phylogenetic  significance 

The  cranium  of  Aulopyge  is  shown  in  Figs  2  and  3.  In  general  appearance  it  is  depressed  and 
elongate.  The  ethmoid  region  is  narrow  and  shallow,  the  supraethmoid  bearing  a  sloped,  valley- 
like  depression  and  anteriorly  having  slight  lateral  expansions  and  a  short  rostral  extension  which 
is  medially  indented  (srp,  Fig.  2).  The  kinethmoid  (Fig.  4d)  is  of  the  rod-shaped  type  considered  by 
Howes  (1978;  1981)  as  plesiomorphic  for  cyprinoids.  Each  lateral  ethmoid  is  extended  medially 
along  the  parasphenoid  and  contacts  its  partner,  their  being  no  anterior  myodome.  Laterally,  each 
bone  extends  a  narrow,  posteriorly  pointing  wing  which  ventrally  bears  a  well-developed  round 
facet  against  which  the  entopterygoid  facet  articulates  (lef,  Fig.  2).  This  is  an  unusual  feature  and  is 
discussed  further  below. 

The  frontals  are  narrowed  anteriorly  and  nasal  bones  are  absent;  the  supraorbital  bones  are 
small  but  not  excessively  reduced.  Otherwise,  the  cranium  of  Aulopyge  exhibits  no  features  which 
may  be  regarded  as  anything  but  plesiomorphic  among  cyprinoids,  viz.:  the  prootic  is  elongate 
with  a  long  lateral  commissure,  the  subtemporal  fossa  is  round  and  deep,  there  is  no  posttemporal 
fossa,  and  the  basioccipital  has  a  short  posterior  process  and  small,  round  masticatory  plate 
(Figs  2  &  3). 

Likewise  the  jaws  and  elements  of  the  suspensorium  (Fig.  4),  other  than  the  entopterygoid 
(discussed  below),  show  no  departure  from  the  'generalised'  cyprinoid  morphology  (see  Howes, 
1978, 1981, 1984). 

The  lateral  ethmoid  and  its  articulation  with  the  entopterygoid 

The  presence  in  Aulopyge  of  a  facet,  ventrally  on  the  lateral  ethmoid,  apposing  an  entopterygoid 
facet  is  a  feature  which  has  a  restricted  distribution  amongst  the  Cyprinidae.  Ramaswami  (1955) 
drew  attention  to  a  mesial  entopterygoid  facet  articulating  with  the  lateral  ethmoid  in  Labeo 


170 


G.  J.  HOWES 


so  Pa 


fc 


epo 


se 


exo 


1C 


loc 


fm 


csi 


Fig.  3    Aulopyge  huegelii,  neurocranium  in  lateral  (above)  and  posterior  (below)  views.  Scale  bar  in  mm. 


macrostoma  and  Cyprinus  carpio.  Howes  ( 1 976: 46)  noted  that  such  a  facet  was  variously  developed 
in  cyprinids,  supposing  it  best  developed  in  those  species  with  a  long  ethmoid  region  and  least  in 
those  with  a  short  ethmoid.  However,  further  investigation  has  not  endorsed  this  claim  and  it 
appears  that  the  presence  of  an  entopterygoid  facet  is  not  positively  correlated  with  the  length 
of  the  ethmoid.  Its  presence  seems  to  require  a  purely  phylogenetic  rather  than  a  functional 
explanation.  Thus,  entopterygoid-lateral  ethmoid  facets  occur  only  in  taxa  included  in  the 
Cyprininae,  being  absent,  but  for  a  single  exception  (Tinea;  see  below),  in  the  Leuciscinae,  (i.e.  all 
non-barbelled  cyprinids).  The  most  highly  developed  form  of  this  articulation  occurs  in  some 
species  of  Barbus,  Cyprinus  and  in  the  schizothoracin  genus  Diptychus  (Figs  5  &  6). 

In  Cyprinus,  the  ventral  surface  of  the  lateral  ethmoid  wing  is  broadly  triangular  with  the  ventral 
articular  facet  situated  antero-medially  (Fig.  5b);  the  facet  is  sloped  posteriorly  and  articulates 
against  a  round  facet  on  the  dorso-anterior  border  of  the  entopterygoid,  just  posterior  to  that 
bone's  articulation  with  the  palatine. 

In  Barbus  barbus,  B.  nasus,  B.  plebejus,  B.  bocagei,  B.  meridionalis  and  B.  barbulus  the  lateral 
ethmoid  facet  is  a  large  triangular  platform  (Fig.  5a).  In  some  'large'  Barbus  species,  e.g.  the  Asian, 
B.  grahami,  Barbus  ( =  Tor)  tor  and  the  North  African,  B.  setivemensis  the  articular,  boss-like 
facet  is  situated  at  the  midpoint  of  the  lateral  ethmoid  wing  (Figs  6c-e).  In  all  these  species  the 
entopterygoid  facet  is  moderately  developed.  In  yet  other  African  and  Asian  'large'  Barbus  species 
the  lateral  ethmoid  facet  lies  along  the  posterior  margin  of  the  wing  and  in  some  taxa,  e.g.  the 
majority  of  'large'  African  Barbus  and  Varicorhinus  species,  a  distinct  facet  is  barely  developed, 
there  being  only  a  bevelling  of  the  posterior  border  of  the  wing  (Figs  5d  &  6f).  In  these  taxa  an 
entopterygoid  articulatory  surface  is  feebly  developed  also  (Fig.  5d).  However,  in  the  majority  of 
African  and  Asian  Barbus  examined  lateral  ethmoid  and  entopterygoid  facets  are  lacking.  This 
appears  to  be  the  condition  in  all  the  so-called  'small'  African  Barbus  species. 

Amongst  schizothoracins  a  lateral  ethmoid  facet  is  variously  developed  (Figs  6k-m),  but  in 


CYPRINID  FISH  GENUS  AULOPYGE 

.hyo        met  enf 


171 


Fig.  4     Aulopyge  huegelii.  (Above)  suspensorium  in  lateral  view;  (below),  (a)  palatine;  (b)  maxilla;  (c) 

premaxilla;  (d)  kinethmoid.  Scale  bar  in  mm. 


enf 


Fig.  5  Articular  facets  on  the  ventral  surface  of  the  lateral  ethmoid  wing  and  antero-dorsal  surface  of 
the  entopterygoid  in:  (a)  Barbus  barbus;  (b)  Cyprlnus  carpio;  (c)  Tor putitora',  (d)  Barbus  oxyrhynchus; 
(e)  Tinea  tinea;  (f)  Labeo  coubie,  entopterygoid  facet  also  shown  in  lateral  view.  In  (a)  dashed  outline 
represents  attachment  area  of  lateral  ethmoid  ligament.  Anterior  to  the  left.  Scale  bar  =  5  mm. 


172 


G.  J.  HOWES 


n 


Fig.  6  Lateral  ethmoid  facets  of:  (a)  Diptychus  dybowski;  (b)  Barbus  nasus;  (c)  B.  grahami;  (d)  B. 
setivemensis;  (e)  B.  lithopides;  (f)  Varicorhinus  tanganicae;  (g)  Barbus  callensis;  (h)  B.  serra;  (i)  B. 
progeny  s;  (j)  B.  canis;  (k)  Schizothorax  grahami;  (1)  S.  taliensis;  (m)  S.  intermedius;  (n)  S.  esocinus  and  S. 
richardsoni.  Semi-diagrammatic;  all  drawn  to  same  scale;  anterior  to  the  left. 


none,  apart  from  Diptychus  (Fig.  6a)  is  there  a  condition  approaching  that  in  the  Eurasian  Barbus 
species  cited  above,  and  an  entopterygoid  facet  is  rarely  present. 

In  the  squaliobarbins  (Squaliobarbus ,  Ctenopharyngodon  and  Mylopharyngodori),  a  group  con- 
sidered as  primitive  cyprinines  (see  Howes,  1981,  and  Fig.  21),  the  lateral  ethmoid  articular  surface 
is  elongate,  with  a  bevelled  anterior  margin  against  which  abuts  the  posterior  edge  of  the  palatine. 
The  entopterygoid  articulates  only  with  the  posterior  rim  of  the  lateral  ethmoid  wing  as  in  some 
'large'  African  Barbus  described  above. 

In  labeins,  Labeo  (sensu  Reid,  1 985)  has  an  extensive  lateral  ethmoid  whose  ventral  surface  bears 
a  fossa  which  cups  an  entopterygoid  condyle  (Fig.  5f).  Garra,  on  the  other  hand,  has  a  narrow 
lateral  ethmoid  wing,  which  is  only  connected  ligamentously  with  the  entopterygoid. 

Lateral  ethmoid  and  entopterygoid  facets  are  also  lacking  in  Cyprinion,  Gibelion  and  Capoeta; 
whether  this  condition  represents  secondary  loss  or  a  plesiomorphic  state  is  uncertain  in  the 
absence  of  recognised  synapomorphies  indicating  the  relationships  of  these  taxa. 

That  there  is  a  phylogenetic  rather  than  a  functional  basis  for  the  various  types  of  articulatory 
surfaces  among  cyprinines  is  seemingly  supported  by  the  following  observations. 

In  those  taxa  where  there  is  a  well-developed  articulation  between  the  two  bones,  e.g.  Cyprinus 
and  some  Eurasian  Barbus  species,  the  anterior  portion  of  the  entopterygoid  is  almost  horizontal 
(Figs  8a  &  b),  and  it  is  also  horizontal  in  those  taxa  which  have  only  a  moderate  articulation 
between  these  bones,  e.g.  some  'large'  African  Barbus  and  Varicornhinus  species  (Fig.  8c).  In 
Aulopyge,  where  there  are  well-developed  lateral  ethmoid  and  entopterygoid  facets,  the 
entopterygoid  slopes  at  an  angle  similar  to  that  in  taxa  which  lack  such  close  articulation,  e.g. 
Schizothorax  esocinus  (Fig.  8d).  Thus,  whilst  the  slope  of  the  entopterygoid  is  correlated  with 
cranial  width  (being  horizontal  in  those  taxa  with  the  widest  crania)  there  is  apparently  no  corre- 
lation between  slope  (both  in  the  horizontal  and  vertical  planes)  and  the  presence  or  absence  of 


CYPRINID  FISH  GENUS  AULOPYGE 


173 


lien 


Fig.  7     Connection  between  the  lateral  ethmoid  and  entopterygoid  in,  (a)  Cyprininae;  Barbus  barbus, 
and  (b)  Leuciscinae;  Raiamas  loati.  Scale  bars  =  3  mm. 


articulatory  surfaces.  Even  if  one  accepts  this  as  evidence  for  the  apomorphic  status  of  articulatory 
facets,  there  is  nothing  to  suggest  which  type  of  facet  morphology  is  the  more  derived,  be  the 
extensive  well-developed  articulation  of  the  Eurasian  Barbus  and  Cyprinus  or  the  less  intimate 
connection  of  the  African  Barbus  and  Varicorhinus  species. 

It  was  noted  above  that  all  but  one  leuciscine  taxon  lack  an  articular  connection  between  the 
lateral  ethmoid  and  entopterygoid.  Instead,  the  two  bones  are  ligamentously  connected  and  often 
the  entopterygoid  extends  anterior  to  the  lateral  ethmoid  (Fig.  7b).  Tinea  is  the  exception  amongst 
leuciscines,  in  that  the  entopterygoid  bears  a  distinct  and  deep  fossa  which  articulates  with  a  lateral 
ethmoid  facet  (Fig.  5e). 

That  articulatory  lateral  ethmoid  and  entopterygoid  facets  occur  only  amongst  cyprinines 
further  supports  an  internal  division  of  the  Cyprinidae,  but  whether  this  represents  the  derived 
condition,  and  if  so,  whether  it  is  synapomorphic  for  those  taxa  in  which  the  articulation  occurs  is 
problematic  (see  remarks  above). 

The  types  of  ligamentous  connection  between  the  lateral  ethmoid  and  the  entopterygoid 
support  the  subfamilial  division  of  the  Cyprinidae  (see  p.  166  and  Appendix  2).  The  widespread 
ostariophysan  condition  is  for  there  to  be  a  strong  ligament  running  from  the  upper  medial  face 
of  the  lateral  ethmoid  wing  to  the  dorsolateral  surface  of  the  entopterygoid  (Fig.  7a;  see  also 
Vanderwalle,  1977,  Fig.  4  of  Barbus  barbus). 

In  all  members  of  the  subfamily  Cyprininae  investigated,  apart  from  Ctenopharyngodon,  there  is 
a  single,  slender  ligament  connecting  the  bones;  in  Ctenopharyngodon  a  broad  ligamentous  band 
connects  the  bones.  In  the  subfamily  Leuciscinae  a  ligament  of  the  type  found  in  the  Cyprininae  is 
absent  and  connection  between  the  lateral  ethmoid  and  entopterygoid  is  via  undifferentiated 
connective  tissue.  Vandewalle  (1977)  showed  in  Leuciscus  leuciscus  a  ligament  (labelled  Li  18) 
running  from  the  edge  of  the  lateral  ethmoid  to  the  entopterygoid.  I  find  no  such  discrete  ligament, 
but  instead  thickened  connective  tissue  running  to  the  lateral  edge  of  the  entopterygoid  (Fig.  7b). 

The  widespread  occurrence  and  constant  position  of  the  lateral  ethmoid-entopterygoid  ligament 
amongst  ostariophysans  suggests  it  is  plesiomorphic  and  thus  its  absence  in  the  Leuciscinae  is 
considered  a  derived  loss.  It  is  interesting  to  note  in  this  group,  as  compared  with  the  Cyprininae, 
what  appears  to  be  an  anterior  shift  of  the  entopterygoid  head,  and  its  somewhat  looser  connection 
with  the  palatine,  features  which  may  be  associated  with  the  absence  of  a  ligamentous  connection. 


174 


G.  J.  HOWES 

.enf 


Fig.  8  Anterior  views  of  the  right  suspensorium  in;  (a)  Cyprinus  carpio;  (b)  Barbus  barbus;  (c) 
Varicorhinus  tanganicae;  (d)  Schizothorax  esocinus;  (e)  Aulopyge  huegelii.  Scale  bar  for  a-d  =  5  mm, 
for  e  =  1  mm. 


Sensory  canals  and  their  associated  bones 

Aulopyge  possesses  the  pattern  of  supraorbital  canals  corresponding  to  Illick's  (1956)  group 
IVAA,  where  a  marked  gap  separates  the  supraorbital  and  infraorbital  canal  systems;  the  frontal 
and  parietal  canals  are  distant  and  the  parietal  canals  are  separated  from  one  another  by  a  midline 
gap.  The  dorso-cranial  canals  are  bony  tubes  lying  on  the  surface  of  their  respective  bones;  the 
frontal  canal  contains  9-10  pores.  The  infraorbital  series  is  reduced  to  bony  tubes,  the  first  and  last 
infraorbitals  being  fragmented  into  several  elements  (Fig.  9a). 

The  most  unusual  feature  of  the  Aulopyge  infraorbital  canal  is  its  disassociation  from  the 
lachrymal  (1st  infraorbital,  lac,  Fig.  9a).  In  an  alizarin  stained  specimen  of  52mm,  the  weakly 
ossified  canal  lies  somewhat  ventral  to  the  well-developed  elongate  'lachrymal'  bone.  This  situ- 
ation is  evident  in  all  the  specimens  of  Aulopyge  examined  (60-127  mm  SL)  with  the  exception  of  an 
84  mm  SL  female,  where  a  bony  canal  tube  is  attached  to  the  face  of  the  lachrymal  (Fig.  9b).  Those 
portions  of  the  canal  posterior  and  anterior  are  epidermal. 

The  mandibular-preopercular  canal  is  incomplete.  Only  a  single,  small  tube  lies  below  and 
separate  from  the  dentary  (Fig.  4).  There  is  a  short  groove  along  the  ventro-lateral  border  of  the 
dentary,  but  no  sign  of  a  canal  associated  with  the  anguloarticular.  The  canal  reappears  as  a  series 
of  weakly  ossified,  epidermal  tubes  along  the  posterior  part  of  the  preoperculum;  at  the  point  of 
curvature,  the  canal  runs  through  the  bone  (poc,  Fig.  4),  then  continues  in  three  or  four  epidermal 
tubes,  the  last  terminating  close  to  the  dorsal  tip  of  the  preoperculum. 

The  development  of  cyprinoid  sensory  canals  was  studied  by  Lekander  ( 1 949)  who  summarised 
the  results  and  theories  of  previous  authors.  Lekander  showed  that  the  sensory  canals  can,  from  the 
earliest  ontogenetic  stages  either  be  united  with  their  respective  bone,  later  fuse  with  it,  or  remain 
separate  from  it.  He  drew  particular  attention  to  the  'antorbital'  ( =  lachrymal  of  most  authors) 
noting  that  in  some  cypriniforms,  the  canal  remains  separate  from  its  lamellar  portion.  Such  is  the 
case  in  the  Cobitidae,  where  there  is  apparently  an  antorbital,  i.e.  a  bone  lying  antero-dorsal  to  the 
1st  infraorbital,  while  the  elongate  lachrymal  is  by-passed  ventro-laterally  by  the  subcutaneous 
sensory  canal  (Lekander,  1949;  Parshall,  1983). 

As  in  the  adult  Aulopyge,  the  developing  infraorbital  canals  in  some  cyprinids  often  appear 
irregularly  spaced  and  remain  unfused  to  one  another;  a  'splenial'  bone  may  be  present  (Lekander, 
1 949:  8 1 ),  and  the  preopercular  latero-sensory  canal  tubes  remain  separated  from  one  another  and 
from  the  preoperculum  (Lekander,  1949:  95;  102;  1 12). 


CYPRINID  FISH  GENUS  AULOPYGE 


175 


lac 


let 


Fig.  9  Infraorbital  bones  of  Aulopyge  huegelii;  (a)  complete  series  of  52  mm  SL  specimen;  (b)  the 
lachrymal  of  an  84  mm  SL  specimen  with  canal  attached  to  the  bone,  (c)  Barbus  barbulus,  showing 
disassociated  lachrymal  canal.  Scale  bars  in  mm. 


Lekander  (1 949:  1 1 3)  makes  the  point  that  sensory  canals  in  cyprinids  develop  later  than  in  most 
other  teleosts  he  examined.  Whether  this  is  so  or  not,  I  observe  a  temporal  difference  in  the 
development  of  the  infraorbital  canals  between  two  species  of  Barbus.  In  specimens  of  Barbus  cf. 
paludinosus  of  17  mm  SL  the  sensory  canal  of  the  1st  infraorbital  (lachrymal)  is  present  in  the  bone 
although  it  does  not  become  completely  enclosed  until  24  mm  SL.  However,  in  Barbus  barbus  of 
25  mm  SL,  the  canal  is  subcutaneous  and  well-separated  from  the  membranodermic  part  of  the 
lachrymal.  These  species  are  respectively,  tropical  and  temperate,  and  small  and  large  sized.  Thus, 
the  variation  in  canal  development  may  reflect  the  different  temperature  and  hormonally  con- 
trolled growth  rates.  In  an  adult  specimen  of  Barbus  barbulus,  the  posterior  part  of  the  canal  lies 
subcutaneously,  whereas  the  anterior  part  is  attached  to  the  lachrymal  (Fig.  9c). 


176 


Fig.  10  Lachrymal  (1st  infraorbital)  bones  of;  (a)  Barbus  barbus  of  66  mm  SL;  (b)  B.  barbus,  adult;  (c)  B. 
comiza;  (d)  B.  plebejus;  (e)  B.  capita  (also  in  B.  sclateri);  (f)  B.  bocagei  (also  in  B.  albanicus);  (g) 
B.  longiceps;  (h)  B.  grypus  (also  in  B.  canis,  B.  sharpeyi,  B.  reinii);  (i)  B.  intermedius  intermedius;  (j)  B. 
trimaculatus;  (k)  B.  altus;  (1)  B.  callipterus;  (m)  Labeo  coubie;  (n)  5.  mursa;  (o)  5.  semz;  (p)  5.  andrewi. 
Scale  bars  =  1  mm. 


If  Lekander  (1949)  is  correct  in  recognising  three  distinct  types  of  association  between  the 
laterosensory  and  membranodermic  parts  of  the  canal  bones  (at  least  amongst  cypriniforms),  then 
it  may  be  that  these  represent  arrest  at  successive  ontogenetic  stages.  In  this  case,  that  exhibited  by 
Aulopyge  and  some  cobitids  where  the  sensory  and  membranodermic  components  are  separate 
represented  the  earliest,  whilst  that  in  which  they  are  united,  as  in  Leuciscus,  would  represent  the 
most  advanced  ontogenetic  stage. 

The  lachrymal  in  Aulopyge  is  virtually  oblong  in  lateral  view  being  somewhat  tapered  anteriorly. 
In  most  cyprinids  the  lachrymal  is  a  deep,  triangular  or  pentagonal  bone,  as  in  Cyprinus,  Labeo  and 
the  majority  of  Barbus  species  (Figs  lOi-m).  In  some  Eurasian  Barbus  species,  however,  the 
lachrymal  has  the  same  oblong  shape  as  in  Aulopyge,  and  the  sensory  canal  also  runs  in  the  ventral 
part  of  the  bone.  In  this  latter  respect  the  Eurasian  species  also  differ  from  other  African  and  Asian 
Barbus  where  the  canal  runs  centrally  through  the  lachrymal  (Fig.  10k).  In  Barbus  barbus,  one  of 


C YPRINID  FISH  GENUS  A  ULOPYGE  1 77 

the  species  with  an  oblong  lachrymal  in  adults,  there  is  a  marked  ontogenetic  change  in  the  bone's 
shape.  In  a  specimen  of  66  mm  SL,  it  is  almost  square  with  a  short,  dorsally  curved  sensory  canal 
(Fig.  lOa).  In  adults,  the  bone  is  elongated,  with  a  greatly  lengthened  canal  (Fig.  lOb),  the  anterior 
part  of  the  canal  having  become  more  deeply  forked  and  an  additional  pore  developing  at  the 
posterior  elongation  of  the  canal. 

The  adult  lachrymal  morphology  of  Barbus  barbus  resembles  that  characteristic  of  certain  other 
Eurasian  species  (Figs  9c,  lOc,  d,  f  &  g).  A  variant  of  this  condition  is  found  in  the  Middle-eastern 
species  B.  canis,  B.  sharpeyi,  B.  grypus,  B,  reinii  and  the  Asian  Barbus  (=Tor)  tor,  where  the 
anterior  part  of  the  sensory  canal  runs  close  to  the  anterior  border  of  the  bone,  and  the  dorsal 
border  is  concave  (Fig.  lOh). 

It  is  difficult  to  evaluate  the  shape  of  the  lachrymal  as  a  phylogenetic  character.  Skelton  (1980) 
pointed  out  that  the  South  African  West  Cape  species  Barbus  andrewi  and  B.  serra  possess  a 
lachrymal  of  the  same  elongate  form  as  that  of  the  Eurasian  species.  However,  the  lachrymal  of 
these  two  species  differs  from  that  in  the  Eurasian  taxa  in  having  the  ventral  border  convex  rather 
than  straight  (Figs  lOo  &  p);  see  also  p.  195. 

My  own  comparisons  permit  the  following  generalisations: 

*in  all  Leuciscinae  the  lachrymal  has  a  square  or  even  rounded,  never  elongate  shape,  even  in 
those  species  with  a  relatively  long  ethmoid  region  (e.g.  Elopichthys  bambusa;  Fig.  12A  in  Howes, 
1978). 

*in  Barbus  there  is  some  degree  of 'intermediacy'  in  shape  between  such  forms  as  B.  trimaculatus 
and  B.  altus  (Figs  lOj  &  k)  and  the  B.  canis  type  (Fig.  lOh)  exemplified  by  B.  oxyrhynchus  and  B. 
intermedius  (Fig.  10i). 

*there  is  a  distinct  (?apomorphic)  type  characterising  a  group  of  Eurasian  Barbus  species;  see 
above. 

Vertebral  column,  dorsal  and  anal  fins 

The  general  morphology  of  the  Weberian  ossicles  and  centra  ofAulopyge  resembles  that  ofBarbus 
barbus.  In  both  taxa  the  neural  complex  is  low,  with  a  concave  anterior  border.  Its  posterior  border 
is  irregular  and  widely  separated  from  the  4th  neural  spine,  which  is  almost  half  the  height  of  the 
neural  complex  and  is  inclined  posteriorly. 

Neural  complex.  The  comparative  morphology  of  the  cyprinid  neural  complex  has  not  been 
subject  to  any  detailed  treatment  and  from  the  following  perfunctory  observations  appears 
worthy  of  closer  study.  The  so-called  'neural  complex'  in  cypriniforms  is  a  supraneural  having 
synchondral  contact  with  the  3rd  and  4th  neural  arches.  There  is  usually  a  long  gap  between  the 
supraoccipital  and  the  neural  complex  and  only  rarely  are  they  in  close  contact  (see  Reid,  1985). 
Within  the  Cyprinidae,  two  morphotypes  of  neural  complex  are  recognisable  (briefly  described  in 
Howes,  1981:  29-30;  see  also  Chen  et  al,  1984);  these  can  be  correlated  with  the  subfamily  division 
already  recognised  as  Cyprininae  and  Leuciscinae  (see  above  and  Appendix  2). 

In  Cyprininae,  the  neural  complex  is  most  often  tall,  axe-shaped  and  lamellate,  with  a  vertical  or 
forwardly  inclined  anterior  border  and  without  a  grooved  dorsal  surface.  The  4th  neural  spine  is 
rarely  as  high  as  the  neural  complex,  most  often  being  half  or  less  than  half  its  height  and  narrowly 
separated  from  it.  The  first  free  supraneural  never  articulates  directly  with  the  neural  complex. 

In  Leuciscinae,  the  neural  complex  is  most  often  low,  oblong  or  square,  vertically  or  backwardly 
inclined;  its  dorsal  surface  contains  a  groove,  and  in  some  taxa,  the  neural  complex  is  deeply 
forked  (Howes,  1981,  Fig.  22);  the  1st  free  supraneural  articulates  with  the  groove  (Howes,  1978: 
19;  Fig.  13).  The  4th  neural  spine  is  most  often  as  tall  as  the  neural  complex  and  may  be  widely 
separated  from  it. 

The  morphology  of  the  neural  complex  is  variable  within  the  Cyprininae,  but  from  the  data  at 
hand  it  is  possible  to  make  a  broad  and  tentative  classification.  Within  Barbus,  the  'small'  species 
examined  (B.  radiatus,  B.  paludinosus,  B.  perince,  B.  leonensis,  B.  hulstaerti)  and  some  Asian  taxa 
(including  B.  ( =  Puntius]  sophore}  possess  a  tall,  oblong  neural  complex,  either  vertical  or  sloping 
backward  and  narrowly  separated  from  the  4th  neural  spine  which  is  the  same  height  as  the  neural 
complex  (Figs  1  lf-1). 


178 


G.  J.  HOWES 

nc 


ns4 


Fig.  11  Neural  complex  (shaded)  and  position  of  4th  neural  spine  in  (a)  Aulopyge  huegelii;  (b)  Barbus 
barbus;  (c)  B.  plebejus;  (d)  B.  altianalis  radcliffi;  (e)  Cyprinus  carpio;  (f)  Barbus  paludinosus',  (g) 
B.  perince;  (h)  Puntius  sophore;  (i)  Barbus  marequensis  ('long-head  morph');  (j)  Varicorhinus 
steindachneri;  (k)  V.  ensifer;  (1)  Schizopygopsis  stoliczkae.  Drawings  made  from  radiographs,  all  to 
approximately  the  same  scale. 


CYPRINID  FISH  GENUS  A  ULOPYGE  1 79 

Within  the  'large'  Barbus  species,  as  in  other  Cyprininae,  the  neural  complex  is  tall  and  axe- 
shaped.  Its  relationship  to  the  4th  neural  spine  is  variable.  In  some  taxa  the  spine  is  short  and 
curved  forward,  e.g.  B.  intermedius ,  B.  arabicus,  B.  altianalis  (Fig.  lid),  B.  (=Tor)  putitora, 
Carassius  auratus,  or  long  and  curved  forward,  e.g.  Cyprinus  carpio  (Fig.  1  le),  Cydocheilichthys, 
or  short,  vertical  or  sloping  backward,  e.g.  majority  of 'large'  African  Barbus  (Fig.  Hi),  and  some 
Asian  Barbus.  In  some  Varicorhinus  species  the  spine  is  minute  and  barely  developed  as  is  also  the 
case  in  Cyprinion  species  (see  Howes,  1982).  In  all  these  taxa,  however,  the  4th  neural  spine  is 
closely  apposed  to  the  posterior  border  of  the  neural  complex  (Fig.  1  Ij). 

As  noted  above,  the  neural  complex  of  Aulopyge  huegelli  and  Barbus  bar  bus  exhibit  another 
morphotype  (Figs  1  la  &  b),  being  squat  to  oblong  with  a  concave  anterior  border  and  an  indented 
posterior  border  leaving  a  wide  gap  between  it  and  the  4th  neural  spine.  Other  taxa  with  this 
morphology  are  the  Eurasian,  Middle-eastern  and  Chinese  Barbus  species  plebejus,  nasus, 
meridionalis ,  barbulus,  schejch,  subquincunciatus  and  grahami.  An  exaggerated  variant  of  this 
condition  occurs  among  the  schizothoracin  genera  Schizocypris  (Fig.  1 11),  Diptychus,  Gymnocypris 
and  Schizothorax,  where  the  neural  complex  is  irregularly  shaped  and  widely  separated  from  a 
small  4th  neural  spine. 

From  this  limited  survey  it  cannot  be  said  which  of  these  is  a  derived  morphotype.  That 
characteristic  of  Aulopyge,  some  Eurasian  and  Middle-eastern  Barbus  and  schizothoracins  may 
simply  be  a  correlate  of  the  generally  elongate  and  depressed  bodies  of  those  taxa.  There  is  also  a 
degree  of  intraspecific  and  ontogenetic  variability.  For  example,  the  ontogenetic  sequence  of 
neural  complex  development  in  the  'large'  African  Barbus  intermedius  is  at  21  -5  mm  SL  (Fig.  12a) 
that  of  the  adult  morphology  (almost  identical  to  that  of  B.  altianalis,  shown  in  Fig.  1  Id)  in  which 
the  complex  is  narrowly  separated  from  the  4th  neural  spine.  At  25  mm  SL  the  neural  complex  is 
tilted  forward,  is  relatively  taller  and  has  a  large  gap  separating  it  from  the  neural  spine.  At  3 1  mm 
SL  the  neural  complex  is  upright  and  the  4th  neural  spine  is  tall  and  narrowly  separated  from  it. 

The  four  ontogenetic  stages  shown  in  Fig.  12  of  specimens  21-5, 23-5, 25-0  and  31  -0  mm  SL  seem 
to  reflect  four  of  the  similar  adult  morphotypes  described  above. 

To  summarise  the  conditions  of  the  neural  complex  among  cyprinines: 

*tall  and  oblong  with  long  4th  neural  spine — in  'small'  African  Barbus  and  (?all)  Asian  Puntius 

*tall  and  axe-shaped  with  4th  neural  spine  closely  apposed — in  'large'  African  and  Asian  Barbus 
and  most  other  cyprinines,  subgrouped  as: 

4th  neural  spine  short — some  African  Barbus  and  other  cyprinines 
4th  neural  spine  long — most  African  and  Asian  Barbus 
4th  neural  spine  minute — Varicorhinus  and  Cyprinion 

*low,  oblong  or  square  with  irregular  anterior  and  posterior  borders  and  with  4th  neural  spine 
widely  separated  posteriorly — in  Aulopyge,  Eurasian  Barbus  and  schizothoracins 

Vertebral  number.  Aulopyge  has  a  total  of  37-38  vertebrae,  of  which-  10  (including  the  four 
Weberian  vertebrae)  are  pre-dorsal,  i.e.  the  neural  spine  of  the  last  vertebra  in  the  set  lies  in  front 
of  the  1st  dorsal  pterygiophore.  This  total  vertebral  number  lies  within  the  modal  range  for 
Cyprininae. 

In  a  sample  of  46  'large'  African  Barbus  species  the  range  is  36-42,  of  which  20  species  have  a 
range  of  9-11  pre-dorsal  vertebrae,  4  species  have  11-12  (oxyrhynchus ,  somereni,  mariae  and 
ethiopicus)  and  the  remaining  22  species  have  13-17.  These  latter  species,  apart  from  the  South 
African  Cape  B.  serra  and  B.  andrewi,  are  European  and  Middle-eastern  species  (Table  1). 
Schizothoracin  genera  have  both  higher  total  (46-48)  and  pre-dorsal  (13-17)  vertebral  numbers 
(Table  2).  In  other  Cyprininae,  the  numbers  of  pre-dorsal  vertebrae  rarely  exceed  10;  in  Cyprinion 
there  are  8-12,  in  Cyprinus  9-10,  Gibelion  8  and  Catlacarpio  8-9.  In  labeins,  Garra  has  9-12,  and 
Labeo  has  8-9.  Squaliobarbin  taxa  also  have  a  high  pre-dorsal  number,  10-12. 

Skelton  (1976)  recorded  the  vertebral  numbers  in  four  groups  of  African  Barbus,  groupings 
made  on  the  basis  of  scale  striae  pattern  and  degree  of  ossification  in  the  last  unbranched  dorsal  fin 
ray.  He  found  higher  counts  in  the  group  with  parallel  striated  scales  and  with  the  dorsal  fin  ray 
ossified  and  smooth,  a  group  to  which  belong  the  'large'  African  Barbus  species  cited  above. 


180 


G.  J.  HOWES 


Fig.  12     Barbus  intermedius  Ontogenetic  development  of  the  neural  complex,  at  (a)  21-5  mm  SL;  (b) 
23-5  mm;  (c)  25  mm;  (d)  31  mm.  Scale  bars  =  0-5  mm. 


Following  Lindsey's  (1976)  broader  discussion  of  pleomerism,  Skelton  (1980)  pointed  out  that 
Jordan's  rule  (the  correlation  of  increased  vertebral  number  with  higher  latitudes)  may  be  a  factor 
when  considering,  for  example,  the  endemic  high-latitude,  high-altitude  redfin  'Barbus'  which 
have  a  more  frequently  occurring  range  (36-38)  than  species  of  'small'  African  Barbus  (31-38). 
Skelton  argues  that  such  specialisation  signifies  that  the  higher  vertebral  number  represents  a 
synapomorphy,  one  he  uses  to  recognise  the  redfin  'Barbus'  as  a  monophyletic  group. 

In  the  Cyprininae,  the  total  vertebral  number  never  exceeds  48,  and  the  modal  range  is  38-40;  in 
the  Leuciscinae  the  total  range  is  greater,  being  33-61,  as  is  the  modal  range  of  40-45  (see  Howes, 
1978,  Table  1).  Perhaps  more  significant  is  the  consistently  higher  range  of  pre-dorsal  vertebrae  in 
Leuciscinae,  10-19  versus  9-14  in  Cyprininae.  Howes  (1978;  1984)  considered  a  high  number  of 
vertebrae  as  a  synapomorphy  for  the  aspinin  group  of  leuciscine  cyprinids,  since  the  range  for  this 
group  exceeds  that  of  other  leuciscines  in  both  abdominal  and  caudal  vertebrae. 


CYPRINID  FISH  GENUS  AULOPYGE 


181 


Table  1  Vertebral  and  lateral  line  counts  in  Barbus  species  having  high  total  and  pre-'dorsal  numbers  of 
vertebrae  and  lateral  line  scales,  and  having  a  serrated  last  unbranched  dorsal  fin  ray.  In ' Barbus'  species  with 
a  pre-dorsal  vertebral  count  of  9-1 1,  the  total  count  rarely  exceeds  43. 


Species 

Total 

Pre-dorsal 

Lateral  line  scales 

albanicus 

44 

14 

57 

andrewi 

38^0 

14-16 

38-40 

barbulus 

44 

13 

52-54 

barbus 

46 

14 

55-63 

bocagei 

42^4 

14-15 

45-49 

brachycephalus 

47 

11 

63 

capita 

42-45 

13 

57-65 

(including  specimens  labelled  as  kersiri) 

comiza 

43 

12 

48-50 

(syntypes) 

esocinus 

48 

14 

76-78 

graellsi 

42^3 

14 

47-52 

grypus 

44-47 

13-14 

40 

lacerta 

43 

13 

55-63 

longiceps 

43^44 

13-14 

51-60 

meridionalis 

40 

13 

48-60 

mursa 

43 

14 

90-97 

nasus 

43^4 

13-14 

49-78 

plebejus 

41^2 

13-14 

49-78 

rajanorum 

45-46 

13-14 

57-65 

(including  specimens  labelled  as  schejch) 

sclateri 

42 

12 

46-47 

(syntypes) 

serra 

39^1 

14-17 

42^3 

sharpeyi 

40-42 

13-14 

30-31 

(lacks  serrated  last  dorsal  spine) 

subquincunciatus 

45 

13-14 

80-84 

xanthopterus 

44 

13 

58-60 

Table  2    Vertebral  counts  in  a  selection  of  schizothoracins. 


Species 


Total 


Pre-dorsal 


Diptychus  dybowski 
Diptychus  maculatus 
Gymnocypris  sp. 
Schizothorax  dipogon 
Schizothorax  esocinus 
Schizothorax  chrysochlorus 
Schizothorax  grahami 
Schizothorax  intermedius 
Schizothorax  richardsoni 
Schizothorax  prenanti 
Schizothorax  sinuatus 
Schizothorax  yunnanensis 
Schizopygopsis  stoliczkae 


48 

49 

46 

49 

46-47 

42 

47 

48 

46 

46 

48 

46 

48 


182 


G.  J.  HOWES 


dfs 
2-3 


Fig.  13 


rp 


Unbranched  dorsal  fin  rays  of,  (a)  Aulopyge  huegelii,  1 1 5  mm  SL;  (b)  Barbus  barbus,  24  mm  SL 
(cartilage  stippled);  scale  =  1  mm. 


It  is  difficult  to  assign  polarity  to  vertebral  number  for  other  groups  of  cyprinids  because, 

*there  is  a  continuum  from  the  relatively  low  numbers  in  Cyprininae  to  the  higher  numbers  in 
Leuciscinae 

*there  is  the  phenomenon  of  pleomerism  (see  Lindsey,  1975) 

*vertebral  numbers  may  be  influenced  by  latitudinal  position  and  temperature  changes  (see 
Lindsey,  1975;  Lindsey  &Arnason,  1981). 

Lindsey  (1975)  commented  that  the  Catostomidae  display  significant  pleomerism  among  its 
species,  but  not  in  the  family  as  a  whole.  The  same  observation  can  be  applied  to  the  Cyprinidae, 
where  deep-bodied  genera  such  as  Cyprinion  and  Megalobrama  have  similar  maximum  lengths  to 
those  of  cylindrical,  depressed  or  compressed  and  slender  forms,  but  possess  lower  vertebral 
numbers. 

Dor  sal  fin  and  serrated  unbranched  dorsal  fin  ray.  In  Aulopyge  the  first  (reduced)  dorsal  fin  ray  lies 
on  a  vertical  just  anterior  to  the  base  of  the  pelvic  fin  and  above  the  15th  vertebra. 

In  those  Barbus  with  a  high  number  of  pre-dorsal  vertebrae  the  1st  dorsal  fin  ray  lies  above  the 
16th-18th  vertebra  and  above  or  somewhat  anterior  to  the  origins  of  the  respective  fins.  In  the 
majority  of  the  Cyprininae,  the  dorsal  fin  lies: 

*above  or  anterior  to  the  origin  of  the  pelvic  fins. 

*rarely  posterior  to  the  pelvic  fin  origin,  (e.g.  'Labeo1  stoliczkae,  Barbus  paludinosus,  B.  serra, 
some  Puntius  species). 

In  the  Leuciscinae,  however,  only  exceptionally  does  the  dorsal  fin  originate  in  advance  of  the 
pelvics,  (e.g.  Pogonichthys).  In  both  Cyprininae  and  Leuciscinae  there  are  several  taxa  where  the 
dorsal  fin  origin  is  immediately  above  the  pelvic  fin  insertion.  Such  a  situation  occurs  in  both  basal 
leuciscines  with  short,  cylindrical  bodies,  (e.g.  Opsariichthys,  Zacco)  and  those  with  elongate, 
compressed  bodies,  (e.g.  Salmostoma,  Macrochirichthys).  In  cyprinines,  this  generalised  position 
of  the  dorsal  fin  is  present  in  many  'large'  and  'small'  African  Barbus  species.  Skelton  (1980) 
considered  a  dorsal  fin  posteriorly  placed  in  relation  to  the  pelvics  as  a  synapomorphy  uniting 
serrated-dorsal  fin  rayed  redfin  Barbus  species.  However,  if  one  assumed  the  Cyprininae  to  be  the 
derived  sister-group  of  the  Leuciscinae,  such  a  posterior  dorsal  fin  position  may  indicate  the 


CYPRINID  FISH  GENUS  AULOPYGE 


183 


ah 


hb1 


ph 


ih 


bb1-3 


p2-3 


cb4-5 


Fig.  14     Aulopyge  huegelii,  branchial  arches  of  left  side  in  dorsal  view.  Scale  =  2  mm. 


plesiomorphic  condition.  The  forward  placement  of  the  dorsal  fin  in  the  Cyprininae,  seen  in  its 
most  extreme  form  amongst  labeins,  is  more  likely  to  be  the  derived  state. 

An  analysis  of  the  position  of  the  1st  dorsal  fin  ray  in  relation  to  the  vertebral  column  again 
reflects  the  major  taxonomic  grouping  of  the  Cyprininae  and  Leuciscinae.  In  the  majority  of 
cyprinines,  the  1st  dorsal  ray  lies  above  the  ll-18th  vertebra  (14— 15th  in  Aulopyge),  whereas  in 
leuciscines,  it  may  lie  above  any  from  the  16th  to  the  31st  vertebra  (modally  between  the  18th  and 
2 1  st).  The  furthest  posterior  position  of  the  1  st  dorsal  fin  ray  occurs  in  the  chelin  assemblage,  where 
it  lies  above  the  21st-26th  vertebra  in  Salmostoma  and  the  30th-31st  in  Macrochirichthys.  In  the 
schizothoracins  the  1st  dorsal  ray  lies  above  the  17th-21st  vertebra. 

In  Aulopyge  there  are  3  unbranched  dorsal  fin  rays,  the  last  being  moderately  serrated  along  its 
distal  posterior  border;  there  are  7-8  branched  dorsal  fin  rays. 

The  number  of  unbranched  dorsal  fin  rays  preceding  the  1st  branched  ray  varies  in  the 
Cyprinidae  from  2-6.  Gosline  (1978)  found  some  significance  in  the  numbers  of  unbranched  dorsal 


184 


G.  J.  HOWES 


cl 


scp 


Fig.   15    Aulopyge  huegelii,  right  pectoral  girdle  in  lateral  view.  The  (medial)  positions  of  the 
mesocoracoid  and  part  of  the  scapula  are  indicated  by  dashed  lines. 


fin  rays,  believing  a  modal  count  of  4  to  be  representative  of  the  Cyprininae  whilst  3  was  present  in 
'. . .  other  cyprinid  subfamilies'.  Although  Gosline's  subfamily  concept  differs  from  that  presented 
here,  I  find  his  statement  justified.  A  possible  reason  for  there  being  a  high  number  of  unbranched 
dorsal  fin  rays  in  cyprinines  may  be  correlated  with  the  often  marked  ossification  of  the  last  such 
element.  A  large  heavy  spine-like  ray,  in  order  to  remain  rigid  may  require  some  anterior  bracing  in 
the  form  of  several  and  strong  elements  in  the  fin. 

Highly  ossified  dorsal  rays  rarely  occur  in  the  Leuciscinae,  (e.g.  Capoetobramd)  and  never  bear 
serrations. 

In  the  Cyprininae  the  last  unbranched  ray  is  always  the  largest  but  varies  from  flexible  to  heavily 
ossified,  and  may  be  smooth  or  serrated  along  its  posterior  margin.  When  present,  a  serrated  ray 
may  bear  serrae  over  its  entire  or  partial  proximal  length.  A  serrated  dorsal  ray  occurs  only  in  some 
species  of  the  genera  Barbus,  'Puntius\  Schizothorax  and  Mystacoleucus,  while  in  other  genera, 
such  as  Acrossocheilus ,  Cyclocheilichthys  and  Cyprinus,  all  species  possess  a  serrated  last 
unbranched  dorsal  ray. 

Based  principally  on  the  classification  of  Boulenger  (1911),  Skelton  (1976)  recognised  four 
group  of  Barbus,  of  which  only  one  (Group  III)  contained  species  with  a  serrated  dorsal  fin  ray. 
Within  this  group,  the  subgroup  (IIIA)  comprises  the  'large'  African  Barbus  and  contains  those 
species  which  also  have  relatively  high  total  and  pre-dorsal  vertebral  counts  (see  above,  p.  1 79  and 
Table  1).  The  ranking  of  serrated  dorsal  fin  rays  as  a  synapomorphy  is  dubious  since  the  feature 
has  an  irregular  distribution  among  genera  recognised  as  monophyletic,  (e.g.  Cyprinion;  see 
Howes,  1982).  However,  it  would  be  possible  to  test  for  the  plesiomorphic  nature  of  dorsal  fin  ray 
serrations  by  observing  their  presence  in  some  ontogenetic  stage  of  those  taxa  whose  adults  lack 
them.  In  Barbus  barbus,  in  which  the  last  unbranched  dorsal  ray  bears  serrations,  they  begin  to 
appear  at  23-5  mm  SL  when  that  ray  is  still  segmented  (Fig.  1 3b). 


CYPRINID  FISH  GENUS  AULOPYGE 


185 


hyp  6 


Fig.  16  Aulpyge  huegelii,  (above)  caudal  fin  skeleton  of  52  mm  SL  specimen.  Scale  =  1  mm;  (below) 
variation  on  second  neural  spine  (dark  shading)  on  PU2  of  specimens  (a)  106mm,  (b)  112mm,  (c) 
127mmSL. 


Other  osteological  features 

Aulopyge  is  conservative  in  its  other  skeletal  elements. 

The  gill  arches  are  of  a  generalised  cyprinid  type  except  for  the  complete  absence  of  gill-rakers  on 
the  outer  margin  of  the  1st  ceratobranchial  and  only  3  or  4  rakers  on  the  1st  epibranchial.  The 
pharyngeal  bone  (5th  ceratobranchial,  Fig.  14)  is  broad  and  bears  a  single  row  of  four  teeth,  the 
first  somewhat  globular  with  a  prominent  cusp,  the  others  having  bevelled  or  chisel-like  crowns. 

The  pectoral  girdle  has  a  tall,  upright  cleithral  limb  and  a  short  horizontal  limb  with  a  narrow 
lamina  (Fig.  1 5).  The  cleithral-coracoid  foramen  is  minute  and  the  coracoid  is  small.  The  size  of  the 
cleithral-coracoid  foramen  is  variable  amongst  cyprinids,  both  intra-  and  interspecifically  (see 
Howes,  1979:  180),  and  appears  to  have  little  worth  as  a  phyletic  character.  There  is  a  single,  long 
postcleithrum  in  Aulopyge. 

The  caudal  fin  skeleton  is  of  a  generalised  type  with  6  hypurals,  a  well-developed  hypurapophysis, 
paired  uroneurals  and  a  long,  proximally  expanded  epural  (Fig.  16).  There  is,  however,  variable 
development  in  the  neural  arch  on  PU2.  In  the  smallest  specimen  available  (52  mm  SL)  there  are 
two  neural  arches  on  PU2,  the  posterior  arch  having  only  a  short  spine  (Fig.  16).  In  a  specimen  of 
106  mm  SL  there  are  two  arches  with  fully  developed  spines,  and  in  the  largest,  127  mm  SL,  the 
condition  resembles  that  of  the  smallest  specimen,  namely,  the  second,  posterior  arch  having  a 
small  neural  spine  (Figs  16a-c). 


186 


Fig.  17    Aulopyge  huegelii,  anal  tube  and  anal  fin  of  female.  Drawn  from  dissection  and  X-Radiograph 

of  1 16  mm  SL  specimen. 

Radiographs  of  a  wide  range  of  cyprinines  reveal  the  presence  of  a  second  neural  arch  on  PU2  to 
be  of  not  infrequent  occurrence,  although  when  it  does  occur,  the  neural  spine  is  usually  fully 
developed,  (e.g.  Barbus plebejus ,  Barbuscanis,  Barbusmicropogon,  Barbus  barbulus ,  Acrossocheilus 
yunnanensis,  Carassius  auratus).  A  reduced  second  PU2  neural  spine  is  found  in  Barbus  comiza. 

The  significance,  phyletic  or  otherwise,  of  a  second  neural  arch  and  spine  on  PU2  is  unknown.  Its 
mosaic  and  wide  distribution  in  cyprinines  make  polarity  assignment  impossible.  It  is  of  interest  to 
note,  however,  that  in  leuciscines,  it  is  the  3rd  preural  centrum  which  bears  a  double  neural  arch 
rather  than  the  2nd  as  in  cyprinines  (see  Howes,  1984:  296).  Variability  of  neural  arches  on  the 
posterior  caudal  centra  may  be  a  plesiomorphic  feature  of  teleosts;  Greenwood  (1970:  134)  noted 
such  variability  in  Elopiformes. 

Sexual  dimorphism  and  genitalia 

Seeley  (1 886)  pointed  out  the  marked  sexual  dimorphism  of  Aulopyge  exhibited  in  the  morphology 
and  position  of  the  anal  and  genital  openings  and  in  the  smaller  body  size  of  the  male. 

In  the  male  Aulopyge,  the  anus  and  genital  opening  are  separated,  the  genital  orifice  being 
posterior  in  position  and  lying  in  front  of  the  first  unbranched  anal  fin  ray.  In  the  female,  both 
openings  and  their  respective  ducts  are  contained  in  a  fleshy  tube  which  is  adnate  to  the  2nd 
unbranched  anal  fin  ray.  The  oviduct  is  firmly  joined  to  the  flexible  2nd  ray  for  part  of  its  length 
(Fig.  1 7).  In  both  males  and  females  the  1  st  unbranched  anal  fin  ray  is  vestigial  and  does  not  project 
from  the  body  surface.  The  genital  morphology  of  Aulopyge  is  unique  among  cypriniforms. 


Discussion 
Aulopyge  relationships  and  barbin  classification 

Aulopyge  exhibits  a  condition  well  known  to  cyprinid  systematists,  namely  the  possession  of 
several  unique  features  (autapomorphies)  and  few,  if  any,  recognisable  synapomorphies  with 
other  cyprinid  taxa.  Too  few  published  comparative  anatomical  data  exist  for  barbelled  carps 
(Cyprininae)  and  the  comparisons  made  during  this  study  are  of  limited  taxonomic  scope. 
However,  some  information  has  emerged  which  may  signpost  useful  characters  for  determining 
subgroups  amongst  barbins.  The  phylogenetic  position  of  Aulopyge  can  only  be  discussed  in  the 
context  of  these  wider  issues. 

It  was  stated  in  the  Introduction  that  Aulopyge  is  a  member  of  the  Cyprininae.  This  subfamily 
was  one  of  the  divisions  recognised  by  Howes  (1981)  on  the  basis  of: 


CYPRINID  FISH  GENUS  A  ULOPYGE  1 87 

*a  maxillary  barbel  associated  with  a  foramen  in  the  maxillary  bone  through  which  the  barbel  is 
supplied  by  a  branch  of  the  VII  facial  nerve. 

*a  rostrally  extended  supraethmoid  with  a  laterally  convex  border. 

At  present  only  two  monophyletic  assemblages  have  been  identified  within  the  Cyprininae,  viz. 
the  squaliobarbin  group  (Howes,  1981)  and  the  labein  group  (Reid,  1982;  1985).  The  former  is  a 
group  of  three  seemingly  plesiomorphic  genera  (Ctenopharyngodon,  Mylopharyngodon  and 
Squaliobarbus)  having  a  native  distribution  restricted  to  China.  The  labeins  are  a  speciose 
assemblage  of  c.  16  genera  with  an  Afro- Asian  distribution.  Aulopyge  shares  none  of  those  derived 
characters  listed  by  Howes  (1981)  and  Reid  (1982;  1985)  as  defining  either  group. 

The  Cyprininae  may  be  subdivided  on  the  basis  of  the  morphology  of  the  dilatator  fossa  (Howes, 
1981:15).  There  are  two  conditions  of  the  fossa;  1 )  it  indents  the  dorso-lateral  cranial  surface,  or  2) 
it  is  a  foramen  in  the  ventral  lamella  of  the  frontal,  and  in  the  case  of  the  labeins,  the  sphenotic  as 
well. 

It  is  assumed  from  its  widespread  occurrence  in  teleosts,  and  its  universal  presence  in  all  non- 
barbelled  cyprinids  (Leuciscinae)  and  other  cyprinoids,  that  the  dorsal  cranial  dilatator  fossa 
represents  the  pleisiomorphic  condition.  That  the  ventral,  foraminate  dilatator  fossa  is  a  derived 
condition  is  reinforced  by  its  ontogenetic  history. 

The  development  of  the  foraminate  fossa  was  traced  in  a  series  ofBarbus  intermedius  specimens 
20-55  mm  SL.  In  the  smallest  specimens  the  fossa  is  of  the  plesiomorphic  type,  (i.e.  dorso-laterally 
placed  and  indenting  the  surfaces  of  the  sphenotic  and  frontal);  the  dilatator  operculi  muscle  is  a 
narrow  band-like  element.  At  29  mm  SL  there  is  a  lateral  process  on  both  the  frontal  and  sphenotic; 
the  indentation  for  the  muscle  in  the  frontal  has  deepened.  By  31  mm  SL  the  spheiiotic  process  has 
curved  downward  and  the  frontal  lamella  is  perforated;  the  anterior  part  of  the  dilatator  muscle 
runs  through  the  foramen  and  fibres  also  originate  from  its  lateral  rim  and  the  sphenotic  process 
(Figs  1 8a-c).  By  55  mm  SL  the  foramen  is  well-formed  and  increased  in  size  by  medial  attrition  of 
the  frontal  lamella. 

In  the  smallest  specimens  ofBarbus  barbus  available  (25  mm  SL)  there  is  no  sign  of  a  foraminate 
dilatator  fossa  and  the  condition  resembles  that  in  the  smallest  specimen  ofBarbus  intermedius.  It  is 
reasonable  to  assume  that  the  development  of  the  fossa  in  this  species  proceeds  along  much  the 
same  course  as  that  described  in  B.  intermedius.  In  the  two  closely  related  species  Barbus  litamba 
from  Lake  Malawi  and  B.  mattozi  from  the  Limpopo,  the  fossa  is  foraminate  only  in  specimens 
above  103  mm  SL,  and  then  only  has  a  small  opening. 

Although  it  may  be  argued  that  a  foraminate  dilatator  fossa  could  have  been  derived  inde- 
pendently within  different  cyprinine  lineages,  it  will  be  accepted  as  a  working  hypothesis  that  it  is 
the  principal  synapomorphy  for  one  group  of  Cyprininae.  Since  Aulopyge  lacks  a  foraminate 
dilatator  fossa  it  must  be  included  with  the  squaliobarbins,  schizothoracins,  several  Barbus  species 
and  other  taxa  listed  in  Table  3.  Of  these,  the  most  likely  candidate  for  sister-group  relationship  to 
Aulopyge  is  the  schizothoracin  assemblage.  Some  schizothoracin  genera  lack  scales,  possess  a 
narrow  ethmoid,  serrated  last  unbranched  dorsal  fin  ray  and  have  a  well-developed  ventral  facet  on 
the  lateral  ethmoid,  all  derived  characters  shared  with  Aulopyge.  However,  these  characters  are 
mosaically  distributed  amongst  schizothoracin  species,  no  one  taxon  possessing  all  together,  and 
so  a  relationship  between  Aulopyge  would  involve  only  certain  species,  thus  making  the  schizo- 
thoracins a  paraphyletic  group.  Previous  authors,  in  recognising  the  subfamily  Schizothoracinae 
sensu  Berg,  1912,  have  tacitly  assumed  monophyly.  Such  an  assumption  is  based  on  the  possession 
by  all  included  taxa  of 'tile'  scales,  i.e.  a  row  of  regular,  oblong  scales  at  the  base  of  the  anal  fin.  This 
synapomorphy  is  supported  by  another,  namely  the  presence  of  a  bony  strut  extending  from  the 
parasphenoid  to  contact  the  prootics  in  the  midline  and  thus  dividing  the  posterior  myodome.  As 
such,  this  feature  resembles  the  basisphenoid  present  in  other  teleosts,  but  which  is  absent  in 
ostariophysans.  These  two  characters  indicate  the  monophyly  of  the  schizothoracins  and  as  such 
exclude  Aulopyge,  which  lacks  both  of  them. 

Aulopyge  also  shares  the  character  of  well-developed  lateral  ethmoid  and  entopterygoid  facets 
with  some  species  of  Barbus.  This  character  distribution  immediately  raises  the  question;  what  is 
meant  by  Barbus] 

At  the  present  time  Barbus  includes  c.  800  nominal  species  distributed  in  Eurasia  and  Africa, 


188 


G.  J.  HOWES 


Fig.  18  Barbus  intermedius  intermedius,  ontogenetic  development  of  dilatator  fossa;  (a)  at  21  -5  mm;  (b) 
at  25  mm;  (c)  at  3 1  mm  SL.  Dashed  line  indicates  position  of  dilatator  operculi  muscle;  all  in  ventro- 
lateral  view.  Scales  =  0-5  mm. 


many  of  which,  even  to  the  non-specialist,  bear  scant  resemblance  to  the  type  species  of  the  genus, 
the  European  Barbus  barbus  (Linn.).  Some  authors  have  opted  to  recognise  separate  genera,  (e.g. 
Puntius,  Tor)  for  Indian  and  South  East  Asian  species,  a  solution  which  does  little  to  elucidate 
relationships  since  these  'genera'  are  not  defined  on  derived  characters.  The  definition  of  Barbus 
can  only  be  approached  through  an  adequate  anatomical  comparison  of  the  Eurasian  and  African 
species. 

Comparisons  and  character  analyses  made  during  this  study  have  demonstrated  that  a  'group' 
including  the  type  species  Barbus  barbus  and  other  Eurasian  species  can  be  defined  on  a  suite  of  five 
characters: 

1)  shield-shaped  supraethmoid  (Fig.  19a) 

2)  oblong  lachrymal  with  ventral  sensory  canal  (Fig.  lOb) 

3)  enlarged  lateral  ethmoid  facet  articulating  with  a  well-developed  entopterygoid  facet  (Fig.  5a) 

4)  13-15  pre-dorsal  vertebrae 


CYPRINID  FISH  GENUS  AULOPYGE 
Table  3     Distribution  of  the  dilatator  fossa  morphotypes  amongst  examined  Cyprininae. 


189 


Foramen  present: 

Single  foramen 

Acrossocheilus 

Barbichthys 

Barbus  (part;  see  Table  4) 

Capoeta 

Carassius 

Cyprinus 

Probarbus 

Varicorhinus 


Foramen  absent: 

Ageniogarra 

Aulopyge 

Barbus  (part;  see  Table  4) 

Cyprinion 

Mystacoleucus 

Onychostoma 

Prolabeo 


Double  foramen 

Catla 

Cirrhina 

Crossocheilus 

Garra 

Labeo 

Labiobarbus 

Lobocheilos 

Osteocheilus 

Semilabeo 

Tylognathus  (sensu  Reid,  1985) 

Typhlogarra 


and  in  the  squaliobarbins 

Squaliobarbus 

Mylopharyngodon 


Fig.  19     Ethmoid  region  in  dorsal  view  of,  (a)  Barbus  bar  bus;  (b)  B.  altianalis  altianalis;  (c)  B.  leonensis 
(scale  bar  =  0-5  mm);  (d)  B.  serra;  (e)  Tor putitora. 


190  G.J.HOWES 

Table  4     Condition  of  the  dilatator  fossa  in  80  species  of  'Barbus\ 


Foramen  present 

African  species:  altianalis  (all  subspecies),  andrewi,  biscarensis,  callensis,  camp  I acanthus,  fritschi,  guirali, 

intermedius  (all  subspecies),  jacksonijohnstoni,  litamba  (some,  see  text),  macrolepis,  marequensis  (all  morphs), 

natalensis,  nigrodor sails,  oxyrhynchus,progenys,  reinii,  rothschildi,  serra,  setivemensis,  somerini,  trimaculatus , 

tropidolepis. 

Eurasian  species:  barbus,  barbulus,  bocagei,  canis,  comiza,  douronensis,  graellsii,  grahami,  hexagonolepis, 

longiceps,  meridionalis ,  nasus,  plebejus,  sharpeyi,  tambroides,  tor,  xanthopterus. 

Foramen  absent 

African  species:  ablabes,  amphigramma,  argenteus,  aspilus  aurantiacus,  dorsolineatus,  eutaenia,  holotaenia, 
hospes,  hypsolepis,  kersteni,  leonensis,  lineomaculatus,  macrops,  mimus,  neglectus,  neumayeri,  paludinosus, 
paytoni,  perince,  poechi,  profundus,  radiatus,  tenuis,  thalamakanensis. 

Asian  species:  aurilius,  bimaculatus,  binotatus,  burmanicus,  chola,  collingwoodi,  conchonius,  filamentosus, 
pentazona,  lithopides,  orphoides,  sarana,  sophore,  titteya. 


5)  low  neural  complex  widely  separated  from  the  4th  neural  spine  (Fig.  1  Ib) 
Of  these  only  characters  1  and  2,  because  of  their  restricted  distribution,  can  be  treated  as 
synapomorphies  (character  2  is  also  shared  with  Aulopyge;  see  below).  Characters  3-5,  when 
viewed  in  the  context  of  cyprinoid  distribution  are  apparently  derived.  Their  disparity  among 
cyprinines,  however,  does  not  make  them  highly  corroborated  synapomorphies.  Nonetheless  they 
are  congruent  with  characters  1  and  2. 

If,  on  the  basis  of  this  character  suite,  Barbus  is  restricted  to  only  some  Eurasian  species  (see 
Appendix  1)  then  it  remains  to  be  determined  how  closely  related  it  is  to  those  African  and  Asian 
species  presently  included  in  Barbus,  Tor  and  Puntius.  From  the  distribution  of  the  foraminate 
dilatator  fossa  (see  above)  it  is  clear  that  African  and  Asian  barbins  do  not  constitute  a  mono- 
phyletic  assemblage.  Of  80  African  and  Asian  'Barbus'  species  selected  at  random,  virtually  50% 
possess  the  character  (4 1  with,  39  without;  Table  4).  Also  emerging  from  this  analysis  is  that  almost 
none  of  the  'small'  African  Barbus  examined  possess  a  foraminate  fossa.  Thus,  on  the  basis  of  the 
synapomorphic  foraminate  fossa,  some  Barbus  species  are  more  closely  related  to  labein  and  other 
cyprinine  genera  such  as  Capoeta,  Cyprinus,  Varicorhinus  and  Acrossocheilus  than  to  other  Barbus 
species. 

Immediate  relationships  ofAulopyge 

Although  it  lacks  a  foraminate  dilatator  fossa,  Aulopyge  shares  with  some  Eurasian  Barbus  species 
(termed  from  hereon  Barbus  sensu  stricto)  well-developed  lateral  ethmoid  and  entopterygoid 
articular  facets  and  an  oblong  lachrymal  with  a  ventral  sensory  canal  (although  in  Aulopyge  the 
canal  is  not  fused  with  the  bone;  see  p.  1 74.  The  lack  of  a  foraminate  fossa  may  be  interpreted  either 
as  a  loss  or  as  a  plesiomorphic  condition,  in  which  case  the  lateral  ethmoid-entopterygoid  facets 
and  oblong  lachrymal  must  be  viewed  as  having  been  independently  derived.  Fewer  assumptions 
are  required  to  support  the  'loss'  of  the  derived  dilator  fossa  in  Aulopyge  than  are  demanded  by 
other  schemes  of  relationship  (Figs  20A-C).  Support  for  'loss'  is  that  Aulopyge  exhibits  heter- 
ochrony  in  the  late  development  and  fusion  of  infraorbital  sensory  canals  and  in  the  absence  of 
scales.  It  may  be  that  the  dorso-lateral  dilatator  fossa  is  also  the  retention  of  an  early  ontogenetic 
stage  (see  p.  187).  Outright  dismissal  of  independent  origin  on  grounds  of  parsimony  must  be 
treated  with  caution,  however,  since  it  is  noted  that  amongst  the  schizothoracins  a  lateral  ethmoid 
facet  is  present  in  some  taxa  (p.  1 70).  Since  the  schizothoracins  are  almost  certainly  a  monophyletic 
group  within  the  Cyprininae  (see  p.  1 87)  it  follows  that  this  feature  has  been  derived  independently 
from  that  in  Aulopyge,  Barbus,  sensu  stricto  and  other  barbins  (including  Cyprinus). 
Whether  Aulopyge  is  recognised  as  the  sister-group  to  Barbus  sensu  stricto,  or  to  Barbus  sensu 


CYPRINID  FISH  GENUS  AULOPYGE 


191 


Fig.  20  Three  possible  schemes  of  relationship  between  Aulopyge  and  other  cyprinines.  A,  the  most 
parsimonious,  involves  the  loss  of  character  2  in  Aulopyge.  B.  involves  the  loss  of  characters  3  and  4  in 
'Barbins'  and  Labeins.  C,  involves  independent  derivation  of  characters  3  and  4  in  Aulopyge  and 
Barbus  sensu  stricto. 

Character  1.  lateral  ethmoid  articular  facet;  2.  foraminate  dilatator  fossa;  3.  expansion  of  lateral 
ethmoid  facet  and  presence  of  entopterygoid  facet;  4.  oblong  lachrymal;  5.  double-foraminate  fossa 
(additional  characters  defining  Labeins  are  given  in  Reid,  1985).  'Barbins'  include  those  taxa  listed  in 
Table  3,  under  'foramen  present'. 


stricto  plus  other  barbins  and  labeins  cannot  be  resolved  on  those  characters  considered  here. 
However,  no  synapomorphies  have  been  identified  that  would  suggest  Aulopyge  is  more  closely 
related  to  any  cyprinine  taxon  lacking  a  foraminate  dilatator  fossa,  including  the  schizothoracins. 
Karaman's  (1971)  hypothesis  of  an  intermediate  taxonomic  position  for  Aulopyge  between 
'barbine'  and  'schizothoracine'  subgroups  (see  above,  p.  165)  is  not  supported  by  this  study. 
Schizothoracins  do  share  with  Barbus  sensu  stricto  high  total  and  pre-dorsal  numbers  of  vertebrae 
(Table  2),  but  the  polarity  of  this  character  is  uncertain  (see  p.  182)  and  if  treated  as  a  synapomor- 
phy  in  a  scheme  of  relationship  involving  Aulopyge,  Barbus  and  'other  cyprinines'  it  is  incongruent 
with  other  synapomorphies. 


192  G.J.HOWES 

I  have  found  no  evidence  in  support  of  Arai's  (1982)  contention  that  Aulopyge  possesses  some 
gobionine  characters. 

From  the  data  analysed  two  hypotheses  are  available: 

*Aulopye  is  a  derived  member  of  the  Barbus  sensu  stricto  lineage,  with  specialisation  through 
reduction  and  'loss'  (Fig.  20A) 
*  Aulopyge  is  a  member  of  the  'stem-group'  of  Eurasian  plus  African  barbins  (Figs  20B  &  C). 


Acknowledgements 

I  am  most  grateful  to  my  colleagues  Keith  Banister,  Humphry  Greenwood,  Alwyne  Wheeler  and  Peter 
Whitehead  for  their  critical  comments  on  the  manuscript,  and  for  their  many  enlivened  discussions  over  the 
years  on  the  taxonomy  of  Barbus.  To  Gordon  Reid  of  the  Horniman  Museum,  I  am  particularly  indebted  for 
his  critical  refereeing,  his  sound  advice  and  continuing  encouragement.  Paul  Skelton  of  the  J.  L.  B.  Smith 
Institute  of  Ichthyology  has  my  gratitude  for  providing  specimens  and  so  much  stimulating  discussion. 

My  sincerest  thanks  are  due  to  Primoz  Zupancic  of  Dolsko,  Yugoslavia,  for  obtaining  the  specimens  of 
Aulopyge  upon  which  most  of  this  work  is  based,  and  for  providing  me  with  photographs  and  information  on 
their  habitat;  and  to  Keith  Easton  of  the  Severn  Trent  Water  Authority  for  providing  juvenile  specimens  of 
Barbus  barbus. 

Special  thanks  go  to  my  colleague  Mandy  Holloway  for  preparing  the  radiographs. 

After  this  paper  had  been  submitted  for  publication,  Dr  Friedhelm  Krupp  of  the  Senckenberg  Museum 
generously  provided  me  with  additional  information  which  has  been  added  to  the  text. 


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Manuscript  accepted  for  publication  1 1  April  1986 


Appendix  1 
The  genus  Barbus  sensu  stricto 

Definition  and  included  species 

Banister  (1980)  placed  the  majority  of  Middle-eastern  Barbus  into  two  groups — the  'European' 
and  'Afro-Indian',  which  he  characterised  on  overall  morphology,  scale  type  and  serrated  or 
smooth  last  dorsal  fin  ray.  Banister  stated  that  both  groups  might  be  monopyletic  and  so  tacitly 
restricted  Barbus  to  the  'European'  group. 

Leveque  &  Daget  (1984)  stated  that  'Strictly  speaking  the  generic  name  Barbus  shall  be  restricted 
to  European  and  some  north- African  species'.  These  authors'  remarks  are  supported  by  this  study 
and  reference  has  been  made  in  the  text  to  Barbus  sensu  stricto.  Only  a  thorough  comparative 
anatomical  study  of 'barbins'  will  provide  a  strict  diagnosis  (based  on  synapomorphies)  of  Barbus. 
The  definition  of  Barbus  used  here  is  based  on  the  characters  analysed  above  and  forms  a  working 


194  G.J.HOWES 

hypothesis  for  a  more  critical  evaluation.  Those  taxa  not  embraced  by  this  definition  are  referred  to 
as  'Barbus'  or  Barbus  sensu  lato;  in  the  case  of  the  Asian  species,  the  generic  names  Puntius  and  Tor 
are  already  widely  used  (see  for  example,  Jayaram,  1981).  For  African  'Barbus'  several  generic 
names  are  available  (see  synonymy  in  Leveque  &  Daget,  1984). 

Barbus  sensu  stricto  is  defined  on  the  basis  of  its  members  having  a  total  vertebral  count  of  40-48 
of  which  13-15  are  pre-dorsal  vertebrae;  a  well-developed,  centrally  to  anteriorly  situated  ventral 
lateral  ethmoid  facet  articulating  with  a  well-developed  anterodorsal  entopterygoid  facet;  a 
'shield'-shaped  supraethmoid  with  (usually)  a  prominent  rostral  process  (Fig.  19a);  neural  com- 
plex low  with  a  deeply  indented  anterior  border,  its  posterior  border  (usually)  well-separated  from 
the  4th  neural  spine  which  is  at  least  half  the  height  of  the  neural  complex;  lachrymal  elongate, 
often  oblong  with  tapered  anterior  tip  and  sometimes  an  indented  posteroventral  border,  sensory 
canal  running  through  the  lower  half  of  the  bone;  49-90  scales  in  the  lateral  line,  (cf.  20-55  in 
African  and  Asian  'Barbus'). 

Almaca  (1981)  distinguished  three  groups  of  Eurasian  Barbus  on  the  basis  of  lateral  line  scale 
counts  but  he  pointed  out  the  lability  of  this  character  due  to  influences  of  temperature  and 
latitudinal  variation  (see  similar  remarks  under  'vertebrae',  p.  182). 

The  following  species  are  considered  to  constitute  Barbus  sensu  stricto: 
Barbus  albanicus  Steindachner,  1870  (including  B.  graecus  (Steindachner,  1895)) 
Distribution:  Albania-Greece 

Barbus  barbus  (Linnaeus,  1758),  type  species  of  the  genus. 

Distribution:  Europe  (see  Almaca,  1981  for  detailed  distributional  data  and  recognition  of 
subspecies) 

Barbus  barbulus  Heckel,  1846 
Distribution:  Tigris 
Barbus  bocagei  Steindachner,  1 865 
Distribution:  Iberia 
Barbus  capita  (Giildenstadt,  1 773) 
Distribution:  Caspian  and  Aral  Sea  basins;  Amu  Darya 
Barbus  comiza  Steindachner,  1865 
Distribution:  Iberia 

This  species  is  included  in  Barbus  on  the  basis  of  its  possessing  a  high  vertebral  number,  and  a 
typical  oblong  lachrymal  (Fig.  lOc).  However,  it  differs  from  other  species  in  its  longer  and 
narrower  head  (see  Almaca,  1967;  1972),  concave  dorsal  profile,  lower  number  of  pre-dorsal 
vertebrae  (1 2,  cf.  1 3-15),  tall  neural  complex  narrowly  separated  from  the  4th  neural  spine,  and  the 
absence  of  a  fleshy  overhanging  upper  lip.  In  its  striking  dorsal  and  lateral  head  profiles,  and 
narrow  ethmoid  B.  comiza  greatly  resembles  Aulopyge.  However,  no  synapomorphies  have  been 
identified  that  would  suggest  these  features  are  anything  other  than  homoplasies. 
Barbus  esocinus  (Heckel,  1843) 
Distribution:  Tigris-Euphrates 
Barbus  graellsi  (Steindachner,  1 866) 
Distribution:  Portugal 
Barbus  lacerta  Heckel,  1843 
Distribution:  Tigris-Euphrates  and  Qwarq  rivers 
Barbus  longiceps  Valenciennes,  1 842 
Distribution:  Jordan  River  system 
Barbus  lorteti  Sauvage,  1882 
Distribution:  Orontes  R. 
Barbus  microcephalus  Almaca,  1 967 
Distribution:  Iberia 

Barbus  meridionalis  Risso,  1826  (including  B.  peloponnesius  Val.,  1842). 
Distribution:  NE  Spain — S.  France — Yugoslavia — Greece 
Barbus  nasus  Giinther,  1874 
Distribution:  Morocco 
Barbus  pectoralis  Heckel,  1843 


CYPRINID  FISH  GENUS  AULOPYGE  195 

Distribution:  Orontes  R. 

BarbusplebejusEonaparle,  1839  (including  the  subspecies  recognised  by  Almac. a,  1981;  1983) 

Distribution:  N.  Italy-Greece 

Barbus  rajanorum  Heckel,  1 843 

Distribution:  Tigris-Euphrates 

Karaman  (1971)  considers  B.  schejch  (Heckel,  1843)  and  B.  barbulus  Heckel  (listed  here  as  a 
separate  species),  to  be  synonyms  of  B.  rajanorum.  This  synonymy  is  doubtful  and  the  'rajanorum 
complex'  requires  a  taxonomic  reappraisal.  In  Dr  F.  Krupp's  opinion  (pers.  com.)  B.  rajanorum  is  a 
hybrid  between  B.  pectoralis  and  Capoeta  damascinus. 
Barbus  sclateri  Giinther,  1 868 
Distribution:  Iberia 
Barbus  subquincunciatus  Giinther,  1 868 
Distribution:  Tigris-Euphrates 
Barbus  steindachneri  Almac.  a,  1967 
Distribution:  Iberia 
Barbus  xanthopterus  Heckel,  1843 
Distribution:  Tigris-Euphrates 

Although  the  Middle-eastern  species  Barbus grypus  and  B.  sharpyeihave  relatively  high  vertebral 
numbers  (Table  1),  they  lack  the  elongate  lachrymal  of  the  other  species  listed  and  possess,  in 
common  with  B.  canis  and  B.  reinii  what  is  considered  to  be  another  derived  form  of  lachrymal  in 
which  the  sensory  canal  runs  along  the  anterodorsal  border  (see  p.  177).  Barbus  canis  and  B.  reinii 
both  have  low  vertebral  numbers,  respectively  38 — 39  and  37  (totals)  and  10  and  10-1 1  predorsal 
elements.  Barbus  sharpy ei  differs  from  other  species  of  this  group  in  having  only  30-31  lateral  line 
scales. 


The  systematic  positions  of  Barbus  brachycephalus  Kessler,  1872  and  B.  mursa  (Guldenstadt,  1773) 

The  generic  placements  of  these  two  south  Central  Asian  species  (respectively  the  Aral  and 
Caspian  Seas  and  the  Kura  system)  are  problematical.  Both  species  although  having  high  vertebral 
counts  differ  in  several  ways  from  Barbus  sensu  stricto  and  other  species  of 'Barbus'. 
Barbus  brachycephalus  has  rather  slender  barbels,  unlike  the  thick,  often  papillate  barbels  of 
'typical'  species  of  the  genus,  and  7  branched  dorsal  fin  rays,  cf.  8  in  the  majority  of  Barbus  sensu 
stricto  and  also  in  Barbus  sensu  lato.  The  cranium  is  broad  and  flat,  lacking  the  transverse  convexity 
of  most  barbins.  There  are  a  total  of  47  vertebrae,  but  only  1 1  are  predorsal,  cf.  13-15  in  Barbus 
sensu  stricto. 

Barbus  mursa  has  a  deep  lachrymal  with  an  anterior  branching  pattern  resembling  that  of  Barbus 
canis  and  related  species  discussed  above  (Fig.  lOn).  However,  it  possesses  a  series  of  preanal  scales 
and  a  prominent  genital  papilla  more  reminiscent  of  schizothoracins. 


The  systematic  positions  of  Barbus  andrewi  Barnard,  1937  and  B.  serra  Peters,  1864 

These  two  species  are  restricted  to  the  South  African  Western  Cape.  On  the  basis  of  the  characters 
given  for  Barbus  sensu  stricto  both  species  should  be  included.  Both,  however,  have  a  higher  pre- 
dorsal vertebral  count  than  other  Barbus  sensu  stricto,  viz.:  14-17,  cf.  13-15,  but,  a  relatively  low 
total  vertebral  number,  viz.:  38-41,  cf.  40-47.  The  supraethmoid  has  the  same  'shield'-shaped 
appearance  as  in  Barbus  sensu  stricto  (Fig.  19d),  but  the  vomer  is  broader  anteriorly  and  extends 
further  dorsally  in  B.  serra  and  B.  andrewi.  Again,  the  lachrymal  bones  of  the  two  species,  while 
having  the  same  overall  appearance  of  that  bone  in  the  Eurasian  Barbus  have  a  sloped,  rather  than 
a  perpendicular  posterior  margin  and  more  convex  ventral  border.  Because  of  these  differences  I 
am  hesitant  to  include  the  Cape  species  in  Barbus  sensu  stricto.  According  to  Skelton  (1 980),  Barbus 
serra  and  B.  andrewi  are  sister-taxa,  not  closely  related  to  any  African  'Barbus'  he  examined. 


196 


G.  J.  HOWES 


Appendix  2 
Characteristics  of  the  subfamilies  Cyprininae  and  Leuciscinae 


CYPRININAE 
(including  'Schizothoracinae') 

Maxillary  barbel  present 
Maxillary  foramen 

Supraethmoid    with    medially    indented    rostral 
process 


4  Articular  facets  present  on  lateral  ethmoid  and 
entopterygoid  in  some  taxa 

5  Single,  strong  ligament  connecting  posterior  face 
of  lateral  ethmoid  with  dorsomedial  surface  of 
entopterygoid 

6  Lachrymal  (1st  infraorbital)  sometimes  elongate 
and  oblong 

7  Dilatator  operculi  muscle  sometimes  originating 
from  ventral  surface  of  the  frontal  and  passing 
through  a  frontal-sphenotic  foramen 

8  Neural  complex  lacking  grooved  dorsal  surface; 
sometimes  close  to,  or  even  contacting  the  cranium 
(supraoccipital) 

9  1  st  free  supraneural  not  contacting  neural  complex 


10  Dorsal  fin  origin  lies  above  or  anterior  to  that  of  the 
pelvics  (rarely  posterior) 

11  1st  unbranched  dorsal  fin  ray  lies  above  the 
llth- 18th  vertebra 

1 2  Last  unbranched  dorsal  fin  ray  often  serrated 


LEUCISCINAE 

No  maxillary  barbel 

No  maxillary  foramen 

No  supraethmoid  rostral  process 

Articular  facets  absent  (except  in  Tinea) 

Connection  between  lateral  ethmoid  via  connec- 
tive tissue  sheet,  sometimes  thickened  laterally 

Lachrymal  never  elongate  or  oblong 


Dilatator  operculi  muscle  confined  to  cranial 
surface;  fossa  never  foraminate 


Neural  complex  with  grooved  dorsal  surface, 
never  contacting  the  cranium 


1st  free  supraneural  articulating  with  neural 
complex 

Dorsal  fin  origin  rarely  above  or  in  advance  of  the 
pelvics 

1st  unbranched  dorsal  fin  ray  lies  above  16th-31st 
vertebra 

Last  unbranched  dorsal  fin  ray  never  serrated 


1 3     Modal  number  of  vertebrae  38^40  (never  exceeding      Modal  number  of  vertebrae  40-45  (range  33-61 ) 
48) 

Note:  The  genus  Tinea  is  included  here  in  the  Leuciscinae,  although  possessing  a  cyprinine  feature 
(character  4).  Chen  et  al.  (1984)  consider  Tinea  to  be  the  sister-group  of  the  Cyprininae  (their 
'Barbines').  Although  for  the  most  part,  these  authors  appear  to  base  their  hypothesis  on  differ- 
ences rather  than  on  shared  homologies,  their  cladogram  requires  serious  consideration  and  offers 
a  much  needed,  testable  classification  of  the  Cyprinidae. 


British  Museum  (Natural  History) 

The  birds  of  Mount  Nimba,  Liberia 

Peter  R.  Colston  &  Kai  Curry- Li ndahl 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Tring,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1 986,  1 29pp.  Hardback.  0  565  00982  6  £  1 7.50. 


Titles  to  be  published  in  Volume  52 


Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophora  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barhus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata). 

By  P.  J.  Chimonides 


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


Bulletin  of  the 

British  Museum  (Natural  History) 


Revision  of  the  genera  Acineria,  Trimyema 
and  Trochiliopsis  (Protozoa,  Ciliophora) 


H.  Augustin,  W.  Foissner  &  H.  Adam 


Zoology  series    Vol  52    No  6    25  June  1987 


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 
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The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 

Keeper  of  Zoology  :  MrJ.  F.  Peake 
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ISBN  0  565  05030  3 

ISSN  0007- 1 498  Zoology  series 

Vol52  No.  6  pp  197-224 

British  Museum  (Natural  History) 

Cromwell  Road 

London  SW7  5BD  Issued  25  June  1987 


(NAT 

26 
Revision  of  the  genera  Acineria,  Trimyema  and 

Trochiliopsis  (Protozoa,  Ciliophora) 

H.  Augustin,  W.  Foissner  &  H.  Adam 

Zoologisches  Institut  der  Universitat  Salzburg,  Hellbrunnerstrasse  34,  Salzburg,  Austria,  A-5020 


Conte 


Synopsis . 
Introduction    . 
Materials  and  Methods 
Genus  Acineria 
Genus  Trimyema 
Genus  Trochiliopsis . 
References 


Synopsis 

The  genera  Acineria,  Trimyema  and  Trochiliopsis  are  reviewed.  The  revision  is  based  on  an  investigation  of 
each  of  the  type-species,  namely  Acineria  incurvata  Dujardin,  Trimyema  compressa  Lackey,  and  Trochiliopsis 
opaca  Penard,  which  were  found  in  a  sewage-treatment  plant.  Acineria  comprises  three  species;  A.  incurvata, 
A.  nasuta,  and  A.  uncinata.  A.  acuta  is  a  synonym  of  A.  incurvata.  Trimyema  comprises  eight  species; 
T.  alfredkahli,  T.  claviformis,  T.  compressa,  T.  echinometrae,  T.  kahli,  T.  marina,  T.  minuta  and  T.  pleuri- 
spiralis  but  T.  alfredkahli  and  T.  claviformis  are  perhaps  synonyms  of  T.  marina.  Trochiliopsis  is  monotypic 
and  new  for  the  fauna  of  Austria.  This  genus  is  apparently  closely  related  to  the  autochthonous  soil  ciliate 
Stammeridium  kahli. 


Zusammenfassung 

Die  Gattungen  Acineria,  Trimyema  und  Trochiliopsis  werden  revidiert.  Die  Revision  basiert  auf  der 
Untersuchung  der  Typusarten,  namlich  Acineria  incurvata  Dujardin,  Trimyema  compressa  Lackey  und 
Trochiliopsis  opaca  Penard,  die  in  einer  Klaranlage  gefunden  wurden.  Die  Gattung  Acineria  umfasst  drei 
Arten;  A .  incurvata,  A .  nasuta  und  A .  uncinata.  A .  acuta  ist  ein  Synonym  von  A .  incurvata.  Von  Trimyema  sind 
acht  Arten  beschrieben;  T.  alfredkahli,  T.  claviformis,  T.  compressa,  T.  echinometrae,  T.  kahli,  T.  marina,  T. 
minuta  und  T.  pleurispiralis.  T.  alfredkahli  und  T.  claviformis  sind  moglicherweise  Synonyme  von  T.  marina. 
Trochiliopsis  ist  monotypisch  und  neu  fur  die  Fauna  Osterreichs.  Diese  Gattung  ist  hochstwahrscheinlich 
nahe  verwandt  zum  autochthonen  Boden-Ciliaten  Stammeridium  kahli. 


Introduction 

Only  few  activated-sludge  ciliates  have  been  characterized  by  silver-staining  techniques  which  is 
sometimes  necessary  for  their  correct  identification.  To  overcome  this  deficiency,  a  project  to 
redescribe  the  most  frequently  occurring  species  was  begun.  During  these  studies  the  poorly  known 
type-species  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  were  found.  They  have  been 
reinvestigated  using  modern  techniques  which  provide  a  base  upon  which  to  revise  these  genera. 

Supported  by  the  'Fonds  zur  Forderung  der  wissenschaftlichen  Forschung,  Projekt  Nr.  P  5889'. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  52(6):  197-224  Issued  25  June  1987 

197 


1 98  H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 

Materials  and  Methods 

Acineria  incurvata,  Trimyema  compressa,  and  Trochiliopsis  opaca  were  obtained  from  activated 
sludge  of  the  sewage-treatment  plant  at  Aspach,  Upper  Austria. 

Small  samples  of  activated  sludge  were  placed  in  glass  petri-dishes  where  they  remained  without 
additional  aeration.  In  such  cultures  a  surprising  succession  and  enrichment  of  ciliates  often 
occurred.  Acineria  incurvata  could  also  be  cultured  in  tap  water  enriched  with  a  crushed  wheat 
grain  which  supported  the  growth  of  many  small  prey  ciliates  (Dexiotricha,  Uronemd). 

The  infraciliature  was  revealed  with  a  protargol  silver-staining  method  (Foissner,  1982).  The 
silverline  system  was  studied  in  specimens  impregnated  by  a  modified  'dry'  silver-impregnation 
technique  (Foissner,  1976).  The  oral  structures  of  Trimyema  compressa  were  impregnated  by  the 
pyridinated  silver  carbonate  method  of  Fernandez-Galiano  (1976)  as  improved  by  Augustin  et  al. 
(1984). 

For  scanning  electron  microscopy  Acineria  cells  were  fixed  for  10  minutes  in  Parducz's  solution 
(2%  OsO4  and  concentrated  Hg-sublimate  solution,  4:1),  rinsed  in  0-05  M  sodium  cacodylate 
buffered  at  pH  6-3,  dehydrated  in  an  isopropyl  alcohol  series  (60%,  70%,  80%,  90%,  100%,  100%, 
five  minutes  each)  and  put  into  a  mixture  of  isopropyl  alcohol  (100%)  and  frigen  1 1  (2  :  1,  1  :  1, 
1 :  2,  five  minutes  each).  Finally,  cells  were  transferred  into  pure  frigen  1 1  and  critical  point  dried, 
using  frigen  13.  Specimens  were  gold-sputtered  three  times  for  six  minutes  each. 

Each  species  was  drawn  from  life  as  well  as  from  impregnated  specimens  using  a  camera  lucida 
for  the  latter.  The  drawings  are  only  slightly  diagrammatic.  All  statistical  procedures  follow 
methods  described  in  Sokal  &  Rohlf  (1981). 

Genus  ACINERIA  Dujardin,  1841 

DIAGNOSIS.  Amphileptidae  Biitschli,  1889  with  (1)  compressed  oral  slit  anteriorly  rolled  up  and 
overlapping  to  the  left  side  forming  (together  with  the  anterior  dorsal  margin)  an  oblique  spoon- 
like  excavation,  (2)  three  perioral  kinetics  (one  left  and  two  right  of  the  cytostome,  (3)  somatic 
kinetics  on  the  right  side  successively  shortened  along  the  cytostome,  (4)  oral  slit  measuring  less 
than  half  of  body  length  being  located  at  the  convex  side  of  the  tapering  anterior.  Freshwater  and 
marine,  prefers  polysaprobic  conditions. 

TYPE-SPECIES.  Acineria  incurvata  Dujardin,  1841 

REMARKS.  Acineria  was  mentioned  for  the  first  time  by  Dujardin  (1840)  in  the  family  'Trichodiens' 
but  without  any  valid  characterization.  In  1 84 1  he  gave  a  rather  vague  diagnosis  of  the  genus  and  of 
two  species.  Maupas  (1883)  criticized  the  unsatisfactory  diagnosis  and  gave  a  better  description 
of  Acineria  incurvata  even  noting  the  overlapping  anterior  end  which  is  the  main  character  of 
Acineria;  nevertheless,  he  did  not  include  this  character  in  the  diagnosis.  How  Maupas  (1883) 
arrived  at  the  conclusion  that  his  species  was  the  same  as  that  described  by  Dujardin  remains, 
however,  inexplicable.  It  was  only  Kahl  (1926)  who  used  the  real  character  of  the  overlapping 
dorsal  end  of  the  mouth  to  the  left  side  to  distinguish  Acineria  from  the  most  closely  related  genus 
Litonotus.  But  there  is  no  indication  in  the  infraciliature  that  the  dorsal  margin  and  the  left  side 
coalesce  as  supposed  by  Kahl  (1926,  1931).  Thus,  Kahl's  interpretation  that  a  part  of  the  ciliated 
right  side  of  the  genus  Litonotus  has  shifted  over  to  the  left  side  in  Acineria  is  not  supported  by  our 
investigations.  We  consider  the  rolled  up  anterior  part  of  the  mouth  to  be  the  reason  for  the 
anterior  overlapping  of  the  dorsal  margin.  The  occurrence  of  somatic  kinetics  on  the  left  side,  as 
stressed  by  Kahl  (1926)  is  a  weak  distinctive  character  because  this  happens  also,  more  or  less 
pronounced,  in  the  genera  Litonotus  and  Amphileptus  (Foissner,  1984). 

Key  to  the  species 

la    Single  spherical  macronucleus A.nasuta 

Ib    Macronucleus  in  two  parts  with  a  single  micronucleus  between  them 2 

2a    Cytostome  restricted  to  the  rolled  up  anterior  pole,  right  side  with  3  somatic  kinetics,  left  side 

unciliated A.uncinata 

2b    Cytostome  about  one  third  of  body  length,  10-12  normally  ciliated  somatic  kinetics  A.  incurvata 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS  199 

Descriptions  of  species 
Acineria  incurvata  Dujardin,  1841 

?Trachelius  anaticula  Ehrenberg,  1 833 

Acineria  acuta  Dujardin,  1841 

?Amphileptus  anaticula  Claparede  &  Lachmann,  1859 

Lionotus  reversus  Kahl,  1926 

Amphileptus  incurvatus  Lepsi,  \926a 

Litonotus  lamella  Fryd-Versavel  et  al.,  1975 

NEOTYPE-SPECIMENS.  Slide  (protargol  silver  impregnated)  of  neotype-specimens  has  been  deposited 
in  the  British  Museum  (Natural  History)  in  London,  reference  number  1986:5:30:1 . 

REDESCRIPTION  (Figs  1-3,  12-58,  Table  1).  Type  species  of  the  genus.  Freshwater  and  marine. 
About  45-200  urn  (Dujardin,  1841;  Maupas,  1883;  Kahl,  1926,  1928,  1931,  1933;  Horvath  & 
Kuhn,  1941;  Bick,  1972;  Foissner,  1977/78).  Abnormal,  giant  individuals  up  to  500  jam  showing 
most  organelles  duplicated  observed  by  Foissner  (1977/78)  and  probably  by  Lepsi  (1965)  (Figs 
28-30).  Body  oblong,  slightly  contractile,  laterally  compressed,  rounded  posteriorly,  narrowing 
anteriorly  to  a  blunt  point.  Rather  variable  in  shape  (slender  to  wide  and  plump)  depending  on 
nutritional  condition  (Figs  3 1-34).  Ventral  side  more  or  less  convex,  dorsal  side  straight  or  concave 
in  the  anterior,  convex  in  the  posterior  region.  Excavated  region  conspicuous,  shining  brightly. 
Anterior-most  dorsal  top  somewhat  refractive,  due  to  the  rolled  up  oral  slit.  Macronucleus  in  two 
spherical  to  ovoid  parts  with  a  single  micronucleus  between  them.  1-3  micronuclei  according  to 
Maupas  (1883).  Macronuclear  parts  fuse  during  bipartition  (Horvath  &  Kuhn,  1941)  (Figs  38-42) 
and  divide  in  the  later  fission  stages  (Kahl,  1926).  Single  contractile  vacuole  at  the  posterior  pole, 
diameter  about  7  um,  with  5-8  pores  on  the  right  lateral  side  (Horvath  &  Kuhn,  1941)  (Fig.  43) 
which  could  not  be  seen  in  our  slides.  Cytoproct  terminal,  a  slightly  laterally  located  slit  (Maupas, 
1883;  Kahl,  1926).  Pellicle  soft,  flexible,  with  longitudinal  furrows  in  which  the  cilia  and  bristles 
originate.  Furrows  disappear  in  well-fed  individuals.  Extrusomes  straight  to  slightly  fusiform 
(arrow-shaped  according  to  Foissner,  1977/78),  thin,  about  4  um  long  (2  urn  according  to  Horvath 
&  Kuhn,  1941),  located  along  the  cytostome,  a  small  accumulation  of  them  in  the  ventral  side  of  the 
posterior  end  and  even  a  few  scattered  throughout  the  body  (Figs  48,  49).  Cytoplasm  of 
normally-fed  specimens  rather  clear,  containing  some  small  colourless  spheres.  Carnivorous,  feeds 
on  small  hymenostome  ciliates,  e.g.  Colpidium,  Cyclidium,  Glaucoma,  Pseudocohnilembus,  Loxo- 
cephalus,  Uronema  (Maupas,  1883;  Lepsi,  1926a;  Kahl,  1926,  1931;  Buck,  1961;  Struhal,  1969). 
Starved  individuals  feed  even  on  'cysts'  ofEuglena  viridis  (Horvath  &  Kuhn,  1941)  and  perhaps  on 
bacteria  (Lepsi,  1926a).  Ingestion  vacuoles  rather  large,  dividing  quickly  into  smaller  food 
vacuoles  (Horvath  &  Kuhn,  1941).  Movement  moderately  quick,  gliding  on  the  bottom  of  the 
petri-dish  or  swimming  in  rotation  along  its  longitudinal  axis.  Bipartition  by  transverse  fission 
(Lepsi,  1926a;  Horvath  &  Kuhn,  1941)  (Figs  38-42).  Opisthe  almost  spherical  when  it  separates 
from  the  proter  (Kahl,  1926;  Horvath  &  Kuhn,  1941)  (Fig.  40).  Very  small  degenerative  forms  tend 
to  conjugate;  during  this  process  the  mouth  of  an  individual  fuses  with  the  back  of  another  (Kahl, 
1926)  (Fig.  20).  Encystment  frequently  occurring  when  food  is  depleted  (Horvath  &  Kuhn,  1941). 
Endocyst  forms  within  an  hour,  later  the  macronuclear  parts  fuse  to  a  worm-shaped  product.  Wall 
of  ectocyst  without  visible  structure.  Cysts  surrounded  by  some  material  which  sticks  them  to  the 
bottom  of  the  culture  dishes  or  to  the  bacterial  film  on  the  surface  of  the  culture  medium  (Horvath 
&Kuhn,  1941)  (Fig.  35). 

Three  different  types  of  cilia:  (1)  normal  cilia,  about  10  um,  (2)  short  bristles,  about  0-5-1-0  um, 
(3)  club-shaped  bristles,  up  to  2-0  um.  Eleven  longitudinal  kinetics  with  cilia  type  1,  about  8-9  of 
them  on  the  right  and  about  3  on  the  left  side.  This  is  in  accordance  with  the  numbers  given  by  Kahl 
( 1 926),  Horvath  &  Kuhn  ( 1 94 1 ),  and  Fryd-Versavel  et  al.  ( 1 975).  In  addition  to  the  normal  somatic 
kinetics  the  following  are  found  on  the  more  differentiated  left  side:  (1)  a  single  kinety  with  cilia 
type  2  located  to  the  left  of  the  brosse  kinety  and  often  extending  only  to  the  middle  of  the  body,  its 
posterior  basal  bodies  less  closely  spaced,  (2)  one  brosse  row  of  obliquely  arranged,  paired  bristles 
(cilia  type  3)  being  posteriorly  continued  by  a  row  of  unciliated  kinetosomes  (or  by  kinetosomes 


200 


H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS  201 

with  very  short  bristles  only),  (3)  one  kinety  consisting  apically  of  2-3  cilia  of  type  3  (probably 
constituting  a  rudimentary  brosse  row)  and  being  continued  by  a  few  unciliated  kinetosomes 
(about  5  in  the  anterior  third  and  about  3  kinetosomes  in  the  middle  of  the  body).  Kahl  (1926, 
1931)  described  the  brosse  as  being  built  up  of  3  rows  of  bristles  (Fig.  21).  Foissner  (1977/78) 
observed  only  a  file-shaped  structure  there,  most  probably  suggested  by  the  single  row  of  paired 
brosse-bristles. 

Cytostome  more  or  less  curved,  anteriorly  overlapping  to  the  left  side  but  not  to  the  right  as 
described  by  Lepsi  (\926a, b,  1928).  Perioral  kinety  1  left  of  cytostome,  with  paired  basal  bodies 
along  the  mouth,  however,  only  the  anterior  basal  body  each  bearing  cilia  of  type  2.  Perioral  kinety 
2  and  3  to  the  right  of  the  oral  slit  showing  closely  spaced  basal  bodies  and  constituting  the  so-called 
'mane',  a  conspicuous  compact  ciliature.  Perioral  kinety  2  with  paired  basal  bodies  along  the  oral 
slit,  the  anterior  basal  body  bears  a  cilium  of  type  1 .  This  kinety  appears  unciliated  post-orally. 
Perioral  kinety  3  with  single  basal  bodies  but  ciliated  along  the  whole  body  with  cilia  type  1. 
Horvath  &  Kuhn  (1941)  misinterpreted  the  perioral  kinetics  2  and  3  as  left  and  right  perioral 
kinetics.  Their  drawing,  however,  shows  the  correct  situation,  that  is  to  say  also  perioral  kinety  1 
(Figs  43,  44).  Fryd-Versavel  et  al.  (1975)  overlooked  the  perioral  kinety  3  (Figs  45-47). 

The  silverline  system  is  a  linearly  orientated  fine-meshed  lattice  (Foissner,  1977/78)  (Fig.  50a). 

OCCURRENCE  AND  ECOLOGY.  Dujardin  (1841)  found  this  species  in  a  20-day-old  infusion  of  material 
from  the  Mediterranean  Sea.  Later  it  was  recorded  from  the  brackish  waters  of  Oldesloe  and  Kiel 
(Kahl,  1928, 1933),  from  the  Roumanian  littoral  of  the  Black  Sea  (Lepsi,  \926a,b,  1928;Tucolesco, 
1962a)  and  from  the  periphyton  of  brackish  and  marine  waters  of  Konigshafen  near  List  (Sylt, 
Germany)  (Kiisters,  1974). 

Some  authors  mentioned  also  terrestrial  habitats  (Radu  &  Tomescu,  1972;  Tomescu,  1978),  but 
a  reliable  record  is  not  available  (Foissner,  1987).  The  drawing  made  by  Stella  (1948),  who  claimed 
to  have  found  Acineria  incurvata  in  a  pine  forest,  indicates  that  it  was  (probably)  a  member  of  the 
genus  Spathidium  (Fig.  22). 

Acineria  incurvata  has  been  frequently  found  in  strongly  saprobic  freshwater  habitats,  such  as 
different  sewage-loaded  watercourses  (Horvath  &  Kuhn,  1941;  Buck,  1961;  Bick,  1972;  Madoni  & 
Ghetti,  1977;  Foissner,  1977/78),  in  Sphaerotilus  tufts  (Vasicek,  1964;  Struhal,  1969),  on  the 
bottom  of  the  river  Elbe  upstream  from  Hamburg  (Grimm,  1968),  in  a  cesspool  (Kahl,  1926),  and 
in  sewage-treatment  plants  (trickling  filters  in  good  working  order,  aeration  tanks)  (Buck,  1961; 
Weninger,  1971;  Madoni,  1981).  Fryd-Versavel  et  al.  (1975)  found  their  'Litonotus  lamella'  in  a 
pond  in  the  year  1962.  Sramek-Husek  (1956,  1958)  noted  it  as  a  true  member  of  the  'Colpidietum 
colpodae'.  Weninger  (1971)  found  a  decreasing  abundance  when  nitrate  or  ammonium  was  added 
to  sewage,  whereas  phosphate  strongly  increased  its  number. 

The  above  data  suggest  that  Acineria  incurvata  is  a  widely  distributed  poly  saprobic  eury  hyaline 
indicator  species  with  a  rather  high  tolerance  of  lack  of  oxygen  and  high  concentrations  of  NH  4. 


Figs  1-23     Acineria. 

Figs  1, 2     Acineria  incurvata  after  Dujardin  (1841). 

Fig.  3     Acineria  acuta  after  Dujardin  (1841). 

Fig.  4    Acineria  nasuta  after  Lepsi  ( 1 962). 

Figs  5-11     Acineria  uncinata  after  Tucolesco  (\962a).  5  Anterior  pole.  6  Posterior  pole,  7,  8  Right  and 
left  side.  9  Mouth  and  anterior  pole  overlapping  towards  the  left  side.  10  Ventral  view.  11  Dorsal  view. 

Figs  12-23  Acineria  incurvata.  12-15  After  Maupas  (1883).  12  An  individual  swallowing  an  Uronema. 
13  Right  side  (Maupas  called  it  dorsal  view).  14  Left  side  (Maupas  called  it  ventral  view).  15  Pellicle.  16 
After  Kahl  (1931),  left  side.  17-21  After  Kahl  (1926).  17, 18  Left  side  of  different  individuals.  19  Right 
side.  20  Conjugants.  21  Left  anterior  region.  22  After  Stella  (1948),  probably  a  Spathidium.  23  After 
Buck  (1961). 


202  H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 

Table  1     Biometrical  characterization  of  Acineria  incurvata 


Character1 


M       SD      SE      CV     Min    Max     n 


Body,  length 
Body,  width 

Number  of  macronucleus  parts 
Macronucleus  part,  length 
Macronucleus  part,  width 
Number  of  micronuclei 
Micronucleus,  length 
Micronucleus,  width 
Cytostome,  length  (measured  as  chord) 
Distance  from  apex  to  posterior  end  of  brosse 
Number  of  brosse-bristles 
Brosse-bristles,  maximal  length 
Number  of  left  perioral  kinetics 
Number  of  right  perioral  kinetics 
Number  of  normally  ciliated  kinetics  (cilia  type  1), 
perioral  kinetics  excluded 


56-25    54-5    7-50    1-67    13-3    46-0    75-0    20 
15-50     16-0    2-01    0-45    13-0    12-0    19-0    20 


2-00 
9-65 
7-35 
1-00 
2-42 
2-12 
22-55 
16-90 
41-20 
1-72 
1-00 
2-00 

2-0 
10-0 
7-5 
1-0 
2-2 
2-0 
22-0 
19-5 
40-0 
1-8 
1-0 
2-0 

0-00 
1-60 
0-83 
0-00 
0-66 
0-47 
3-50 
2-66 
4-18 
0-24 
0-00 
0-00 

0-00 
0-36 
0-18 
0-00 
0-15 
0-11 
0-78 
0-60 
0-93 
0-05 
0-00 
0-00 

0-0 
16-6 
13-3 
0-0 
27-1 
22-3 
15-5 
13-6 
10-1 
14-3 
0-0 
0-0 

2-0 
7-0 
6-0 
1-0 
1-8 
1-6 
15-0 
14-0 
34-0 
1-2 
1-0 
2-0 

2-0 
13-0 
9-0 
1-0 
4-0 
3-6 
28-0 
25-0 
48-0 
2-0 
1-0 
2-0 

20 
20 
20 
20 
20 
20 
20 
20 
20 
20 
20 
20 

10-85     11-0    0-59    0-13      5-4    10-0    12-0    20 


'All  data  are  based  on  protargol  silver  impregnated  specimens.  All  measurements  in  |im.  Legend:  x,  mean;  M,  median;  SD, 
standard  deviation;  SE,  standard  error  of  mean;  CV,  coefficient  of  variation  in  %;  Min,  minimum;  Max,  maximum;  n, 
sample  size. 

REMARKS.  Trachelius  anaticula  Ehrenberg,  1833  is  an  older  but  unreliable  synonym  of  this  species. 
Acineria  acuta  Dujardin,  1841,  which  was  observed  in  the  water  of  a  wheel-track  in  1838,  has  been 
very  insufficiently  described  and  therefore  cannot  be  discriminated  from  Acineria  incurvata.  Thus, 
Acineria  acuta  is  here  treated  as  synonym.  Amphileptus  anaticula  perhaps  is  a  synonym,  too,  but 
the  figure  given  by  Claparede  &  Lachmann  (1859)  shows  an  unidentifiable  individual  with  a 
voluminous  ingestion  vacuole.  The  synonym  Lionotus  reversus  Kahl,  1926  results  par  lapsus,  since 
Kahl  mentioned  in  a  footnote  that  he  had  found  Maupas'  good  description  of  Acineria  incurvata 
just  after  having  finished  the  manuscript.  The  synonym  Litonotus  lamella  results  from  an  obvious 
misidentification  by  Fryd-Versavel  et  al.  (1975). 

Acineria  nasuta  Lepsi,  1962 

DIAGNOSIS  (Fig.  4).  Marine.  About  90-100  urn  long,  rather  wide.  Only  one  single  macronucleus. 
Pellicle  with  5-6  distinct  stripes.  Postapical,  to  the  right  of  the  so-called  'nose'  a  peculiar  line 
(perhaps  the  mouth)  which  is  said  to  be  characteristic  of  this  species. 

OCCURRENCE  AND  ECOLOGY.  Only  a  few  individuals  were  found  in  a  raw  culture  of  putrefying 
marine  algae.  In  the  same  culture  Holophrya  torquabilis  occurred  in  large  numbers,  probably 
serving  as  food  for  Acineria. 


Figs  24-^47  Acineria  incurvata.  24-30  After  Lepsi  (1965).  24  Normal  aspect.  25  Trachelius-\ike  form. 
26  Slender  form,  resembling  Spathidium.  27  Degenerated  individual  resembling  Litonotus.  28-30 
Abnormal,  degenerated  forms.  31^14  After  Horvath  &  Kuhn  (1941).  31-34  Outlines  of  well-fed  and 
starved  specimens.  35  Cyst.  36  Left  side,  with  extrusomes  along  the  cytostome.  37  Right  side,  location 
of  contractile  vacuole  and  of  kinetics.  38-42  Bipartition.  43,  44  Infraciliature  (right  and  left  side) 
revealed  by  Bresslau's  opalblue-technique.  45-47  After  Fryd-Versavel  et  al.  (1975),  misidentified  as 
Litonotus  lamella.  45, 46  Infraciliature  of  left  and  right  side.  47  Diagram  of  different  types  of  cilia  and 
bristles  in  the  anterior  region. 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS 


203 


46 


47 


204 


H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 


49 


Figs  56-58  Acineria  incurvata,  scanning  electron  micrographs.  56  Total  view  of  left  side.  57  Anterior 
part  with  dorsal  oral  region  rolled  up  forming  a  spoon-like  excavation.  Note  the  club-shaped  brosse- 
bristles,  the  short  bristles  of  the  perioral  kinety  1 ,  and  the  long  cilia  of  perioral  kinetics  2  and  3  (arrows). 
58  Detail  of  anterior  third  with  brosse-bristles,  short  bristles  and  normal  cilia  (arrows). 


Figs  48-55  Acineria  incurvata,  originals.  48  Left  side  from  life  and  according  to  scanning  electron 
microscopic  observations,  scale  =  20  um.  49  Extrusome,  length  about  4  um.  50  Reconstructed  cross- 
sections  in  different  regions  of  body.  50a  Silverline  system  in  the  oral  region,  dry  silvered,  after 
Foissner  (1977/78).  51  Right  side,  infraciliature  of  a  protargol  silver  stained  specimen.  P2,  P3,  perioral 
kinetics  2  and  3.  52  Left  ventro-lateral  view  of  a  protargol  silver  stained  specimen  with  different  types 
of  cilia  and  bristles  according  to  SEM-observations.  53, 54  Infraciliature  of  the  left  ventro-lateral  and 
the  right  dorso-lateral  side  of  a  protargol  silver  impregnated  specimen.  Pl-3,  perioral  kinetics  1-3;  Br, 
srale  =  30  um  55  Ventral  view. 


206  H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 

REMARKS.  Lepsi  (1962)  assumed  that  this  species,  which  has  remained  unmentioned  since 
original  description,  could  be  a  form  of  A.  incurvata  and  mentioned  some  relationship  with  the 
genera  Chilophrya  and  Plagiocampa.  His  figure  and  description  are  so  incomplete  that  it  is  at 
present  impossible  to  find  any  reliable  affinity.  The  single  macronucleus  suggests  that  it  is  not  an 
Amphileptidae,  although  he  could  have  observed  a  dividing  stage  with  fused  macronucleus. 

Acineria  uncinata  Tucolesco,  19620 

DIAGNOSIS  (Figs  5-1 1).  Brackish  and  freshwater.  About  35-55  urn.  Body  lanceolate  without  lateral 
edge.  Anterior  pole  overlapping  towards  the  left  side.  Two  spherical  macronuclei  showing  a  clearer 
zone  at  their  central  region.  Sometimes  a  single,  elongated,  tapered  nucleus.  Contractile  vacuole 
terminal,  often  surrounded  by  a  group  of  smaller  vacuoles.  Cytostome  a  straight  and  short  slit 
restricted  to  the  rolled  up  anterior  pole.  Can  therefore  feed  only  on  small  prey  (flagellates).  Three 
somatic  kinetics  on  the  right  side  with  20-22  cilia  each.  Cilia  at  the  ventral  margin  of  the  anterior 
third  transformed  to  regularly  curved  crotchets. 

OCCURRENCE  AND  ECOLOGY.  This  species  was  found  in  summer  1954  in  a  small  dirty  brackish 
puddle  near  Lake  Tekirghiol  and  in  mesosaprobic  freshwaters  of  Bucarest. 

REMARKS.  Tucolesco  (1962a)  separated  this  species  from  A.  incurvata  by  the  non-overlapping 
post-oral  dorsal  margin.  However,  in  A  incurvata  the  situation  is  rather  similar  (page  199).  Thus, 
we  propose  the  following  characters  for  discrimination  from  A.  incurvata:  the  presence  of  only 
three  somatic  kinetics  on  the  right  side,  the  (probably)  unciliated  left  side,  and  the  short  oral  slit 
being  restricted  to  the  anterior  pole.  Unmentioned  since  description.  Note  after  proof  reading: 
This  is  a  valid  species  which  we  rediscovered  recently!  Redescription  is  in  preparation. 


Genus  TR1MYEMA  Lackey,  1925 
Sciadostoma  Kahl,  1926 

DIAGNOSIS.  Trimyemidae  Kahl,  1933  (syn.  Sciadostomatidae  Kahl,  1926)  with  vestibulum  and 
cytostome  near  apical  end.  Vestibular  ciliature  consisting  of  three  rows  of  cilia,  two  rather  long 
ones  arranged  approximately  in  a  semicircle  at  the  left  margin  of  the  vestibulum  and  an  inner  rather 
short  third  row  located  near  the  cytostome  at  the  posterior  left  of  the  vestibulum.  Somatic  ciliature 
in  longitudinal  kineties  but  arranged  in  a  way  that  a  more  or  less  wide  band  of  oblique  spirals  is 
formed.  Prominent  caudal  cilium.  Body  small,  mostly  tapered  at  both  ends.  Free-living  and 
endocommensally,  freshwater  and  marine,  polysaprobic. 

TYPE-SPECIES.  Trimyema  compressa  Lackey,  1925 

REMARKS.  There  is  much  confusion  about  the  exact  orientation  of  the  cell:  dorsal,  lateral,  and 
ventral  sides  are  often  mixed  up  in  descriptions.  In  addition  some  authors  have  given  incorrect 
figures  focusing  the  microscope  on  the  lower  surface  of  their  specimens.  Thus,  they  attained 
inverted  figures  (see  explanations  to  figures).  Most  species  of  the  genus  Trimyema  are  only  super- 
ficially described.  The  oral  structures  are  known  exactly  only  of  T.  compressa  (Figs  83,  107)  and 


Figs  59-76  Trimyema  compressa.  59  After  Lackey  ( 1 925)  (inverted).  60  After  Wang  &  Nie  ( 1 935),  left 
lateral  view.  61-65  After  Liebmann  (1936).  61  Left  ventro-lateral  view.  62-65  Defecation  by  the  aid  of 
the  caudal  cilium.  66-69  After  Kahl  (1926).  66  Left  lateral  view.  67  Oral  region  during  progressed 
bipartition.  68  Ventral  side.  69  Bipartition.  70  After  Kahl  (1931),  constant  marine  form,  rather  similar 
to  T.  claviformis  described  later.  71  After  Kahl  (1933),  left  lateral  view.  72  After  Pennak  (1953) 
(inverted).  73  After  Bick  ( 1 972),  left  lateral  view.  74  After  Sladecek  ( 1 972),  left  lateral  view.  75, 76  After 
Schmall  (1976),  infraciliature  of  protargol  silver  stained  specimens  (inverted),  ventral  and  dorsal  view 
(Schmall  called  it  dorsal  and  ventral  view). 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS 


207 


76 


208  H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 

partly  of  T.  pleurispiralis  (Fig.  96)  and  T.  echinometrae  (Fig.  93).  From  the  descriptions  and  our 
investigations  we  deduced  the  basic  structure  of  the  oral  apparatus  as  described  above  (compare 
Fig.  1 07).  Faure-Fremiet  ( 1 962)  and  Borror  ( 1 972)  obviously  overlooked  the  short  third  vestibular 
kinety.  Borror  (1972)  described  only  an  inner  and  an  outer  'polykinety'.  Detcheva  el  al.  (1981), 
however,  showed  in  T.  compressa  electronmicroscopically  that,  despite  their  polykinetal  appear- 
ance, the  vestibular  ciliary  systems  are  not  separate  polykineties  but  are  the  anterior  parts  of  the 
somatic  kinetics  that  are  preceded  by  parasomal  sacs  and  retain  the  same  fibrillar  systems  as  the 
somatic  kinetosomes.  Jankowski  (\964a, b)  gave  no  evidence  for  his  statement  that  there  were  four 
vestibular  kinetics  in  T.  compressa. 

Encystment  is  unknown  in  this  genus.  Czapik  (19750)  noted  that  even  starved  specimens  (of  T. 
compressa)  die  without  forming  cysts.  Morphogenesis  has  not  yet  been  exactly  studied.  However, 
the  oral  apparatus  is  supposed  to  reduce  before  cell  division,  because  during  division  both  proter 
and  opisthe  show  the  same  state  of  development  of  the  oral  apparatus  (Kahl,  1926)  (Figs  67,  69). 

The  silverline  system  has  been  demonstrated  only  in  T.  compressa  (Klein,  1930;  Faure-Fremiet, 
1962;  Jankowski,  \964a, b;  Czapik,  19750).  Klein  (1930)  gave  the  description  that  best  agrees  with 
our  observations  (Figs  106,  1 10).  But  he  did  not  draw  the  transverse  silver  lines  connecting  the 
longitudinal  lines  in  the  region  of  the  ciliary  spirals.  The  granules  located  at  and  in  the  silverlines 
(Fig.  1 10)  have  been  said  to  be  mucocysts  ('Relationskorner')  or  rudimentary  basal  bodies  (Klein, 
1930).  However,  the  electronmicroscopic  investigation  shows  only  mucocysts  (Detcheva  et  al., 
1981). 

The  exact  taxonomic  position  of  the  genus  is  still  unclear.  Kahl  (1926)  created  a  new  family  for 
the  rather  special  helical  ciliature.  This  author,  Corliss  (1979),  and  Curds  (1982)  included  the 
family  in  the  order  Trichostomatida  Biitschli,  1889.  Faure-Fremiet  (1962)  noted  that  the  family 
Trimyemidae  indeed  presents  one  of  numerous  possibilities  existing  in  the  order  to  use  the  anterior- 
most  somatic  kinetics  for  building  up  a  vestibular  ciliature.  In  addition,  he  indicated  possible 
affinities  of  Trimyema  with  Mycterothrix  and  Maryna,  which  are  now  'good'  colpodids  (Foissner, 
1985a).  Jankowski  (1980)  erected  the  new  order  Trimyemida  (incertae  sedis)  giving  no  reasons  for 
this  decision.  On  the  contrary,  Detcheva  et  al.  (1981)  stated  that  Trimyema  is  a  member  of  the 
Vestibulifera  and  that  the  Trimyemidae  show  the  same  general  type  of  vestibular  architecture  as 
the  Plagiopylidae  and  the  Coelosomidae.  However,  a  more  reasonable  classification  demands 
further  investigations  especially  on  the  morphogenetic  processes. 

Ruinen  (1938)  is  wrong  in  transferring  Palmarium  salinum  Gajevskaja,  1925  to  the  genus 
Trimyema,  since  Palmarium  is  illustrated  as  having  an  adoral  zone  of  membranelles  (Figs  97-101) 
(Borror,  1972). 

Trimyema  pur  a  (Ehrenberg)  is  listed  by  Curds  (1975)  as  a  species  occurring  in  percolating 
filters  and  in  activated  sludge.  We  suppose  that  this  species  has  been  described  as  Trichoda  pura 
Ehrenberg,  1831,  which  according  to  Corliss  &  Dougherty  (1967)  is  a  synonym  of  Tetrahymena 
pyriformis. 

Lackey  (1925)  classified  Trimyema  as  female  using  the  latin  ending  -a  for  his  species  T. 
compressa.  Since  we  could  not  find  any  greek  word  comparable  to  'myema'  from  which  the  name  of 
the  genus  and  its  sex  could  be  derived  we  accept  Lackey's  proposal  of  the  sex.  This,  however, 
requires  the  endings  of  T.  claviforme,  T.  marinum,  T.  minutum,  and  T.  pleurispirale  to  be  emended 
(see  below). 


Key  to  the  species 

la  3  somatic  ciliary  spirals 2 

Ib  Usually  more  than  3  somatic  ciliary  spirals 3 

2a  Posterior  end  of  body  tapered,  length  25-65  um T.  compressa 

2b  Posterior  end  of  body  broadly  rounded,  prominent  beak-like  pharynx  opening,  length  c.  20  um  . 

.   T.  minuta 

3a  Body  broadly  oval,  width  c.  half  length  of  body 4 

3b  Body  rather  slender,  fusiform  or  oblong,  width  much  less  than  half  length  of  body      ...         5 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCH1LIOPSIS  209 

4a    4(-6)  somatic  ciliary  spirals  restricted  to  the  anterior  half  of  body,  length  c.  20-45  um . 

T.pleurispiralis 

4b    7  somatic  ciliary  spirals  restricted  to  the  anterior  half  of  body,  length  c.  25^0  jam,  endocommen- 

sally  in  sea-urchins T.  echinometrae 

5a     Body  club-shaped,  thickened  in  the  anterior  region  and  slender  in  the  posterior  region,  length  c. 

40  urn,  (not  totally  reliable  species!) T.  claviformis 

5b    Body  not  club-shaped       .        .  6 

6a    Shape  of  body  obviously  asymmetric,  tapered  at  both  ends,  anterior  pole  bent  to  the  right, 

posterior  pole  bent  to  the  left,  peristome  measures  c.  one  third  of  cell  length        .        .      T.kahli 

6b    Shape  of  body  symmetrical,  slender  fusiform  to  slender  oblong 7 

7a     Body  length  c.  40  urn,  peristome  measures  c.  one  fourth  of  body T.  marina 

7b    Body  length  c.  60  um,  peristome  measures  less  than  one  fourth  of  body  (not  totally  reliable 

species!)  ..............     T.  alfredkahti 


Descriptions  of  species 
Trimyema  compressa  Lackey,  1925 

Sciadostoma  difficile  Kahl,  1926 
Trimyema  compressum  Kahl,  1933 
Trimyema  marinum  Faure-Fremiet,  1962 

NEOTYPE-SPECIMENS.  Slides  (dry  silvered  and  protargol  silver  impregnated)  of  neotype-specimens 
have  been  deposited  in  the  British  Museum  (Natural  History)  in  London,  reference  numbers 
1986:5:30:2-3. 

REDESCRIPTION  (Figs  59-86,  104-1 13,  Table  2).  Type  species  of  the  genus.  Freshwater  and  marine. 
In  vivo  about  25-50(-60)  x  15-20(-35)  urn  (Lackey,  1925;  Kahl,  1926,  1928,  1931,  1933;  Wang  & 
Nie,  1935;  Liebmann,  1936;Czapik,  1975a;  Schmall,  1976;  Detcheva  6>f0/.,  1981).  Body  fusiform  to 
plump  S-shaped,  laterally  slightly  flattened,  anterior  and  posterior  end  slightly  tapered.  Dorsally 
and  ventrally  an  inconspicuous  ectoplasmatic  ridge,  the  so-called  keel  (unrecognized  by  us)  (Kahl, 
1926;  Wang  &  Nie,  1935).  Macronucleus  spherical  to  slightly  oval,  located  centrally  in  most 
specimens.  Schmall  (1976)  found  it  to  be  more  variable,  also  located  posteriorly.  Micronucleus 
closely  attached  to  the  macronucleus.  In  protargol  impregnated  specimens  often  a  second,  weakly 
stained  macronucleus-like  structure,  probably  a  large  ingestion  vacuole  (Fig.  109).  Macronucleus 
usually  heavily  stained,  surrounded  by  dark,  slightly  curved  rods  measuring  c.  2  um  in  length  and 
0-5  um  in  width.  These  aggregated  rods  look  like  bacteria.  Detcheva  et  al.  (1981),  however, 
consider  them  to  be  mitochondria,  which  is  not  supported  by  recent  studies  on  other  sapropelic 
ciliates  (Van  Bruggen  et  al.,  1984).  Contractile  vacuole  and  its  pore  located  in  the  region  of  the  last 
ciliary  spiral  on  the  right  ventro-lateral  side.  Cytoproct  a  slit  circa  5-10  um  long,  located  in  the 
right  dorso-lateral  surface  (Figs  104,  105,  108).  Pellicle  thin,  flexible  and  deformable,  with  very 
slight  ridges  paralleling  the  longitudinal  kinetics.  In  protargol  impregnated  specimens  these  ridges 
appear  darkly  stained  and  produce  a  negative  image  of  the  silverline  system.  Cytoplasm  rather 
transparent,  contains  a  lot  of  refractive  long-oval  (length  about  0-5-1-5  um)  granules  which  are 
also  visible  in  protargol  stained  specimens.  They  are  most  probably  the  mucocysts  described  by 
Detcheva  et  al.  (1981).  Food  vacuoles  about  5  um  in  diameter.  Cyclosis  pronounced  (Lackey, 
1925).  Feeds  on  bacteria  but  is  not  dependent  on  sulphur  bacteria  (Liebmann,  1947).  Moves  slowly 
and  slightly  tremblingly  in  a  straight  line  or  in  the  arc  of  a  circle  rotating  on  its  longitudinal  axis 
(Lackey,  1925).  Reproduction  by  transverse  fission  (Lackey,  1925;  Kahl,  1926,  1931;  Czapik, 
1975a). 

Somatic  cilia  7-9  um,  strongly  beating,  arranged  in  about  50-60  longitudinal  kinetics  but  more 
commonly  viewed  as  3  oblique  spirals.  In  the  anterior  region  of  these  spirals  the  third,  fourth,  and 
fifth  kinetosomes  are  paired,  constituting  the  compact  field  of  cilia,  consisting  of  3  x  4  and  2x2 
cilia,  described  by  Schmall  (1976)  (Figs  75, 1 13).  A  short  row  of  about  5-10  cilia  on  the  ventral  side 
extends  obliquely  from  the  posterior  end  of  the  anteriormost  somatic  spiral  to  the  right.  Posterior 


Figs  77-86  Trimyema  compressa.  77-79  After  Jankowski  ( 1 964).  77  From  life  (inverted).  78  Left  lateral 
view  of  a  dry  silver  impregnated  specimen.  79  Lateral  view  of  a  dry  silvered  specimen  (inverted).  80-82 
After  Czapik  (1975a),  specimens  stained  by  Chatton's  method  as  modified  by  Corliss.  80  Scheme 
representing  the  disposition  of  the  ciliary  spirals.  81  Ventro-apical  region.  82  Ventral  side  (Czapik 
called  it  right  side).  83  After  Detcheva  et  al.  ( 1 98 1 ),  dorsal  view  (inverted).  84-86  After  Faure-Fremiet 
(1962)  who  identified  it  erroneously  as  T.  marina.  84  From  life.  85  Apical  view  of  Chatton-Lwoff 
impregnated  specimen.  86.  Infraciliature  and  silver  lines  of  left  dorso-lateral  side  of  a  Chatton-Lwoff 
impregnated  specimen  (Faure-Fremiet  interpreted  it  as  left  ventro-lateral  view). 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS 


211 


95 


212  H.  AUGUSTIN,  W.  FO1SSNER  &  H.  ADAM 

third  of  body  unciliated  apart  from  the  caudal  cilium  measuring  about  one  third  to  one  half  of  body 
length  (Lackey,  1925;  Kahl,  1931;  Wang  &  Nie,  1935);  it  is  perhaps  involved  in  the  process  of 
defecation  (Liebmann,  1936)  (Figs  62-65). 

Vestibulum  circa  one  third  of  body  length,  funnel-shaped.  Left  half  of  the  oral  depression  more 
excavated  than  the  right  one  and,  as  a  consequence,  the  left  margin  becomes  a  thin,  transparent 
layer  of  ectoplasm  and  forms  a  cap  or  hood-like  process  bordering  the  vestibulum  (Kahl,  1926; 
Wang  &  Nie,  1935).  Cytopharyngeal  fibres  inconspicuous,  rectangular  to  the  entrance  of  the 
vestibulum.  Vestibular  kinety  1  a  bit  longer  than  vestibular  kinety  2.  At  their  anterior  ends  4  to  5 
pairs  of  basal  bodies  or  single  basal  bodies  with  parasomal  sacs.  Vestibular  kinety  3  consists  of  only 
6-7  cilia  (Figs  107,  111,  112).  In  stained  specimens  somatic  as  well  as  vestibular  kinetosomes 
appear  to  be  paired  (Figs  106-1 1 3)  but  in  fact,  the  anterior  granule  is  a  parasomal  sac  (Detcheva  et 
al.,  1981),  probably  with  the  exception  of  the  above  mentioned  compact  field. 

About  60  longitudinal  silver  lines  (Czapik,  1975a  mentioned  52  lines),  connected  by  transverse 
lines  which  are  located  between  the  somatic  ciliary  spirals.  In  front  of  the  anteriormost  ciliary  spiral 
a  circumoral  silver  line  from  which  a  few  longitudinal  lines  extend  to  the  vestibulum  forming 
square-like  fields  at  its  rim.  The  longitudinal  silver  lines  fuse  at  the  posterior  third  forming  rough 
meshes  (Figs  106,  110). 

OCCURRENCE  AND  ECOLOGY.  First  recorded  from  the  sewage  disposal  of  Imhoff  tanks  in  New  Jersey 
and  later  listed  as  an  obligate  anaerobe  (Lackey,  1925,1938;  Noland  &  Gojdics,  1 967).  Very  similar 
habitats  were  reported  by  Liebmann  (1936,  1947,  1951),  who  found  T.  compressa  regularly  in 
waters  containing  a  lot  of  organic  matter  and  H2S,  such  as  in  over-loaded  percolating  filters,  in 
Imhoff  tanks  (S-Sind.ml"1  and  40  hid. ml"1),  in  sewers,  and  at  the  outfalls  of  communal  waste 
waters. 

Further  habitats  are  the  sapropel  of  ponds  near  Leningrad  (Jankowski,  \964a,b),  ponds  used  for 
the  treatment  of  sugar  factory  wastes  (Grabacka,  1973),  the  plankton  of  the  eutrophic  pond 
Toppelsdorfer  Weiher'  in  Bonn  (Wilbert,  1969),  a  small  eutrophic  lake  at  Uttendorf/Salzburg 
(Foissner,  unpublished),  and  an  arctic  tundrapond  at  Barrow/Alaska  (Fenchel,  1975).  Detcheva 
( 1 972)  and  Czapik  ( 1 915a,b)  listed  up  Bulgarian  and  Polish  habitats  like  ponds,  lakes,  ditches,  and 
polluted  rivers.  Wang  &  Nie  (1935)  observed  some  individuals  among  decaying  organic  substances 
taken  from  Lake  Ho  Hu.  Kahl  ( 1 926, 1931,1933)  found  it  in  the  sapropel,  in  a  cesspool,  in  sewage, 
and  more  rarely  in  the  brackish  waters  of  Oldesloe  (Kahl,  1928)  thus  considering  it  to  be  of 
freshwater  origin.  Faure-Fremiet  (1962)  found  it  in  a  rock  pool  on  the  French  Atlantic  coast. 
Tucolesco  (\962b)  recorded  it  from  the  Black  Sea  and  from  the  saliferous,  para-marine  Lake 
Tekirghiol  in  Roumania.  According  to  Sladecek  (1972)  T.  compressa  developed  in  great  numbers 
(up  to  10,000  ind.ml"  *)  in  a  sample  of  industrial  waste  water  from  a  textile  factory. 


Figs  87-103     Trimyema. 

Figs  87-89     Trimyema  marina.  87, 88  After  Kahl  (1933).  89  After  Kahl  (1931). 

Figs  90, 91     Trimyema  minuta  after  Kahl  (1931),  dorsal  and  left  lateral  view. 

Fig.  92     Trimyema  claviformis  after  Kahl  ( 1 933). 

Figs  93, 94     Trimyema  echinometrae  after  Groliere  et  al.  ( 1 980),  protargol  silver  impregnated  specimens 
(inverted),  lateral  view  and  ventral  view  (the  latter  designated  as  lateral  view,  too). 

Figs  95,  96     Trimyema  pleurispiralis  after  Borror  (1972).  95  Left  ventro-lateral  view.  96  Anterior  pole 
with  anteriormost  somatic  ciliary  spiral  (at  the  left)  and  with  vestibular  ciliature. 

Figs  97-101     Palmarium  salinum  Gajevskaja  after  Ruinen  (1938).  97,  98  Normal  form,  ventral  and 
dorsal  view.  99  Slender  (abnormal?)  form.  100, 101  Peristome,  ventral  and  lateral  view. 

Fig.  102     Trimyema  alfredkahli  after  Tucolesco  ( 1 962a),  left  lateral  view. 
Fig.  103     Trimyema  kahli  after  Tucolesco  ( 1 962o),  left  lateral  view. 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS 


213 


107 


Figs  104-109  Trimyema  compressa,  originals,  scale  =  20  um  each.  104  Left  ventro-lateral  view,  from 
life.  105  Right  dorso-lateral  view  of  an  S-shaped  individual.  CP,  cytoproct.  106  Dorsal  view  of  a  dry 
silvered  specimen.  107  Vestibular  ciliature  revealed  by  Fernandez-Galiano's  method.  The  shape  of  the 
vestibular  kinetics  has  been  slightly  deformed  by  preparation;  they  are  less  curved  in  life.  Vl-3, 
vestibular  kinetics  1-3.  108,  109  Ventral  and  dorsal  view  of  a  protargol  silver  impregnated  specimen 
amended  with  details  from  individuals  impregnated  with  Fernandez-Galiano's  method.  CVP, 
contractile  vacuole  pore. 


214 


H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 


^^Y**  l**^s.  '  V 

X.    •*  *»i*    ^T^k-  •    •     ' 


110 


111 


•••*•»  -^  *» .  •  *  * 

. 


112 


113 


Figs  110-113  Trimyema  compressa.  110  Silverline  system  revealed  by  the  dry  silver  impregnation 
technique,  dorsal  view.  111-113  Specimens  stained  by  Fernandez-Galiano's  impregnation  technique. 
Ill  Left  side  with  the  three  ciliary  spirals  and  apex  with  vestibular  ciliature  consisting  of  two  long  rows 
and  one  short  row  of  cilia  (arrow).  112  Dorsal  view,  arrow  indicates  the  three  vestibular  kinetics.  113 
Ventral  view,  arrow  indicates  the  isolated  basal  bodies  at  the  anteriormost  region  of  the  vestibular 
kinetics. 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS  2 1  5 

Bick  (1968,  1972)  gave  the  most  detailed  ecological  characterization.  T.  compressa  is  an 
outstanding  indicator  of  polysaprobity  (Liebmann,  1951),  isosaprobity  and  even  metasaprobity 
(Sladecek,  1973)  and  occurs  in  waters  receiving  fresh  manure  and  sewage,  or  waste  waters  contain- 
ing cellulose  material  (paper  mill  outlets,  etc.).  The  species  seems  to  prefer  conditions  with  low 
ammonia  content,  i.e.  conditions  prevailing  during  the  decay  of  cellulose  and  other  material  poor 
in  nitrogenous  compounds.  The  saprobiological  evaluation  is  indicated  by  Sladecek  (1972):  x  =  0, 
o  =  0,  0  =  0,  a  =  0,  p=  10,  G  =  5,  s  =  5-3  (E,  H2S). 

REMARKS.  This  species  differs  from  T.  minuta  particularly  by  the  tapered  posterior  end.  It  can  easily 
be  distinguished  from  the  other  species  by  its  having  only  three  somatic  ciliary  spirals.  Faure- 
Fremiet  (1962)  observed  an  abundant  population  of  Trimyema  (Figs  84-86)  and  identified  it  as  T. 
marina  although  it  was  of  an  ovoid  and  stocky  form  which  was  not  described  by  Kahl  (1931,  1933, 
1935).  On  the  contrary  this  author  later  stated  that  T.  marina  is  usually  one  third  to  one  half  more 
slender  than  he  drew  it  in  1 93 1  (Kahl,  1931,  Fig.  89;  Kahl,  1933,  Figs  87,88).  Thus  we  suppose  that 
Faure-Fremiet  worked  on  T.  compressa. 

Table  2     Biometrical  characterization  of  Trimyema  compressa 

Character1  x         M      SD      SE      CV     Min    Max     n 


Body,  length 

39-05 

39 

5 

4-58 

1-03 

11-7 

32-0 

47-0 

20 

Body,  width 

22' 

30 

23' 

0 

3-01 

0-67 

13-5 

17-0 

26-0 

20 

Macronucleus,  length 

11 

05 

11 

0 

1-57 

0-35 

14-2 

9 

•0 

14-0 

20 

Macronucleus,  width 

9 

35 

9 

'5 

1-50 

0-34 

16-0 

6-0 

12-0 

20 

Number  of  vestibular  ciliary  rows 

3' 

00 

3 

0 

0-00 

0-00 

0-0 

3 

0 

3-0 

20 

Number  of  somatic  ciliary  rows 

3 

00 

3 

0 

0-00 

0-00 

0-0 

3 

0 

3-0 

20 

Number  of  caudal  cilia 

1 

00 

1 

0 

0-00 

0-00 

0-0 

1 

0 

1-0 

20 

Distance  from  apex  to  posterior  end  of  vestibulum 

9 

80 

10-0 

1-88 

0-42 

19-2 

7 

•0 

15-0 

20 

Distance  between  posterior  end  of  body  and  posterior 

end  of  ciliary  spirals 

11 

00 

11 

0 

1-78 

0-40 

16-2 

7 

•0 

15-0 

20 

1  See  footnote  Table  1 


Trimyema  a#ra/£«M  Tucolesco,  \962a 

DIAGNOSIS  (Fig.  102).  Marine.  About  60  jim.  Body  oblong  and  slender,  slightly  tapering  anteriorly 
and  posteriorly.  Oral  apparatus  particularly  small,  bounded  at  the  right  margin  by  a  conspicuous 
dilatation.  Macronucleus  spherical.  Cilia  long  and  fine.  According  to  Tucolesco's  figure  ciliary 
spirals  cover  nearly  the  whole  body,  which  contrasts  his  description.  Caudal  cilium  longer  than  half 
body  length. 

OCCURRENCE  AND  ECOLOGY.  Found  in  an  abundant  population  in  a  mixed  polysaprobic  culture 
taken  from  the  Black  Sea  in  March  1955. 

REMARKS.  This  species  has  remained  unmentioned  since  description.  It  can  perhaps  be  dis- 
tinguished from  T.  marina  by  its  oblique  orientation  of  the  oral  apparatus,  which  is  stressed  by 
Tucolesco  (1962#),  and  by  its  larger  size.  However,  synonymy  cannot  be  excluded. 

Trimyema  claviformis  Kahl,  1933 
Trimyema  claviforme  Kahl,  1933 

DIAGNOSIS  (Fig.  92).  Marine.  Circa  40  urn.  Body  club-shaped.  Posterior  third  of  body  unciliated. 
OCCURRENCE  AND  ECOLOGY.  Found  in  sapropelic  habitats  of  Sylt  and  Kiel  (Germany). 


216  H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 

REMARKS.  Very  insufficiently  described.  With  exception  of  the  unciliated  tapering  posterior  third  of 
body  identical  with  T.  marina.  Even  Kahl  (1935)  noted  that  he  established  this  species  with  some 
doubt.  Thus,  synonymy  cannot  be  excluded. 

Trimyema  echinometrae  Groliere,  Puy torac  &  Grain,  1 980 

DIAGNOSIS  (Figs  93,  94).  Marine.  Living  endocommensally  in  sea-urchins.  About  31  (27-40)  x  17 
(1 3-20)  urn.  Body  peg-top  like.  Macronucleus  spherical,  5-7-5  urn  in  diameter,  posteriorly  located. 
Micronucleus  not  visible.  60  to  70  longitudinal  somatic  kinetics.  Cilia  distributed  in  7  parallel 
spirals  in  the  anterior  half  of  body.  Three  vestibular  kinetics  very  similarly  arranged  as  in  T. 
compressa. 

OCCURRENCE  AND  ECOLOGY.  Found  in  the  sea-urchins  Diadema  antillarum  and  Echinometra 
lucunter  from  the  Gulf  of  Mexico  and  the  Gulf  of  Guadeloupe.  Housing  together  with  other 
commensal  species  like  Biggaria  echinometris,  Metanophrys  elongata  and  Metopus  circumlabens 
(Groliere  et  al.,  1980).  Perhaps  already  Profant  (1966)  observed  this  species,  since  he  mentioned 
Trimyema  sp.  to  be  a  ciliate  inhabiting  echinoids  in  the  Eastern  Pacific  Ocean. 

REMARKS.  T.  echinometrae  is  a  reliable  species.  It  differs  from  the  other  members  of  the  genus  in  the 
number  of  ciliary  spirals.  The  figures,  however,  are  obviously  inverted,  because  in  the  genus 
Trimyema  the  spirals  run  the  other  way  round.  Furthermore,  the  identification  is  impeded  by  the 
missing  drawing  from  life. 

Trimyema  kahli Tucolesco,  \962a 

DIAGNOSIS  (Fig.  103).  Para-marine.  About  36-40  urn.  Body  conspicuously  asymmetric,  inverted 
S-shaped.  Peristome  in  the  anterior  third  of  body.  Macronucleus  spherical,  usually  located  in  the 
middle  of  the  cell.  Contractile  vacuole  close  behind  the  middle  of  body.  Cilia  long  and  fine.  Ciliary 
spirals  extending  to  the  posterior  pole.  Caudal  cilium  almost  rigid,  bent  to  the  left. 

OCCURRENCE  AND  ECOLOGY.  Polysaprobic,  found  constantly  in  the  para-marine  Roumanian  Lake 
Tekirghiol  (Tucolesco,  \962a,b). 

REMARKS.  This  species  has  remained  unmentioned  since  1962.  However,  from  its  general 
appearance  it  seems  to  be  a  reliable  but  insufficiently  described  species. 

Trimyema  marina  (Kahl,  1931) 

Sciadostoma  marinum  Kahl,  1931 
Trimyema  marinum  Kahl,  1933 

DIAGNOSIS  (Figs  87-89).  Marine.  About  40  urn.  Slender  fusiform  to  slender  oblong  (4 :  1).  In  the 
original  figure  (Fig.  89)  similar  to  T.  compressa  but  later  figured  and  redescribed  with  5-6  ciliary 
spirals  (Figs  87,  88). 

OCCURRENCE  AND  ECOLOGY.  Repeatedly  observed  in  putrid  water  of  the  North  and  East  Sea  (Sylt, 
Kiel)  and  in  salt-water  from  Oldesloe  (Kahl,  1931,  1933,  1935). 

REMARKS.  Kahl  (1931)  considered  T.  marina  to  be  a  separable  species  because  he  never  found 
similar  forms  among  numerous  populations  of  the  freshwater  form  of  T.  compressa.  Later  he 
thought  that  two  forms  of  this  species  probably  exist  and  erected  the  species  T.  claviformis  (Kahl, 
1933)  which,  however,  is  not  a  totally  reliable  species  (Kahl,  1935).  We  consider  this  species  and 
T.  alfredkahli  perhaps  to  be  junior  synonyms  of  T.  marina. 

Trimyema  minuta  nov.  comb. 
Sciadostoma  minutum  Kahl,  1931 

DIAGNOSIS  (Figs  90,  91).  Freshwater  and  marine.  About  20  um.  Rounded  posterior  and  a 
prominent  beak-like  pharynx-opening.  Ectoplasmatic  ridge  (keel)  more  pronounced  than  in  T. 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS  217 

compressa,  extending  from  the  beak-like  pharynx-opening  over  the  back  to  the  posterior.  Cilia 
longer  and  more  rigid  than  in  T.  compressa. 

OCCURRENCE  AND  ECOLOGY.  This  species  was  found  together  with  T.  compressa  and  was  first 
considered  as  a  modification,  but  once  an  abundant  population  occurred  in  a  ditch  contaminated 
with  liquid  manure  (Kahl,  1931).  Wenzel  (1961)  observed  T.  minuta  in  the  sponge  Halichondria 
panicea  from  the  Gulf  of  Naples.  Tucolesco  (19626)  recorded  it  twice  from  old,  mixed  infusions  of 
the  para-marine  Roumanian  Lake  Tekirghiol. 

REMARKS.  Kahl  (1931)  doubted  the  species  status  of  this  form  and  did  not  mention  it  again  in  his 
publication  of  the  year  1935.  Further  investigations  are  necessary. 

Trimyema  pleurispiralis  Borror,  1972 

DIAGNOSIS  (Figs  95,  96).  Marine.  About  20-44  x  16-23  (usually  less  than  20)  urn.  Shape  of 
prepared  individuals  egg-like,  circular  in  cross  section  (Fig.  95).  Macronucleus  spherical,  central. 
Micronucleus  not  observed.  Cytoproct  an  elongated  (approximately  8  urn)  slit  near  posterior  pole, 
lying  in  the  same  latitude  as  cytostome  and  suture  at  ends  of  ciliary  spirals.  Contractile  vacuole 
pore  not  observed.  Except  for  elongated  caudal  cilium,  all  somatic  cilia  restricted  to  anterior  half 
of  cell,  arranged  in  at  least  four  spirals  (a  few  individuals  possess  a  partial  or  even  complete 
fifth  spiral,  and  even  a  few  cilia  of  a  sixth  spiral).  Outer  vestibular  kinety  in  a  semicircle 
dipping  posteriorly  into  vestibulum  and  terminating  near  cytostome.  Inner  vestibular  kinety  with 
three  regions:  (1)  anteriormost  two  isolated  tufts  of  approximately  five  cilia  each,  (2)  a  row  of 
kinetosomes  closely  paralleling  the  outer  kinety,  extending  from  the  tufts  down  to  cytostome,  (3) 
posteriormost  a  J-shaped  field  of  cilia.  As  already  mentioned,  this  interpretation  of  the  oral 
structure  is  a  little  erroneous  and  incomplete. 

OCCURRENCE  AND  ECOLOGY.  Like  the  other  species  of  this  genus  T.  pleurispiralis  is  bacterivorous 
and  occurred  only  irregularly  in  New  Hampshire  tidal  salt  marshes  (Borror,  1972). 

REMARKS.  This  species  differs  from  the  other  members  of  the  genus  in  number  and  location  of 
ciliary  spirals,  which  are  restricted  to  the  anterior  half  of  body.  Unfortunately,  Borror  (1972)  did 
not  give  a  drawing  from  life.  Thus,  the  real  body  shape  is  unknown.  Redescription  is  needed. 


Genus  TROCHILIOPSIS  Penard,  1922 

DIAGNOSIS.  Microthoracidae  Wrzesniowski,  1870  with  cytostome  in  the  anterior  third  of  body. 
Three  preoral  kinetics  subapically  on  the  left  body  side.  Somatic  kinetics  from  either  side  terminate 
near  the  pointed  beak-like  region  formed  by  the  oral  structures.  Apex  smooth.  Rightmost  somatic 
kinety  of  the  right  side  interrupted.  Contractile  vacuole  located  almost  centrally.  Freshwater, 
polysaprobic. 

TYPE-SPECIES.  TrochiliopsisopacaPenard,  1922. 

REMARKS.  Trochiliopsis  shows  many  characters  which  are  very  likely  homologous  to  genera  of  the 
family  Microthoracidae  Wrzesniowski  1870  according  to  the  classification  of  Foissner  (19856). 
Thus,  a  separation  of  Trochiliopsis  at  the  familial  level  as  suggested  by  Jankowski  (1975)  is  not 
justified  (Compare  Corliss,  1979;  Curds,  1982).  On  the  contrary,  the  organization  of  Trochiliopsis, 
especially  the  general  appearance  of  the  infraciliature  and  the  location  and  structure  of  the  oral 
apparatus,  allows  a  classification  close  to  the  genus  Stammeridium.  These  similarities  might  have 
induced  Kahl  (1931)  to  synonymize  Trochiliopsis  with  Trichopelma  Levander  and  Leptopharynx 
Mermod.  There  are  just  sufficient  differences  in  the  location  of  the  preoral  kinetics,  the  paroral 
membrane,  and  the  shape  of  the  anteriormost  region  for  separating  these  two  genera.  Further- 
more, by  a  trivial  twist  of  some  organelles  of  Trochiliopsis,  the  typical  organization  of  the  genus 
Stammeridium  can  be  achieved  (Figs  126,  127):  The  preoral  kinetics  move  to  the  apex  between 
serrated  processes,  the  paroral  membrane  gets  located  obliquely  to  the  longitudinal  axis  and  the 
contractile  vacuole  moves  close  to  the  ventral  side. 


218  H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 

Key  to  the  genera  of  Microthoracina  Jankowski  1967  (based  on  Foissner  1985&) 

la     Microthoracina  with  somatic  cirri-like  organelles,  fusiform  extrusomes,  and  wide-meshed  silver- 
line  system       ..........        (Discotrichidae)  Discotricha 

1  b    Microthoracina  with  normal  cilia,  anchor-like  extrusomes,  and  granular  or  fine-meshed  silverline 
system     ................. 

2a     Microthoracina  with  more  than  10  uninterrupted  somatic  kinetics 

(Pseudomicrothoracidae)  Pseudomicrothorax 

2b    Microthoracina  with  fewer  than  1 0,  usually  6  partly  interrupted  somatic  and  three  preoral  kinetics 

(Microthoracidae) 

3a    Oral  apparatus  ventrally  in  the  posterior  third  of  body Microthorax 

3b    Other       ....  

4a    Oral  apparatus  ventrally  between  middle  and  posterior  third  of  body,  body  more  or  less  oblong  . 

Drepanomonas 

4b    Other .        . 

5a     Oral  apparatus  between  middle  and  anterior  third  of  body,  rightmost  somatic  kinety  or  right  side 
uninterrupted,  preoral  kinetics  run  in  distinct  furrows  from  the  ventral  to  the  right  body  side  and 

form  a  keel Leptopharynx 

5b    Other       .  ... 

6a     Rightmost  somatic  kinety  of  right  side  interrupted,  preoral  kinetics  run  anterior-posteriorly  on  the 
left  side  of  the  body,  paroral  membrane  circa  half  body  length    ....       Trochlliopsis 

single  species:  Trochiliopsis  opaca 
6b    Preoral  kinetics  apically  in  furrows,  apex  distinctly  serrated,  paroral  membrane  shorter  than  a 

third  of  body  running  obliquely  to  the  longitudinal  axis Stammeridium 

single  species:  Stammeridium  kahli 


Description  of  species 
Trochillopsis  opaca  Penard,  1 922 

Trichopelma  opaca  Kahl,  1931 
Leptopharynx  opaca  Detcheva,  1972 

NEOTYPE-SPECIMENS.  Slides  (protargol  silver  impregnated  and  dry  silvered)  of  neotype-specimens 
have  been  deposited  in  the  British  Museum  (Natural  History)  in  London,  reference  numbers 
1986:5:30:4-5. 

REDESCRIPTION  (Figs  114-131,  Table  3).  Type  species  of  the  genus.  Freshwater.  In  vivo  circa 
30-40(-50)  x  17-20  um.  Body  outline  oval,  anteriorly  curved  slightly  to  the  ventral  side  terminat- 
ing in  a  pointed  beak-like  region  (peak).  Body  strongly  compressed  laterally  (circa  2:1).  Somatic 
kinetics  in  deep,  crenelated  furrows,  which  terminate  near  the  oral  peak.  Macronucleus  spherical, 
more  or  less  centrally  located,  in  vivo  hardly  discernible.  Micronucleus  closely  attached  to  the 
macronucleus.  Contractile  vacuole  centrally  located,  close  to  the  right  lateral  surface,  diameter 
about  4  um;  contractile  vacuole  pore  at  the  end  of  the  paroral  membrane.  Cytoproct  slightly 
posterior  to  the  contractile  vacuole  pore,  visible  as  black  line  in  dry  silvered  specimen  (Fig.  130). 
Pellicle  rigid,  colourless,  opaque.  Extrusomes  about  3  um,  fusiform,  scattered  over  the  whole  body 
in  the  ribs  between  the  furrows,  show  four  anchor-like  processes  at  the  distal  end  in  the  exploded 
phase.  Probably  feeds  on  bacteria,  but  no  food  vacuoles  were  found.  Slow,  trembling  and  swaying 
movements. 

Length  of  cilia  8-10  um.  Six  somatic  kinetics  (Kl-6),  three  preoral  kinetics  (Pl-3),  and  a  short 
x-kinety  (Figs  124, 125).  Kl  anterior  with  8-10,  posterior  with  4,  K2  anterior  with  2,  posterior  with 
5-6,  K3  anterior  with  12-16,  posterior  with  5-8,  K4  (anterior)  with  6-8,  K5  anterior  with  3-4, 
posterior  with  2,  K6  with  10-12  kinetosomes.  At  the  end  of  K4  and  in  the  middle  of  K5  sometimes  a 
single  unciliated  kinetosome,  respectively.  Basal  bodies  of  Kl-5  mostly  paired,  K6  always  with 
single  kinetosomes.  Preoral  kinety  1  with  4-5  pairs,  preoral  kinety  2  constantly  with  5  singles,  and 
preoral  kinety  3  constantly  with  7  singles,  x-kinety  with  1-2  paired  basal  bodies  located  left  of  the 
posterior  end  of  the  paroral  membrane  (Figs  120-126). 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS 


219 


116 


114 


115 


117 


118 


119 


120 


121 


122 


123 


K1 


K3 


125 


126 


127 


Figs  114-126  Trochiliopsisopaca,  scale  =  10  urn  each.  114-117  After  Penard(  1922).  114, 115  Right  and 
left  lateral  view.  116  View  from  the  apex.  117  Extrusomes  with  2,  3,  and  4  processes.  118, 119  After 
Kahl  (1931),  right  and  left  side.  120-123  Originals,  from  life  and  protargol  silver  stained  specimens, 
right  and  left  side  respectively.  CP,  cytoproct.  124,  125  Schematized  organization  of  T.  opaca,  right 
and  left  lateral  view.  Kl-6,  somatic  kinetics  1-6;  M,  adoral  membranelles;  PM,  paroral  membrane; 
Pl-3,  preoral  kinetics  1-3;  x-K,  x-kinety  126  Probable  evolution  of  Stammer idium  from  Trochiliopsis. 

Fig.  127     Schematic  organization  of  the  genus  Stammeridium  (after  Foissner,  1985£). 


220 


H.  AUGUSTIN,  W.  FOISSNER  &  H.  ADAM 


/•'  '     i 


128 


129 


130 


131 


Figs  128-131     Trochiliopsis  opaca.  128, 129  Protargol  silver  impregnated  specimens,  right  and  left  side. 
130, 131  Dry  silver  impregnated  specimens,  right  and  left  side. 


Probably  only  two  adoral  membranelles,  located  at  the  oral  peak.  Anterior  adoral  membranelle 
most  likely  built  up  by  two  rows,  posterior  one  probably  by  three  rows  of  kinetosomes.  Paroral 
membrane  with  8-9  paired  basal  bodies  (Figs  120,  122,  124).  Cyrtos  invisible  in  life  even  with 
interference  contrast,  but  slightly  impregnated  with  protargol  silver. 

Silverline  system  granular  or  very  fine-meshed. 

OCCURRENCE  AND  ECOLOGY.  Penard  (1922)  found  few  individuals  between  dead  leaves  of  the  'swan 
pond'  at  Ariana  ('. . .  a  1'etang  des  Cygnes',  Switzerland).  Kahl  (1931)  noted  T.  opaca  sporadically 
in  the  sapropel  and  sometimes  numerously  in  sapropelic  infusions  of  Glyceria.  Lackey  (1938) 
recorded  it  once  from  a  polluted  stream,  twice  from  a  trickling  filter,  and  five  times  from  an 
activated-sludge  chamber.  Noland  &  Gojdics  (1967)  mentioned  that  T.  opaca  occurs  when  the 
sludge  has  reached  the  finely  particulate  stage  and  the  bacteria  in  it  are  well  distributed.  Detcheva 


THE  GENERA  ACINERIA,  TRIMYEMA  AND  TROCHILIOPSIS 


221 


(1972)  listed  some  Bulgarian  habitats,  namely  a  pond  in  the  surroundings  of  the  village  Bosnek  in 
the  Witoscha  mountains,  a  marshy  meadow  in  the  vicinity  of  the  village  Kasitschene  near  Sofia, 
and  a  river  in  the  Wrabniza  quarter  of  Sofia.  Apart  from  in  activated  sludge,  we  found  this  species 
once  in  the  polysaprobic  zone  of  a  heavily  polluted  river  (Ager  near  Lenzing,  Upper  Austria). 
These  localities  suggest  T.  opaca  to  be  a  good  indicator  of  heavily  polluted  (polysaprobic) 
conditions.  It  might  also  have  some  tolerance  of  H2S. 


Table  3     Biometrical  characterization  of  Trochiliopsis  opaca 


Character1 


M       SD      SE      CV     Min    Max     n 


Body,  length 

25' 

66 

26-0 

1-12 

0-37 

4-4 

24-0 

27-0 

9 

Body,  width 

13 

'22 

13-0 

0-83 

0-28 

6-3 

12-0 

15-0 

9 

Macronucleus,  length 

6 

33 

6-5 

0-35 

0-12 

5-6 

6-0 

7-0 

9 

Macronucleus,  width 

6 

11 

6-0 

0-42 

0-14 

6-8 

5-5 

7-0 

9 

Distance  from  apex  to  the  beginning  of  macronucleus 

12 

•22 

12-0 

1-30 

0-44 

10-6 

10-0 

14-0 

9 

Micronucleus,  length 

1 

62 

1-6 

0-30 

0-10 

18-7 

1-2 

2-0 

9 

Micronucleus,  width 

1 

51 

1-5 

0-31 

0-10 

20-8 

1-0 

1-8 

9 

Number  of  kinetosomes  of  paroral  membrane 

17 

•78 

18-0 

0-67 

0-22 

3-7 

16-0 

18-0 

9 

Number  of  kinetosomes  of  anterior  kinety  1 

8 

•22 

8-0 

0-67 

0-22 

8-1 

8-0 

10-0 

9 

Number  of  kinetosomes  of  posterior  kinety  1 

4-00 

4-0 

0-00 

0-00 

0-0 

4-0 

4-0 

9 

Number  of  kinetosomes  of  anterior  kinety  2 

2' 

00 

2-0 

0-00 

0-00 

0-0 

2-0 

2-0 

9 

Number  of  kinetosomes  of  posterior  kinety  2 

5 

89 

6-0 

0-33 

0-11 

5-7 

5-0 

6-0 

9 

Number  of  kinetosomes  of  anterior  kinety  3 

13 

11 

12-0 

1-45 

0-48 

11-1 

12-0 

16-0 

9 

Number  of  kinetosomes  of  posterior  kinety  3 

6 

11 

6-0 

0-93 

0-31 

15-2 

5-0 

8-0 

9 

Number  of  kinetosomes  of  kinety  4 

6 

•67 

6-0 

0-87 

0-29 

13-0 

6-0 

8-0 

9 

Number  of  kinetosomes  of  anterior  kinety  5 

3 

11 

3-0 

0-33 

0-11 

10-7 

3-0 

4-0 

9 

Number  of  kinetosomes  of  posterior  kinety  5 

2 

•00 

2-0 

0-00 

0-00 

0-0 

2-0 

2-0 

9 

Number  of  kinetosomes  of  kinety  6 

10 

•33 

10-0 

0-71 

0-24 

6-8 

10-0 

12-0 

9 

Number  of  kinetosomes  of  the  x-kinety 

3 

•78 

4-0 

0-67 

0-22 

17-6 

2-0 

4-0 

9 

Number  of  kinetosomes  of  preoral  kinety  1 

9 

•89 

10-0 

0-33 

0-11 

3-4 

9-0 

10-0 

9 

Number  of  kinetosomes  of  preoral  kinety  2 

5 

•00 

5-0 

0-00 

0-00 

0-0 

5-0 

5-0 

9 

Number  of  kinetosomes  of  preoral  kinety  3 

7 

•00 

7-0 

0-00 

0-00 

0-0 

7-0 

7-0 

9 

'See  footnote  Table  1 


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Manuscript  accepted  for  publication  26  May  1986 


British  Museum  (Natural  History) 

The  birds  of  Mount  Nimha,  Liberia 

Peter  R.  Colston  &  Kai  Curry-Lindahl 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Tring,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1986,  129pp.  Hardback.  0  565  00982  6  £17.50. 


Titles  to  be  published  in  Volume  52 


Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophora  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata). 

By  P.  J.  Chimonides 


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


Bulletin  of  the 

British  Museum  (Natural  History) 


The  baculum  in  the  Vespertilioninae 
(Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus 
and  Eptesicus,  and  the  descriptions  of  a  nev* 
genus  and  subgenus 


J.  E.  Hill  &  D.  L.  Harrison 


Zoology  series    Vol52    No  7    30  July  1987 


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 
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©  Trustees  of  the  British  Museum  (Natural  History),  1987 


The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 

Keeper  of  Zoology  :  Mr  J.  F.  Peake 
Editor  of  Bulletin  :  Dr  C.  R.  Curds 
Assistant  Editor  :  Mr  C.  G.  Ogden 


ISBN  0565  05031  1 

I SSN  0007- 1 498  Zoology  series 

Vol  52  No.  7  pp  225-305 
British  Museum  (Natural  History) 
Cromwell  Road 
London  SW7  5BD  Issued  30  July  1 987 


(NATURAL  HISTORY) 

,-;c? 
The  baculum  in  the  Vespertilioninae  (Chiroptera: 

Vespertilionidae)  with  a  systematic  review,  a  synopsis 
of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of  a 
new  genus  and  subgenus 

J.  E.  Hill 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7  5BD 

D.  L.  Harrison 

Harrison  Zoological  Museum,  Bowerwood  House,  St  Botolph's  Road,  Sevenoaks,  Kent 
TN133AL 


Contents 

Synopsis 

Introduction 

Functional  and  systematic  significance  of  the  baculum 

Nature  and  scope  of  this  study 

Materials  and  methods 

Authorship  and  responsibility 

The  baculum  of  Pipistrellus 

The  baculum  of  Eptesicus 

The  baculum  in  other  Vespertilioninae 

Systematic  considerations         ..... 

Genus  Pipistrellus  Kaup,  1829 

Genus  Eptesicus  Rafi nesque,  1 820 

The  status  of  the 'Nycticeini'. 

The  classification  of  the  Vespertilioninae 
Zoogeographical  considerations 

Conclusions 

Addendum 

References 

Appendix  1 .  Specimens  examined 

Tables  1-3 

Figures  1-22 


Synopsis 

Current  classification  of  the  Vespertilioninae  rests  chiefly  on  a  suite  of  mainly  adaptive  characters,  among 
which  facial  shortening  throughout  the  subfamily  with  consequent  changes  in  the  structure,  size,  relative 
position  and  number  of  the  incisive  and  premolar  teeth  features  prominently.  Such  characters  may  not 
necessarily  reflect  relationships  or  phyletic  diversity,  and  sometimes  do  not  serve  properly  to  distinguish  the 
genera  that  they  purport  to  define,  as  in  the  distinction  of  Pipistrellus  and  Eptesicus,  where  generic  boundaries 
remain  unclear.  The  search  for  possibly  less  strongly  adaptive  features  suggested  the  possibility  that  the 
morphology  of  the  os  penis  or  baculum  might  prove  valuable  in  the  study  of  the  systematics  of  these  genera 
and  perhaps  in  the  subfamily  as  a  whole. 

This  paper  reviews  earlier  studies  of  the  baculum  in  the  Chiroptera  and  their  relevance  to  systematics  in  the 
Order,  with  an  examination  of  its  gross  morphology  throughout  the  Vespertilioninae,  especial  attention  being 
given  to  species  currently  allocated  either  to  Pipistrellus  or  to  Eptesicus.  A  synoptic  review  of  the  species 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  52  (7):  225-305 


Issued  30  July  1987 


225 


226  J.  E.  HILL  &  D.  L.  HARRISON 

content  of  these  genera  is  presented,  with  the  recognition  and  definition  of  subgenera  and  included  species 
groups:  three  such  (pumilus,  capensis  and  tenuipinnis)  currently  referred  to  Eptesicus  on  dental  grounds  seem 
instead  more  closely  related  to  Pipistrellus  to  which  they  are  here  transferred.  One  subgenus  of  Pipistrellus  is 
described  as  new  (p.  250). 

The  Vespertilioninae  as  a  whole  display  a  wide  range  of  bacular  variation,  which  falls  into  two  major  and 
several  minor  groups.  This  has  suggested  a  revision  of  the  current  classification  of  the  subfamily,  combining 
bacular  features  with  those  conventionally  in  use.  Bacular  morphology  provides  a  clear  indication  that 
the  'Nycticeini'  (or  'Nycticeiini')  is  an  artificial  grouping  and  that  the  genus  Nycticeius  as  presently  under- 
stood is  composite.  Currently  it  is  held  to  include  two  species,  the  North  American  humeralis  and  the  African 
schlieffenii:  these  are  here  thought  to  be  sufficiently  characterised  to  justify  generic  separation  and  a  new 
generic  name  is  proposed  for  schlieffenii  (p.  254). 

A  suggested  classification  of  the  subfamily  is  presented,  with  a  tabulated  review  of  earlier  classifications; 
possible  relationships  between  the  constituent  genera  are  discussed  and  the  zoogeography  of  the  bacular  types 
within  the  subfamily  is  examined. 


Introduction 

A  penial  bone  is  known  to  occur  among  mammalian  Orders  in  the  Insectivora,  Chiroptera, 
Primates,  Rodentia  and  Carnivora.  Variously  called  the  os  penis,  os  priapi  or  os  glandis,  it  was  first 
named  the  baculum  by  Thomas  (191 5a),  the  corresponding  structure  in  the  female,  the  os  clitoridis, 
being  later  called  the  baubellum  by  Shortridge  (1934:  327,  footnote).  The  features  of  the  baculum 
have  been  used  extensively  in  attempts  to  determine  phyletic  relationships  at  various  systematic 
levels  (Patterson  &  Thaeler,  1982).  Thomas  (loc.  cit.),  for  example,  suggested  that  the  baculum 
might  provide  evidence  valuable  in  the  subfamilial  classification  of  the  Sciuridae  and  indeed 
pointed  out  that  in  this  connection  there  were  no  bacular  features  to  support  the  association  of  the 
dwarf  squirrels  in  a  separate  subfamily,  the  Nannosciurinae.  More  commonly,  bacular  features 
have  been  used  to  indicate  or  determine  relationships  within  genera  in  the  Sciuridae,  among  New 
World  rodents,  and  in  the  Mustelidae.  Such  characteristics  have  been  employed  in  species  descrip- 
tions, especially  where  bacular  variation  is  pronounced,  and  also  for  age  determination.  Numerous 
examples  of  these  uses  of  the  baculum  are  summarised  by  Patterson  &  Thaeler  (loc.  cit.)  while  Burt 
(1960)  gave  an  account  of  the  earlier  of  such  studies.  Similar  early  accounts  of  the  baculum  in  the 
Chiroptera  are  reviewed  by  Hamilton  (1949). 

The  presence  of  a  baculum  in  some  at  least  of  the  Chiroptera  has  been  long  established. 
Daubenton  (1760)  described  and  illustrated  (in  part)  the  baculum  of  Nyctalus  noctula  and 
Blainville  (1840)  similarly  studied  the  baculum  of  Rhinolophus  fermmequinum,  R.  hipposideros, 
Vespertilio  murinus  and  again  of  Nyctalus  noctula,  the  latter  author  providing  perhaps  the  first 
accurate  and  quite  detailed  drawings  of  this  structure.  Later  workers  such  as  Ercolani  (1868), 
Robin  (1881),  Gilbert  ( 1 892),  Rauther  ( 1 903),  Gerhardt  ( 1 905)  and  Chaine  ( 1 926)  provided  further 
details  of  penial  and  bacular  morphology  in  the  Chiroptera,  Chaine  in  particular  discussing  and 
illustrating  the  baculum  in  several  species  and  to  some  extent  summarising  earlier  work  in  the  field. 
However,  none  attempted  to  use  the  structure  of  the  baculum  for  systematic  purposes. 

The  first  use  of  the  baculum  in  chiropteran  systematics  appears  to  be  by  Thomas  (\9\5b)  who 
employed  bacular  characteristics  in  defining  the  species  of  Nyctophilus.  This  worker  clearly  fore- 
saw the  value  of  bacular  features  in  the  definition  of  some  at  least  of  the  species  of  bats,  beginning 
from  that  time  a  collection  of  vespertilionid  bacula  at  the  British  Museum  (Natural  History) 
although  subsequently  making  little  use  of  the  material  that  was  accumulating,  except  in  1928 
employing  bacular  characters  to  separate  Indo-Chinese  species  of  Pipistrellus  (Thomas,  19280,  K). 
Since  then  the  baculum  has  been  utilised  in  a  variety  of  taxonomic  studies  of  bats,  for  example  by 
Krutzsch  (1959,  1962)  and  Lanza  (1969)  to  examine  its  value  in  indicating  relationships  in  the 
Megachiroptera,  by  Topal  (19700)  in  determining  the  affinities  of  la,  or  by  Heller  &  Volleth  (1984) 
as  an  indicator  of  relationship  among  the  species  of  Pipistrellus  and  Eptesicus.  The  baculum  of 
Plecotus  was  found  valuable  by  Lanza  (1960)  in  discriminating  between  P.  auritus  and  P.  wardi 
(  =  P.  austriacus):  the  subsequent  use  of  the  baculum  in  distinguishing  these  species  is  summarised 
by  Corbet  (1964).  Genoways  &  Jones  (1969)  found  that  bacular  features  distinguished  closely 


VESPERTILIONINE  SYSTEMATICS  227 

related  species  of  North  American  Myotis,  LaVal  (1973#)  employing  bacular  characters  for  the 
same  purpose  among  the  Neotropical  species  of  this  genus. 

The  emphasis  placed  on  bacular  characters  in  chiropteran  systematics  is  perhaps  best  illustrated 
by  the  number  of  studies  devoted  chiefly  to  bacular  structure,  often  on  a  regional  or  faunal  basis,  as 
for  instance  the  work  by  Hamilton  (1949)  and  Krutzsch  &  Vaughan  (1955)  on  North  American 
species,  by  Brown  et  al.  (1971)  on  Neotropical  bats,  by  Topal  (1958)  on  central  European  species, 
by  Bhatnagar  (1967),  Agrawal  &  Sinha  (1973),  Sinha  (1976)  and  Khajuria  (1979,  1980,  1982)  on 
Indian  bats,  or  by  Wassif  &  Madkour  (1972)  and  Wassif,  Madkour  &  Soliman  (1984)  on  Egyptian 
bats.  Bacula  are  sometimes  studied  in  discrete  taxonomic  groupings,  as  for  example  those  of  New 
World  molossids  by  Brown  (1967)  or  of  Malaysian  Hipposideros  by  Zubaid  &  Davison  (in  press). 
Thus  among  the  Chiroptera  the  baculum  has  been  employed  as  a  source  of  taxonomic  features  at 
several  systematic  levels,  but  primarily  to  indicate  degrees  of  relationship  or  for  separation  at  the 
specific  and  sometimes  the  generic  grades,  or  especially  for  distinguishing  closely  related,  often 
sympatric  species  whose  conventional  morphological  characters  are  otherwise  very  similar,  as  in 
Myotis  and  Plecotus. 


Functional  and  systematic  significance  of  the  baculum 

Conflicting  hypotheses  for  bacular  variation  were  reviewed  in  detail  by  Patterson  &  Thaeler  (1982). 
These  authors  proposed  that  among  rodents  at  least  the  probability  was  that  the  baculum  has  a 
precise  reproductive  purpose  and  functions  primarily  as  a  device  contributing  to  species  isolation. 
Bacular  differences  among  closely  related  taxa  might  well  then  take  an  exaggerated  form.  As  such, 
the  baculum  would  be  therefore  a  poor  basis  for  supra-specific  classification,  but  an  excellent 
structure  for  species  diagnosis.  Thus  they  would  not  consider  a  phyletic  basis  for  bacular  variation 
to  be  appropriate.  They  admit,  however,  that  while  in  some  rodent  groups  there  are  patterns  of 
bacular  morphology  that  do  not  agree  with  phyletic  divergence  as  indicated  by  other  morpho- 
logical features,  there  exist  also  gross  patterns  of  bacular  variation  in  other  groups  that  do  in  fact 
conform  with  accepted  phyletic  relationships.  Indeed,  they  remarked  that  there  can  be  little  doubt 
that  the  baculum  exhibits  phyletic  weight  and  consequently  may  serve  as  a  valuable  taxonomic 
tool.  Moreover,  taxa  that  differ  in  external  and  cranial  characters  may  have  similar  bacula,  while 
others  that  are  similar  in  such  features  may  exhibit  highly  distinctive  genitalia.  Patterson  & 
Thaeler  (loc.  cit.)  suggested  that  although  bacular  morphology  reflects  phyletic  history  on  a  gross 
scale,  discordance  between  patterns  of  bacular  and  phyletic  divergence  supports  a  functional 
interpretation  of  bacular  variation,  especially  at  the  species  level. 

Similarly,  opinions  vary  as  to  the  value  of  bacular  morphology  in  chiropteran  systematics. 
Hamilton  ( 1 949)  examined  the  baculum  in  North  American  vespertilionids  and  concluded  that  in 
this  family  the  baculum  was  useful  in  defining  relationships  when  considered  with  skull  and  other 
skeletal  characteristics.  Thus  he  was  able  to  suggest  that  the  close  similarity  between  the  bacula  of 
Myotis  (Fig.  19i,  j)  and  Pizonyx  (Fig.  19k)  indicated  their  close  relationship,  and  that  the  dissimi- 
larity between  the  bacula  of  Pipistrellus  subflavus  (Fig.  2d)  and  P.  Hesperus  (Fig.  8d)  suggested 
generic  or  at  least  subgeneric  difference.  This  author  also  noted  that  in  most  instances  among 
North  American  vespertilionids  there  were  marked  generic  differences  in  the  baculum.  He  con- 
sidered that  further  study  was  needed  to  determine  the  usefulness  of  the  baculum  in  chiropteran 
systematics  and  that  with  time  and  sufficient  material  the  bone  might  be  utilised  in  classification. 
These  conclusions  were  reinforced  by  Krutzsch  &  Vaughan  (1955)  who  examined  the  bacula  of 
further  North  American  species.  They  remarked  that  in  the  case  of  those  that  are  closely  related 
the  baculum  can  serve  as  a  criterion  in  judging  relationship  when  other  clear  cut  distinguishing 
characters  are  lacking.  These  authors  found  bacular  variation  in  closely  related  bat  species  to  be 
chiefly  in  shape,  detail  of  outline,  and  gross  size:  their  study  led  to  the  belief  that  in  at  least  some 
superficially  similar  species  well  marked  and  consistent  bacular  differences  reinforced  the  more 
subtle  external  and  cranial  dissimilarities. 

Krutzsch  (1959)  accepted  the  view  that  the  baculum  can  provide  additional  evidence  for  classifi- 
cation, or,  in  the  absence  of  other  clearly  defined  characters,  can  serve  as  a  criterion  in  judging 


228  J.  E.  HILL  &  D.  L.  HARRISON 

relationship.  He  added  in  relation  to  the  Pteropodidae  that  by  virtue  of  its  relative  simplicity  and 
structural  stability  the  baculum  might  well  serve  to  help  place  entities  of  doubtful  relationship  in 
their  natural  position,  although  they  might  be  otherwise  morphologically  contradictory.  Among 
pteropodids  he  found  that  infrageneric  differences  in  the  baculum  involved  minor  details  of  shape, 
outline  and  size.  Genera,  however,  might  be  separated  by  more  profound  differences.  His  study 
suggested  that  although  within  the  genus  well  marked  and  consistent  differences  existed  between 
the  bacula  of  individual  species,  there  was  nevertheless  a  basic  similarity  in  pattern  throughout  the 
genus,  leading  to  the  suggestion  that  marked  variants  from  this  morphological  standard  in  a  single 
genus  might  provide  grounds  for  a  reappraisal  of  the  affinities  of  the  variant.  A  further  study 
(Krutzsch,  1962)  confirmed  these  opinions,  especially  in  the  broad  agreement  of  bacular  variation 
in  pteropodids  with  the  taxonomic  arrangement  of  this  family  by  Andersen  (1912)  and  by  Tate 
(1942ft).  Krutzsch  concluded  that  strong  intrageneric  similarities  exist  among  the  bacula  of  ptero- 
podids, but  that  representative  bacula  of  different  genera  differ  distinctly:  although  serving  well  as 
a  source  of  diagnostic  features  for  the  genus,  the  baculum  does  not  seem  to  offer  exceptional  insight 
into  suprageneric  relationships.  The  large  genus  Pteropus,  however,  to  some  extent  proves  to  be 
an  exception,  with  various  of  its  many  species  demonstrating  considerable  variation  in  bacular 
structure:  on  occasion  differences  between  species  equal  those  between  some  megachiropteran 
genera. 

Lanza  ( 1 969)  examined  the  baculum  of  Pteropus  in  detail  and  found  that  its  bacular  morphology 
did  not  conform  to  the  classification  proposed  by  Andersen  (1912),  a  conclusion  also  reached  by 
Davis  ( 1 947)  who  examined  only  five  species.  Lanza  found  that  in  many  cases  bacula  of  an  identical 
size  and  shape  could  be  found  among  species  belonging  to  the  same  group  as  well  as  to  different 
groups;  or  that  the  baculum  could  be  extremely  different  among  forms  apparently  otherwise  very 
closely  related.  Thus  in  this  genus  he  found  the  baculum  to  be  of  limited  value  in  phyletic  analysis. 
Similarly,  LaVal  (1973ft)  found  that  with  one  exception  the  bacula  of  the  various  species  of  the 
vespertilionid  Rhogeessa  are  not  sharply  differentiated  from  each  other:  although  in  shape  they 
show  substantial  geographic  and  individual  variation  within  species  they  seem  nevertheless  to 
differ  between  species  in  areas  of  sympatry  or  near  sympatry.  Harrison  &  Brownlow  (1978)  found 
that  individual  variation  in  the  baculum  of  adults  of  another  vespertilionid,  Scotophilus,  was  such 
that  it  rendered  this  structure  of  little  or  no  value  in  species  diagnosis  in  this  difficult  genus. 

Martin  (1978)  discussed  the  adaptive  value  of  the  baculum  in  bats,  having  found  a  wider  range  of 
structural  variation  among  several  pteropodid  species  than  was  previously  thought.  He  considered 
that  the  baculum  may  have  a  number  of  roles  of  varying  adaptive  significance  in  supporting  the 
penis,  as  a  stimulatory  structure,  or  in  preventing  urethral  closure  during  the  pressures  of  copula- 
tion. Although  these  might  allow  the  baculum  to  maintain  morphological  stability  within  certain 
taxonomic  units,  this  possible  variability  of  function  he  thought  tended  to  reduce  its  value  in 
classification  at  the  specific  and  subspecific  levels. 

Despite  these  possible  limitations,  many  authors  admit  at  least  the  species-specificity  of  bacular 
variation  among  bats,  using  the  baculum  to  provide  additional  characters  to  separate  species  that 
sometimes  otherwise  closely  resemble  each  other.  Some  examples  have  been  mentioned:  others 
include  Wallin  (1969)  who  drew  attention  to  bacular  differences  in  Japanese  Pipistrellus  and  who 
used  such  differences  to  define  two  species  groups  in  Vesper tilio,  or  Baag0e  (1973)  who  utilised 
bacular  characters  in  comparing  sibling  species  of  European  Myotis.  Zubaid  &  Davison  (in  press) 
found  the  baculum  to  be  specifically  diagnostic  among  Malaysian  Hipposideros.  In  some  genera 
authors  have  routinely  described  and  illustrated  the  baculum  of  new  species:  for  instance  Sinha 
(1969)  in  describing  Pipistrellus  peguensis  compared  its  baculum  with  the  bacula  of  the  related 
species.  Similarly,  McKean  et  al.  (1978)  described  and  illustrated  the  baculum  of  'Eptesicus' 
sagittula,  comparing  it  with  the  bacula  of  other  Australian  'Eptesicus',  while  Kitchener  (1976) 
employed  the  baculum  of ' Eptesicus'  douglasorum  in  the  same  way.  Bacular  characters  sometimes 
form  an  essential  part  of  revisionary  study,  as  by  Kitchener  et  al.  (1986)  in  defining  and  keying  the 
Australo-Papuan  representatives  of  Pipistrellus  and  Falsistrellus.  The  baculum  has  also  featured 
in  generic  revision,  Hill  (1966a)  for  example  describing  and  illustrating  that  of  Philetor  in  the 
course  of  such  a  study,  or  (1976)  that  of  the  majority  of  the  species  of  Hesperoptenus. 

Bacular  variation  has  also  been  employed  for  generic  and  subgeneric  distinction  within  the 


VESPERTILIONINE  SYSTEMATICS  229 

Vespertilioninae.  Wallin  (1969)  used  penial  characters  in  establishing  Vespertilio  as  a  genus  distinct 
from  Eptesicus  and  in  recognising  Hypsugo  as  a  subgenus  within  Pipistrellus,  while  Topal  (1970a) 
noted  that  bacular  morphology  allied  the  aberrant  genus  la  more  closely  to  Eptesicus  than  to 
Pipistrellus  with  which  it  had  been  associated  by  some  authors.  Heller  &  Volleth  (1984)  summar- 
ised published  illustrations  of  the  bacula  of  Pipistrellus,  Eptesicus  and  some  of  their  associated 
genera,  drawing  attention  to  their  taxonomic  implications.  At  a  further  systematic  level,  Pine  et  al. 
(1971)  discussed  the  penial  and  bacular  morphology  of  Antrozous  and  Bauerus  in  relation  to  the 
presumed  affinities  of  these  North  American  genera  to  the  Australian  and  New  Guinea  genera 
Nyctophilus  and  Pharotis  with  which  they  have  been  associated  in  the  subfamily  Nyctophilinae. 

It  is  clear  from  the  foregoing  account  that  the  baculum  is  regarded  as  a  valuable  source  of 
diagnostic  information  by  many  students  of  chiropteran  systematics.  This  seems  especially  true  in 
the  Vespertilioninae,  a  subfamily  in  which  diagnosis  and  definition  at  both  specific  and  generic 
levels  is  sometimes  difficult  if  only  the  orthodox  morphological  characters  of  external,  cranial  and 
dental  structure  are  to  be  relied  upon. 


Nature  and  scope  of  this  study 

The  basis  of  the  current  classification  of  the  Vespertilioninae  was  first  set  out  in  detail  by  Miller 
(1907),  who  recognised  a  total  of  thirty-two  genera  in  the  group,  with  diagnoses  and  short  descrip- 
tive accounts.  The  classification  of  Miller  was  based  chiefly  on  external,  cranial  and  dental  features. 
Tate  (1942a)  reviewed  the  characters  used  for  diagnosis  in  some  detail,  dividing  the  subfamily 
into  four  main  (tribal)  aggregations,  and  attempting  to  quantify  the  interrelationships  of  its 
many  genera.  The  major  outlines  of  his  classification  have  since  been  followed,  sometimes  with 
local  modification  as  for  instance  by  Koopman  (19840,  b,  1985)  who  subsumed  the  subfamily 
Nyctophilinae  into  the  Vespertilioninae.  Hill  (1966)  pointed  out  that  the  subfamily  comprises  a 
complex  of  closely  interrelated  genera  separated  in  some  instances  by  comparatively  slender  or 
even  rather  arbitrary  distinctions,  the  patterns  of  relationship  often  obscured  by  parallelism  or 
convergence. 

The  narrowness  of  the  orthodox  distinctions  that  define  many  of  the  constituent  genera  of  the 
Vespertilioninae  has  led  to  much  taxonomic  combination  and  recombination  since  Tate  wrote. 
This  situation  is  exemplified  by  the  more  extreme  variants  of  classification  that  have  been  pro- 
posed. For  example,  Kuzyakin(  1944, 1950, 1 965)  included  Pipistrellus  and  Eptesicus  in  Vespertilio 
while  Simpson  (1945)  included  Glischropus,  Scotozous,  Nyctalus  and  la  in  Pipistrellus  and  Rhinop- 
terus,  Hesperoptenus,  Tylonycteris,  Mimetillus,  Philetor,  Histiotus  and  Laephotis  in  Eptesicus.  A  yet 
more  extreme  viewpoint  was  adopted  by  Sokolov  (1973)  who  considered  that  Vespertilio  should 
include  not  only  Pipistrellus  and  Eptesicus  as  was  thought  by  Kuzyakin,  but  also  all  of  the  other 
above  mentioned  genera  except  Nyctalus.  Horacek  &  Hanak  (19850,  b)  commented  that  the 
concepts  of  Kuzyakin  and  Sokolov  (with  the  inclusion  of  Nyctalus}  might  be  provisionally 
accepted,  at  least  until  factual  proof  of  paraphylly  in  the  group  was  forthcoming.  Nevertheless, 
they  considered  this  to  be  a  retrograde  solution  since  it  expresses  nothing  of  the  factual  diversity  of 
the  group,  proposing  instead  that  the  problematic  taxa  should  be  arranged  in  separate  genera,  their 
diagnoses  then  making  their  content  clearer  though  narrower.  Both  Simpson  and  Sokolov  also 
included  Scotoecus  and  Scotomanes  in  Nycticeius  as  then  understood,  Baeodon  in  Rhogeessa, 
Glauconycteris  in  Chalinolobus  and  Dasypterus  in  Lasiurus  to  produce  a  heavily  'lumped'  classifica- 
tion. The  status  of  some  such  as  Scotoecus,  Dasypterus  and  Idionycteris  has  varied  from  one  author 
to  another  for  decades:  in  the  Australian  region  Scoteanax  and  Scotorepens  have  recently  achieved 
generic  rank  after  many  years  as  nominal  subgenera  (Kitchener  &  Caputi,  1985)  while  la  has  once 
again  reverted  to  Pipistrellus  (Koopman  1984a,  b,  1985)  after  a  brief  spell  with  Eptesicus.  The 
major  variants  of  vespertilionine  classification  are  summarised  in  Table  1. 

Many  of  the  characters  used  to  define  taxa  and  relationships  among  the  Vespertilioninae  appear 
strongly  adaptive  and  of  equivocal  value  in  generic  and  suprageneric  systematics.  Most  concern 


230  J.  E.  HILL  &  D.  L.  HARRISON 

ear  size  and  shape,  tragal  structure,  the  architecture  of  the  skull,  and  the  number  and  formation  of 
the  teeth.  Zima  &  Horacek  (1985)  pointed  out  that  the  use  of  the  morphological  characters 
employed  hitherto  in  the  classification  of  the  Vespertilionidae  as  a  whole  might  not  lead  invariably 
to  correct  taxonomic  conclusions,  their  degree  of  differentiation  perhaps  reflecting  the  orientation 
and  intensity  of  selection  pressure  rather  than  actual  phyletic  relationships.  These  authors  indi- 
cated an  urgent  need  for  new,  sufficiently  reliable  and  taxonomically  useful  criteria  based  on 
features  that  did  not  possess  a  direct  adaptive  significance,  including  among  these  the  morphology 
of  the  reproductive  organs  and  the  baculum. 

Much  weight  has  been  placed  in  the  past  upon  the  progressive  shortening  of  the  muzzle  apparent 
throughout  the  Vespertilioninae  with  concomitant  reduction  and  loss  of  the  incisors  and  pre- 
molars  (Tate,  1942a).  In  the  incisive  dentition  the  first  upper  tooth  (i1)3  is  absent,  as  in  all  bats. 
Reduction  results  in  the  remaining  inner  tooth  (i2)  becoming  peg-like  and  unicuspid,  although 
sometimes  quite  massive,  in  a  reduction  in  size  of  the  outer  tooth  (i3),  its  displacement  inwards 
or  outwards,  or  in  its  eventual  obsolescence  or  loss.  In  the  mandible,  the  first  (ij  and  second 
(i2)  incisor  teeth  are  invariably  present,  but  exceptionally  the  third  (i3)  may  be  absent.  The  process 
also  involves  the  reduction  and  loss  of  the  second  upper  and  lower  premolars  (pm})  and  then 
of  the  anterior  upper  premolar  (pm2):  thus  the  premolar  formula  ranges  from  pm  rf  y£  to  pm 
5  i  5  f  the  first  upper  and  lower  premolars  (pm})  being  presumed  to  be  those  that  are  absent  from  all 
bats.  Seven  different  combinations  of  incisors  and  premolars  occur  in  the  subfamily,  if  Antrozous 
and  Bauerus  are  included.  The  full  complement  is  usually  taken  as  the  primitive  condition,  the 
reduction  and  disappearance  of  teeth  as  derived.  These  are  summarised  in  Table  2,  which  gives  the 
incisive  and  premolar  formulae  usually  attributed  to  each  of  the  various  genera.  However,  Myotis 
occur  in  which  pm3  or  pmj  are  absent  (Hill  &  Topal,  1973),  thus  in  the  first  instance  producing 
the  formula  typified  by  Lasionycteris  or  Plecotus,  in  the  second  the  formula  for  Pipistrellus  or 
Nyctalus;  pm2  may  be  absent  from  Pipistrellus  to  give  the  arrangement  for  Eptesicus,  or  may  be 
present  in  Eptesicus  to  produce  the  formula  for  Pipistrellus  (Hill  &  Topal,  loc.  cit.);  i3  is  variable  in 
Scotozous  (of  45  examined,  present  in  34,  absent  from  one  side  or  the  other  in  8,  completely  absent 
from  3),  when  totally  absent  to  produce  the  incisive  formula  that  usually  characterises  Nycticeius 
and  its  associates  (but  Thomas  &  Wroughton  ( 1 908)  report  an  example  of  'Nycticeius '  schlieffenii 
in  which  the  left  i3  is  present);  pm2  is  variable  in  Scotoecus  (Hill,  1974)  and  in  Chalinolobus  (Ryan, 
1966;  Koopman,  1971),  and  very  rarely  may  be  present  in  'Nycticeius'  schlieffenii  (Dobson,  1878; 
Thomas,  1890). 

Most  genera  of  Vespertilioninae  can  be  defined  by  other  features  besides  those  of  the  incisive  and 
premolar  dentition,  although  sometimes  only  in  differing  combinations.  Thus  although  some 
species  exist  that  combine  the  external  features  of  Myotis  with  the  dental  formula  of  Pipistrellus  to 
the  extent  that  initially  they  (annectans,  ridleyi)  were  described  in  the  latter  genus,  other  characters 
such  as  the  form  and  structure  of  the  tragus  and  the  structure  of  the  incisors  enable  them  to  be 
referred  confidently  to  Myotis  (Topal,  19706;  Hill  &  Topal,  1973).  Another  (rosseti)  was  first 
described  in  Glischropus,  subsequently  removed  to  Pipistrellus  by  Hill  (1969)  and  finally  (with 
ridleyi)  to  Myotis  by  Hill  &  Topal  (loc.  cit.).  However,  the  genera  Pipistrellus  and  Eptesicus  do  not 
offer  further  conventional  characters  in  this  way  and  are  separated  for  the  most  part  by  the 
presence  of  pm2  in  the  former  and  its  absence  in  the  latter.  Wallin  (1969)  and  Hill  &  Topal  (loc.  cit.) 
discussed  the  variability  of  this  tooth  in  Pipistrellus  and  Eptesicus  in  detail,  the  latter  authors 
concluding  that  the  presence  or  absence  of  pm2  can  have  no  universal  validity  in  defining  the  two 
genera.  Heller  &  Volleth  (1984)  also  examined  the  relevance  of  pm2  in  separating  Pipistrellus  and 
Eptesicus  and  concluded  that  it  does  not  seem  to  be  a  reliable  characteristic,  a  classification  based 
on  it  perhaps  misrepresenting  true  relationship.  Tate  (\942a)  recognised  this  difficulty  but  adhered 
to  the  conventional  practice  of  separating  the  two  genera  by  this  feature,  and  indeed  the  majority  of 
authors  have  retained  the  distinction  as  a  matter  of  convenience,  often  using  the  extent  of  reduction 
and  degree  of  displacement  of  pmf  from  the  line  of  the  toothrow  as  a  diagnostic  feature  between  the 
species  of  Pipistrellus. 


"Dental  notation  of  Miller  (1907) 


VESPERTILIONINE  SYSTEMATICS  23 1 

Koopman  (1975)  has  commented  upon  this  problem.  This  author  examined  African  species 
allocated  variously  to  Pipistrellus  and  Eptesicus  in  an  attempt  to  find  some  other  character  that 
would  divide  this  large  assemblage  into  two  major  groups.  He  could  find  none  among  the  usual 
suite  of  morphological  features.  Although  he  found  no  African  Pipistrellus  species  that  closely 
resembled  the  hottentotus,  tenuipinnis  orfloweri  groups  of  Eptesicus  and  no  African  member  of  the 
latter  genus  that  was  similar  to  thepipistrellus,  Hesperus,  savii  or  rueppellii  groups  of  Pipistrellus  as 
he  defined  them,  he  did  find  a  resemblance  amounting  in  some  cases  to  virtual  identity  (if  pm2  was 
ignored)  between  the  kuhlii  group  of  Pipistrellus  and  the  Eptesicus  capensis  group.  Expanding  a 
view  first  expressed  by  Tate  (19420)  in  his  account  of  Eptesicus,  Koopman  commented  that  it  is 
probable  that  the  anterior  upper  premolar  has  been  lost  more  than  once  here,  and  that  true  phyletic 
relationships  run  across  the  'generic'  line.  He  thought  that  it  is  even  possible  that  in  some  cases  a 
'Pipistrellus  species'  and  an  'Eptesicus  species'  are  actually  conspecific,  but  was  of  the  opinion 
that  the  available  material  was  insufficient  to  establish  this  with  certainty  for  any  such  pair  at 
the  present  time.  Although  retaining  Pipistrellus  and  Eptesicus  as  separate  genera  since  he 
believed  that  the  problem  should  be  attacked  on  a  cosmopolitan  basis,  he  remarked  that  such  an 
arrangement  is  almost  certainly  wrong.  More  recently,  Horacek  &  Hanak  (1985-1986)  have 
offered  further  definitions  of  Pipistrellus,  Hypsugo  and  Eptesicus. 

Many  varying  interpretations  can  be  placed  upon  external,  cranial  and  dental  morphology  or  on 
karyological  data  in  the  Vespertilioninae.  These  range  from  the  relationship  of  one  species  to 
another  to  suprageneric  relationships,  even  to  the  view  that  Pipistrellus  and  Eptesicus  may  be 
polyphyletic.  Menu  (1984),  for  example,  remarked  that  an  exhaustive  odontological  study  of 
the  Vespertilioninae  indicated  that  Pipistrellus  includes  species  wrongly  associated  by  reason 
of  their  identical  dental  formula,  but  which  are  not  related.  Williams  &  Mares  (1978)  discussed  the 
karyology  of  Eptesicus,  which  as  currently  defined  they  thought  seemed  to  be  a  composite  taxon, 
encompassing  perhaps  several  phyletic  lines  of  pipistrelloid  species  with  reduced  numbers  of 
premolars.  Heller  &  Volleth  (1984)  suggested  that  Pipistrellus  may  be  a  very  heterogeneous 
assemblage  and  after  reviewing  the  relevance  of  pm2  in  separating  this  genus  from  Eptesicus 
considered  the  baculum  to  be  a  more  reliable  guide  to  the  phylogeny  of  the  species  of  Pipistrellus 
and  Eptesicus,  using  its  features  to  supplement  their  findings  from  karyological  data.  Many  years 
before  this  Tate  (19420)  remarked  that  it  seemed  probable  that  study  of  the  baculum  in  the 
Microchiroptera  would  yield  valuable  results,  with  the  implication  that  this  might  have  signifi- 
cance in  the  classification  of  the  Vespertilionidae.  Indeed,  Tate  records  that  G.  M.  Allen  had 
gathered  together  a  number  of  bacula  representing  many  of  the  species  of  Pipistrellus  which  he 
intended  to  employ  in  revising  the  genus.  Moreover,  relatively  few  species  of  the  nominal  genera 
Pipistrellus  and  Eptesicus  have  so  far  been  studied:  the  impression  gained  from  the  literature  is 
that  Pipistrellus  as  currently  understood  is  dignified  chiefly  by  a  long,  slender  shafted  baculum  and 
most  Eptesicus  as  it  is  presently  classified  by  a  small,  triangular  structure,  which  we  have  found  not 
to  be  the  case. 

Initially  our  intention  was  to  examine  the  bacula  of  as  many  species  of  Pipistrellus  as  possible  to 
establish  the  pattern  of  bacular  variation  within  the  genus,  and  to  compare  it  with  the  species 
groupings  proposed  by  Tate  (19420)  and  by  Koopman  (1973,  1975).  As  the  work  progressed, 
however,  it  became  increasingly  apparent  that  its  implications  extended  far  beyond  the  limits  of 
this  nominal  genus  and  that  it  was  necessary  in  addition  to  study  the  bacula  of  Eptesicus  so  far  as  we 
were  able,  and  to  examine  the  boundary  between  these  two  conventional  groupings.  Finally,  to 
place  our  findings  in  proper  perspective,  we  have  surveyed  the  bacula  of  most  of  the  remaining 
genera  of  the  Vespertilioninae  and  have  attempted  to  assess  the  generic  significance  of  bacular 
variation  in  the  subfamily.  We  have  also  examined  the  bacula  of  Nyctophilus  and  Pharotis,  both 
usually  referred  to  the  closely  related  subfamily  Nyctophilinae.  This  has  been  united  recently  with 
the  Vespertilioninae  (Koopman,  19840,  b,  1985)  and  is  thus  relevant  to  our  study. 

We  have  made  no  detailed  examination  of  the  gross  morphology  of  the  chiropteran  penis  except 
insofar  as  it  is  reflected  by  bacular  structures.  Nor  have  we  attempted  to  study  its  histomorphology. 
These  features  are  discussed  by  Smith  &  Madkour  (1980)  in  an  effort  to  elucidate  their  relevance  to 
interordinal  and  infraordinal  phylogenetic  relationships,  and  who  review  earlier  studies  of  penial 
morphology. 


232  J.  E.  HILL  &  D.  L.  HARRISON 

Materials  and  methods 

We  have  been  able  to  examine  bacula  from  the  majority  of  species  currently  listed  in  Pipistrellus 
and  from  most  of  those  presently  assigned  to  Eptesicus.  In  a  few  instances  we  have  relied 
upon  illustrations  and  descriptions  from  the  literature.  Similarly,  for  the  other  genera  of  the 
Vespertilioninae  our  study  material  has  been  drawn  chiefly  from  specimens  and  to  a  much  lesser 
extent  from  the  published  works  of  others.  The  specimens  that  we  have  examined  are  listed  in 
Appendix  1 .  Our  aim  as  far  as  genera  other  than  Pipistrellus  and  Eptesicus  are  concerned  has  been 
to  provide  illustrations  of  representative  bacula,  but  in  those  instances  where  bacular  structure  has 
not  before  been  studied  we  have  endeavoured  to  examine  as  many  species  within  each  genus  as  the 
available  specimens  permitted.  Clearly,  the  material  available  to  us  has  been  quite  inadequate  to 
establish  the  extent  of  individual  variation  in  any  one  species  or  subspecies.  While  only  adult 
specimens  (wing  epiphyses  fully  fused)  have  been  used,  we  have  necessarily  had  to  accept  that  for 
the  majority  of  species  our  data  is  limited.  We  have  concentrated  therefore  on  studying  and 
comparing  the  gross  morphology  (size,  shape,  gross  structure)  of  the  bacula  that  we  have  examined 
in  an  attempt  to  identify  similarities,  differences  and  general  trends.  The  finer  details  perhaps  more 
valuable  in  species  distinction  have  received  much  less  attention,  although  where  it  is  known  that 
species  are  difficult  to  separate  by  conventional  means  attention  has  been  drawn  to  bacular  features 
that  may  assist  in  identification. 

The  specimens  used  in  this  study  have  been  drawn  almost  entirely  from  the  collections  of  the 
British  Museum  (Natural  History),  London  (BM(NH))  and  the  Harrison  Zoological  Museum, 
Sevenoaks,  Kent  (HZM).  Apart  from  these  we  have  been  able  to  examine  one  from  the  Natur- 
historisches  Museum,  Wien  (NMW),  by  courtesy  of  Dr  K.  Bauer,  and  one  from  the  Carnegie 
Museum  of  Natural  History,  Pittsburgh  (CMNH),  an  anomalous  specimen  loaned  for  identifica- 
tion by  Dr  D.  A.  Schlitter,  while  Dr  K.  F.  Koopman  generously  brought  to  London  an  example  of 
Nycticeius  humeralis  from  the  American  Museum  of  Natural  History,  New  York  (AMNH)  from 
which  a  much  needed  baculum  was  obtained.  Specimens  prepared  many  years  ago  at  the  British 
Museum  (Natural  History)  are  dry,  sometimes  mounted  on  card:  the  remainder  have  been  pre- 
pared in  the  course  of  this  study.  This  has  been  accomplished  by  maceration  for  a  short  period  in  a 
5%  solution  of  potassium  hydroxide  to  which  a  small  quantity  of  alizarin  red  has  been  added,  after 
which  the  grosser  macerated  tissue  was  removed  by  dissection,  the  specimen  then  being  cleared  and 
stored  in  glycerin. 

Drawings  have  been  prepared  using  either  a  stereoscopic  microscope  with  graticule  scale  and 
attached  camera  lucida,  or  freehand  using  a  similar  instrument.  A  few  were  drawn  freehand  using  a 
stereo  projection  microscope  with  travelling  micrometer  stage.  The  wide  range  of  size  variation 
among  vespertilionine  bacula  (for  example  from  a  length  of  1  mm  or  less  to  as  much  as  9  or  10  mm 
in  Pipistrellus)  has  necessitated  the  use  of  several  scales  of  magnification.  So  far  as  possible  all 
drawings  on  any  one  page  of  figures  are  at  the  same  magnification,  with  an  appropriate  scale:  to 
facilitate  comparison  the  varying  magnifications  used  follow  an  arithmetic  progression  whereby 
each  successive  larger  value  is  twice  its  predecessor.  It  has  not  always  been  possible  to  conform  to 
this  arrangement,  especially  where  drawings  have  been  prepared  from  published  illustrations.  As  a 
rule  dorsal  (D)  and  right  lateral  (RL)  views  of  each  baculum  are  provided:  rarely  through  damage 
the  left  lateral  (LL)  aspect  is  given.  Occasionally  where  it  is  of  especial  interest  a  half  ventral  (RVL 
or  LVL)  drawing  has  been  made,  and  in  a  few  instances  where  drawings  have  been  taken  from  the 
literature  it  has  been  necessary  to  give  the  ventral  (V)  rather  than  the  dorsal  aspect. 

Authorship  and  responsibility 

We  take  joint  responsibility  for  the  results  and  opinions  put  forward  and  expressed  in  this  paper, 
and  for  the  new  names  proposed  therein. 

The  baculum  of  Pipistrellus 

Four  bacular  types  have  been  identified  within  the  nominal  genus  Pipistrellus.  With  some  excep- 
tions, modification  and  combinations,  these  are  in  broad  agreement  with  the  groupings  of  species 


VESPERTILIONINE  SYSTEMATICS  233 

proposed  by  Tate  (19420)  and  Koopman  (1973,  1975).  The  classifications  of  Tate  and  Koopman 
are  summarised  in  Table  3. 

(1)  An  elongate  structure  (Fig.  la)  with  a  slender  shaft  and  paired  basal  flanges  (e.g.  Figs  2a-c, 
3,  4,  5),  the  ventral  surface  of  the  proximal  part  of  the  shaft  transversely  concave,  its  distal  part 
cylindrical  or  nearly  so;  in  profile  the  base  in  line  with  the  shaft  or  more  or  less  deflected  downward 
at  an  angle  to  it;  the  shaft  may  be  more  or  less  straight,  flexed  or  variously  curved  in  the  vertical 
plane,  while  the  tip  is  generally  bifid  or  forked  and  may  be  directed  ventrally  to  a  greater  or  lesser 
extent. 

Species  aggregations  in  which  this  type  of  baculum  is  found  include  the  abramus,  pipistrellus, 
coromandra  and  tennis  groups  of  Tate  (19420);  Koopman  (1973)  amalgamated  these  to  form  a 
pipistrellus  group  to  which  he  added  (1975)  the  African  nanus  and  permixtus.  However,  nanus 
(Fig.  6b)  proves  to  have  a  very  different  baculum,  as  does  imbricatus  (Fig.  9a),  included  by  Tate  in 
the  coromandra  group  and  thus  by  Koopman  ( 1 973)  in  the  pipistrellus  group.  Pipistrellus  babu  (Fig. 
4a),  provisionally  placed  by  Tate  in  the  kuhlii  group,  also  has  the  long,  relatively  straight  baculum 
characteristic  of  this  part  of  the  division,  as  do  endoi  (Imaizumi,  1959)  (Fig.  3b)  and  peguensis 
(Sinha,  1969)  (Fig.  15c),  both  described  since  Tate  wrote.  The  more  recently  described  westralis 
(Koopman,  1984c)  (Fig.  lOd),  adamsi  (Fig.  lOc)  and  wattsi  (Fig.  lOg)  (Kitchener  et  al,  1986)  also 
belong  with  tenuis  (Fig.  9d)  and  its  allies  in  this  grouping.  Taxa  referred  to  the  ceylonicus  group  by 
both  Tate  and  Koopman  (1973)  prove  to  have  this  bacular  structure,  as  do  those  that  have  been 
examined  of  the  rueppellii  group  (Figs  7e,  f,  lOa,  b)  of  Koopman  (1975).  Pipistrellus  kuhlii  and  its 
associates  (Figs  5a-d,  6c)  also  belong  in  this  division.  In  these,  however,  the  basal  lobes  of  the 
baculum  are  sharply  angled  to  the  shaft  in  the  vertical  plane,  and  this  bacular  profile  is  very 
characteristic  of  kuhlii  and  its  relatives.  The  shaft  is  straight,  without  flexion,  and  the  tip  is  usually 
bifid  and  not  directed  ventrally.  Koopman  (1975)  included  anchietae  (Fig.  6e)  in  the  kuhlii  group, 
but  this  proves  to  have  a  very  different  bacular  configuration. 

A  long-shafted  baculum  of  this  type  occurs  with  little  modification  in  the  majority  of  the 
Australian  species  (Figs  lla-f,  12k)  currently  referred  to  Eptesicus,  in  Nyc talus  (Fig.  lOf),  in 
Scotozous  (Fig.  16d)  (to  which  rueppellii  and  its  immediate  associates  have  sometimes  been 
referred)  and  in  Scotoecus  (Fig.  20a-e),  in  which  the  'horns'  of  the  bifid  tip  extend  in  some  instances 
almost  to  form  a  ring,  a  condition  foreshadowed  in  Pipistrellus  pater culus  (Fig.  3c).  The  Australian 
Scoteanax  (Fig.  16i)  and  Scotorepens  (Figs  16g,  h,  21e,  0  also  share  this  bacular  type:  in  Scoteanax 
the  'horns'  at  the  tip  have  become  a  transverse  bar,  but  the  species  of  Scotorepens  retain  the  bifid 
or  slightly  bifid  tip.  A  similar  long-shafted  baculum  but  with  a  simple  tip  occurs  in  the  genera 
Hesperoptenus  (Fig.  21a-c,  g)  and  Chalinolobus  (Fig.  17a-e).  The  baculum  of  Glischropus  (Fig. 
18a),  although  very  small,  is  also  of  this  type,  with  paired  basal  lobes,  a  slender  shaft,  and  bifid  tip. 

(2)  A  very  small  structure  (Fig.  Ib),  consisting  of  a  broad  base  with  two  basal  lobes  (e.g.  Figs  2d, 
e,  9c,  h),  supporting  a  short,  very  slightly  hollowed  shaft.  This  bacular  type  is  found  in  subflavus 
(Fig.  2d),  circumdatus  (Fig.  2e),  societatis  (Fig.  9c)  and  the  more  recently  described  cuprosus  (Hill  & 
Francis,  1984)  (Fig.  9h). 

(3)  A  relatively  short,  stout  shafted  baculum  (Fig.  Ic),  sometimes  with  expanded  base  and  tip 
(e.g.  Figs  6a,  b,  7a,  h,  8e,  f),  the  base  on  occasion  divided  into  paired  lobes,  sometimes  angled 
vertically  to  the  line  of  the  shaft,  which  is  fluted  ventrally  rather  than  mostly  cylindrical;  tip  when 
expanded  having  its  anterior  edge  sometimes  divided  into  several  irregular  serrations  and  on 
occasion  downwardly  directed. 

Such  bacula  are  found  in  the  savii  group  of  Tate  (19420)  and  Koopman  (1973,  1975)  but  not  in 
maderensis  (Fig.  5b)  which  was  put  into  the  savii  group  by  both  authors.  Its  baculum  is  however 
quite  different  and  is  like  that  of  kuhlii  and  its  associates.  Pipistrellus  anchietae  (Fig.  6e),  referred  to 
the  kuhlii  group  by  Koopman  (1975)  also  belongs  with  savii,  and  the  same  bacular  type  occurs  in 
nanus  (Fig.  6b),  allocated  with  permixtus  to  the  pipistrellus  group  by  the  same  author.  We  have  been 
unable  to  examine  the  baculum  of  permixtus  but  that  of  nanus  and  ofhelios  (Fig.  6d)  is  of  the  type 
characteristic  of  this  division,  with  its  basal  part  quite  sharply  flexed  to  the  shaft  although  not 
especially  deep,  and  with  an  expanded,  downwardly  directed  distal  part.  Pipistrellus  eisentrauti 


234  J.  E.  HILL  &  D.  L.  HARRISON 

(Fig.  9g),  referred  to  the  rueppellii  group  by  Koopman  (1975)  also  shares  this  bacular  type.  The 
North  American  Hesperus  (Fig.  8d)  was  placed  by  this  author  in  a  Hesperus  group,  with  the  African 
musciculus.  It  has  a  robust  baculum  of  the  type  found  in  this  division,  somewhat  flattened,  without 
basal  lobes  but  broadened  just  beyond  the  base,  the  shaft  narrowing  towards  the  tip;  the  ventral 
surface  is  shallowly  fluted  throughout  its  length.  Unfortunately,  no  baculum  has  been  available  for 
musciculus  but  provisionally  it  is  referred  to  this  division  on  other  grounds. 

This  bacular  class  also  includes  imbricatus  (Fig.  9a),  referred  to  the  coromandra  group  by  Tate 
(1942a)  and  by  Koopman  (1973),  and  pulveratus  (Fig.  8c),  lophurus  (Fig.  8f),  and  kitcheneri  (Fig. 
8e)  which  Tate  placed  in  the  qffinis  group  (vide  infra).  Koopman  (loc.  cit.)  followed  this  lead  with 
respect  to  kitcheneri  but  did  not  include  pulveratus  and  lophurus  since  these  were  extralimital  to 
his  study.  Pipistrellus  bodenheimeri  (Fig.  9f),  described  (Harrison,  1960)  since  Tate  wrote  and 
extralimital  to  Koopman  (loc.  cit.,  1975)  also  belongs  in  this  group.  Tate  referred  macrotis,  vorder- 
manni  and  curtatus  to  the  savii  group  on  account  of  their  reduced  pm2  but  indicated  that  this 
allocation  might  not  be  tenable:  however,  the  baculum  of  macrotis  shows  that  it  should  be  placed  in 
this  group.  This  author  also  created  the  joffrei  group  to  include  joffrei,  anthonyi,  brachypterus,  and 
stenopterus:  brachypterus  has  since  proved  to  be  a  Philetor  (Hill,  1971).  We  have  been  unable  to 
examine  the  baculum  in  either  joffrei  or  anthonyi  but  that  of  stenopterus  (Fig.  7h)  indicates  that  it 
belongs  here.  Both  joffrei  and  stenopterus  have  been  referred  variously  to  Nyctalus  (Chasen,  1940; 
Ellerman  &  Morrison-Scott,  1951)  or  to  Pipistrellus  (Tate,  19420;  Hill,  19660)  but  the  baculum  of 
stenopterus  has  no  resemblance  to  the  long-shafted  structure  of  the  former  genus. 

(4)  A  relatively  large,  short  but  strong  baculum  (Fig.  Id),  broad,  with  little  or  no  proximal  or 
distal  expansion  (e.g.  Figs  8a,  b,  g,  lOh),  the  ventral  surface  transversely  deeply  concave  so  that  it  is 
strongly  arched  or  fluted  throughout  its  length.  This  grouping  includes  affinis  (Fig.  8a)  andpetersi 
(Fig.  8b),  placed  in  the  qffinis  group  by  Tate  (19420)  and  in  the  case  ofpetersi  in  the  same  group  by 
Koopman  (1973),  qffinis  being  extralimital  to  his  study,  together  with  the  Australian  tasmaniensis 
(Fig.  8g)  for  which  Tate  maintained  a  tasmaniensis  group.  A  further  Australian  form,  mackenziei 
(Kitchener  et  al.,  1986)  (Fig.  lOh)  is  very  like  tasmaniensis  and  also  belongs  here.  To  some  extent 
this  grouping  is  linked  to  the  previous  division  by  pulveratus,  imbricatus,  lophurus,  kitcheneri  and 
their  immediate  associates:  Tate  allocated  all  except  imbricatus  to  the  affinis  group. 

The  baculum  of  Eptesicm 

We  have  been  able  to  identify  three  bacular  types  among  the  species  currently  referred  to  the 
nominal  genus  Eptesicus.  There  is  no  single  reference  for  species  groupings  in  this  aggregation  of 
species,  but  for  African  forms  these  bacular  types  agree  almost  exactly  with  the  species  groups 
defined  by  Koopman  (1975). 

(1)  A  more  or  less  triangular  structure  (Fig.  le),  its  apex  occasionally  drawn  out  into  a  slight, 
short  shaft,  the  base  widened  and  sometimes  slightly  lobed  but  the  tip  not  expanded,  usually  more 
or  less  pointed  or  gently  rounded  (e.g.  Figs  13, 14a,  c).  This  type  of  baculum  is  flattened,  with  little 
ventral  fluting  or  concavity:  there  is  little  vertical  flexion  either  of  the  base  or  of  the  more  distal  part, 
and  the  tip  is  not  deflected  downwards.  So  far  as  we  have  been  able  to  establish,  this  bacular  class 
occurs  in  all  of  the  forms  that  are  currently  referred  to  Eptesicus  from  the  New  World,  Europe  and 
Asia,  and  in  the  African  forms  that  Koopman  (1975)  included  in  the  serotinus  andfloweri  groups. 

(2)  The  structure  in  a  small  group  of  species,  wholly  Australian,  in  which  the  baculum  has 
usually  a  long  cylindrical  or  slightly  fluted  shaft  with  paired  expanded  basal  lobes  (Fig.  If)  and 
usually  a  blunt  tip  (e.g.  Fig.  1  la-e),  very  similar  in  fact  to  the  first  of  the  bacular  types  that  we  have 
described  for  Pipistrellus.  This  grouping  includes pumilus  (Figs  1  la,  b,  12k),  vulturnus  (Fig.  1  le), 
douglasorum  (Fig.  1  Id)  and  regulus  (Fig.  1  le):  in  pumilus  and  douglasorum  the  shaft  and  base  may 
be  flexed  rather  like  those  of  Pipistrellus  kuhlii  and  its  immediate  relatives,  while  in  regulus  the  shaft 
has  an  expansion  just  behind  the  tip  (McKean  et  al.,  1978;  Kitchener,  1976).  A  further  species, 
sagittula  (Fig.  llf),  also  appears  to  belong  here,  its  baculum  being  perhaps  a  shorter-shafted 
version  of  this  type. 


VESPERTILIONINE  SYSTEMATICS  235 

(3)  A  slender-shafted  baculum  (Fig.  Ig),  usually  with  distinct  paired  basal  lobes,  sometimes 
angled  or  flexed  to  the  line  of  the  shaft,  which  is  cylindrical,  with  variously  expanded  tip,  the  distal 
expansion  varying  from  a  downwardly  directed  spatulate  plate  to  a  large,  anteriorly  directed, 
downwardly  deflected  lobed  structure  (e.g.  Figs  12a-j,  14b).  Taxa  with  this  type  of  baculum  are 
wholly  African  and  the  grouping  comprises  those  forms  referred  to  the  capensis  and  tenuipinnis 
group  by  Koopman  (1975). 


The  baculum  in  other  Vespertilioninae 

Before  considering  the  implications  of  bacular  morphology  in  relation  to  the  systematics  of 
Pipistrellus  and  Eptesicus,  a  brief  review  of  bacular  types  in  the  remaining  genera  of  the 
Vespertilioninae  will  serve  to  place  these  nominal  genera  in  the  perspective  of  bacular  structure  in 
the  subfamily  as  a  whole. 

My otis  (Fig.  19i,  j).  The  baculum  of  Myotis  has  been  figured  and  described  by  numerous 
authors.  Palaearctic  species  have  been  studied  by  Topal  (1958),  Hanak  (1965,  1970,  1971),  Wallin 
(1969),  Atallah  (1970)  and  Baag0e  (1973),  among  others.  Nearctic  and  Neotropical  species  have 
been  examined  by  Hamilton  (1949),  Wimsatt  &  Kallen  (1952),  Krutzsch  &  Vaughan  (1955),  Davis 
&  Rippy  (1968),  Genoways  &  Jones  (1969),  LaVal  (19730)  and  Warner  (1982).  In  this  genus  the 
baculum  is  much  like  a  small  saddle.  In  profile  the  base  and  tip  are  slightly  elevated,  the  baculum 
ventrally  slightly  concave.  In  dorsal  aspect  the  baculum  is  more  or  less  triangular  or  projectile- 
shaped,  anteriorly  bluntly  or  sometimes  more  sharply  pointed,  the  base  divided  to  a  greater  or 
lesser  extent  into  two  lobes,  the  ventral  surface  deeply  fluted.  In  some  instances  at  least  there  is 
evidently  distinctive  variation  between  species  and  it  is  possible  that  some  clear  infrageneric 
division  into  bacular  types  might  be  made.  There  is  also  apparently  considerable  individual  varia- 
tion in  some  species,  leading  LaVal  (19730)  in  the  case  ofnigricans  to  suggest  the  possibility  that 
the  material  that  he  studied  was  a  composite  of  sibling  species.  However,  for  so  large  a  genus 
there  seems  to  be  a  surprising  degree  of  broad  homogeneity  in  gross  bacular  structure.  We  have 
examined  relatively  few  bacula  from  such  a  well  known  genus,  but  it  is  of  some  interest  to  note  that 
the  baculum  ofridleyi  (Fig.  1 9i),  described  and  retained  for  many  years  in  Pipistrellus  on  account  of 
its  dentition  which  corresponds  to  that  genus  (Hill  &  Topal,  1973)  is  undeniably  of  the  type 
characteristic  of  Myotis. 

Pizonyx  (Fig.  19k).  Figured  and  described  by  Hamilton  (1949).  The  baculum  is  similar  to  that  of 
Myotis,  but  lacks  much  of  the  saddle-like  appearance,  and  is  more  triangular  in  dorsal  aspect, 
tapering  distally  to  a  flattened,  slightly  elevated  tip;  it  is  also  elevated  proximally  so  that  in  profile 
the  base  inclines  slightly  upwards.  Ventrally  the  baculum  is  slightly  fluted. 

Lasionycteris  (Fig.  17f).  Figured  and  described  by  Hamilton  (1949).  The  baculum  has  a  large, 
swollen,  bilobed  base,  a  long  cylindrical  shaft,  and  a  slight  distal  enlargement,  the  tip  and  base 
elevated  dorsally.  Proximally,  there  is  sometimes  a  flattened  dorsal  prominence  on  the  base,  its 
bilobed  extremity  projecting  beyond  the  main  bulbous  part. 

Plecotus  (including  Corynorhinus).  The  Old  World  forms  (Plecotus,  Figs  14d,  19g,  h)  are  figured 
and  described  by  Topal  (1958),  Lanza  (1960)  and  Ibanez  &  Fernandez  (1986),  American  taxa 
(Corynorhinus,  Fig.  15f-h)  by  Nader  &  Hoffmeister  (1983).  In  most  the  baculum  is  arrow-head- 
shaped,  slightly  saddle-like,  with  basal  lobes  and  broad,  short  distal  part,  the  base  elevated 
dorsally,  the  ventral  surface  deeply  fluted:  in  two  taxa  (auritus,  Fig.  19g,  teneriffae,  Fig.  14d)  it  has  a 
longer,  more  slender  shaft  with  paired  basal  lobes  and  is  less  saddle-like. 

Idionycteris  (Fig.  15e).  Figured  and  described  by  Nader  &  Hoffmeister  (1983).  An  elongate 
baculum,  with  triangular  basal  plate,  its  apex  directed  posteriorly,  and  narrow  shaft,  curved 
dorsally  and  ventrally  shallowly  grooved. 

Euderma.  The  baculum  in  this  genus  is  so  far  apparently  unknown. 


236  J.  E.  HILL  &  D.  L.  HARRISON 

Barbastella  (Fig.  18j).  Figured  and  described  by  Topal  (1958).  A  small,  saddle-like  baculum 
similar  in  many  ways  to  that  of  Plecotus,  with  elevated  base,  narrowed  distal  part  which  is 
upwardly  curved,  and  with  slightly  raised,  elevated  tip. 

Rhogeessa  (Fig.  18k).  Figured  and  described  by  LaVal  ( 1 9736).  A  small  baculum  with  expanded, 
bilobed  base  and  short,  stubby  shaft  lacking  any  distal  modification,  the  shaft  ventrally  fluted. 

Baeodon  (Fig.  15b).  Figured  and  described  by  Brown  et  al.  (1971)  and  LaVal  (19736).  Baculum 
very  like  that  of  Rhogeessa  but  with  shorter  shaft. 

Nycticeius.  Australian  forms  hitherto  referred  to  Nycticeius  have  been  recognised  as  Scoteanax 
and  Scotorepens  by  Kitchener  &  Caputi  (1985).  As  therefore  it  is  currently  understood,  Nycticeius 
includes  two  species,  humeralis  from  North  America  and  schlieffenii  from  Africa.  These  have 
widely  differing  bacula.  In  humeralis  (Fig.  1 7k)  the  baculum  is  blade-like,  with  short,  narrow  shaft, 
the  base  thickened,  proximally  forming  a  prominent  angle  which  inclines  towards  the  ventral 
surface,  the  distal  portion  deep,  with  convex  walls  which  terminate  in  an  ascending  point.  This 
structure  is  figured  by  Hamilton  (1949)  who  remarked  that  it  differed  markedly  from  the  bacula  of 
other  [North  American]  genera:  in  fact  it  is  not  closely  approached  by  any  other  vespertilionine. 
The  baculum  of  schlieffenii  (Fig.  16e),  by  contrast,  has  a  broad  bilobed  base  with  tapering,  fluted 
shaft,  its  tip  unmodified,  bluntly  pointed,  and  unexpanded.  Moreover,  the  bacular  morphology  of 
the  Australian  species  formerly  referred  to  Nycticeius  supports  their  separation  from  this  genus. 
This  matter  is  discussed  more  fully  below. 

Otonycteris  (Fig.  16a).  Figured  and  described  by  Wassif  &  Madkour  (1972),  Fairon  (1980)  and 
Wassif,  Madkour  &  Soliman  (1984).  An  unusual  baculum,  mostly  a  more  or  less  parallel-sided 
narrow  shaft,  the  base  and  tip  not  expanded,  both  strongly  elevated  dorsally,  the  shaft  tapering 
distally  to  a  raised  tip. 

Lasiurus  (Fig.  191).  Figured  and  described  by  Hamilton  (1949).  A  slipper-like  baculum  with 
broad,  dorsally  elevated  base,  a  short  shaft,  fluted  ventrally,  and  with  slightly  expanded  and 
elevated  tip. 

Dasypterus  (Fig.  18f).  Figured  and  described  by  Brown  et  al.  (1971)  and  Hamilton  (1949). 
Baculum  like  that  of  Lasiurus  but  tip  as  a  rule  not  upturned. 

Antrozous  (Fig.  18b).  Figured  and  described  by  Krutzsch  &  Vaughan  (1955)  and  Pine  et  al. 
(1971).  Baculum  broadly  triangular  in  dorsal  view,  tapering  to  broad,  blunt  point,  fluted  ventrally, 
and  with  the  base  elevated  dorsally.  It  is  very  different  from  the  baculum  in  Nyctophilus  and 
Pharotis,  and  from  that  of  Otonycteris,  with  which  genera  Antrozous  has  been  associated  in  the 
past. 

Bauerus  (Fig.  15i).  Figured  and  described  by  Pine  et  al.  (1971).  Baculum  like  that  of  Antrozous 
but  smaller  and  narrower,  the  distal  part  not  upcurved. 

Scotomanes  (Fig.  18g).  A  short  baculum  with  broad,  bilobed  base  merging  into  a  very  narrow, 
short  cylindrical  shaft  with  no  distal  expansion,  lacking  any  upward  deflection  either  proximally  or 
distally. 

Scotophilus  (Fig.  17g-j).  Figured  and  described  by  Harrison  &  Brownlow  (1978).  Baculum 
irregularly  sub-rectangular  or  sub-triangular,  flattened,  anteriorly  usually  bluntly  rounded, 
slightly  concave  in  ventral  transverse  section,  with  slight  basal  lobes. 

Vespertilio.  Figured  and  described  by  Topal  (1958)  and  Wallin  (1969).  In  two  species  (murinus 
and  orientalis)  the  baculum  is  situated  at  the  base  of  the  penis,  which  is  supported  by  a  cartilaginous 
pseudobaculum.  In  the  third  (superans)  the  baculum  is  situated  not  at  the  base  of  the  penis  but 
midway  along  the  shaft,  and  there  is  no  pseudobaculum.  The  baculum  in  orientalis  (Fig.  21j)  and 
superans  is  flattened  and  triangular,  with  a  broad,  bilobed  base,  tapering  anteriorly  to  a  narrow 
point  and  with  slight  vertical  flexion.  The  baculum  of  murinus  (Fig.  21i)  is  broad  but  less  triangular 
in  outline,  and  has  a  wide,  bluntly  rounded  distal  part.  The  bacula  of  orientalis  and  superans  in 
particular  are  similar  in  many  respects  to  those  of  the  Eptesicus  serotinus  group  (vide  supra). 


VESPERTILIONINE  SYSTEMATICS  237 

Histiotus  (Fig.  18c-e).  A  very  small  baculum,  with  expanded  bilobed  base  and  short,  narrow 
cylindrical  shaft,  its  tip  unexpanded,  the  base  and  tip  deflected  slightly  upwards. 

la  (Fig.  2 Id).  Figured  and  described  by  Topal  (19700).  A  large,  flattened,  triangular  baculum 
similar  to  those  of  the  Eptesicus  serotinus  group  (vide  supra). 

Tylonycteris  (Fig.  18h,  i).  Baculum  small,  similar  to  that  of  Histiotus  or  to  those  of  the  Eptesicus 
serotinus  group  (vide  supra),  but  with  the  distal  part  extended  into  a  narrowed  shaft  and  with 
relatively  wider,  expanded  base  with  a  slight  trace  of  basal  lobes. 

Glauconycteris  (Fig.  19a-f).  Baculum  very  small  and  somewhat  variable  within  the  genus,  but 
mostly  more  or  less  triangular,  with  some  modification,  usually  as  reduction,  to  a  deeply  lobed  base 
with  a  short,  blunt  distal  portion.  However,  on  occasion  the  base  is  slightly  or  considerably 
expanded  and  the  distal  portion  lengthened  to  a  short  shaft. 

Mimetillus.  We  have  been  unable  to  establish  the  presence  of  a  baculum  in  this  monotypic  genus, 
from  which  it  appears  to  be  lacking. 

Eudiscopus.  The  baculum  ofEudiscopus  (if  present)  is  apparently  unknown. 

Nyctalus  (Fig.  100-  Figured  and  described  by  Topal  (1958)  and  Lanza  (1959).  A  long,  slender 
baculum  with  narrow  basal  lobes,  a  long  cylindrical  shaft,  and  slightly  bifurcated  tip. 

Laephotis  (Fig.  160-  Baculum  with  expanded,  bilobed  base,  narrow  fluted  shaft  and  broadly 
expanded  tip  with  slight  downward  deflection,  a  small  protuberance  on  its  upper  surface.  Similar  in 
many  respects  to  the  baculum  in  the  Eptesicus  capensis  and  E.  tenuipinnis  groups  (vide  supra). 

Glischropus  (Fig.  18a).  A  very  small  baculum,  with  paired  basal  lobes,  narrow  cylindrical  shaft 
and  slightly  expanded,  bifid  tip. 

Scotozous  (Fig.  16d).  Figured  and  described  by  Sinha  (1976).  A  long  baculum  with  slight  basal 
lobes,  a  narrow,  fluted  shaft,  and  slightly  bifid  tip,  the  shaft  slightly  flexed. 

Scoteanax  (Fig.  1 6i).  Figured  and  described  by  Kitchener  &  Caputi  ( 1 985).  A  long  baculum  with 
strong,  expanded  bilobed  base,  a  slender  cylindrical  shaft,  and  with  the  tip  embellished  into  a  short, 
transverse  bar. 

Scotorepens  (Figs  16g,  h,  21e,  f)-  Figured  and  described  by  Kitchener  &  Caputi  (1985).  A  long 
baculum  with  expanded,  bilobed  base  and  slender,  cylindrical  shaft,  the  tip  slightly  expanded  and 
bifid,  the  'horns'  deflected  ventrally. 

Scotoecus  (Figs  20a-e,  21h).  A  long,  slender  baculum  with  slightly  expanded  and  bilobed 
base,  long  cylindrical  shaft  and  an  expanded,  bifurcated  tip,  the  'horns'  extending  ventrally  and 
sometimes  curving  to  form  an  almost  complete  ring. 

Philetor  (Fig.  16b).  Figured  and  described  by  Hill  (19660).  A  strong  but  relatively  short  baculum 
with  paired  basal  lobes,  a  short,  fluted  shaft,  and  expanded  rugose  tip,  the  base  and  tip  elevated  and 
deflected  upwards. 

Hesperoptenus  (Fig.  21a-c,  g).  Figured  and  described  by  Hill  (1976)  and  Hill  &  Francis 
(1984).  Baculum  long  and  slender,  with  paired  basal  lobes,  a  flattened,  ventrally  fluted  shaft,  and 
unmodified,  rounded  tip. 

Chalinolobus  (Fig.  17a-e).  Baculum  long,  with  clearly  defined  basal  lobes,  a  long  cylindrical 
shaft,  and  an  expanded  tip,  the  expansion  sometimes  slight,  considerable,  or  bifid  with  two  obtuse 
projections. 

Although  the  two  genera  are  commonly  referred  to  a  separate  subfamily,  the  Nyctophilinae,  for 
purposes  of  comparison  we  have  also  examined  the  baculum  in  Nyctophilus  and  Pharotis. 

Nyctophilus  (Figs  16c,  22a-g).  Figured  by  Churchill  et  al.  (1984).  A  long,  rather  broad  baculum 
with  scarcely  expanded,  bilobed  base  and  a  broad  shaft  tapering  distally  to  a  blunt  point,  or  wider 


238  J.  E.  HILL  &  D.  L.  HARRISON 

terminally  with  a  median  emargination  to  produce  a  shallowly  bifid  tip;  shaft  ventrally  deeply 
fluted. 

Pharotis  (Fig.  22h).  Baculum  similar  to  that  of  Nyctophilus  but  shaft  narrower,  tapering  to 
slightly  expanded  tip. 


Systematic  considerations 

The  majority  of  genera  in  the  Vespertilioninae  have  bacula  which  overall  display  a  wide  range  of 
variation  in  their  gross  morphology.  Most  have  a  distinctive  baculum:  where  closely  similar  bacula 
occur  in  genera  currently  recognised  as  distinct,  as  for  example  in  Scotozous  and  Pipistrellus ,  then 
close  relationship  has  been  presumed  on  other  morphological  grounds.  We  are  thus  persuaded  that 
in  this  subfamily  the  baculum  can  be  used  as  a  guide  to  infrageneric  and  intergeneric  classification, 
although  it  seems  that  its  value  as  a  suprageneric  indicator  may  be  less.  In  the  same  way,  although 
we  have  not  explored  the  point  in  detail,  it  has  become  apparent  that  in  many  genera  the  minor 
details  of  bacular  morphology  can  be  used  to  assist  in  species  distinction.  These  considerations 
have  led  us  to  the  view  that  the  very  dissimilar  bacular  types  that  we  have  been  able  to  identify  and 
define  within  Pipistrellus  and  Eptesicus  do  indeed  reflect  natural  groupings  and  show  that  Eptesicus 
as  it  is  currently  defined  is  a  composite.  Certainly  it  seems  true  to  say  that  the  current  classifi- 
cation of  both  nominal  genera  does  not  properly  reflect  the  relationships  that  we  believe  bacular 
morphology  suggests  exist  within  and  between  them. 

Genus  Pipistrellus  Kaup,  1829 

Pipistrellus  Kaup,  1829:98.  Vesper tilio  pipistrellus  Schreber. 

Romicia  Gray,  1838:  495.  Romicia  calcarata  Gray  =  Vespertilio  kuhlii  Kuhl. 

Rotnicius  Blyth,  1840:  75.  Variant  of  Romicia  Gray. 

Hypsugo  Kolenati,  1956:   131.  Included   Vespertilio  maurus  Blasius  =  Vespertilio  savii  Bonaparte,  and 

Vespertilio  krascheninnikowii  Eversmann.  Type  species  fixed  as  Vespertilio  savii  Bonaparte  by  Wallin 

(1969).  Valid  as  a  subgenus. 
Nannugo  Kolenati,  1856:  131.  Included  Vespertilio  nathusii  Keyserling  &  Blasius,  Vespertilio  kuhlii  Kuhl 

and  Vespertilio  pipistrellus  Schreber. 
Alobus  Peters,  1868:  707.  Vespertilio  temminckii  Cretzschmar  =  Vespertilio  ruppellii  Fischer.  Preoccupied  by 

Alobus  Le  Conte,  1856  (Coleoptera). 

Euvesperugo  Acloque,  1899:  35.  Included  six  species,  one  being  Vespertilio  pipistrellus  Schreber. 
Eptesicops  Roberts,  1926:  245.  Scotophilus  rusticus  Tomes. 
Neoromicia  Roberts,  1926:  245.  Eptesicus  zuluensis  Roberts.  Valid  as  a  subgenus. 
Vansonia  Roberts,  1946:  304.  Pipistrellus  vernayi  Roberts  =  Vespertilio  ruppellii  Fischer. 
Vespadelus  Iredale  &  Troughton,  1934:  iii,  95.  Scotophilus  pumilus  Gray.  Nomen  nudum. 
Vespadelus  Troughton,  1943:  348.  Scotophilus  pumilus  Gray.  Valid  as  a  subgenus. 
Registrellus  Troughton,  1943:  349.  Pipistrellus  regulus  Thomas  (see  Hill,  19666). 
Falsistrellus  Troughton,  1943:  349.  Vespertilio  tasmaniensis  Gould.  Valid  as  a  subgenus. 
Perimyotis  Menu,  1984: 409, 415.  Vespertilio  subflavus  F.  Cuvier.  Valid  as  a  subgenus. 
Parastrellus  Horacek  &  Hanak,  1985a:  unpaginated;  19856:  62;  1985-1986:  15,  fig.  4.  Pipistrellus  Hesperus 

H.  Allen.  Nomen  nudum. 

The  genus  Pipistrellus  cannot  be  diagnosed  by  conventional  morphological  characters  that  are 
individually  exclusive.  Its  current  definition  rests  on  Miller  (1907)  who  based  his  diagnosis  on  the 
structure  of  i2  which  is  simple  or  has  a  well  developed  secondary  cusp;  on  the  reduction  of  i3  which 
is  smaller  than  i2  but  nevertheless  extends  beyond  the  cingulum  of  that  tooth;  on  rather  short 
canines,  c1  often  but  not  invariably  with  incipient  secondary  cusp  on  its  posterior  edge;  and  on  the 
absence  of  pmf  to  give  the  dental  formula  iff  |,  c},  pm^f  |,  m}2^  =  34,  with  pm2  barely  or  not  in  the 
toothrow.  He  remarked  that  the  members  of  the  genus  were  recognisable  by  their  dental  formula, 
large  i3,  unmodified  skull  and  ears,  and  the  normally  long  fifth  finger. 

The  definition  of  the  genus  is  briefly  discussed  by  Tate  (1942a),  Ellerman  &  Morrison-Scott 
(1951)  and  Kitchener  et  al.  (1986).  All  recognised  the  unreliability  of  the  presence  or  absence  of 
pm2  as  a  prime  diagnostic  character,  Ellerman  &  Morrison-Scott  also  remarking  that  'strictly 


VESPERTILIONINE  SYSTEMATICS  239 

speaking  Pipistrellus  is  not  more  than  a  subgenus  of  Eptesicus,  which  itself  might  be  referred  to 
Vespertilio',  but  for  convenience  they  and  most  other  recent  authors  have  followed  the  conven- 
tional distinction.  It  is  clear  from  the  foregoing  account  of  the  baculum  in  the  Vespertilioninae  that 
the  species  allocated  to  Pipistrellus  can  be  separated  from  most  other  vespertilionine  genera  by 
their  bacular  morphology:  those  genera  which  have  bacula  similar  to  those  of  some  Pipistrellus 
species  (e.g.  Nyctalus,  Scotozous)  can  be  defined  by  other  morphological  features  of  the  skull  and 
dentition,  as  they  were  by  Miller  (loc.  cit.). 

Species  groups  in  Pipistrellus  are  difficult  and  in  some  instances  almost  impossible  to  define  on 
external,  cranial  and  dental  characters:  most  (Tate,  19420;  Koopman,  1973,  1975)  are  brought 
together  by  combinations  of  characters  with  few  or  sometimes  no  exclusive  features.  Some  species 
of  Pipistrellus,  moreover,  appear  difficult  to  separate  from  some  of  Eptesicus  (Koopman,  1975; 
Heller  &  Volleth,  1984)  except  by  the  presence  or  absence  of  pm2  which  is  itself  evanescent.  Bacular 
morphology  appears  to  offer  at  least  a  partial  solution  to  this  difficulty,  at  the  same  time  indicating 
that  the  genus  as  currently  understood  is  a  composite  of  several  different  groups  of  species,  as 
suggested  by  Heller  &  Volleth  (loc.  cit.)  on  the  basis  of  its  known  karyology. 

Chromosomal  features  so  far  as  they  have  been  established  in  Pipistrellus  are  reviewed  by  Heller 
&  Volleth  (1984)  and  Zima  &  Horacek  (1985).  Their  summaries  demonstrate  that  karyologically 
Pipistrellus  as  currently  constituted  is  a  very  heterogeneous  and  diverse  group,  with  2N  varying 
from  26-44  and  FN  from  44-60.  At  this  stage  we  have  been  unable  to  find  any  consistent  correla- 
tion between  the  chromosome  formulae  that  these  authors  quote  for  various  species  and  the 
groupings  that  we  recognise  on  bacular  and  other  grounds.  However,  many  species  remain  to 
be  studied  karyologically  and  it  appears  from  Zima  &  Horacek  that  for  the  present  karyotype 
variability  in  the  Vespertilioninae  may  be  only  of  limited  value  as  a  taxonomic  criterion. 

While  at  present  we  would  not  support  the  generic  division  of  Pipistrellus  as  has  been  indicated 
or  suggested  by  Menu  (1984),  Horacek  &  Hanak  (19850,  b}  or  Kitchener  et  al.  (1986)  since  besides 
bacular  features  there  appear  to  be  few  or  no  characters  reported  for  its  constituent  groups  that 
would  support  this  wider  separation,  we  consider  that  the  divisions  apparent  within  the  genus 
justify  subgeneric  recognition.  Wallin  (1969)  has  already  anticipated  this  view  to  some  extent, 
employing  Hypsugo  Kolenati,  1856  for  P.  savii,  in  part  on  bacular  grounds.  Horacek  &  Hanak 
(1985-1986)  recognised  Hypsugo  as  a  distinct  genus.  The  gross  morphology  of  the  baculum 
also  indicates  that  the  Australian  taxa  formerly  referred  to  Eptesicus  should  be  transferred  to 
Pipistrellus,  as  Heller  &  Volleth  (1984)  suggested,  and  that  the  African  forms  hitherto  allocated  to 
the  capensis  and  tenuipinnis  groups  of  Eptesicus  also  represent  Pipistrellus  as  these  authors  inferred 
on  account  of  their  known  karyology.  Thus  we  would  classify  Pipistrellus  in  the  following  manner, 
listing  included  taxa  without  distinction  as  to  taxonomic  rank:  some  are  not  necessarily  valid 
species  or  subspecies  and  for  obvious  reasons  we  have  been  unable  to  examine  every  named  form  in 
the  genus. 

Subgenus  Pipistrellus  (Pipistrellus) 

Baculum  long,  with  strong,  extended  shaft,  well  developed  basal  lobes,  nearly  always  with  a  bifid 
tip.  Braincase  high,  rounded,  not  flattened,  sometimes  globose;  postorbital  region  usually  wide; 
cranial  profile  generally  straight  or  nearly  straight  from  occiput  to  nares;  interdental  palate  longer 
than  wide;  maxillary  toothrows  parallel  or  only  slightly  convergent  anteriorly;  i2  generally 
bicuspid;  pm2  usually  large,  pm2  not  greatly  reduced,  usually  about  3/4  crown  area  of  pm4. 
Pipistrellus  kuhlii  and  its  associates  differ  slightly  in  almost  unicuspid  i2;  greatly  reduced  i3,  much 
reduced  pm2  and  more  reduced  pm2  although  some  of  these  features  occur  in  isolation  in  other 
species  of  the  subgenus. 

(a)  pipistrellus  group 

Basal  lobes  of  baculum  more  or  less  in  line  with  the  bacular  shaft  in  the  vertical  plane;  i2  bicuspid, 

but  not  strongly  so,  cusps  not  deeply  divided. 

(a)  (i)  pipistrellus  subgroup.  Braincase  high,  rounded;  postorbital  region  wide;  supraorbital 
region  not  widened  or  swollen;  rostrum  long,  not  greatly  broadened,  with  shallow  median  rostral 


240  J.  E.  HILL  &  D.  L.  HARRISON 

depression;  cranial  profile  almost  straight  from  occiput  to  nares,  slightly  depressed  over  anterior 
part  of  orbit;  premaxillae  not  shortened;  zygomata  slender,  lacking  any  jugal  eminence;  interdental 
palate  longer  than  wide;  maxillary  toothrows  parallel  for  most  of  their  length,  anteriorly  slightly 
convergent;  short  bony  post-palate;  slight  basial  pits;  i2  bicuspid,  posterior  cusp  1/2-3/4  the  height 
of  anterior  cusp;  i3  about  the  same  in  crown  area  or  a  little  larger  than  i2,  about  1/2  or  a  little  more 
its  height,  with  larger  central  and  smaller  lateral  accessory  cusps,  lying  postero-externally  to  that 
tooth,  separated  from  c1  by  a  small  diastema;  pm2  large,  unreduced,  its  crown  area  similar  to  that 
or  i2  or  a  little  less,  slightly  intruded  but  separating  c1  and  pm4;  i  l  _  3  not  much  imbricated,  i3  about 
twice  the  bulk  of  i1  _2;  pm2  not  usually  much  reduced,  about  1/2-3/4  or  more  the  crown  area  of 
pm4. 

Included  taxa:  aladdin,  bactrianus,  lacteus,  nathusii  (Fig.  2b);  mediterraneus,  (?)  permixtus, 
pipistrellus  (Fig.  2a). 

Among  African  Pipistrellus  we  have  been  unable  to  examine  the  baculum  of  permixtus  (Aellen, 
1957)  compared  by  its  describer  chiefly  with  nathusii.  Its  dentition,  with  bicuspid  i2,  the  posterior 
cusp  2/3  the  height  of  the  anterior  cusp,  i3  with  lateral  accessory  cusps,  its  main  cusp  equal  in  height 
to  the  posterior  cusp  of  i2,  large,  slightly  intruded  pm2  which  is  about  as  big  as  i3,  and  unreduced 
pm2,  its  crown  area  about  3/4-4/5  the  crown  area  of  pm4  suggests  that  it  should  be  referred  to  the 
pipistrellus  subgroup.  Koopman  (1975)  referred  it  to  the  pipistrellus  group. 

(a)  (ii)  javanicus  (abramus)  subgroup.  Braincase  slightly  globular,  elevated  posteriorly;  post- 
orbital  region  wide;  supraorbital  region  distinctly  broadened  to  produce  abruptly  incurving  lateral 
margins  to  the  anterior  part  of  the  postorbital  area;  rostrum  broad,  dorsally  flattened,  with  no 
more  than  an  indication  of  a  median  rostral  depression;  cranial  profile  almost  straight  from 
occiput  to  nares,  slightly  flattened  over  the  occiput  and  a  little  depressed  over  the  anterior  part  of 
the  orbits;  premaxillae  not  shortened;  zygomata  slender  but  not  weak,  lacking  any  jugal  eminence; 
interdental  palate  only  little  longer  than  wide;  palate  strongly  domed  with  broad  anterior  emargi- 
nation;  maxillary  toothrows  more  or  less  parallel,  scarcely  convergent  anteriorly;  short  bony 
post-palate;  shallow  basial  pits;  i2  well  developed,  bicuspid,  posterior  cusp  sometimes  small, 
usually  about  3/4  height  of  anterior  cusp;  i3  similar  in  size  to  i2  or  slightly  larger,  about  as  high  as  its 
posterior  cusp,  with  larger  central  and  smaller  lateral  accessory  cusps,  lying  postero-externally  to 
that  tooth,  separated  from  c1  by  a  narrow  diastema;  pm2  little  reduced,  equal  to  or  rather  less  than 
i3  in  crown  area,  in  recess  between  c1  and  pm4  which  approach  but  do  not  touch;  ij_3  scarcely 
imbricated,  i3  as  a  rule  similar  in  size  to  i2,  both  a  little  more  massive  than  i^  pm2  about  1/2-3/4  the 
size  of  pm4,  very  slightly  intruded  from  toothrow. 

Included  taxa:  abramus  (Fig.  3a),  akokomuli,  babu  (Fig.  4a),  bancanus,  camortae  (Fig.  15d), 
endoi  (Fig.  3b),  irretitus,  javanicus  (Fig.  lOe),  meyeni,  peguensis  (Fig.  15c),  paterculus  (Fig.  3c), 
pumiloides. 

Current  treatments  of  Asian  Pipistrellus  usually  include  abramus  in  P.  javanicus  (tralatitius , 
Laurie  &  Hill,  1954)  as  a  valid  subspecies.  There  appear  to  be  few  conventional  features  that  clearly 
separate  javanicus  from  abramus  but  their  bacula  differ  quite  sharply  in  the  high  degree  of  vertical 
flexion  of  the  shaft  evident  in  the  latter.  This  difference  was  used  by  Thomas  (1928a)  who  examined 
Indo-Chinese  Pipistrellus  and  differentiated  abramus  from  raptor,  javanicus  (as  tralatitius)  and 
coromandra  by  virtue  of  the  double  curvature  of  its  baculum,  the  others  being  straight.  Van  Peenen 
et  al.  (1969)  recorded  coromandra,  javanicus  and  mimus  from  Vietnam  but  the  baculum  that  they 
illustrate  for  javanicus  is  clearly  that  of  abramus.  This  bacular  difference  suggests  thatjavanicus  and 
abramus  should  be  considered  specifically  distinct  even  although  there  seem  to  be  few  cranial  and 
dental  characters  to  separate  them.  The  braincase  in  javanicus  is  slightly  more  inflated  than  in 
abramus  and  its  rostrum  narrower,  the  palate  is  usually  a  little  wider  in  relation  to  its  length  and  is 
slightly  more  excavated  and  domed,  while  pm2  is  a  little  less  reduced  and  less  intruded,  tending 
rather  more  to  separate  c1  and  pm4.  Both  occur  in  Vietnam  (Thomas,  1928a;  specimens  listed 
below).  It  seems  likely  that  bancanus  and  camortae,  which  has  an  unflexed  baculum,  are  more 
closely  related  to  javanicus  than  to  abramus. 

Soota  &  Chaturvedi  (1980)  remarked  that  Thomas  (191 5c)  had  pointed  out  that  the  baculum  of 
abramus  is  doubly  curved  and  that  in  paterculus  it  is  straight,  but  they  stated  further  that  material  of 


VESPERTILIONINE  SYSTEMATICS  241 

paterculus  in  the  collections  of  the  Zoological  Survey  of  India  revealed  that  its  baculum  is  doubly 
curved.  However,  specimens  in  the  collections  of  the  British  Museum  (Natural  History)  referred  to 
paterculus  (some  the  original  material  seen  by  Thomas)  have  relatively  straight  bacula  when 
compared  with  the  sinuous  baculum  of  abramus.  We  have  found  this  sinuous  baculum  to  be 
characteristic  of  abramus,  to  which  perhaps  the  specimens  seen  by  Soota  &  Chaturvedi  should  be 
referred. 

The  very  elongate  baculum  of  paterculus,  with  its  strongly  bifid  tip,  the  'horns'  of  which  are 
deflected  ventrally  and  extend  to  some  extent  to  form  a  ring  (Thomas,  191 5c)  is  reminiscent  of  the 
baculum  of  Scotoecus.  A  very  long  baculum  is  also  found  in  endoi,  but  in  this  species  the  tip  is 
less  strongly  bifid  and  the  'horns'  are  deflected  dorsally.  Both,  however,  are  clearly  referable  to 
Pipistrellus  on  cranial  and  dental  characters,  Scotoecus  being  distinguished  especially  by  a  massive 
unicuspid  i2,  the  loss  of  i3,  a  grooved  c1,  and  usually  by  the  absence  of  pm2. 

(a)  (iii)  coromandra  subgroup.  Small,  with  small,  rounded  braincase,  elevated  posteriorly  and 
slightly  so  frontally;  postorbital  region  wide;  rostrum  short,  relatively  narrow;  no  median  rostral 
depression;  cranial  profile  straight  or  nearly  so  from  occiput  to  tip  of  rostrum;  premaxillae  excep- 
tionally short;  zygomata  slender,  without  jugal  projection;  interdental  palate  about  as  long  or  a 
little  longer  than  wide;  short  bony  post-palate:  no  basial  depressions;  i2  usually  bicuspid,  posterior 
cusp  sometimes  very  small  or  rarely  absent,  when  present  about  1/2  or  a  little  more  the  height  of  the 
anterior  cusp;  i3  equal  or  greater  than  i2  in  crown  area,  reaching  to  tip  of  its  posterior  cusp,  with 
larger  principal  cusp  and  smaller  lateral  accessory  cusps,  lying  postero-externally  to  the  inner 
tooth;  pm2  not  much  reduced,  nearly  as  great  or  as  great  in  crown  area  as  i3,  with  well  developed, 
slightly  inwardly  directed  pointed  cusp,  in  recess  between  c1  and  pm4;  ^  _  3  not  much  imbricated,  i3 
a  little  larger  than  i:  _2;  pm2  about  1/2  crown  area  and  height  of  pm4,  slightly  extruded. 

Included  taxa:  adamsi  (Fig.  lOc),  afghanus,  angulatus,  collinus  (Fig.  4b),  coromandra  (Fig.  7c), 
glaucillus,  mimus  (Fig.  7g),  murrayi  (Fig.  4c),  nitidus,  papuanus  (Fig.  2c),  ponceleti  (Fig.  4d), 
portensis,  principulus,  sewelanus,  sturdeei;  possibly  subulidens  which  may  however  represent 
javanicus;  tenuis  (Fig.  9d),  tramatus  (Fig.  7b),  wattsi  (Fig.  lOg),  westralis  (Fig.  lOd). 

(a)  (iv)  ceylonicus  subgroup.  Large,  with  rather  short,  broad  braincase;  wide  postorbital  region; 
some  degree  of  supraorbital  expansion;  rostrum  broad,  rather  long;  weak,  diffuse  median  rostral 
depression;  cranial  profile  slightly  convex,  raised  over  the  frontal  region;  premaxillae  normal,  not 
shortened;  zygomata  moderate,  without  jugal  eminence  or  process,  interdental  palate  longer  than 
wide;  maxillary  toothrows  parallel;  short  bony  post-palate;  slight  basial  pits;  i2  large  and  massive, 
bicuspid  to  almost  unicuspid,  with  moderate  to  small  posterior  cusp  about  2/3  height  of  anterior 
cusp;  i3  massive,  as  large  or  larger  than  i2,  extending  to  or  a  little  beyond  posterior  cusp  of  that 
tooth,  with  large  principal  cusp  and  smaller  lateral  accessory  cusps,  lying  postero-laterally  to  i2, 
narrowly  separated  from  c1;  pm2  large,  nearly  as  great  or  greater  in  crown  area  than  i3,  usually 
filling  the  recess  between  c1  and  pm4  into  which  it  is  intruded,  these  almost  in  contact  labially;  i:  _  3 
slightly  imbricated,  i3  a  little  larger  than  i1-2'  pm2  almost  as  large  in  crown  area  as  pm4,  very 
slightly  extruded  from  the  toothrow. 

Included  taxa:  borneanus,  ceylonicus  (Fig.  7d),  chrysothrix,  indicus,  (?)  minahassae,  raptor  (Fig. 
3d),  shanorum,  subcanus. 

An  account  of  minahassae  is  given  by  Tate  (\942a)  who  referred  it  to  a  minahassae  group  of 
which  it  was  the  sole  member.  The  skull  of  the  holotype  has  never  been  described  and  Tate's 
remarks  are  based  on  a  referred  specimen  in  the  American  Museum  of  Natural  History,  New  York 
(AMNH  102359).  It  has  a  short,  high  braincase  with  rudiments  of  a  sagittal  crest,  prominent 
supraorbital  tubercles  and  slender  zygomata;  i2  is  long,  with  well  developed  posterior  cusp,  c1 
slender,  lacking  an  accessory  cusp,  pm2  only  slightly  intruded,  its  crown  area  greater  than  that  of  i3, 
and  i^-j  scarcely  imbricated.  These  features  suggest  that  if  this  specimen  represents  minahassae 
the  taxon  should  be  allocated  to  Pipistrellus  (Pipistrellus)  and  provisionally  we  place  it  in  the 
ceylonicus  subgroup  of  the  pipistrellus  group,  but  clearly  these  decisions  can  only  be  speculative. 

(b)  rueppellti  group 

Baculum  as  in  pipistrellus  group;  braincase  high,  broadened,  rounded  and  globose;  postorbital 


242  J.  E.  HILL  &  D.  L.  HARRISON 

region  wide;  supraorbital  region  slightly  expanded;  rostrum  short;  with  shallow,  ill-defined  median 
depression;  cranial  profile  almost  straight,  a  little  raised  over  frontal  region,  a  little  depressed  over 
rostrum;  premaxillae  not  shortened;  zygomata  slender,  without  jugal  projection;  interdental  palate 
a  little  longer  than  wide;  maxillary  toothrows  slightly  convergent;  short  bony  post-palate;  no  basial 
pits,  instead  a  shallow  depression;  i2  strongly  bicuspid,  posterior  cusp  about  3/4  height  of  anterior 
cusp;  i3  usually  very  small  or  minute,  its  crown  area  less  than  1/2  that  of  i2,  its  tip  sometimes  barely 
rising  above  the  cingulum  of  the  inner  tooth,  on  occasion  (e.g.  nanulus)  larger,  equal  to  or  slightly 
exceeding  i2  in  crown  area,  about  1  /2  or  a  little  more  the  height  of  that  tooth;  i3  lying  sublaterally  to 
i2,  separated  from  c1  by  a  wide  diastema;  pm2  not  usually  greatly  reduced,  its  crown  area  similar  to 
that  of  i2,  with  strong  cusp,  separating  c1  and  pm4,  occasionally  (crassulus)  much  reduced,  similar 
in  size  to  i3  in  its  much  reduced  condition,  or  (crassulus,  nanulus)  recessed  between  these  teeth;  it  _  3 
little  imbricated,  i3  slightly  the  largest  as  a  rule;  pm2  about  3/4  or  more  as  large  in  crown  area  as 
pm4  and  about  3/4  its  height,  rarely  (coxi,  crassulus)  more  reduced,  about  1/2  crown  area  and 
height  of  pm4. 

Included  taxa:  Probably  coxi',  crassulus  (Fig.  le),fuscipes,  leucomelas,  nanulus  (Fig.  lf),pulcher 
(Fig.  lOa),  rueppellii  (Fig.  \0b),senegalensis,  vernayi. 

Vansonia  Roberts,  1946  is  available  should  further  separation  of  the  rueppellii  group  be  thought 
justified:  an  earlier  name,  Alobus  Peters,  1867  is  preoccupied. 

(c)  kuhlii  group 

Baculum  of  moderate  length  with  narrow  cylindrical  shaft  and  paired  basal  lobes  as  in  pipistrellus 
and  rueppellii  groups  but  basal  lobes  strongly  angled  to  line  of  shaft  in  vertical  plane;  braincase  low 
but  not  flattened,  rounded,  only  slightly  elongate;  postorbital  region  wide;  supraorbital  region  not 
widened  or  swollen;  rostrum  long,  unwidened,  with  very  slight  median  flattening;  cranial  profile 
almost  straight  from  occiput  to  nares,  slightly  raised  over  frontal  region,  slightly  depressed  over 
front  of  orbits;  premaxillae  slightly  shortened;  zygomata  slender,  weak,  without  jugal  eminence; 
interdental  palate  longer  than  wide;  maxillary  toothrows  almost  parallel;  short  bony  post-palate; 
small,  narrow  basial  pits;  i2  usually  unicuspid,  at  best  only  slightly  bicuspid;  i3  small,  its  crown  area 
1/2  or  less  that  of  i2,  its  tip  extending  only  slightly  beyond  the  cingulum  of  that  tooth,  to  which  it  lies 
laterally  or  sublaterally,  separated  from  c1  by  a  moderate  or  narrow  diastema;  pm2  small,  similar  in 
crown  area  to  i3,  intruded  to  lie  in  recess  between  c1  and  pm4,  these  more  or  less  in  contact;  it  _3 
moderately  imbricated,  i3  slightly  the  largest;  pm2  reduced,  about  1/2  or  less  the  crown  area  and 
height  of  pm4. 

Included  taxa:  Probably  aero;  deserti(aegyptius,  Qumsiyeh,  1985)  (Fig.  5c),fuscatus,  ikwanius; 
probably  inexspectatus;  kuhlii  (Fig.  5a),  maderensis  (Fig.  5b),  marrensis,  rusticus  (Figs  5d,  6c). 

We  have  been  unable  to  examine  the  baculum  of  inexspectatus  (Aellen,  1959)  but  this  taxon  was 
placed  in  the  kuhlii  group  by  Koopman  (1975)  who  also  referred  maderensis  to  the  savii  group. 
However,  an  example  of  maderensis  in  the  collections  of  the  British  Museum  (Natural  History)  has 
a  baculum  clearly  of  the  kuhlii  type. 

Romicia  Gray,  1838  is  available  for  the  kuhlii  group  should  this  be  thought  worthy  of  further 
separation. 

Subgenus  Pipistrellus  (  Vespadelus) 

Baculum  usually  with  long  cylindrical  or  ventrally  slightly  fluted  shaft,  paired  basal  lobes  and  a 
blunt  tip;  shaft  shorter  and  wider  in  sagittula;  basal  lobes  sometimes  flexed  to  line  of  shaft  in 
vertical  plane;  braincase  slightly  broadened,  flattened  and  elongated;  postorbital  region  wide; 
supraorbital  region  slightly  broadened;  rostrum  short  but  not  greatly  widened;  shallow  median 
rostral  depression;  cranial  profile  almost  straight  from  occiput  to  nares,  a  little  depressed  over 
rostrum;  premaxillae  not  shortened;  zygomata  slender,  without  jugal  process;  interdental  palate  a 
little  longer  than  wide;  maxillary  toothrows  slightly  convergent  anteriorly;  short  bony  post-palate; 
no  basial  pits:  i2  bicuspid,  posterior  cusp  almost  as  high  as  anterior  cusp;  i3  much  reduced,  its 
crown  area  1  /2  or  less  that  of  i2,  its  tip  barely  extending  beyond  the  cingulum  of  that  tooth,  to  which 
i3  lies  postero-laterally,  separated  from  c1  by  a  narrow  diastema;  pm2  almost  invariably  absent, 


VESPERTILIONINE  SYSTEMATICS  243 

when  present  a  small  spicule  in  recess  between  c1  and  pm4:  i{  _3  moderately  imbricated,  i3  slightly 
the  largest,  pm2  greatly  reduced,  in  crown  area  about  1/2  or  more  usually  less  the  crown  area  of 
pm4,  and  1/2  its  height. 

Included  taxa:  caurinus (Fig.  lib), douglasorum (Fig.  1  ld),pumilus (Figs  1  la,  12k),  regulus (Fig. 
1  le),  sagittula  (Fig.  1 1 0,  vulturnus  (Fig.  1  Ic). 

Formerly  referred  to  Eptesicus,  the  transfer  of  these  taxa  to  Pipistrellus  was  first  suggested  by 
Heller  &  Volleth  (1984),  purely  on  bacular  grounds.  The  bacular,  cranial  and  dental  features  of  this 
group  suggest  that  it  represents  P.  (Pipistrellus)  in  Australia,  the  few  members  of  this  subgenus 
(adamsi,  westralis  and  perhaps  javanicus)  that  also  occur  there  being  possibly  slightly  less  differen- 
tiated by  virtue  of  their  relatively  slightly  less  shortened  skulls  and  their  retention  of  pm2.  The 
pipistrellus  group  of  the  subgenus  extends  widely  through  the  islands  of  Indo- Australia  to  New 
Guinea,  the  Solomon  Islands  and  Australia,  chiefly  as  the  coromandra  subgroup,  to  which  adamsi 
and  westralis  belong.  Thejavanicus  subgroup  reaches  at  least  to  Java  and  Sulawesi  and  may  extend 
to  Australia  (Hill,  1983)  but  the  Australian  record  ofjavanicus  is  based  on  two  old  examples  and 
has  never  been  confirmed.  Possibly  the  slightly  differentiated  pumilus  and  it  allies  result  from  a 
further  perhaps  earlier  invasion  of  Australia.  Bacular  differences  in  this  subgenus  (Figs  11,  12k) 
suggest  that  it  may  consist  of  two  groups:  it  has  been  possible  to  examine  only  pumilus. 

There  has  been  hitherto  a  wide  geographical  hiatus  in  the  Indo-Australian  distribution  of 
Eptesicus  as  formerly  understood.  Beyond  these  Australian  forms,  no  other  taxon  attributed  to 
this  nominal  genus  has  been  reported  further  east  in  Indo-Australia  than  southern  Thailand,  other 
than  an  unconfirmed  record  from  Sarawak  of  Eptesicus  sp.  (Pirlot,  1968)  which  provided  no 
details. 

Subgenus  Pipistrellus  (Perimyotis) 

Baculum  very  small,  Y-shaped,  with  paired  basal  lobes  and  very  short  shaft;  braincase  slightly 
elongate,  rounded,  almost  globose;  postorbital  region  wide;  supraorbital  region  slightly  broad- 
ened; rostrum  long,  elevated,  slightly  widened;  shallow  median  frontal  depression;  a  very  slight 
lateral  depression  on  each  side  just  anterior  to  the  orbital  rim;  cranial  profile  sinuous,  raised 
over  frontal  region,  a  little  depressed  over  front  of  orbits;  premaxillae  not  shortened;  zygomata 
moderate,  a  slight  jugal  eminence;  interdental  palate  longer  than  wide  with  wide  anterior  palatal 
emargination;  maxillary  toothrows  convergent  anteriorly;  very  short  bony  post-palate;  slight 
basial  depressions;  i2  bicuspid  with  well  developed  posterior  cusp  about  3/4  height  of  anterior  cusp; 
i3  massive,  its  crown  area  exceeding  that  of  i2,  in  height  reaching  or  exceeding  the  height  of  anterior 
cusp  of  that  tooth,  with  larger  principal  cusp  and  smaller  lateral  accessory  cusps,  lying  postero- 
externally  to  inner  tooth,  separated  from  c1  by  a  wide  diastema;  pm2  large,  its  crown  area  equal  to 
that  of  i3,  in  toothrow,  sometimes  separated  from  pm4  by  a  slight  diastema;  i2"3  and  pm2  almost 
identical  to  those  of  P.  nathusii;  i^_3  not  imbricated,  i3  only  slightly  bulkier  than  i:_2;  pm2  not 
greatly  reduced  or  compressed  in  toothrow,  its  crown  area  about  1/2  or  more  that  of  pm4,  about 
1/2-3/4  its  height;  tragus  myotine,  about  1/2  height  of  ear,  tapering  to  blunt  point. 

Included  taxon:  subflavus  (Fig.  2d). 

Menu  (1984)  proposed  the  genus  Perimyotis  for  P.  subflavus,  chiefly  on  account  of  the  features  of 
the  canine  and  post-canine  dentition  in  which  he  believed  this  species  to  approach  Myotis.  How- 
ever, Hill  &  Topal  (1973)  in  discussing  Myotis  rosseti  and  M.  ridleyi  which  also  combine  the  tragal 
features  of  Myotis  with  the  Pipistrellus  dentition  (pm|  absent)  noted  that  in  Myotis  i2  is  short  and 
broad,  its  posterior  cusp  wider  basally  than  the  anterior  cusp,  while  in  Pipistrellus  this  tooth  is 
linear,  often  narrower  posteriorly  than  anteriorly.  Also,  in  Myotis  the  principal  cusp  of  i3  is  equal 
to  or  exceeds  that  of  i2  in  height  and  the  tooth  is  often  hooked  to  produce  a  caniniform  apperance 
while  in  Pipistrellus  it  is  lower  and  is  not  hooked.  In  Myotis  i3  is  usually  much  larger  than  il  _ 2  but 
in  Pipistrellus  there  is  as  a  rule  no  such  great  distinction  in  size.  The  incisive  dentition  of  subflavus 
corresponds  closely  with  that  of  Pipistrellus. 

The  baculum  of  subflavus  is  of  a  type  not  found  in  Myotis.  Menu  (1984)  stated  on  the  basis  of 
published  figures  that  the  baculum  approached  that  of  certain  Myotis  and  more  particularly  that  of 
Plecotus  auritus.  We  find  no  significant  resemblance  to  the  morphologically  rather  stable,  saddle- 


244  J.  E.  HILL  &  D.  L.  HARRISON 

like  baculum  ofMyotis  and  although  there  are  some  similarities  with  the  bacula  ofPlecotus  auritus 
(Fig.  19g)  and  P.  teneriffae  (Fig.  14d),  that  of  P.  austriacus  (Fig.  19h)  is  nearer  in  structure  to  the 
myotine  baculum.  The  bacular  type  found  in  subflavus  occurs  in  a  similar  form  in  Pipistrellus 
circumdatus  (Fig.  2e),  P.  societatis  (Fig.  9c)  and  P.  cuprosus  (Fig.  9h).  There  are  considerable 
differences,  however,  between  subflavus  and  circumdatus  and  its  allies,  not  least  in  the  degree  of 
reduction  of  pm2,  this  tooth  in  these  three  species  being  very  small  or  absent. 

The  unshortened  rostrum  and  the  dental  features  of  subflavus  suggest  that  it  is  nearest  to  P. 
(Pipistrellus),  which  it  appears  to  represent  in  North  America.  We  find  hesperus,  the  other  North 
American  species  of  Pipistrellus,  to  belong  on  bacular  and  dental  grounds  to  P.  (Hypsugo).  Thus 
we  do  not  support  Menu's  view  (p.  410,  footnote)  that  Pipistrellus  is  limited  to  the  Old  World  and 
that  the  lines  leading  to  this  genus  did  not  enter  the  North  American  continent.  The  marked 
differences  between  subflavus  and  hesperus  indicate  two  quite  different  pipistrelline  groups,  as 
Hamilton  (1949)  remarked  in  relation  to  their  bacula,  but  bacular  and  dental  evidence  suggests 
alliance  to  established  Old  World  groupings,  the  baculum  of  subflavus  being  perhaps  a  reduced 
form  of  the  shafted  structure  found  in  P.  (Pipistrellus) ,  that  of  hesperus  a  modification  of  the  type 
found  in  P.  (Hypsugo) . 


Subgenus  Pipistrellus  (Hypsugo) 

Baculum  usually  short,  stout,  sometimes  with  expanded  base  and  tip;  base  rarely  bilobed,  some- 
times dorsally  elevated;  shaft  generally  flattened  dorso-ventrally,  sometimes  wide,  its  underside 
transversely  concave  or  fluted;  tip  ventrally  hollowed  as  an  extension  of  ventral  fluting  of  shaft, 
when  expanded  anteriorly  sub-square  or  slightly  rounded,  its  anterior  edge  sometimes  irregularly 
serrated,  tip  sometimes  downwardly  directed,  its  lateral  margins  on  occasion  forming  two  broadly 
based,  ventrally  directed  projections;  pm2  generally  much  reduced,  small,  minute,  or  rarely  absent. 
Wallin  ( 1 969)  considered  Hypsugo  a  valid  subgenus  within  Pipistrellus  but  included  only  P.  savii: 
Horacek  &  Hanak  (1985#,  b)  added  cadornae  and  pulveratus  and  suggested  the  elevation  of 
Hypsugo  to  generic  rank,  subsequently  (1985-1986)  widening  its  possible  content  and  considering 
it  generically  distinct. 

(a)  savii  group 

Postorbital  region,  supraorbital  region  and  rostrum  not  greatly  widened;  supraorbital  tubercles  if 

present  small  and  undeveloped. 

(a)  (i)  pulveratus  subgroup.  Braincase  elongate,  inflated;  postorbital  region  wide,  supraorbital 
area  not  broadened;  rostrum  long,  not  widened;  shallow  frontal  depression;  no  median  rostral 
depression;  broad,  shallow  lateral  depressions  above  anterior  part  of  orbit;  cranial  profile 
somewhat  sinuous,  depressed  over  front  of  orbits;  premaxillae  not  shortened;  zygomata  robust, 
with  very  slight  jugal  eminence;  interdental  palate  longer  than  wide;  maxillary  toothrows  almost 
parallel;  moderate  bony  post-palate;  no  basial  pits;  i2  bicuspid,  posterior  cusp  about  3/4  height  of 
anterior  cusp;  i3  large,  wide,  its  crown  area  equal  to  or  slightly  exceeding  that  of  i2,  reaching  to  tip 
of  the  posterior  cusp  of  that  tooth,  with  moderate  lateral  accessory  cusps,  lying  postero-externally 
to  the  inner  tooth,  separated  from  c1  by  a  moderate  diastema;  pm2  about  equal  or  nearly  equal  to 
i3  in  crown  area,  in  recess  between  c1  and  pm4  which  are  closely  approximated;  ^.3  slightly 
imbricated,  i3  a  little  the  largest;  pm2  a  little  less  than  1/2  the  crown  area  of  pm4, 1/2-3/4  its  height. 
Included  taxon:  pulveratus  (Fig.  8c). 

(a)  (ii)  nanus  subgroup.  Braincase  elevated,  slightly  inflated,  more  or  less  globose  but  a  little 
elongated;  postorbital  region  wide;  supraorbital  area  slightly  widened  with  small  supraorbital 
swellings;  rostrum  not  especially  shortened  or  broadened;  shallow  median  rostral  depression; 
slight  lateral  depressions  just  anterior  to  supraorbital  region;  cranial  profile  sinuous,  strongly 
depressed  and  concave  over  rostrum;  premaxillae  not  shortened;  zygomata  slender,  lacking  jugal 
projection;  interdental  palate  longer  than  wide;  maxillary  toothrows  slightly  convergent;  short 
bony  post-palate;  no  basial  pits;  i2  unicuspid  or  with  small  posterior  cusp  extending  to  about  3/4  of 


VESPERTILIONINE  SYSTEMATICS  245 

its  height;  i3  wide,  its  crown  area  slightly  exceeding  that  of  i2,  about  1/2-3/4  the  height  of  that  tooth, 
extending  almost  to  the  tip  of  its  posterior  cusp,  with  slight  lateral  cusps,  lying  postero-externally 
to  the  inner  tooth,  separated  from  c1  by  a  wide  diastema;  pm2  about  1/2-2/3  or  a  little  more  the 
crown  area  of  i3,  intruded  into  recess  between  c1  and  pm4,  these  sometimes  in  contact  or  nearly  so; 
i:  _  3  not  or  only  very  slightly  imbricated,  i3  slightly  the  largest;  pm2  about  1/2  the  crown  area  and 
height  of  pm4. 

Included  taxa:  arabicus  (Fig.  7a),  culex,  Helios  (Fig.  6d);  probably  musciculus;  nanus  (Fig.  6b), 
stampflii. 

Current  listings  (i.e.  Hayman  &  Hill,  1971;  Koopman,  1975)  unite  Helios  with  P.  nanus  as  a 
synonym  or  possibly  as  a  valid  subspecies.  However,  the  bacular  features  of  this  pale  form  suggest 
that  it  may  represent  a  species  distinct  from  nanus  with  which  it  may  be  sympatric  in  northern  and 
eastern  Kenya  and  in  the  Sudan.  No  baculum  has  been  available  for  musciculus,  which  was  placed 
in  a  Hesperus  group  by  Koopman  (loc.  cit.).  Although  its  incisive  and  premolar  dentition  agrees 
with  the  nanus  subgroup  its  placement  here  remains  speculative. 

(a)  (in)  savii  subgroup.  Braincase  rather  low  and  flat,  elongate  rather  than  globose;  postorbital 
region  not  especially  widened;  supraorbital  region  unwidened  or  only  slightly  widened;  rostrum  of 
moderate  length;  a  shallow  median  rostral  depression;  usually  slight  lateral  rostral  depressions  just 
anterior  to  supraorbital  and  anterior  orbital  rim;  cranial  profile  straight  or  slightly  concave; 
premaxillae  not  shortened;  zygomata  robust,  often  with  slight  jugal  process;  interdental  palate  a 
little  longer  than  wide;  maxillary  toothrows  more  or  less  parallel;  short  bony  post-palate;  shallow 
or  no  basial  pits;  i2  unicuspid  or  with  posterior  cusp,  when  present  about  3/4  height  of  anterior 
cusp;  i3  similar  to  or  exceeding  i2  in  crown  area,  about  1/2-3/4  the  height  of  i2,  with  strong 
central  cusp  flanked  by  smaller  lateral  accessory  cusps,  lying  postero-externally  or  more  laterally 
(anchietae)  to  the  inner  tooth,  separated  from  c1  by  a  strong  diastema;  pm2  much  reduced,  minute 
or  absent,  when  present  crown  area  less  than  1/2  that  of  i3,  in  recess  between  c1  and  pm4,  these  as  a 
rule  in  contact;  \^  _  3  slightly  or  more  strongly  imbricated,  similar  in  size  or  i3  slightly  the  bulkiest; 
pm2  reduced,  about  1/2  or  less  in  crown  area  than  pm4  and  about  2/3  its  height. 

Included  taxa:  anchietae  (Fig.  6e);  probably  ariel',  probably  austenianus;  bodenheimeri  (Fig.  9f), 
caucasicus,  darwini,  maurus,  savii  (Fig.  6a). 

We  have  been  unable  to  examine  the  baculum  of  ariel.  The  baculum  of  a  small  Pipistrellus  from 
the  Naturhistorisches  Museum,  Wien  (from  Sayala,  Upper  Egypt)  tentatively  identified  as  ariel  is 
illustrated  by  Gaisler  et  al.  (1972)  but  is  evidently  of  the  kuhlii  type.  Qumsiyeh  (1985)  employs  the 
description  of  this  baculum  in  his  account  of  ariel.  However,  Dr  K.  Bauer  informs  us  (in  litt.)  that 
the  specimen  (NHW  10351)  of  which  the  baculum  is  figured  by  Gaisler  et  al.  (loc.  cit.)  is  not 
referable  to  ariel  but  is  instead  a  small  deserti,  an  identification  clearly  supported  by  its  bacular 
structure.  Moreover,  Dr  Bauer  has  loaned  three  similarly  small  specimens,  one  male,  the  others 
female  (NHW  27501-3)  (length  of  forearm  29-2,  28-9,  28-2;  condylobasal  length  11-0,  10-5.  10-9; 
c-m3  4-0,  3-8,  3-9)  apparently  from  Upper  Egypt,  that  also  represent  deserti:  a  baculum  from  this 
sample  is  again  exactly  of  the  kuhlii  type.  The  cranial  (narrow  braincase,  unexpanded  rostrum, 
short  broad  narial  and  anterior  palatal  emarginations,  narrow  basioccipital)  and  dental  (long  i3, 
minute  pm2)  features  of  ariel  clearly  indicate  that  it  belongs  with  savii,  to  which  group  Koopman 
(1975)  referred  it. 

\syntype  of  Eptesicusbicolor(Bocage,  1889)(BM(NH)  89.5. 1.3)  (Fig.  9e)  proves  to  be  identical 
cranially,  dentally  and  in  bacular  morphology  with  Pipistrellus  anchietae  (Seabra,  1900)  (vide 
infra,  p.  249).  However,  the  point  needs  confirmation  or  otherwise  by  examination  of  the  other 
syntype  in  the  Museu  Nacional  de  Lisboa.  It  should  be  noted  that  bicolor  is  the  prior  name 
(HonackietaL,  1982). 

The  relationship  between  the  pulveratus ,  nanus  and  savii  subgroups  is  illustrated  by  arabicus  and 
bodenheimeri,  the  bacula  of  which  are  compared  directly  by  Harrison  (1982).  The  baculum  of 
arabicus  (Fig.  7a)  approaches  that  of  anchietae  (Fig.  6e)  yet  cranially  and  dentally  this  species  is 
nearer  to  nanus  (Fig.  6b),  while  that  of  bodenheimeri  (Fig.  9f)  is  like  the  baculum  of  pulveratus  (Fig. 
8c)  but  cranially  and  dentally  the  species  is  close  to  savii  (Fig.  6a).  These  combinations  of  features 
link  the  three  subgroups. 


246  J.  E.  HILL  &  D.  L.  HARRISON 

(a)  (iv)  Hesperus  subgroup.  Baculum  a  fluted  structure,  much  like  that  ofpulveratus  or  bodenhei- 
meri.  Braincase  low  but  broad,  elongated;  postorbital  region  wide;  supraorbital  area  slightly 
widened;  rostrum  short,  not  greatly  broadened;  a  shallow  median  frontal  depression;  slight  lateral 
rostral  depressions  just  above  anteorbital  foramina;  cranial  profile  almost  straight,  slightly 
depressed  above  anterior  root  of  zygomata;  premaxillae  not  shortened;  zygomata  slender,  a  little 
widened  anteriorly,  lacking  any  jugal  eminence;  interdental  palate  about  as  wide  as  long;  maxillary 
toothrows  convergent;  short  bony  post-palate;  no  basial  pits;  cochlear  bullae  inflated  with  narrow 
basioccipital;  i2  unicuspid;  i3  slightly  greater  in  crown  than  i2  but  about  1/2  its  height,  with  little 
trace  of  lateral  accessory  cusps,  lying  postero-externally,  separated  from  c1  by  moderate  to  small 
diastema;  pm2  small  to  minute,  at  best  about  1/2  or  less  in  crown  area  than  i3,  in  recess  between  c1 
and  pm4  which  are  closely  approximated;  ij_3  scarcely  or  not  imbricated,  similar  in  size;  pm2 
reduced,  about  1/2  the  crown  area  of  pm4,  a  little  less  than  1/2  its  height. 

Included  taxon:  Hesperus  (Fig.  8d). 

Horacek  &  Hanak  (1985a,  b,  1985-1986)  have  indicated  that  they  intend  to  propose  generic 
status  for  Hesperus  and  indeed  have  suggested  that  it  be  referred  to  Parastrellus  which  they  offer  as  a 
new  name.  It  is  however  a  nomen  nudum  in  these  publications.  There  seem  good  grounds  for 
considering  Hesperus  the  North  American  representative  of  P.  (Hypsugo)  to  which  its  bacular, 
cranial  and  dental  features  ally  it.  Like  bodenheimeri  (Fig.  9f)  its  baculum  approaches  that  of 
pulveratus  (Fig.  8c)  but  cranially  and  dentally  it  is  nearer  to  savii&nd  its  immediate  allies.  Koopman 
(1975)  referred  Hesperus  to  a  Hesperus  group  in  which  he  also  included  the  African  musciculus,  here 
provisionally  allocated  to  the  nanus  subgroup. 

(a)  (v)  eisentrauti  subgroup.  Braincase  broad,  elevated  and  globular;  inflated  frontally;  post- 
orbital  region  wide;  supraorbital  region  broadened,  with  small  supraorbital  tubercles;  rostrum 
short,  deep,  wide  and  massive;  slight  median  rostral  depression;  cranial  profile  straight  or  slightly 
convex;  premaxillae  not  shortened;  zygomata  strong,  lacking  any  jugal  projection;  interdental 
palate  very  slightly  longer  than  wide;  maxillary  toothrows  almost  parallel;  short  bony  post-palate; 
slight  basial  pits  usually  present;  i2  long,  narrow,  bicuspid,  posterior  cups  about  3/4  height  of 
anterior  cusp;  i3  short,  wide,  similar  to  or  slightly  greater  in  crown  area  than  i2,  about  1/2  or  a  little 
more  its  height,  with  larger  central  cusp  and  smaller  lateral  accessory  cusps,  lying  laterally  and 
slightly  posteriorly  to  the  inner  tooth,  separated  from  c1  by  a  moderate  diastema;  pm2  small,  about 
the  same  in  crown  area  as  i3,  sandwiched  into  recess  between  c1  and  pm4,  these  almost  in  contact; 
it_3  slightly  imbricated,  i2_3  similar  in  size,  both  larger  than  it;  pm2  about  1/2  crown  area  and 
height  of  pm4. 

Included  taxon:  eisentrauti  (Fig.  9g). 

Koopman  (1975)  places  eisentrauti  in  a  rueppellii  group,  no  doubt  on  account  of  its  elevated, 
inflated  braincase  and  its  bicuspid  i2,  but  its  bacular  features  do  not  associate  it  with  this  species 
and  its  immediate  allies.  Its  baculum  is  very  similar  to  that  of  imbricatus  (Fig.  9a)  or  macrotis 
(Fig.  9b). 

(a)  (vi)  imbricatus  subgroup.  Braincase  inflated,  globular,  raised  posteriorly;  postorbital  region 
Wide;  supraorbital  area  slightly  widened  with  very  small  supraorbital  tubercles;  rostrum  short, 
not  especially  broadened;  no  median  rostral  depression;  cranial  profile  almost  straight;  slightly 
concave  above  supraorbital  region;  premaxillae  not  shortened;  zygomata  moderate  to  strong, 
sometimes  with  a  trace  of  a  jugal  eminence;  interdental  palate  about  as  wide  as  long,  not  domed; 
maxillary  toothrows  almost  parallel;  very  short  bony  post-palate;  well  developed  basial  pits;  i2 
bicuspid,  posterior  cusp  about  3/4  height  of  anterior  cusp;  i3  similar  in  crown  area  to  i2,  about  1/2 
its  height,  with  larger  central  cusp  and  smaller  lateral  accessory  cusps,  lying  laterally  to  the  inner 
tooth,  separated  from  c1  by  a  narrow  diastema;  pm2  greatly  reduced,  1/4  or  less  the  crown  area  of 
i3,  in  recess  between  c1  and  pm4,  these  in  contact;  il  _3  scarcely  imbricated,  i2_3  of  similar  size,  a 
little  larger  than  it;  pm2  about  1/2  the  crown  area  and  height  of  pm4. 

Included  taxa:  curtatus,  imbricatus  (Fig.  9a),  macrotis  (Fig.  9b),  vordermanni. 

(a)  (vii)  lophurus  subgroup.  Braincase  inflated,  rounded,  slightly  elongate,  raised  posteriorly; 
postorbital  region  wide;  supraorbital  area  little  widened;  at  best  only  a  trace  of  supraorbital 


VESPERTILIONINE  SYSTEMATICS  247 

tubercles;  rostrum  moderate  in  length,  longer  than  in  imbricatus  subgroup,  not  broadened;  no 
median  rostral  depression;  cranial  profile  almost  straight,  slightly  depressed  or  concave  above 
supraorbital  region;  zygomata  strong  with  distinct  jugal  eminence;  interdental  palate  a  little  longer 
than  wide;  maxillary  toothrows  slightly  convergent;  moderate  bony  post-palate;  deep  basial  pits; 
incisor  and  premolar  dentition  closely  similar  to  that  of  imbricatus  subgroup  but  i3  lying  more 
postero-laterally  to  i2,  and  pm2  sometimes  (lophurus)  slightly  larger,  about  1/2  crown  are  of  i3. 

Included  taxa:  cadornae,  kitcheneri  (Fig.  8e),  lophurus  (Fig.  80- 

The  baculum  of  kitcheneri  is  unusual  in  the  presence  distally  of  two  anterior  dorso-lateral, 
posteriorly  directed  processes,  with  ventrally  a  more  or  less  tapered  median  gutter.  As  in  lophurus, 
the  tip  is  directed  slightly  ventrally. 

(b)  stenopterus  group 

Braincase  large,  rounded  and  globular;  postorbital  region  very  wide;  supraorbital  region  much 
widened  to  include  well  developed  supraorbital  tubercles;  rostrum  short,  wide;  shallow  median 
rostral  depression  anterior  to  frontal  region;  cranial  profile  slightly  convex,  elevated  over  frontal 
area;  premaxillae  not  shortened;  zygomata  rather  weak,  lacking  jugal  process  but  usually  with 
small  descending  process  external  to  m3;  palate  short  and  broad,  the  interdental  palate  as  wide  as 
long;  maxillary  toothrows  parallel  or  nearly  so;  short  bony  post-palate;  shallow  basial  pits;  i2 
small,  bicuspid,  posterior  cusp  1/2-3/4  height  of  anterior  cusp;  i3  a  little  smaller  in  crown  area  than 
i2,  its  tip  reaching  almost  to  tip  of  the  posterior  cusp  of  that  tooth,  with  large  central  cusp  and 
smaller  lateral  accessory  cusps,  lying  postero-laterally  to  the  inner  tooth,  only  narrowly  separated 
from  c1  or  almost  in  contact  with  it;  c1  with  distinct,  well  defined  posterior  accessory  cusp;  pm2 
small  or  minute,  about  equal  in  crown  area  or  a  little  larger  than  i3  (stenopterus)  or  about  1/3-1/4 
the  crown  area  of  this  tooth  (joffrei,  anthonyi),  in  recess  between  c1  and  pm4,  which  touch;  it  _  3  not 
much  imbricated,  all  of  similar  size;  crown  area  of  pm2  slightly  exceeding  that  of  pm4,  pm2  similar 
in  height  to  the  second  tooth  (stenopterus),  or  crown  area  of  pm2  about  1/2  that  of  pm4,  pm2  almost 
as  high  as  that  tooth  (joffrei,  anthonyi). 

Included  taxa:  anthonyi,  joffrei,  stenopterus  (Fig.  7h). 

The  baculum  of  stenopterus  is  unusual,  although  of  the  savii  type:  it  has  a  narrow  lobed  base, 
hollowed  shaft,  and  expanded  tip  the  lateral  margins  of  which  project  ventrally  as  two  broadly 
based  'horns'.  The  stenopterus  group  as  here  understood  is  the  joffrei  group  of  Tate  ( 1 942a)  and  (in 
part)  of  Koopman  (1973).  Both  joffrei  and  stenopterus  have  been  referred  in  the  past  to  Nyctalus 
but  as  mentioned  above  (p.  234)  the  baculum  of  stenopterus  has  no  resemblance  to  the  long-shafted 
baculum  of  that  genus  (Fig.  100-  Tate  (19420)  referred  both  to  Pipistrellus  with  the  comment  that 
the  group  approached  Oriental  members  of  the  savii  group,  and  might  at  a  later  time  be  accorded 
generic  rank. 

Subgenus  Pipistrellus  (Falsistrellus) 

Baculum  a  broad,  proximally  widened  and  ventrally  deeply  fluted  structure  with  no  distal  expan- 
sion; braincase  elongate;  postorbital  region  wide;  supraorbital  area  not  expanded;  rostrum  long, 
not  broadened;  zygomata  moderate  to  strong;  and  palate  rather  narrow,  the  interdental  palate 
longer  than  wide. 

Pipistrellus  (Falsistrellus)  appears  to  be  related  to  P.  (Hypsugo)  of  which  it  may  be  the  eastern 
representative.  It  is  approached  in  bacular  morphology  by  some  of  the  latter  subgenus  such  as 
imbricatus  (Fig.  9a),  macrotis  (Fig.  9b),  kitcheneri  (Fig.  8e)  and  lophurus  (Fig.  8f),  and  indeed  the 
baculum  in  P.  (Falsistrellus)  appears  to  be  an  extreme  variant  of  the  broad,  ventrally  fluted 
structure  of  many  of  P.  (Hypsugo). 

Kitchener  et  al.  (1986)  raised  Falsistrellus  to  generic  rank  but  did  not  include  affinis  andpetersi, 
confining  their  comparisons  to  the  Australasian  Pipistrellus  (i.e.  adamsi,  angulatus,  collinus, 
papuanus,  wattsi  and  westralis)  here  referred  to  the  coromandra  subgroup  of  P.  (Pipistrellus).  These 
authors  drew  attention  to  its  larger  size;  to  its  small  i3  which  is  anteriorly  displaced  and  swivelled  or 
rotated  outwards  to  lie  alongside  i2,  its  concavity  facing  outwards  (a  feature  which  may  have 
influenced  Iredale  &  Troughton,  1934  in  placing  it  in  Glischropus);  and  to  its  combination  of 


248  J.  E.  HILL  &  D.  L.  HARRISON 

unicuspid  i2,  tiny  pm2  and  pronounced  occipital  crest,  which  asTate  (19420)  noted  gives  the  rear  of 
the  skull  a  'helmeted'  appearance.  Excepting  the  large  size  and  the  presence  of  a  strong  occipital 
crest,  these  features  occur  elsewhere  in  the  various  groups  of  Pipistrellus:  the  extent  of  the  occipital 
crest  may  be  a  function  of  the  large  size  of  the  skull. 

(a)  affinis  group 

Braincase  rather  narrow,  mastoid  width  markedly  less  than  zygomatic  width;  postorbital  region 
wide;  slight,  rather  poorly  developed  supraorbital  ridges;  very  shallow  median  rostral  depression 
just  anterior  to  frontal  region;  dorso-lateral  margin  of  rostrum  more  or  less  straight  from  post- 
orbital  constriction  to  anterior  edge  of  orbit;  cranial  profile  almost  straight,  slightly  elevated 
frontally,  depressed  over  supraorbital  region;  premaxillae  not  shortened;  zygomata  moderate  to 
robust  with  jugal  eminence;  maxillary  toothrows  slightly  convergent;  moderate  bony  post-palate; 
no  basial  pits;  i2  strongly  bicuspid,  posterior  cusp  3/4  or  more  the  height  of  the  anterior  cusp;  i3 
larger  in  crown  area  than  i2,  its  height  about  equal  to  the  height  of  the  posterior  cusp  of  that  tooth, 
with  strong  central  cusp  and  smaller  lateral  accessory  cusps,  lying  postero-laterally  to  the  inner 
tooth,  separated  from  c1  by  a  moderate  diastema;  pm2  almost  as  great  in  crown  area  as  i3  (affinis)  or 
about  3/4  its  crown  area  (petersi),  in  recess  between  c1  and  pm4;  i^  _3  moderately  imbricated,  i3  the 
largest;  pm2  about  1/2  the  crown  area  of  pm4  and  about  2/3-3/4  its  height. 

Included  taxa:  affinis  (Fig.  8a),  (?)  mordax,  petersi  (Fig.  8b). 

It  has  not  been  possible  to  examine  mordax.  Indian  records  of  this  species  appear  to  be  based  on 
specimens  in  the  collections  of  the  British  Museum  (Natural  History)  tentatively  labelled  as  such. 
These,  however,  agree  closely  with  the  description  of  P.  affinis  by  Dobson  (1871)  and  with  the 
account  of  a  specimen  referred  to  this  species  from  Likiang,  Yunnan  by  Tate  (19420).  If  correctly 
allocated,  mordax  (Peters,  1867)  is  the  earliest  name  in  the  group. 

(b)  tasmaniensis  group 

Large  and  distinctive;  braincase  high,  with  well  developed  sagittal  crest;  postorbital  region  wide 
and  strong;  no  median  rostral  depression;  cranial  profile  straight;  premaxillae  slightly  shortened; 
zygomata  strong  with  slight  jugal  process  and  small  inferior  process;  maxillary  toothrows  nearly 
parallel;  short  bony  post-palate;  slight  basial  depressions;  i2  large,  unicuspid;  i3  small,  its  crown 
area  about  1  /4  that  of  i2,  barely  extending  above  the  cingulum  of  that  tooth  to  which  it  lies  laterally, 
its  hollowed  face  outwardly  directed,  separated  from  c1  by  a  moderate  diastema:  pm2  very  small, 
about  1  /3  the  crown  area  of  i3,  in  recess  between  c1  and  pm4,  which  touch;  il  _  3  much  imbricated,  i3 
twice  the  bulk  of  il-2\  pm2  much  reduced,  about  1/4  the  crown  area  of  pm4  and  about  1/2  its 
height. 

Included  taxa:  mackenziei  (Fig.  lOh),  tasmaniensis  (Fig.  8g) 

Subgenus  Pipistrellus  (Neoromicia) 

Baculum  with  distinct  paired  basal  lobes,  slender  cylindrical  shaft  and  variously  expanded  tip; 
braincase  broad,  sometimes  slightly  elongate,  rather  flattened;  postorbital  region  wide;  supra- 
orbital  area  unwidened  or  only  slightly  broadened;  rostrum  moderate  or  slightly  lengthened; 
cranial  profile  straight;  premaxillae  shortened;  zygomata  moderate,  no  jugal  process;  maxillary 
toothrows  only  slightly  convergent;  short  bony  post-palate;  no  basial  pits;  i2  unicuspid  or  with 
small  posterior  cusp  extending  for  about  3/4  its  height;  i3  smaller  than  i2,  its  crown  area  3/4-1/2  or 
less  that  of  the  inner  tooth,  about  1/2  its  height,  with  usually  a  larger  central  cusp  and  slight  lateral 
accessory  cusps,  the  inner  cusp  as  a  rule  very  small,  the  tooth  anteriorly  displaced  to  lie  alongside  or 
almost  alongside  i2,  separated  from  c1  by  a  moderate  to  wide  diastema;  pm2  almost  invariably 
absent,  when  present  very  small,  in  recess  between  c1  and  pm4,  il_3  slightly  to  moderately  imbri- 
cated, of  similar  size  of  with  i2_3  a  little  the  larger;  pm2  reduced,  its  crown  area  1/2  or  less  that  of 
pm4  and  its  height  1/2-3/4  of  the  height  of  that  tooth. 

This  subgenus  is  wholly  African  and  hitherto  its  members  have  been  referred  to  Eptesicus, 
although  there  is  karyological  evidence  (vide  infra)  suggesting  that  one  at  least  should  be  moved  to 
Pipistrellus.  It  incorporates  the  Eptesicus  capensis  and  E.  tenuipinnis  groups  of  Koopman  (1975). 


VESPERTILIONINE  SYSTEMATICS  249 

These  can  be  recognised  readily  by  the  structure  of  the  baculum,  capensis  and  its  allies  (Fig.  12a-d, 
f-i)  having  the  distal  part  of  the  baculum  spatulate  and  ventrally  deflected,  tenuipinnis  and  its 
associates  (Fig.  12e,  j)  having  the  baculum  modified  distally  into  a  lobed,  almost  vertical  plate-like 
structure. 

Published  karyological  data  refers  only  to  capensis,  although  studies  of  other  members  of  the 
subgenus  are  in  progress  (Rautenbach  &  Schlitter,  1985a,  b).  Peterson  &  Nagorsen  (1975)  found 
that  capensis  has  a  diploid  number  of  32  and  a  fundamental  number  of  50:  Williams  &  Mares 
(1978)  discussed  the  possible  composite  nature  of  Eptesicus  as  suggested  by  Koopman  (1975)  and 
pointed  out  that  the  species  fitted  karyologically  within  the  variation  exhibited  by  Pipistrellus.  This 
genus  has  a  diploid  number  varying  from  26  to  44,  and  fundamental  numbers  from  44  to  60,  these 
findings  apparently  supporting  Koopman's  observations.  These  authors  remarked,  however,  that 
the  karyotype  of  capensis  is  more  similar  to  that  of  Pipistrellus  nanus  (2N  =  36,  FN  =  50)  than  to  P. 
kuhlii  (2N  =  44,  FN  =  50),  Koopman  having  thought  capensis  nearer  to  the  kuhlii  group  than  to  the 
pipistrellus  group  in  which  he  placed  nanus.  Williams  &  Mares  (loc.  cit.)  also  found,  in  contrast, 
that  small  Eptesicus  from  the  New  World  (diminutus,furinalis)  have  the  typical  'eptesicoid'  karyo- 
type (2N  =  50,  FN  =  48-50),  and  added  that  the  karyotypic  differences  between  Eptesicus  (sensu 
stricto)  and  Pipistrellus  might  prove  more  useful  for  separating  these  genera  than  other  structural 
features. 

Our  study  of  the  bacula  of  African  'Eptesicus '  confirms  these  observations  and  indicates  the 
isolation  of  capensis,  tenuipinnis  and  their  relatives  from  Eptesicus  sensu  stricto  (vide  infra):  Heller 
&  Volleth  (1984)  also  transferred  capensis  to  Pipistrellus,  entirely  on  account  of  its  published 
karyology.  It  is  interesting  to  note  also  that  the  baculum  of  P.  nanus  indicates  that  this  species 
should  be  referred  to  P.  (Hypsugo)  rather  than  to  P.  (Pipistrellus)  where  Koopman  (1975)  effec- 
tively allocated  it.  The  bacular  morphology  of  capensis,  tenuipinnis  and  their  allies  suggests 
strongly  that  these  former  groupings  of  Eptesicus  are  most  closely  allied  to  P.  (Hypsugo)  as  the 
karyological  similarity  of  capensis  to  P.  nanus  indicates.  The  anterior  upper  premolar  (pm2)  is  very 
small,  vestigial  or  absent  in  P.  savii  and  is  very  small  in  most  other  members  of  P.  (Hypsugo):  very 
rarely  it  is  present  in  capensis  (Wallin,  1969;  Hill  &  Topal,  1973).  On  the  same  point,  we  have  been 
able  to  examine  a  specimen  (MJS  2846)  from  Somalia,  in  the  Carnegie  Museum  of  Natural 
History,  which  has  a  small  pm2  on  both  sides  of  the  jaw,  leading  to  its  erstwhile  identification  as 
Pipistrellus  deserti.  The  baculum,  however,  is  characteristically  that  of  'Eptesicus' somalicus,  which 
in  fact  the  specimen  represents. 

Koopman  (1975)  suggested  that  Vesperus  bicolor  Bocage,  1889  (  =  Eptesicus  bicolor)  and 
Pipistrellus  anchietae  (Seabra,  1900),  both  from  Angola,  may  be  conspecific,  having  examined 
syntypes  of  both  at  the  British  Museum  (Natural  History).  This  author  thought  that  bicolor  might 
be  a  form  of  'Eptesicus '  tenuipinnis  as  Hayman  &  Hill  (1971)  suggested,  or  that  it  might  be  based  on 
a  specimen  of  Pipistrellus  anchietae  with  missing  anterior  upper  premolars.  Bocage  (loc.  cit.)  says 
'pas  de  trace  de  la  premiere  premolaire  a  la  machoire  superieure'.  Further  study  of  the  syntype 
(BM(NH)  89.5. 1 .3)  in  London  shows  it  to  have  a  small  pm2  in  a  recess  between  c1  and  pm4  on  each 
side:  cranially  it  agrees  exactly  with  the  syntype  of  anchietae  (9  BM(NH)  6. 1 .3. 1)  and  its  baculum  is 
exactly  as  in  that  species.  Curiously,  Bocage  states  that  both  original  specimens  of  bicolor  are 
female.  The  specimen  in  London  is  quite  clearly  listed  as  a  'Co-type'  by  Thomas  in  the  relevant 
accession  register. 

(a)  capensis  group 

Tip  of  baculum  flattened,  deflected  ventrally,  sometimes  a  small  sub-apical  dorsal  projection; 
braincase  flattened,  slightly  elongate;  rostrum  not  especially  broadened;  palate  long,  narrow, 
interdental  palate  longer  than  wide;  i3  1/2  or  less  the  crown  area  of  i2. 

Included  taxa:  capensis  (Fig.  12b,  g);  probably  brunneus,  garambae,  grandidieri;  guineensis 
(Fig.  12c),  matroka  (Fig.  12a),  melckorum  (Fig.  12f);  minutus  (?)  (Fig.  12i);  probably  rectitragus; 
somalicus  (Fig.  12h);  probably  vansoni;  zuluensis  (Fig.  12d). 

The  baculum  of  brunneus  sensu  stricto  has  not  been  examined.  That  (Fig.  14b)  of  a  Nigerian 
specimen  (BM(NH)  48.702)  collected  by  I.T.  Sanderson  and  hitherto  referred  to  this  species  is 
very  similar  to  that  of  rendalli  (Fig.  12e),  with  which  this  example  agrees  in  cranial  and  ventral 


250  J.  E.  HILL  &  D.  L.  HARRISON 

characters.  The  series  whence  this  specimen  comes  is  discussed  by  Koopman  ( 1 965)  and  Hayman  & 
Hill  (1971). 

Our  study  of  bacula  in  this  group  shows  clearly  that  mat r oka  belongs  with  capensis:  we  have  been 
unable  to  examine  the  baculum  of  humbloti.  We  find  too  that  capensis  and  somalicus  can  be 
separated  by  bacular  features:  the  baculum  of  capensis  has  distally  a  downwardly  directed,  plate- 
like  expansion,  while  in  somalicus  the  distal  part  of  the  baculum  is  more  spatulate,  depressed  just 
below  the  line  of  the  shaft.  Moreover,  zuluensis  is  very  clearly  of  the  somalicus  type,  and  the  two 
appear  to  be  very  closely  related,  as  Koopman  (1975)  suggested.  Bacular  morphology  also  con- 
firms the  observation  by  this  author  that  melckorum  is  like  a  giant  capensis:  Rautenbach  &  Schlitter 
(19850,  b)  suggested  that  these  are  synonymous. 

(b)  tenuipinnis  group 

Tip  of  baculum  expanded  into  an  almost  vertical,  lobed,  plate-like  structure;  braincase  similar  to 
capensis  group,  but  broader  and  less  elongate;  rostrum  slightly  widened;  palate  short  and  broad, 
interdental  palate  about  as  long  as  wide;  i3  about  1/2-3/4  the  crown  area  of  i2. 

Included  taxa:  Probably  angolensis,faradjius,flavescens,phasma;  rendalli  (Fig.  12e),  tenuipinnis 
(Fig.  12j). 

Pipistrellus  (Arielulus)  subgen.  nov. 

TYPE  SPECIES:  Vespertilio  circumdatus  Temminck,  1840.  Java. 
REFERRED  SPECIES:  Pipistrellus  societatis  Hill,  1 972;  Pipistrellus  cuprosus  Hill  &  Francis,  1 984. 

DISTRIBUTION:  Burma  to  Java  (circumdatus,  Fig.  2e);  Malaya  (societatis,  Fig.  9c);  Borneo 
(cuprosus,  Fig.  9h). 

DIAGNOSIS:  Differs  from  most  other  subgenera  of  Pipistrellus  in  very  small,  Y-shaped  baculum 
which  has  paired  basal  lobes  and  a  short  shaft;  baculum  similar  to  that  of  P.  (Perimyotis)  but 
differing  from  this  subgenus  in  greatly  reduced  i3  and  pm2,  the  former  displaced  anteriorly  to  lie 
alongside  i2,  the  latter  sometimes  absent. 

DESCRIPTION:  Size  small  to  medium  (length  of  forearm  34-7-43-6);  muzzle  short,  broad  and  blunt; 
ears  large,  rounded,  with  blunt  tip,  anterior  margin  with  prominent,  posteriorly  directed  basal 
lobe,  posterior  margin  with  wide  quadrate  lobe  at  insertion  just  behind  angle  of  mouth;  tragal 
margin  concave  anteriorly,  rising  to  anteriorly  directed  point,  upper  margin  of  tragus  nearly 
horizontal,  posterior  margin  strongly  convex;  ears  and  upper  margin  of  tragus  edged  to  a  greater  or 
lesser  extent  with  dull  white  or  yellowish  white;  dorsal  pelage  black  or  blackish  brown,  the  hairs 
tipped  with  yellowish,  orange,  russet,  copper  or  bronze. 

Braincase  high,  inflated,  globose;  postorbital  region  wide;  supraorbital  area  broadened,  with 
small  supraorbital  projections  or  tubercles;  rostrum  short,  widened,  sometimes  a  shallow  median 
rostral  depression;  cranial  profile  almost  straight,  elevated  frontally,  slightly  depressed  behind  and 
above  supraorbital  region;  premaxillae  not  shortened;  zygomata  strong,  no  jugal  eminence;  inter- 
dental palate  longer  than  wide;  short  to  moderate  bony  post-palate;  shallow  to  moderate  basial 
pits;  i2  almost  unicuspid,  posterior  cusp  if  present  insignificant;  i3  very  small,  about  1/4  the  crown 
area  of  i2,  1/3  or  less  its  height,  lying  almost  alongside  this  tooth,  separated  from  c1  by  a  narrow 
to  moderate  diastema;  pm2  very  small  or  absent,  when  present  similar  in  size  to  i3,  recessed  into 
angle  between  c1  and  pm4  which  are  in  contact;  it_3  considerably  imbricated,  \^_2  tricuspid  or 
incipiently  quadricuspid,  i2_3  bulkier,  larger  than  i1,  similar  in  size  to  each  other,  their  cusps 
indistinct;  pm2  1/2-1/4  size  of  pm4,  compressed  in  toothrow. 

ETYMOLOGY:  The  new  subgeneric  name  is  a  diminutive  of  Ariel,  a  little  sprite. 

REMARKS:  Heller  &  Volleth  (1984)  transfer  circumdatus  and  societatis  to  'Eptesicus'  on  karyo- 
logical,  bacular  and  dental  grounds.  However,  the  baculum  in  these  species  does  not  resemble 
closely  any  of  those  found  either  in  Eptesicus  sensu  lato  or  Eptesicus  sensu  stricto.  Possibly  the 
unusual  baculum  in  P.  (Arielulus},  similar  to  that  of  P.  (Perimyotis),  is  a  reduced  form  of  the  P. 


VESPERTILIONINE  SYSTEM ATICS  2  5 1 

(Pipistrellus)  type,  but  the  species  allocated  to  P.  (Arielulus)  differ  widely  cranially  and  dentally 
from  P.  subflavus,  the  sole  species  referred  to  P.  (Perimyotis). 

Genus  Eptesicus  Rafinesque,  1 820 

Eptesicus  Rafinesque,  1820:  2.  Eptesicus  melanops  Rafinesque  =  Vespertilio  fuscus  Palisot  de  Beauvois. 

Cnephaeus  Kaup,  1820:  103.  Vespertilio  serotinus  Schreber. 

Noctula  Bonaparte,  1837:  fasc.  xxi.  Noctula  serotina  Bonaparte. 

Cateorus  Kolenati,  1856:  131.  Vespertilio  serotinus  Schreber. 

Amblyotus  Kolenati,  1858:  252.  Amblyotus  atratus  Kolenati  =  Vespertilio  nilssonii  Keyserling  &  Blasius. 

PachyomusGray,  1866:  90.  Scotophilus  pachyomus  Tomes. 

Nyctiptenus  Fitzinger,  1870:  424.  Vespertilio  smithii  Wagner  =  Vespertilio  hottentota  A.  Smith. 

Rhinopterus  Miller,  1906:  85.  Glauconycteris floweri  De  Winton.  Valid  as  a  subgenus. 

Scabrifer  Allen,  1908:  46.  Substitute  for  Rhinopterus  Allen,  thought  preoccupied  by  Rhinoptera  Kuhl,  1841, 

Pisces. 

Pareptesicus  Bianchi,  1917:lxxvii.  VesperugopachyotisDobson. 
Rhyneptesicus  Bianchi,  1917:  Ixxvii.  Vesperugo  nasutus  Dobson. 
Rhineptesicus  Horacek  &  Hanak,  1985-1986:  16.  Lapsus. 

Baculum  more  or  less  triangular,  its  apex  occasionally  extended  into  a  short  shaft,  basally  rather 
wide,  sometimes  base  expanded  into  small  lobes,  tip  not  expanded,  usually  more  or  less  pointed  or 
gently  rounded.  There  is  little  flexion  in  the  vertical  plane  and  the  tip  is  not  depressed  ventrally; 
transversely  the  base  is  sometimes  slightly  arcuate.  Externally  and  cranially  not  essentially  different 
from  Pipistrellus  but  pm2  invariably  absent,  the  premolar  formula  being  ^  5  f. 

The  karyological  features  of  Eptesicus  are  summarised  by  Heller  &  Volleth  (1984)  and  Zima  & 
Horacek  (1985).  Such  as  have  been  examined  (andinus,  bottae,  brasiliensis,  diminutus ,  furinalis , 
fuscus,  guadeloupensis ,  hottentotus,  japonensis,  lynni,  nilssonii,  parvus,  serotinus,  turcomanus)  are 
homogeneous  in  this  respect,  with  2N  =  50,  FN  =  48-50.  On  present  published  knowledge  only 
capensis  differs  with  2N  =  32,  FN  =  50.  It  is  transferred  to  Pipistrellus  by  Heller  &  Volleth  (loc.  cit.) 
on  this  account  and  in  the  present  paper,  with  others,  on  bacular  grounds.  Pipistrellus  societatis  in 
which  2N  is  also  apparently  50  and  FN  48  is  transferred  to  Eptesicus  by  Heller  &  Volleth  (loc.  cit.) 
on  account  of  its  karyology  and  bacular  structure,  these  authors  considering  it  conspecific  with  P. 
circumdatus  (but  see  Hill  &  Francis,  1984).  Both  species  are  here  retained  in  Pipistrellus,  with  the 
closely  related  P.  cuprosus. 

Subgenus  Eptesicus  (Eptesicus) 

Postorbital  region  not  widened,  evident  postorbital  constriction;  rostrum  not  especially  shortened, 
its  dorsal  margins  not  sharply  angular;  cranial  profile  straight  or  slightly  concave,  not  elevated  over 
frontal  region;  maxillary  toothrows  almost  straight,  only  slightly  convergent;  upper  surface  of 
forearm,  tibia  and  tail  lacking  horny  excrescences. 

(a)  nilssonii  group 

Cranially  large,  the  skull  rather  elongate;  braincase  flattened,  elongate,  no  cranial  crests;  post- 
orbital  region  slightly  widened;  supraorbital  area  unwidened  but  with  very  small  supraorbital 
projections;  margins  of  supraorbital  region  almost  straight  from  postorbital  constriction  to  front 
of  orbit,  no  prominent  supraorbital  ridges  delimiting  upper  surface  of  rostrum;  the  rostrum  long, 
not  widened,  rounded  dorsally,  its  upper  surface  not  flattened  but  transversely  convex  above;  a 
shallow  median  rostral  depression;  slight  lateral  rostral  depressions  on  each  side  just  above  front 
of  orbit;  cranial  profile  straight  or  almost  straight,  slightly  concave  over  supraorbital  region; 
premaxillae  not  shortened;  zygomata  moderate  with  slight  jugal  process;  palate  long,  narrow, 
interdental  palate  longer  than  wide;  maxillary  toothrows  parallel  or  only  slightly  convergent;  very 
short  bony  post-palate;  prominent  basial  pits;  tympanic  bullae  not  enlarged,  not  completely  cover- 
ing cochleae;  i2  bicuspid,  posterior  cusp  about  1/2-3/4  height  of  anterior  cusp;  i3  wide,  as  large  or 
larger  than  i2  in  crown  area,  almost  reaching  tip  of  posterior  cusp  of  inner  tooth,  with  very  small 
lateral  accessory  cusps,  not  displaced  anteriorly,  lying  postero-laterally  to  i2  and  separated  from  c1 
by  a  modei  ate  diastema;  m3  not  greatly  reduced,  with  trace  of  fourth  commissure,  the  tooth  quite 


252  J.  E.  HILL  &  D.  L.  HARRISON 

long;  it  _3  slightly  imbricated,  i3  a  little  the  largest;  pm2  about  1/2  the  crown  area  and  height  of 
pm4,  not  compressed  in  toothrow. 

Included  taxa:  bobrinskoi  (Fig.  \3e),gobiensis,  nilssonii  (Fig.  15a). 

If  subgeneric  recognition  is  thought  justified  for  this  group  then  Amblyotus  Kolenati,  1858  is 
available.  The  nilssonii  group  was  recognised  as  subgenerically  valid  by  Tate  (1942a)  who  however 
included  within  it  a  number  of  taxa  here  allocated  to  the  nasutus  group  (vide  infra).  Strelkov  (1986) 
illustrated  the  bacula  of  nilssonii,  bobrinskoi  and  gobiensis,  considering  the  last  to  be  a  valid  species. 

(b)  nasutus  group 

Cranially  small,  the  skull  not  especially  elongate,  braincase  flattened,  only  slightly  elongate, 
broad;  postorbital  region  relatively  narrower  than  in  nilssonii  group;  supraorbital  area  slightly 
widened;  margins  of  supraorbital  region  nearly  straight  from  postorbital  constriction  to  front  of 
orbit,  supraorbital  ridges  sometimes  prominent;  rostrum  shortened,  its  upper  margins  slightly 
angular,  its  upper  surface  flattened  dorso-ventrally,  transversely  flat,  not  convex  as  in  nilssonii 
group;  a  shallow  or  sometimes  more  pronounced  median  rostral  depression,  slight  lateral  rostral 
depressions  above  front  of  orbit,  small  lateral  rostral  elevations  above  c1 "  ^cranial  profile  straight 
or  nearly  so,  sometimes  slightly  concave  above  supraorbital  region;  premaxillae  sometimes  slightly 
shortened;  zygomata  moderate,  on  occasion  a  slight  jugal  eminence;  interdental  palate  longer  than 
wide;  maxillary  toothrows  slightly  convergent;  short  bony  post-palate;  no  basial  pits;  tympanic 
bullae  very  large,  completely  covering  cochleae;  i2  large,  unicuspid;  i3  small,  about  1/2  crown  area 
and  height  of  i2,  with  larger  main  cusp  and  smaller  lateral  accessory  cusps,  anteriorly  displaced  to 
lie  alongside  or  almost  alongside  the  inner  tooth,  separated  from  c1  by  a  moderate  diastema;  m3 
sometimes  reduced,  usually  with  three  commissures,  no  trace  of  the  fourth,  antero-posteriorly 
rather  short,  compressed,  platelet-like;  i^  _  3  moderately  or  well  imbricated,  similar  in  size  or  with  i3 
slightly  the  largest;  pm2  very  small,  1/3-1/4  the  crown  area  and  1/2-1/3  the  height  of  that  tooth, 
compressed  in  toothrow. 

Included  taxa:  batinensis,  matschiei,  nasutus  (Fig.  I4c),  pellucens,  walli. 

Tate  (1942a)  included  walli,  matschiei  and  pellucens  in  the  nilssonii  group  but  these  agree  more 
appropriately  with  nasutus  as  De  Blase  (1980)  and  Honacki  et  al.  (1982)  recognised:  Ellerman  & 
Morrison-Scott  (1951)  listed  matschiei  and  pellucens  as  subspecies  of  nasutus.  Indeed,  Tate  (loc. 
cit.)  noted  the  large  tympanic  bullae  of  walli  and  the  absence  of  basial  pits  from  this  taxon. 
Rhyneptesicus  Bianchi,  1917  is  available  if  subgeneric  recognition  is  thought  justified  for  this 
group. 

(c)  serotinus  group 

(c)  (i)  serotinus  subgroup.  Cranially  large,  the  skull  elongate;  braincase  flattened,  elongate,  often 
with  lambdoid  and  sagittal  crests  forming  a  distinct  occipital  'helmet';  postorbital  region  slightly 
widened;  supraorbital  area  not  widened  or  only  slightly  so,  with  well  developed  supraorbital  ridges 
in  many  instances;  rostrum  long,  not  broadened,  its  upper  surface  flattened  but  less  so  than  in 
nasutus  group;  very  shallow  or  shallow  median  frontal  depression,  shallow  to  moderate  lateral 
frontal  depressions  just  above  front  of  orbit;  cranial  profile  almost  straight,  a  slight  concavity 
above  front  of  orbits;  premaxillae  sometimes  a  little  shortened;  zygomata  usually  robust  with 
moderate  jugal  projection,  on  occasion  slender  to  moderate,  the  projection  lacking;  palate  long 
and  narrow,  the  interdental  palate  longer  than  wide;  maxillary  toothrows  slightly  convergent; 
short  bony  post-palate;  shallow  basial  pits;  tympanic  bullae  not  covering  cochleae;  i2  bicuspid, 
posterior  cusp  about  3/4  height  of  anterior  cusp;  i3  small  to  very  small,  1/2-1/4  or  less  the  crown 
area  and  height  of  i2,  its  tip  1/2  or  less  the  height  of  the  posterior  cusp  of  that  tooth,  with  very  small 
lateral  accessory  cusps,  the  tooth  displaced  anteriorly  to  lie  alongside  or  almost  alongside  i2, 
separated  from  c1  by  a  moderate  to  small  diastema,  sometimes  almost  in  contact  with  that  tooth; 
m3  sometimes  much  reduced,  its  third  commissure  obsolescent  or  obsolete,  its  second  commissure 
short,  the  tooth  platelet-like;  ^.3  often  massive,  much  imbricated,  i3  the  largest;  pm2  about 
1/3-1/2  the  crown  area  and  1/2  the  height  of  pm4. 
Included  taxa:  andinus  (Fig.  13d),  argentinus,  bottae,  brasiliensis  (Fig.  13k),  dorianus,  fidelis, 


VESPERTILIONINE  SYSTEMATICS  253 

furinalis  (Fig.  \3c),fuscus  (Fig.  13a),  hingstoni,  hispaniolae  (Fig.  13i),  hottentotus,  inca,  innesi(Fig. 
13j),  innoxius,  isabellinus  (Fig.  13h),  megalurus  (Fig.  13b),  melanopterus ,  montosus,  omanensis 
(Fig.  \4a),pachyomus,peninsulae,platyops,punicus,  serotinus  (Fig.  13g),  shirazensis,  sodalis,  tatei, 
turcomanus. 

Tate  (19420)  has  pointed  out  that  the  Old  World  members  of  this  subgroup  fall  into  two 
categories,  one  of  larger  taxa,  the  other  of  smaller  members  of  the  subgroup.  This  is  also  true  of  the 
New  World  taxa:  however,  here  the  larger  forms  are  found  chiefly  in  North  America,  extending 
only  slightly  into  South  America  to  which  the  smaller  taxa  are  entirely  confined  (Thomas,  1920). 
Material  available  to  us  is  quite  inadequate  to  attempt  any  detailed  revision  and  we  have  followed 
the  lead  provided  by  Tate  (loc.  cit.)  in  our  allocation  of  all  to  the  one  category.  Cranial  differences 
between  large  and  small  members  of  the  subgroup  appear  chiefly  to  be  those  associated  with  size. 

The  subgroup  does  not  extend  substantially  into  Africa.  It  is  represented  in  Egypt  by  Eptesicus 
bottae  (innesi)  and  in  northwestern  Africa  by  E.  serotinus  (isabellinus}.  Ibanez  &  Valverde  (1985) 
consider  the  West  African  platyops  to  be  a  subspecies  of  serotinus,  as  may  be  the  South  African 
hottentotus  and  also  loveni  from  Kenya. 

(c)  (ii)  demissus  subgroup.  Eptesicus  demissus  Thomas,  1916  from  Thailand  appears  to  be  known 
only  from  the  holotype,  which  has  a  damaged  skull.  It  is  very  similar  to  the  larger  members  of  the 
serotinus  subgroup  but  has  a  long  bony  post-palate,  prominent  basial  pits,  i3  about  the  same  in 
crown  area  as  i2  and  about  1/2  its  height,  m3  not  especially  reduced,  its  third  commissure  complete, 
and  with  i1_3  moderately  imbricated,  i3  the  largest.  We  follow  Tate  (19420)  in  referring  it  to  a 
separate  subgroup  although  is  likely  that  more  adequate  material  might  enable  its  status  to  be 
determined  more  precisely. 

(c)  (iii)  (?)  pachyotis  subgroup.  We  have  been  unable  to  examine  Eptesicus  pachyotis  (Dobson, 
1871)  from  Assam.  Little  is  known  of  the  species,  of  which  the  holotype  is  in  the  Indian  Museum, 
Calcutta,  and  as  Tate  (19420)  pointed  out,  most  of  the  characters  given  by  Dobson  in  the  original 
description  might  apply  to  almost  any  species  of  Eptesicus.  The  generic  epithet  Pareptesicus 
Bianchi,  1917  was  proposed  for  this  taxon. 

Subgenus  Eptesicus  (Rhinopterus) 

Cranially  small;  braincase  low,  flattened  and  elongate,  inflated  anteriorly;  postorbital  region  wide; 
supraorbital  area  widened  with  very  small  supraorbital  tubercles;  anterior  margin  of  orbit  flange- 
like;  rostrum  short,  flattened  dorso-ventrally,  its  dorsal  margins  angular;  very  shallow  median 
rostral  depression,  shallow  lateral  depressions  just  above  front  of  orbit;  cranial  profile  convex, 
raised  above  frontal  region;  premaxillae  not  shortened;  zygomata  slender,  no  jugal  projection; 
palate  short,  broad,  interdental  palate  about  as  long  as  wide;  maxillary  toothrows  convergent; 
short  to  moderate  bony  post-palate;  no  basial  pits;  i2  bicuspid,  posterior  cusp  3/4  or  more  the 
height  of  the  anterior  cusp;  i3  small  or  minute,  about  1/4  or  less  the  crown  area  of  i2,  about  1/3-1/2 
its  height,  its  lateral  accessory  cusps  very  small  or  obsolete,  lying  postero-laterally  or  almost 
alongside  the  inner  tooth,  separated  from  c1  by  a  moderate  or  small  diastema;  m3  not  much 
reduced,  its  third  commissure  complete;  il_3  strongly  imbricated,  i2  the  smallest,  il  and  i3  of 
similar  size;  pm2  very  small,  about  1/2  crown  area  and  height  of  pm4,  strongly  compressed  in  row; 
horny  excrescences  on  upper  surface  of  forearm,  tibia  and  tail. 
Included  taxa.:floweri  (Fig.  13f),  lowei(Fig.  131). 

The  status  of  the  'NycticeinF 

An  especially  interesting  feature  emerges  from  our  survey  of  bacular  morphology  in  the  Vesperti- 
lioninae.  The  structure  of  the  baculum  suggests  very  strongly  that  the  'Nycticeini'  (or  'Nycticeiini') 
as  presently  accepted  is  not  a  natural  group.  Defined  chiefly  on  dental  characters  (i2  generally 
unicuspid,  i3  and  pm2  absent),  this  group  was  assembled  by  Tate  (19420)  to  include  Baeodon, 
Rhogeessa,  Otonycteris,  Nycticeius  (i.e.  N.  humeralis,  including  cubanus),  'Scoteinus'  (then  includ- 
ing among  others  the  Australian  species  now  referred  to  Scoteanax  and  Scotorepens),  Scotoecus, 
Scotomanes  and  Scotophilus.  Tate,  however,  made  no  mention  of  the  African  species  schlieffenii 


254  J.  E.  HILL  &  D.  L.  HARRISON 

which  was  extralimital  to  his  study  but  which  by  then  had  been  variously  referred  either  to 
Scoteinus  (Miller,  1907)  or  to  Nycticeius  (Hollister,  1918;  Braestrup,  1935).  More  recently,  the 
type  species  of  Scoteinus  (the  Indian  emarginatus)  has  proved  to  be  a  Scotomanes  (Sinha  & 
Chakraborty,  1971)  and  the  other  Indian  species  (pallidus)  formerly  referred  to  it  a  Scotoecus 
(Hill,  1974).  The  Australian  Scoteanax  and  Scotorepens  have  been  considered  to  be  subgenera  of 
Nycticeius  (Laurie  &  Hill,  1954;  Koopman,  1978;  Corbet  &  Hill,  1980)  but  recently  have  been 
accorded  generic  rank  (Kitchener  &  Caputi,  1984;  Corbet  &  Hill,  1986).  Thus  the  current  concept 
of  Nycticeius  is  of  two  species,  N.  humeralis  from  North  America  and  TV.  schlieffenii  from  Africa 
and  southwestern  Arabia. 

The  bacula  ofRhogeessa  (Fig.  1 8k)  and  Baeodon  (Fig.  1 5b)  are  quite  distinctive  and  are  variants 
of  the  saddle-like  or  slipper-like  structure  found  in  Myotis  and  Plecotus  or  their  allies,  as  are  the 
very  characteristic  bacula  of  Otonycteris  (Fig.  16a)  and  Nycticeius  humeralis  (Fig.  17k),  the  type 
species  of  Nycticeius.  Scotomanes  (Fig.  18g)  and  Scotophilus  (Fig.  17g-j)  have  bacula  reminiscent 
of  the  flattened,  triangular  structure  of  Eptesicus  and  its  immediate  associates.  In  contrast,  the 
bacula  of  Scotoecus  (Fig.  20a-e),  Nycticeius  schlieffenii  (Fig.  16e),  Scoteanax  (Fig.  16i),  and 
Scotorepens  (Figs  16g,  h,  21e,  f)  are  closely  similar  to  those  of  Pipistrellus  (Pipistrellus).  Thus  in 
bacular  terms  this  supposed  group  appears  to  be  a  composite  of  different  elements,  so  dissimilar 
among  themselves  that  its  unity  seems  very  unlikely.  Kitchener  &  Caputi  (1984)  contended  on 
the  grounds  of  a  phyletic  analysis  that  Otonycteris  and  Scotophilus  nigrita  fitted  poorly  into  the 
then  current  concept  of  the  'Nycticeini'  and  moreover  on  similar  considerations  that  Nycticeius 
humeralis  and  Nycticeius  schlieffenii  are  not  congeneric.  This  view  contrasts  sharply  with  that  of 
Koopman  ( 1 978)  who  remarked  that  the  latter  are  similar  in  all  important  characters  and  should  be 
retained  together  in  the  subgenus  Nycticeius  (Nycticeius). 

The  sharp  bacular  difference  between  the  American  humeralis  and  the  African  schlieffenii 
suggests  wider  separation  and  indicates  that  their  congeneric  association  is  wrong,  despite  their 
morphological  similarities  in  some  other  ways.  We  propose  therefore  to  dissociate  schlieffenii  from 
Nycticeius  as  generically  distinct.  The  newly  proposed  genus  may  be  called: 


Nycticeinops  gen.  nov. 

TYPE  SPECIES:  Nycticejus  schlieffenii  Peters,  1 860. 
REFERRED  SPECIES:  None. 
DISTRIBUTION:  Mauretania  to  Egypt,  Namibia  and  Mozambique;  SW  Arabia. 

DIAGNOSIS:  Baculum  (Fig.  16e)  distinctive,  with  expanded  base  and  long  fluted  shaft,  very  different 
from  that  of  Nycticeius  humeralis  (Fig.  17k)  which  is  slipper-like,  elevated  proximally  and  distally; 
cranially  similar  to  Nycticeius  sensu  stricto  but  rostrum  shorter,  more  narrowed  anteriorly,  the 
maxillary  toothrows  much  more  convergent,  not  nearly  parallel,  with  correspondingly  narrower 
narial  and  anterior  palatal  emarginations;  mandible  similarly  narrowed  anteriorly,  with  i1_3 
strongly  imbricated,  thrust  further  anteriorly  into  an  arc;  narial  emargination  more  clearly 
U-shaped,  not  prolonged  posteriorly;  anterior  palatal  emargination  extending  further  posteriorly; 
basial  depressions  absent  or  only  very  slight;  pm2  more  reduced.  Similar  to  Australian  Scoteanax 
and  Scotorepens  but  differing  sharply  in  bacular  morphology,  the  baculum  with  a  more  flanged  and 
fluted  shaft  and  lacking  the  modification  of  the  tip  found  in  these  genera,  and  in  less  reduction  of 
mf 

Differs  from  Pipistrellus  in  massive,  unicuspid  i2  which  has  no  trace  of  a  secondary  cusp,  in 
contact  or  nearly  so  with  c1,  the  premaxillae  greatly  shortened,  combined  with  the  almost  invari- 
able absence  of  i3  and  pm2.  Similar  in  some  respects  to  Scotozous  but  differing  in  bacular 
morphology;  in  the  presence  of  a  small,  posteriorly  directed  lobe  at  the  base  of  the  inner  margin  of 
the  ear;  tip  of  tragus  anteriorly  directed;  pm2  almost  invariably  absent;  pm2  more  reduced.  Similar 
also  in  some  ways  to  Scotoecus  but  penis  not  greatly  lengthened,  baculum  similarly  shorter,  its  tip 
not  expanded  and  bifid;  rostrum  narrower,  uninflated;  narial  and  anterior  palatal  emarginations 
not  extensively  deepened;  and  anterior  face  of  c1  rounded,  not  flattened  and  grooved. 


VESPERTILIONINE  SYSTEM ATICS  255 

DESCRIPTION:  Small  (length  of  forearm  about  29-33  mm);  muzzle  flattened,  anteriorly  sparsely 
haired,  nares  opening  obliquely;  ear  rounded  with  broadly  rounded  tip,  anterior  or  medial  margin 
with  small,  posteriorly  directed  basal  lobe,  anterior  margin  slightly  convex  for  most  of  its  length; 
posterior  margin  nearly  straight  distally,  more  convex  proximally  with  well  developed,  thickened 
antitragal  lobe;  tragus  with  bluntly  pointed,  anteriorly  directed  tip,  the  anterior  margin  strongly 
concave  basally,  straight  distally,  upper  margin  nearly  horizontal,  posterior  margin  strongly  con- 
vex, with  prominent  basal  lobe;  calcar  extending  along  a  little  more  than  one  half  of  the  uropatagial 
border;  well  developed,  rounded  post-calacarial  lobe  or  epiblema.  Dorsal  surface  of  head  and  body 
brown  to  pale  brown,  the  pelage  unicolored;  ventral  surface  paler  brown  to  greyish  white,  the 
pelage  usually  unicolored  but  in  the  darker  subspecies  faintly  bicolored,  the  hair  bases  darker  than 
the  tips. 

Skull  low,  with  broad,  flattened  braincase,  not  elevated  frontally;  very  low  cranial  crests  and 
very  slight  occipital  'helmet';  postorbital  region  wide;  supraorbital  area  a  little  broadened;  rostrum 
not  expanded  laterally,  narrow  anteriorly;  cranial  profile  almost  straight,  a  little  depressed  over 
front  of  orbits;  narial  emargination  U-shaped,  extending  posteriorly  one  half  of  distance  from  tip 
of  maxillae  to  a  line  joining  front  of  orbits;  premaxillae  much  shortened;  anteorbital  foramen 
moderate  to  large;  zygomata  slender,  no  jugal  projection;  palate  rather  short,  the  interdental  palate 
little  longer  than  wide,  narrowed  anteriorly,  maxillary  toothrows  convergent,  anterior  palatal 
emargination  narrow,  extending  posteriorly  to  a  line  joining  the  posterior  faces  of  c1"1,  not 
extending  laterally  beyond  the  inner  faces  of  i2"2;  short  to  moderate  bony  post-palate,  a  narrow 
median  post-palatal  spine;  basial  depressions  at  best  only  very  slight. 

Dental  formula  normally  if  23,  prnE^l  m|2f|  =  30.  Upper  incisor  i2  massive,  unicuspid,  usually 
separated  from  c1  by  a  very  short  diastema,  sometimes  in  contact  with  this  tooth;  i3  and  pm2  almost 
invariably  absent  (Thomas,  1 890;  Thomas  &  Wroughton,  1 908);  pm4  in  contact  with  c1 ,  with  small 
protocone;  lingual  shelves  of  m1  ~ 3  widely  separated,  m3  not  reduced,  with  three  commissures  and 
mesostyle,  about  1/2  crown  area  of  m1  or  m2;  ^  _3  strongly  imbricated  to  one  half  of  their  width, 
thrust  forward,  i  l  clearly  tricuspid,  i2  _  3  less  obviously  so,  i  ^  longest,  i2  _  3  more  massive;  pm2  much 
reduced,  1/2-1/4  crown  area  of  pm4  and  1/2  its  height,  compressed  in  toothrow;  m3  slightly 
reduced,  posterior  triangle  smaller  than  anterior  triangle,  hypoconid  and  entoconid  lower  than 
protoconid,  paraconid  and  metaconid. 

Thomas  &  Wroughton  (1908)  reported  a  specimen  (BM(NH)  8.4.3.23)  from  Tette,  Malawi  in 
which  a  well  developed  i3  is  present  in  the  left  side  of  the  jaw.  Dobson  (1878)  remarked  of  two 
specimens  in  the  Museum  National  d'Histoire  Naturelle,  Paris  that  pm2  is  present  on  one  side  in 
one,  on  both  sides  in  the  other,  but  Thomas  (1890)  who  examined  these  noted  that  pm2  is  com- 
pletely absent  from  one  and  present  on  both  sides  in  the  other,  Dobson  having  in  the  first  instance 
perhaps  mistaken  a  grain  of  sand  for  the  tooth.  Allen  (1914)  remarked  of  a  specimen  that  he 
identified  as  schlieffenii  from  Bados,  Blue  Nile  Province,  Sudan  that  pm2  was  present  on  both  sides 
of  the  jaw  and  that  'in  common  with  Scotoecus,  it  has  a  large  penial  bone,  12mm  long'  but 
Koopman  (1965)  pointed  out  that  in  fact  this  specimen  is  a  Scotoecus  (not  Scotophilus  as 
Qumsiyeh,  1985  avers)  and  that  schlieffenii  has  a  very  much  smaller  penis. 

INCLUDED  TAXA:  The  genus  is  monospecific,  its  sole  species  N.  schlieffenii  Peters,  1860.  Taxa 
allocated  to  it  either  as  valid  subspecies  or  synonyms  include  adovanus  Heuglin,  1877;  africanus 
Allen,  1911;  albiventer  Thomas  &  Wroughton,  1908;  australis  Thomas  &  Wroughton,  1908; 
bedouin  Thomas  &  Wroughton,  1908;  cinnamomeus  Wettstein,  \9\6\fitzsimmonsi  Roberts,  1932; 
minimus  Noack,  1887. 

ETYMOLOGY:  The  name  of  the  new  genus  is  derived  from  vu£,  VUKTOCT  or  vuKiioa,  night,  and  oy 
aspect. 

REMARKS:  The  type  species  schlieffenii  has  undergone  a  wide  variety  of  generic  allocations  and 
taxonomic  change  since  Peters  ( 1 860)  first  described  it  as  a  Nycticejus.  Dobson  ( 1 876, 1 878)  placed 
it  in  Scotozous  with  dormeri  while  under  the  impression  that  this  genus  lacked  i3,  and  considered 
(1878)  Scotozous  to  be  a  subgenus  of  Vesperugo.  Noack  (1877)  and  Thomas  (1890)  referred  it  to 
Scotophilus,  the  latter  author  discussing  this  genus  in  relation  to  Scotozous,  which  following 


256  J.  E.  HILL  &  D.  L.  HARRISON 

Dobson  he  thought  to  have  but  one  pair  of  upper  incisors.  Trouessart  (1897)  initially  followed 
Dobs~>n  (1878)  in  allocating  schlieffenii  to  Scotozous  as  a  subgenus  of  Vesperugo,  but  later  (1904) 
changed  this  opinion  to  consider  Scotozous  a  subgenus  of  Scotophilus.  Miller  (1907)  referred 
schlieffenii  to  Scoteinus,  although  in  fact  the  species  does  not  display  the  reduction  of  m|  that  he 
considered  diagnostic  for  this  genus  and  which  occurs  in  the  Australian  species  (balstoni,  greyii, 
now  incorporated  into  Scotorepens)  that  he  allocated  to  it.  Miller's  view  was  adopted  by  Thomas  & 
Wroughton  (1908)  and  in  differing  ways  by  many  subsequent  authors.  However,  Allen  (1911) 
when  describing  africanus  referred  it  to  the  hitherto  American  genus  Nycticeius,  commenting  on  its 
similarity  to  N.  humeralis  and  Hollister  (1918)  remarked  that  Old  World  bats  usually  placed  in  the 
genus  Scoteinus  did  not  seem  to  differ  generically  from  the  American  species  of  Nycticeius,  to  which 
he  also  referred  africanus.  Since  then  africanus  has  been  relegated  to  subspecific  status  or  synonymy 
in  schlieffenii  (Braestrup,  1935;  Allen,  1939;  Aellen,  1952).  Braestrup  (loc.  cit.)  also  employed 
Nycticeius  for  schlieffenii  in  preference  to  Scoteinus,  and  pointed  out  that  its  last  upper  molar  was 
not  reduced  in  the  way  that  Miller  (1907)  had  described  for  that  genus.  This  author  drew  attention 
to  the  affinity  thus  established  between  the  Ethiopian  and  American  faunas,  but  did  not  exclude  the 
possibility  of  convergent  evolution  from  different  Pipistrellus-\ike  forms.  Tate  (1942a)  maintained 
Nycticeius  and  Scoteinus  as  distinct  genera  but  Simpson  (1945)  united  them,  a  lead  followed  by 
many  modern  authors  who  have  considered  Scoteinus  a  subgenus  of  Nycticeius.  Thus  Ellerman  & 
Morrison-Scott  (1951)  and  Ellerman  et  al.  (1953)  referred  schlieffeni  to  Scoteinus  as  a  subgenus 
of  Nycticeius,  while  Laurie  &  Hill  (1954)  listed  the  Australian  species  before  then  allocated  to 
Scoteinus  in  Scoteanax  and  Scotorepens  as  further  valid  subgenera  of  Nycticeius.  On  the  other 
hand,  Rosevear  (1965)  considered  Nycticeius  and  Scoteinus  synonymous.  Koopman  (1965) 
referred  schlieffenii  to  Scoteinus  as  a  subgenus  of  Nycticeius  but  later  (in  litt.  in  Hayman  &  Hill, 
1971)  revised  this  opinion  to  allocate  it  to  Nycticeius  (Nycticeius),  since  then  (1978)  reinforcing  this 
view. 


The  classification  of  the  Vespertilioninae 

Earlier  classifications  of  the  Vespertilioninae  (Miller,  1907;  Tate,  19420)  rely  heavily  on  the  pattern 
of  reduction  of  the  incisor  and  premolar  teeth,  chiefly  on  the  presence  or  absence  of  the  outer  upper 
incisor  (i3),  of  one  or  both  of  the  first  (pm2)  or  second  (pm3)  upper  premolars,  and  on  the  presence 
or  absence  of  the  second  (pm3)  of  the  lower  premolars,  as  Tate's  'phyletic'  diagrams  (loc.  cit.) 
indicate.  These  dental  features  have  been  discussed  in  more  detail  above  (p.  230):  they  reflect 
the  degree  of  shortening  that  forms  an  evident  trend  within  the  subfamily.  When  combined  with 
the  relative  size  of  one  or  more  of  these  teeth  and  the  position  of  the  relevant  tooth  or  teeth  in  the 
toothrow  such  factors  form  an  important  element  in  generic  identification  and  diagnosis  (cf. 
Miller,  loc.  cit.).  The  many  different  combinations  of  incisive  and  premolar  formula  in  the  sub- 
family (Table  2),  the  evanescence  in  some  genera  of  some  of  the  teeth  involved,  the  extreme 
tendencies  towards  reduction  seen  in  some  such  as  Pipistrellus,  and  the  variety  of  positions  within 
the  toothrow  adopted  by  i3  and  pm2  in  particular  reinforce  the  conclusion  that  such  features  reflect 
a  universal  trend  that  may  have  occurred  more  than  once  within  the  group  and  which  as  a  result 
may  not  provide  a  totally  reliable  yardstick  by  which  relationship  may  be  judged. 

In  addition  to  these  dental  features,  Tate  (1942a)  reviewed  a  number  of  other  characters  used  in 
the  classification  of  the  subfamily.  These  include  the  presence  or  absence  of  accessory  canine  cusps; 
the  form  and  shape  of  the  braincase  and  rostrum;  the  degree  of  reduction  of  the  zygomata;  the 
structure  of  the  palate,  its  anterior  emargination  and  accessory  anterior  and  posterior  spines;  the 
presence  or  absence  of  basial  pits;  enlargement  of  the  ears  and  their  associated  bony  structures; 
the  presence  or  absence  of  adhesive  pads  on  the  thumb  or  foot;  and  the  nature  of  other  minor 
structures  such  as  the  calcar.  These  features,  however,  seem  of  greater  value  in  the  distinction  of 
species  and  species  groups,  that  is,  for  infrageneric  classification,  or  for  the  diagnosis  of  individual 
genera. 

The  value  of  such  characters  has  been  discussed  at  some  length  by  Zima  &  Horacek  (1985) 
who  pointed  out  that  there  are  grounds  for  thinking  that  some  of  the  traditional  morphological 


VESPERTILIONINE  SYSTEMATICS  257 

characters  may  not  provide  unequivocally  reliable  criteria  for  the  establishment  of  a  classification 
based  on  presumed  phyletic  relationship,  and  that  their  taxonomic  significance  may  be  limited. 
They  also  remarked  that  such  characters  may  reflect  parallelism  or  convergence,  or  result  from 
selection  pressure  rather  than  relationship.  These  reasons  led  them  to  suggest  that  the  baculum 
might  provide  one  of  several  alternative  sources  of  reliable,  taxonomically  useful  criteria  based  on 
characters  that  do  not  have  a  direct  adaptive  significance. 

The  structure  of  the  baculum  in  the  Vespertilioninae  suggests  some  modifications  to  tribal 
classification  within  the  subfamily,  although  clearly  other  morphological  characters  need  to  be 
given  equivalent  or  greater  weight.  Provisionally,  therefore,  we  offer  an  arrangement  of  the  Vesper- 
tilioninae in  which  bacular  morphology  is  used  in  association  with  the  traditional  diagnostic 
features  to  suggest  possible  relationship.  This  classification  is  presented  in  Table  1 . 

There  appear  to  be  two  major  bacular  types  in  the  Vespertilioninae,  each  with  numerous  varia- 
tions as  might  be  expected  in  such  a  large  and  diverse  subfamily.  A  classification  that  includes  a 
major  consideration  of  bacular  morphology  shows  significant  resemblances  to  earlier  arrange- 
ments based  on  traditional  and  conventional  morphological  features.  However,  there  are  some 
wide  divergences,  as  for  example  the  seemingly  artificial  nature  of  the  'Nycticeini'  or  the  associ- 
ations of  the  various  genera  of  big-eared  bats.  Tate  (19420)  commented  upon  the  latter  and 
pointed  out  that  very  large  ears  and  their  associated  auditory  specialisations  in  the  skull  occurred 
independently  in  three  sections  of  the  subfamily:  indeed,  if  Antrozous  and  Bauerus  are  included, 
these  features  occur  four  times  in  the  group.  In  particular,  both  Miller  (1907)  and  Tate  (loc.  cit.) 
associated  Laephotis  with  Histiotus  on  cranial  and  dental  morphology  but  its  bacular  structure 
shows  a  clear  affinity  with  Pipistrellus  (Neoromicid)  as  here  recognised.  Otonycteris,  another 
big-eared  bat,  was  allied  by  Tate  (loc.  cit.)  to  the  'Nycticeini'  but  proves  to  have  a  baculum  much 
more  like  those  of  the  plecotine  genera. 

One  major  bacular  type  is  'saddle-like'  or  'slipper-like'  and  is  exemplified  by  Myotis  and 
Pizonyx.  Their  bacula  are  very  similar,  emphasising  the  close  relationship  that  is  generally  accepted 
between  these  genera.  The  baculum  of  Lasionycteris  is  somewhat  different  in  the  presence  of  a 
lengthened  shaft.  However,  in  comparison  with  the  long-shafted  bacula  found  in  the  Pipistrellini 
the  baculum  of  Lasionycteris  is  relatively  short,  and  it  retains  indications  of  the  more  characteristic 
myotine  type  in  its  upraised  proximal  and  distal  portions.  The  occasional  presence  of  a  flattened 
dorsal  prominence  on  its  base  also  recalls  the  condition  found  in  Idionycteris.  The  genus,  although 
having  some  specialised  features,  is  allied  firmly  to  Myotis  by  Miller  (1907)  and  Tate  (\942a).  It  has 
slightly  hooked  upper  incisors,  i3  with  a  slightly  caniniform  profile  as  in  Myotis',  pm2  is  in  the  line  of 
the  toothrow;  m3  is  unreduced;  pm2  _  3  are  exactly  as  in  Myotis,  much  smaller  than  pm4,  with  pm3 
not  removed  from  the  line  of  the  other  teeth.  Although  pm3  has  been  lost,  this  appears  to  be  a 
specialisation;  as  Tate  (loc.  cit.)  pointed  out,  pm2  _  3  still  agree  closely  with  those  of  the  less 
specialised  species  of  Myotis  not  only  in  relation  to  each  other  but  also  in  their  proportional  size 
relative  to  pm4.  Although  associated  with  Myotis,  this  genus  is  considerably  specialised  in  other 
ways  (Miller,  loc.  cit.)  and  its  bacular  structure  may  well  reflect  this  divergence.  Its  baculum  might 
be  regarded  as  derived  from  the  more  typical  myotine  structure. 

Bacula  variously  reminiscent  of  the  saddle  shaped  structure  found  in  Myotis  occur  in  a  number 
of  other  genera.  Such  bacula  characterise  Plecotus  (including  Coryhorhinus),  Idionycteris, 
Barbastella,  Rhogeessa,  Baeodon,  Nycticeius,  Otonycteris,  Lasiurus,  Dasypterus,  Antrozous  and 
Bauerus,  and  possibly  may  be  found  in  Euderma.  Tate  (19420)  postulated  the  grouping  'Plecotini' 
for  Plecotus,  (Corynorhinus),  Idionycteris  and  Euderma,  allying  it  to  the  Myotini  but  not  employing 
the  term  in  a  formal  taxonomic  or  systematic  sense.  Bacular  morphology  thus  lends  support  to  his 
hypothesis  that  the  plecotine  genera  should  be  associated  with  Myotis.  Also,  the  baculum  of 
Barbastella  suggests  that  it  too  belongs  here:  Miller  (1907)  postulated  such  a  relationship,  despite 
several  morphological  differences.  Rhogeessa,  Baeodon,  Nycticeius  and  Otonycteris  also  seem 
allied  to  this  grouping.  Tate  (19420)  referred  these  genera  to  the  'Nycticeini'  with  Scotoecus, 
Scotomanes  and  Scotophilus  on  account  of  their  incisive  and  premolar  dentition.  However,  the 
bacula  of  Rhogeessa,  Baeodon,  Nycticeius  and  Otonycteris  are  variants  of  the  saddle-like  type;  that 
of  Scotoecus  is  like  that  of  Pipistrellus  (Pipistrellus),  and  the  bacula  of  Scotomanes  and  Scotophilus 
are  broadly  similar  to  those  ofEptesicus  and  its  allies.  Lasiurus,  Dasypterus,  Antrozous  and  Bauerus 


258  J.  E.  HILL  &  D.  L.  HARRISON 

have  further  variants  of  this  bacular  type,  but  are  quite  distinctive  on  other  morphological 
grounds. 

The  bacula  of  Antrozous  and  Bauerus  are  not  at  all  like  that  of  Otonycteris,  with  which  these 
genera  have  been  tentatively  associated  (Pine  et  al.,  1971),  nor  do  their  bacula  have  any  significant 
resemblance  to  those  of  Nyctophilus  or  Pharotis,  thus  supporting  the  view  (Koopman,  19846, 
1985;  Breed  &  Inns,  1985)  that  these  North  American  genera  should  not  be  associated  with 
the  Australian  Nyctophilus  and  Pharotis  in  the  subfamily  Nyctophilinae.  Bacular  morphology 
suggests  instead  an  association  with  those  genera  that  have  the  myotine  type  of  baculum,  to  which 
the  bacula  of  Antrozous  and  Bauerus  have  many  resemblances.  The  bacula  of  Nyctophilus  and 
Pharotis  (Fig.  22a-h)  are  consistently  homogeneous  and  differ  in  many  ways  from  those  of  the 
genera  usually  referred  to  the  Vespertilioninae.  For  the  present  we  would  place  these  two  genera  in 
a  separate  subfamily,  the  Nyctophilinae,  rather  than  merge  them  into  the  Vespertilioninae  as  is 
done  by  Koopman  (1984a,  19846,  1985). 

A  further  basically  triangular  and  flattened  variant  of  the  saddle-like  baculum  characterises  the 
genera  Eptesicus,  Vesper tilio  (if  the  pseudobaculum  is  ignored),  la  and  Histiotus.  Miller  (1907) 
remarked  that  the  skull  of  Vespertilio  showed  a  strong  likeness  to  that  of  Lasionycteris  but  that 
the  former  was  in  all  respects  a  typical  Eptesicus.  Vespertilio  and  Lasionycteris  are  separated 
by  marked  dental  and  bacular  differences:  the  bacular  morphology  of  Vespertilio  allies  it  with 
Eptesicus  as  Miller  suggested.  It  is  perhaps  not  unreasonable  to  speculate  that  Lasionycteris  which 
has  a  strongly  myotine  dentition  has  diverged  among  the  Myotini  in  the  same  way  as  Vespertilio 
has  diverged  among  the  Vespertilionini,  the  latter  genus  supporting  a  long  penis  either  by  a 
centrally  situated  baculum  or  perhaps  more  effectively  by  the  development  of  a  cartilaginous 
pseudobaculum,  this  function  in  Lasionycteris  by  a  short  shaft.  The  genera  Tylonycteris  and 
Mimetillus  also  belong  here.  The  African  Glauconycteris  has  been  associated  (Ryan,  1966; 
Koopman,  1971)  with  the  Australian  Chalinolobus  but  their  bacula  differ  widely.  Although  struc- 
turally variable  within  the  genus,  the  bacula  of  Glauconycteris  are  more  like  the  vespertilionine 
or  eptesicine  type:  those  of  Chalinolobus  are  long-shafted  and  like  the  bacula  of  Pipistrellus 
(Pipistrellus).  Finally,  the  baculum  ofScotomanes  appears  to  be  a  derivative  of  the  saddle-like  type, 
leading  to  the  distinctive  baculum  of  Scotophilus. 

The  genus  Pipistrellus  seems  to  stand  more  or  less  at  the  centre  of  the  second  major  grouping. 
It  has  broadly  two  divisions  in  bacular  terms,  one  characterised  by  a  long  baculum  with  well 
developed  basal  lobes  and  a  relatively  long,  mostly  cylindrical  shaft,  its  tip  often  bifid  or  with 
similar  elaboration.  The  second  division  includes  those  species  in  which  the  basal  lobes  are  some- 
times small  or  obsolete  and  which  have  a  shorter,  flatter,  ventrally  fluted  shaft,  its  tip  sometimes 
elaborated  into  a  spatulate  or  platelet-like  structure. 

These  groupings  have  been  used  in  this  study  to  support  subgeneric  division  of  this  large  genus. 
The  first  division  includes  Pipistrellus  (Pipistrellus},  P.  (Vespadelus),  P.  (Perimyotis}  and  P. 
(Arielulus}.  Reduction  and  loss  of  pm2  occurs  in  P.  (Arielulus)  and  the  tooth  is  almost  invari- 
ably absent  in  P.  (Vespadelus}.  The  second  division  contains  P.  (Hypsugo)  in  which  pm2  may  be 
very  small  or  absent,  P.  (Neoromicia)  from  which  it  is  again  almost  invariably  absent,  and  P. 
(Falsistrellus}.  Although  primarily  Old  World  in  distribution,  both  of  these  divisions  are  repre- 
sented in  the  New  World,  each  by  a  single  species.  The  Australian  P.  ( Vespadelus}  seems  on  bacular 
features  to  represent  P.  (Pipistrellus);  the  wholly  African  P.  (Neoromicia)  is  apparently  similarly 
related  to  P.  (Hypsugo),  of  which  P.  (Falsistrellus}  appears  to  be  an  eastern  representative. 

The  majority  of  the  genera  here  allocated  to  the  Pipistrellini  show  strong  bacular  affinities  to 
Pipistrellus  (Pipistrellus):  some  such  as  Glischropus  and  Scotozous  have  been  considered  congeneric 
with  Pipistrellus  in  the  past.  Besides  Glischropus  and  Scotozous  these  include  Nycticeinops, 
Scoteanax,  Scotorepens,  Scotoecus,  Nyctalus,  Hesperoptenus  and  Chalinolobus,  all  with  long- 
shafted  bacula.  Of  the  remainder,  Laephotis  in  bacular  structure  is  similar  to  P.  (Neoromicia},  while 
Philetor  has  a  baculum  that  appears  to  be  an  elaboration  of  the  bacular  structure  found  in  some  of 
P.  (Hypsugo).  Tate  (1942a)  postulated  a  relationship  between  Philetor,  Tylonycteris  and  perhaps 
Mimetillus  but  the  bacula  of  the  first  two  are  totally  dissimilar  and  the  structure  is  apparently 
absent  from  Mimetillus:  it  is  very  small  in  Tylonycteris.  Hill  (1966a)  drew  attention  to  the  unusual 
genitalia  of  Philetor  and  following  Tate's  (loc.  cit.)  suggestion  of  affinity  with  Pipistrellus  joffrei  and 


VESPERTILIONINE  SYSTEMATICS  259 

its  associates  allied  Philetor  with  this  group.  Unfortunately,  excepting  for  the  aberrant  species 
stenopterus  the  bacula  of  the  stenopterus  subgroup  (including  P.joffrei)  of  this  present  study  remain 
unknown. 

Bacular  morphology  suggests  that  the  conventional  view  that  Eptesicus  and  its  immediate  allies 
derive  from  or  are  closely  related  to  Pipistrellus  can  be  questioned.  Cranially  and  dentally  there  are 
many  similarities  between  'Eptesicus '  as  formerly  denned  and  Pipistrellus  and  as  Koopman  (1975) 
has  pointed  out,  the  loss  of  pm2  enables  a  species  to  cross  the  boundary  between  the  two  genera  as 
then  understood,  a  process  which  in  his  view  might  have  occurred  more  than  once.  Our  conclusions 
do  not  challenge  this  opinion:  those  'Eptesicus'  species  in  which  pm2  has  been  found  occasionally 
to  occur  prove  on  bacular  grounds  to  be  closer  to  Pipistrellus  than  to  Eptesicus  as  we  understand  it, 
while  Pipistrellus  as  formerly  defined  has  long  been  known  to  include  some  species  from  which  on 
occasion  this  'diagnostic'  tooth  is  absent.  Clearly,  our  findings  support  Koopman's  (loc.  cit.) 
opinion  that  this  process  may  have  occurred  several  times  and  indeed  may  be  occurring  in  some 
species,  but  all  belong  to  the  one  genus,  Pipistrellus. 

As  we  understand  its  composition,  Eptesicus  is  now  a  more  restricted  genus  in  which  the  tri- 
angular, flattened  baculum  is  basically  closer  in  structure  to  the  saddle-like  grouping  than  to  the 
long-shafted  group,  although  some  Eptesicus  do  indeed  have  bacula  that  suggest  the  beginnings 
of  basal  lobulation  or  of  a  very  short  shaft.  We  suggest  therefore  that  in  bacular  terms  the 
Vespertilionini  to  which  we  refer  Eptesicus  may  represent  a  transitional  stage  between  the  saddle- 
like  baculum  and  the  predominantly  basally  lobed  and  long-shafted  type.  Tylonycteris  and 
Glauconycteris  also  show  this  tendency. 

Dental  reduction  proceeds  throughout  both  of  the  major  bacular  groups.  In  the  grouping  with 
broadly  myotine  or  saddle-like  bacula  the  dentition  varies  in  number  of  teeth  from  a  total  of  38 
(Myotis,  Pizonyx)  through  36  (Lasionycteris,  Plecotus  and  allies),  34  (Barbastella,  Eptesicus  and 
allies),  32  (Lasiurus),  30  (Dasypterus,  Rhogeessa,  Baeodon,  Nycticeius,  Otonycteris,  Scotomanes, 
Scotophilus)  to  28  (Antrozous,  Bauerus}.  In  the  second  of  the  two  major  bacular  groups,  dental 
reduction  varies  from  Eudiscopus  with  a  total  of  36  teeth  (its  association  here  is  presumed) 
through  34  (Pipistrellus,  Glischropus,  Scotozous,  Nyctalus,  Chalinolobus),  32  (Laephotis,  Philetor, 
Hesperoptenus)  to  30  (Nycticeinops,  Scoteanax,  Scotorepens,  Scotoecus).  Thus  this  trend  occurs 
concurrently  in  the  two  major  groupings,  taking  the  same  form  in  each  by  increasing  the  size  and 
bulk  of  i2,  the  reduction,  transposition  and  loss  of  i3,  and  the  progressive  reduction,  transposition 
and  loss  of  pm2,  pm3  and  pm2. 


Zoogeographical  considerations 

The  saddle-shaped  or  slipper-like  baculum  characteristic  of  the  Myotini,  Plecotini,  Lasiurini  and 
Antrozoini  as  here  understood  is  cosmopolitan  in  but  one  genus,  Myotis.  It  occurs  in  one  Holarctic 
genus,  Plecotus,  in  one  Palaearctic  genus,  Barbastella,  itself  probably  closely  related  to  Plecotus, 
and  in  one  other  Old  World  genus,  Otonycteris,  that  occurs  in  southwestern  Asia  and  northern 
Africa.  Otherwise  this  bacular  type  is  limited  to  the  New  World.  Lasionycteris,  exclusively  North 
American,  has  a  baculum  apparently  derived  from  this  type,  as  does  Nycticeius,  also  North 
American,  although  in  this  genus  the  baculum  is  considerably  modified  to  the  extent  that  Hamilton 
(1949)  commented  upon  its  unique  character  among  the  genera  that  he  had  examined.  Thus 
although  the  saddle-shaped  baculum  or  its  derivatives  is  represented  about  equally  in  number  of 
species  in  the  Old  and  New  Worlds,  genera  with  bacula  of  this  type  predominate  in  the  latter,  its 
extension  into  the  Old  World  being  primarily  through  the  many  species  of  Myotis,  with  a  lesser 
contribution  from  Plecotus,  Barbastella,  and  Otonycteris. 

A  further  variety  of  this  bacular  type  is  found  in  the  Vespertilionini,  that  is,  in  Eptesicus  and  its 
close  relatives.  In  these,  the  baculum  is  less  strongly  saddle-shaped  or  slipper-like,  flatter,  and  often 
more  triangular  in  outline.  This  bacular  type  is  primarily  Old  World  in  numbers  of  genera  and 
species,  only  Eptesicus  among  Old  World  genera  extending  to  the  New  World  where  there  is  a 
closely  related  genus,  Histiotus.  In  the  Old  World,  Vespertilio  is  also  closely  related  to  Eptesicus. 
Another  Old  World  genus,  la,  is  a  giant  representative  of  this  same  bacular  type.  The  southeastern 


260  J.  E.  HILL  &  D.  L.  HARRISON 

Asian  Tylonycteris  and  the  African  Glauconycteris  have  bacula  that  are  modified  variants  of  this 
type:  Mimetillus,  in  which  no  baculum  has  been  found,  also  appears  to  belong  here.  Two  further 
Old  World  genera,  Scotomanes  and  Scotophilus,  also  have  bacula  that  are  similar  in  many  respects 
to  the  vespertilionine  type. 

The  shafted  or  long-shafted  bacular  type  is  confined  almost  exclusively  to  the  Old  World,  and  is 
represented  in  the  New  World  by  no  more  than  two  species  of  Pipistrellus  in  the  Nearctic  region, 
one  of  these  with  a  highly  modified  baculum.  This  bacular  type  is  restricted  to  the  Pipistrellini  and 
within  that  grouping  to  those  genera  that  for  the  most  part  can  be  shown  on  other  grounds  to 
cluster  around  Pipistrellus.  Indeed,  some  such  as  Scotozous,  Glischropus,  Scoteanax,  Scotorepens 
and  perhaps  even  Nyctalus  might  on  bacular  grounds  be  regarded  as  subgenera  of  this  widespread 
genus.  In  a  reduced  form  this  bacular  type  appears  in  two  of  the  subgenera  of  Pipistrellus,  P. 
(Perimyotis)  and  P.  (Arielulus).  Widespread  in  the  Palaearctic  region  and  in  southeastern  Asia, 
this  bacular  type  is  represented  in  Australia  by  five  distinct  groupings:  Pipistrellus  (Pipistrellus}, 
P.  ( Vespadelus),  Scoteanax,  Scotorepens,  and  Chalinolobus.  This  type  of  baculum  also  occurs  in 
Africa  among  Pipistrellus  kuhlii  and  its  associates,  which  might  in  fact  be  considered  to  warrant 
recognition  as  a  further  subgenus  of  Pipistrellus. 

A  further  variant  of  the  shafted  bacular  type  is  found  in  Pipistrellus  (Hypsugo)  and  P.  (Falsistrel- 
lus). In  these  the  shaft  is  shorter  and  is  ventrally  fluted,  often  with  expansion  of  the  tip.  Pipistrellus 
(Hypsugo)  is  confined  chiefly  to  Asia  and  Africa,  where  in  the  latter  region  it  appears  to  be  closely 
associated  with  P.  (Neoromicia)  in  which  pm2  is  generally  lost.  Thus  as  in  Australia  where  P. 
(Vespadelus)  in  which  pm2  is  also  generally  absent  appears  to  derive  from  P.  (Pipistrellus),  so  in 
Africa  P.  (Neoromicia)  is  apparently  similarly  related  to  P.  (Hypsugo).  Of  the  two  North  American 
pipistrelles,  P.  subflavus  has  a  reduced  form  of  the  P.  (Pipistrellus)  baculum,  the  shaft  very  short 
and  stubby:  this  species  has  a  myotine  tragus  and  has  been  considered  (Menu,  1984)  to  have  a 
myotine  dentition.  However,  on  the  balance  of  features  it  appears  to  be  clearly  referable  to 
Pipistrellus  and  indeed  to  be  cranially  and  dentally  close  to  P.  (Pipistrellus),  which  apparently  it 
represents  in  North  America.  There  do  not  appear  to  be  sufficient  grounds  to  justify  its  generic 
separation  from  Pipistrellus  as  has  been  recently  effected  (Menu,  loc.  cit.),  although  subgeneric 
recognition  within  that  genus  seems  appropriate.  The  second  North  American  species  of  Pipistrel- 
lus, P.  Hesperus,  should  evidently  be  referred  to  P.  (Hypsugo)  with  which  it  has  close  bacular  and 
dental  similarities,  although  recently  generic  separation  (Horacek  &  Hanak,  19850,  b,  1985-1986) 
has  been  proposed  for  it.  Finally,  P.  (Falsistrellus)  is  restricted  to  southeastern  Asia,  Australasia 
and  Tasmania:  the  deeply  ventrally  fluted  baculum  of  this  subgenus,  lacking  basal  and  distal 
modification  but  massive  and  substantial  appears  to  be  an  extreme  of  the  P.  (Hypsugo)  type: 
possibly  P.  (Falsistrellus)  represents  P.  (Hypsugo)  which  seems  to  be  linked  to  it  by  several  of  its 
Asian  species. 

One  corollary  of  the  removal  of  the  African  capensis  and  tenuipinnis  groups  of  'Eptesicus'  to 
Pipistrellus,  and  of  the  similar  transfer  of  the  Australian  species  formerly  referred  to  'Eptesicus' is 
that  in  the  Old  World  Eptesicus  now  becomes  primarily  Palaearctic,  with  outliers,  perhaps  all 
closely  connected  to  E.  serotinus,  in  Africa  while  in  the  New  World  it  extends  over  both  North  and 
South  America.  In  southeastern  Asia  the  genus  becomes  restricted  to  no  further  east  than  southern 
Thailand,  the  former  enormous  hiatus  in  its  distribution  between  this  part  of  southern  Asia  and 
Australia  having  been  removed. 


Conclusions 

(1)  The  current  classification  of  the  Vespertilioninae  is  based  chiefly  on  adaptive  characters  with 
considerable  emphasis  on  facial  shortening  and  concomitant  dental  reduction  and  loss.  Several 
authors  have  drawn  attention  to  the  deficiencies  and  dangers  of  any  classification  that  relies  heavily 
on  such  features.  A  review  of  bacular  morphology  within  the  subfamily  suggests  that  this  structure 
provides  indications  of  relationship  that  in  many  respects  support  the  existing  classification  but 
which  also  indicate  several  changes  to  the  current  arrangement.  In  particular,  bacular  morphology 
suggests  a  number  of  major  and  minor  changes  in  the  systematics  of  the  nominal  genera  Pipistrellus 


VESPERTILIONINE  SYSTEMATICS  261 

and  Eptesicus,  separated  hitherto  only  by  dental  formula,  itself  subject  to  variation  in  both  'genera' 
as  they  are  currently  understood. 

(2)  The  presence  or  absence  of  the  anterior  upper  premolar  (pm2)  in  Pipistrellus  and  Eptesicus, 
used  formerly  as  their  principal  diagnostic  character,  has  little  taxonomic  significance.  The  tooth  is 
variable  in  Pipistrellus  as  here  understood,  being  reduced  or  lost  in  three  of  its  subgenera,  and  is 
absent  from  Eptesicus  as  we  envisage  it. 

(3)  Bacular  morphology  in  Pipistrellus  and  Eptesicus  provides  groupings  that  largely  agree  in 
species  content  with  those  proposed  by  earlier  authors  such  as  Tate  (19420)  and  Koopman  (1973, 
1975)  although  in  basing  their  studies  on  'conventional'  morphological  characters  neither  con- 
sidered these  genera  in  their  entirety.  The  bacular  morphology  of  'Eptesicus'  as  it  is  currently 
understood  provides  a  clear  indication  that  as  such  it  is  not  a  natural  group,  but  that  three  species 
aggregations,  the  Australian  pumilus  group  and  the  African  capensis  and  tenuipinnis  groups, 
should  be  transferred  to  Pipistrellus. 

(4)  It  has  been  possible  to  recognise  and  define  subgenera  for  the  major  species  groups  in  both 
Pipistrellus  and  Eptesicus  and  to  suggest  possible  relationships  between  them.  One  subgenus  is 
described  as  new  as  Pipistrellus  (Arielulus)  for  P.  circumdatus  and  its  allies. 

(5)  There  appear  to  be  clear  links  between  certain  of  the  pipistrelline  subgenera:  Pipistrellus 
( Vespadelus}  in  Australia  seems  to  represent  P.  (Pipistrellus)  in  bacular  terms  while  P.  (Hypsugo)  is 
apparently  represented  in  Indo-Australia  by  P.  (Falsistrellus)  and  is  related  to  the  African  P. 
(Neoromicid).  Although  the  features  of  the  two  Nearctic  species  of  Pipistrellus  have  been  thought 
to  justify  their  recognition  in  separate,  individual  genera  we  consider  that  the  characters  of  one 
(subflavus)  merit  no  more  than  subgeneric  status  as  the  sole  species  of  P.  (Perimyotis),  which  itself 
perhaps  represents  P.  (Pipistrellus),  while  the  other  (Hesperus)  is  perhaps  more  appropriately 
referred  to  P.  (Hypsugo). 

(6)  The  examination  of  bacula  in  Pipistrellus  has  suggested  that  some  taxa  hitherto  ranked  as 
subspecies,  for  example  abramus,  paterculus  or  helios,  might  in  fact  be  distinct  species. 

(7)  As  we  now  understand  the  species  content  of  Pipistrellus  and  Eptesicus  the  former  remains 
primarily  an  Old  World  genus  where  it  is  widespread  and  diverse  in  the  tropics  and  subtropics, 
extending  into  the  temperate  zones  and  just  to  North  America.  In  contrast,  our  concept  of 
Eptesicus  limits  this  genus  to  the  New  World  and  in  the  Old  World  primarily  to  the  Palaearctic, 
with  outlying  representatives  in  Africa.  It  does  not  extend  significantly  into  Australasia. 

(8)  Bacular  morphology  suggests  the  informal  recognition  of  two  major  groupings  within  the 
subfamily  Vespertilioninae.  The  first  includes  the  Myotini,  Plecotini  and  Lasiurini;  Antrozous  and 
Baeurus,  which  in  bacular  terms  have  no  relation  to  Nyctophilus  and  Pharotis  (the  Nyctophilinae); 
the  Scotophilini  to  include  Scotomanes  and  Scotophilus;  and  finally  the  Vespertilionini,  here 
reduced  in  content  to  include  Eptesicus  and  its  close  relatives  Histiotus,  la  and  Vespertilio,  with 
Tylonycteris,  Mimetillus  and  Glauconycteris. 

(9)  The  second  grouping  consists  of  Pipistrellus  and  those  genera  which  cluster  round  it.  All  with 
the  possible  exception  of  Philetor  appear  to  relate  quite  closely  in  bacular  terms  to  one  or  other  of 
the  subgenera  that  we  recognise  in  Pipistrellus,  principally  to  P.  (Pipistrellus).  Laephotis,  formerly 
considered  related  to  Histiotus,  is  instead  in  bacular  terms  closely  associated  with  P.  (Neoromicid). 
The  bacula  of  Chalinolobus  and  Glauconycteris  are  widely  dissimilar  athough  these  genera  have 
been  closely  allied  in  the  past;  Chalinolobus  is  of  the  pipistrelline  type  while  the  baculum  of 
Glauconycteris  apparently  associates  it  more  appropriately  with  Eptesicus  and  its  allies. 

(10)  Bacular  morphology  provides  clear  indications  that  the  'Nycticeini'  of  Tate  (\942a)  and 
Koopman  (1984,  1985)  is  not  a  natural  group,  its  constituent  members  despite  cranial  and  dental 
similarities  having  widely  different  bacula.  Thus  Rhogeessa,  Baeodon,  Nycticeius  sensu  stricto,  and 
Otonycteris  have  been  here  allied  to  the  plecotine  bats  on  bacular  grounds,  while  Scoteanax, 
Scotorepens  and  Scotoecus  are  quite  clearly  associates  in  bacular  terms  of  Pipistrellus.  'Nycticeius', 
at  one  time  thought  to  include  the  Australian  Scoteanax  and  Scotorepens  as  well  as  its  North 


262  J.  E.  HILL  &  D.  L.  HARRISON 

American  type  species  humeralis  and  the  African  schlieffenii,  has  recently  been  restricted  only  to  the 
American  and  African  forms.  These  prove  to  have  widely  different  bacula;  humeralis  has  been 
associated  with  the  plecotine  bats  on  this  account,  while  generic  status  has  been  accorded  to 
schlieffenii  with  the  proposal  of  a  new  generic  name,  Nycticeinops. 

(11)  The  two  broad  bacular  types  that  we  discern  in  the  subfamily  Vespertilioninae  have  definite 
geographical  patterns:  the  saddle-like  baculum  and  its  variants  that  characterise  the  first  group 
noted  above  is  primarily  New  World  and  Palaearctic,  extending  less  significantly  into  the  Old 
World  tropics  or  Australasia,  while  the  shafted  baculum  of  the  second  group  is  chiefly  confined  to 
the  Old  World. 


Addendum 

A  phenetic  analysis  of  the  relationships  of  selected  vespertilionine  species  (chiefly  those  currently 
referred  to  Pipistrellus  and  Eptesicus)  by  Horacek  &  Hanak  (1985-1986)  appeared  while  this  paper 
was  in  press.  These  authors  provided  definitions  of  Pipistrellus,  Hypsugo  (which  they  considered 
generically  valid)  and  Eptesicus,  based  on  the  morphology  of  the  penis  and  baculum,  the  upper 
molars,  the  basisphenoid  pits,  the  pelvic  girdle,  and  the  tibia,  tail  and  epiblema. 

Horacek  &  Hanak  suggested  that  the  classification  of  pipistrelloid  bats  might  be  clarified  by  the 
recognition  of  additional  subgenera  or  genera  for  those  species  or  species  groups  that  do  not 
conform  precisely  with  those  that  they  included  within  these  three  generic  groupings.  To  some 
extent  such  recognition  is  provided  in  several  instances  by  the  classification  here  proposed  and 
although  some  major  differences  exist  between  the  informal  assessments  and  species  groups  of 
Horacek  &  Hanak  and  the  formal  arrangement  put  forward  in  this  paper  there  is  nevertheless 
a  broad  measure  of  agreement.  Horacek  &  Hanak  did  not  attempt  any  classification  of  the 
Vespertilioninae  as  a  whole,  but  'Nycticeius'  schlieffenii,  here  considered  to  represent  a  distinct 
monospecific  genus  (Nycticeinops  gen.  nov.)  was  thought  by  these  authors  to  be  referable  either  to 
Eptesicus  ( Rhyneptesicus),  or  possibly  to  justify  the  establishment  of  a  new  subgenus  within 
Eptesicus. 


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Manuscript  accepted  for  publication  30  June  1986. 


VESPERTILIONINE  SYSTEMATICS  269 

Appendix  1.  Specimens  examined 

AMNH  =  American  Museum  of  Natural  History,  New  York 
BM(NH)  =  British  Museum  (Natural  History),  London 
CMNH  =  Carnegie  Museum  of  Natural  History,  Pittsburgh 
HZM  =  Harrison  Zoological  Museum,  Sevenoaks,  Kent 
NMW  =  Naturhistorisches  Museum,  Wien 

Vespertilioninae 

Myotis  nattereri 

HZM  26.1 1254  Leany  Cave,  Pilis  Heights,  Hungary.  (Fig.  19j) 

Myotis  ridleyi 

BM(NH)  98.3.13.5  Selangor,  Malaya.  Holotype.  (Fig.  19i) 

Pizonyx  vivesi 

HZM  3.10284  Isla  Cordonosa,  Bahia  de  Los  Angeles,  Baja  Norte,  Mexico.  (Fig.  19k) 

Lasionycteris  noctivagans 

BM(NH)  7.7.7.2319  Raleigh,  North  Carolina,  USA.  (Fig.  170 

HZM  2.3708  Delta,  Manitoba,  Canada 

Plecotus  auritus 

HZM  19.1227  Near  Godstone,  Surrey,  England.  (Fig.  19g) 

Plecotus  austriacus 

BM(NH)  91.10.5.4  Duirat,  Tunis 

HZM  3.4867  St.  Pierre  de  Varenne,  Saone  et  Loire,  France. 

HZM  4.8337  Mont  de  Lans,  Les  Deux  Alpes,  Isere,  France. 

HZM  5.8467  Chateau  de  Salse,  Salse,  Rousillon,  France.  (Fig.  19h) 

Barbastella  barbastellus 

HZM  13.1 1222  Kiralyret,  Borzsony  Heights,  Hungary.  (Fig.  18j) 

Rhogeessa  tumida 

HZM  1.12080  Airport  Camp,  Belize.  (Fig.  18k) 

Nycticeius  humeralis 

AMNH  249144  Sierra  de  Tamaulipas,  Acuna,  Tamaulipas,  Mexico,  2890  ft.  (Fig.  17k) 

Otonycteris  hemprichii 

BM(NH)  14.8.17.1  Syrian  Desert. 

HZM  6.8174  17  km  N  of  Hufoof,  Saudi  Arabia.  (Fig.  16a) 

Lasiurus  cinereus 

HZM  1.3695  S  Fork,  Cave  Creek,  near  Portal,  Cochise  County,  Arizona,  USA.  (Fig.  191) 

Dasypterus  argentinus 

BM(NH)  33.6.24.3  Bonifacio,  Argentina.  (Fig.  180 

Antrozous  pallidus 

BM(NH)  50.767  California,  USA.  (Fig.  18b). 

HZM  3.3692  S  Fork,  Cave  Creek,  near  Portal,  Cochise  County,  Arizona,  USA. 

Scotophilus  borbonicus 

BM(NH)  89.1.11.2  E  coast  of  Africa. 

Scotophilus  dinganii 

BM(NH)  79.513  Tokadeh,  Nimba,  Liberia,  600  m. 

Scotophilus  heathii 

BM(NH)  14.7.19.19  Mount  Popa,  Burma. 

BM(NH)  14.7.19.28  Kyauk  Inyaung,  Irrawaddy,  Burma. 

BM(NH)  60.257  Tori,  Pakistan.  (Fig.lTh) 

BM(NH)  70.1488  Bang  Phra,  Cholburi,  Sriracha,  Thailand. 

BM(NH)  76.787-788  Nhatrang,  Annam,  Vietnam. 


270  J.  E.  HILL  &  D.  L.  HARRISON 

Scotophilus  kuhlii 

BM(NH)  75.2955  Chiang  Mai,  Thailand  (Fig.  17i) 

Scotophilus  nigrita  (gigas) 

BM(NH)  22.12.17.55a  Mtondo,  Ruo,  Malawi.  (Fig.  17g) 

Scotophilus  nigritellus 

BM(NH)  78.189  Numan,  Gongola,  Nigeria.  (Fig.  17j) 

Scotomanes  ornatus 

BM(NH)  94.9. 1.21  Foochow,  China.  (Fig.  18g) 

Eptesicus  (Eptesicus) 

Eptesicus  bobrinskoi 

BM(NH)  63.1 187  Outer  Su,  N  of  Mount  Sabalan,  NW  Iran.  (Fig.  13e) 

Eptesicus  nasutus 

HZM  3.4571  Harmul,  10  m  N  of  Sohar,  Oman  (batinensis). 
HZM  12. 1 1 172  Jamma,  near  Rostaq,  Oman  (batinensis).  (Fig.  14c) 
HZM  1.1623  Shaiaba,  Iraq  (pellucens). 

Eptesicus  bottae 

HZM  18.1616  Ser'Amadia,  Kurdistan,  Iraq. 

HZM  5.1628  Basrah,  Iraq  (hingstoni). 

BM(NH)  3.12.8.9  Cairo,  Egypt  (inncsi).  (Fig.  13j) 

HZM  12.8075  Birkat  Sharaf  al  Wadi  Sahtan,  Jebel  al  Akhdar,  Oman  (omanensis).  (Fig.  14a) 

Eptesicus  brasiliensis 

BM(NH)  85.6.26.10  San  Lorenzo,  Rio  Grande  do  Sul,  Brazil.  (Fig.  13k) 

BM(NH)  0.6.29.4  Palmeira,  Parana,  Brazil. 

BM(NH)  98.10.3.32  Valdivia,  Colombia  (andinus).  (Fig.  13d) 

Eptesicus  furinalis 

BM(NH)  4.8.8.5  La  Plata,  Argentina.  (Fig.  13c) 

Eptesicus  fuscus 

BM(NH)  89.6.1.4  Sing  Sing,  New  York,  USA.  (Fig.  13a) 

BM(NH)  52.551  Chinchona,  Jamaica  (hispaniolae).  (Fig.  13i) 

Eptesicus  hottentotus 

BM(NH)  81.7.1 1.1  Drakenburg  Mountains,  Natal,  South  Africa  (megalurus).  (Fig.  13b) 

Eptesicus  serotinus 

BM(NH)  53.555  Blandford,  Dorset,  England.  (Fig.  13g) 

HZM  3.629  Shepreth,  Cambridgeshire,  England. 

BM(NH)  66.1 150  Defilia  Oasis,  Figuig,  Morocco  (isabellinus}.  (Fig.  13h) 

Eptesicus  (Rhinopterus) 

Eptesicus  floweri 

BM(NH)  0.8.6.20  Abu  Zeit,  White  Nile,  Sudan.  (Fig.  130 

BM(NH)  1.5.5.78  Shendy,  Sudan  (lowei).  (Fig.  131) 

Vespertilio  orientalis 

BM(NH)  8.7.25.6,  BM(NH)  8.8.1 1.2  Kuatun,  NW  Fokien,  China 

Histiotus  macrotis 

BM(NH)  71.1 123  Antofagasta,  Lake  Miniques,  Chile,  1450  m.  (Fig.  18e) 

Histiotus  (?)  macrotis 

BM(NH)  6.5.8.3  Jafi,  Tucuman  Province,  Argentina.  (Fig.  18d) 

Histiotus  velatus 

BM(NH)  0.6.29.2  Palmeira,  Parana,  Brazil.  (Fig.  18c) 

Tylonycteris  pachypus 

BM(NH)  9.1. 5.954  Buitenzorg,  Java.  (Fig.  18h) 


VESPERTILIONINE  SYSTEMATICS  271 

Tylonycteris  robustula 

BM(NH)  60.1499  Bukit  Lagong  Forest  Reserve,  Kepong,  Selangor,  Malaya.  (Fig.  18i) 

HZM  3.7444  15th  mile  Ulu  Gombok,  Selangor,  Malaya. 

Mimetillus  moloneyi 

BM(NH)  93. 1 .7.2  Leekie,  Nigeria 

BM(NH)  54.862  Irumu,  Zaire. 

BM(NH)  60.154  Bo,  Sierra  Leone. 

BM(NH)  64.1788  Liwale,  Tanzania. 

HZM  2.7802  Near  Babeke,  River  Isai,  Ituri,  Zaire. 

Glauconycteris  argentata 

BM(NH)  54.863  Banana,  Zaire. 

BM(NH)  59.510  Ikela,  Ikela  Territory,  Zaire.  (Fig.  19d) 

Glauconycteris  beatrix 

BM(NH)  48.713  Eshobe,  Mamfe,  Cameroun.  (Fig.  19c) 

Glauconycteris  humeralis 

BM(NH)  30.11.11.173  River  Wasa,  Semliki  Valley,  Uganda.  (Fig.  1 9e) 

Glauconycteris  poensis 

BM(NH)  96. 1 2.3 1 .2  Sierra  Leone. 

BM(NH)  69.26  Abidjan,  Ivory  Coast.  (Fig.  19a) 

Glauconycteris  variegata 

BM(NH)  76.780  Mole  National  Park,  Ghana.  (Fig.  19b) 

BM(NH)  55.409  Mongue,  near  Inhambane,  Mozambique  (papilio).  (Fig.  190 

Pipistrellus  (  Pipistrellus ) 

Pipistrellus  pipistrellus 

HZM  94.6807  Rabat,  Malta. 

HZM  1 16.8549  Sevenoaks,  Kent,  England.  (Fig.  2a) 

HZM  1 17.8650  Aylesford,  Kent,  England. 

BM(NH)  73.689  Kululai  Rest  House,  Northwest  Frontier  Province,  Pakistan  (bactrianus) 

BM(NH)  14.5.10.19  BM(NH)  14.5.10.22  Djarkent,  Semiretschenskoi,  USSR  (lacteus). 

Pipistrellus  nathusii 

BM(NH)  8.8.4.128  BM(NH)  62.1368  St  Giles,  France  (Fig.  2b) 

Pipistrellus  abramus 

BM(NH)  89.6.17.3-4  Kin  Kiang,  Yangtse  Kiang,  China  (seen  by  Thomas,  1928a). 

BM(NH)  5. 1 .4.8  Tokyo,  Japan. 

BM(NH)  7.7.3.26  Nanking,  China  (seen  by  Thomas,  19280).  (Fig.  3a) 

BM(NH)  14.10.1.1  Lokow,  Hunan,  C  China. 

BM(NH)  26. 1 0.4. 1 8  Hue,  Annam,  Vietnam  (seen  by  Thomas,  1 928a) 

BM(NH)  66.3469-3470  Chihli,  China. 

BM(NH)  86.529  Chusan,  China  (Syntype  irretitus). 

BM(NH)  86.532  Canton,  S  China. 

Pipistrellus  babu 

BM(NH)  45. 1.8.403  Nepal. 

BM(NH)  16.3.25.8  Pashok,  Darjeeling,  India.  (Fig.  4a) 

Pipistrellus  camortae 

BM(NH)  -      -  Car  Nicobar  (Original  No.  3/76).  (Fig.  1 5d) 

Pipistrellus  endoi 

BM(NH)  70.2522  Horobe,  Tayama,  Ajiro-Machi,  Minohe-Gun,  Iwate  Prefecture,  Japan.  (Fig.  3b) 

Pipistrellus  javanicus  (tralatitius) 

BM(NH)  0.8.2.9  Sumatra. 

BM(NH)  9. 1 .5.295  Tjilatjap,  Java. 

BM(NH)  9. 1 .5.997-998  W  Java. 

BM(NH)  16.4.21.3  Sungei  Penoh,  Korinchi,  Sumatra. 


272  J.  E.  HILL  &  D.  L.  HARRISON 

BM(NH)  27.12.1.37  Tarn  Dao,  Tonkin,  Vietnam,  3000  ft  (No.  41 1  of  Thomas,  1928a). 

BM(NH)  28.7. 1 .20  Phu  Qui,  Annam,  Vietnam,  100  ft  (No.  866  of  Thomas,  19286,  who  identified  the  specimen 

as  P.  coromandra  tramatus,  but  with  a  longer  baculum  than  those  previously  examined). 
BM(NH)  83.76  Silau  Silau  Trail,  Mount  Kinabalu,  Sabah,  Borneo.  (Fig.  lOe) 

Pipistrellus  paterculus 

BM(NH)  14.7.8.62  Pyaunggaung,  N  Shan  States,  Burma,  2794  ft. 
BM(NH)  14.7.19.241  Kyauk  Myaung,  Irrawaddy,  W  Burma. 
BM(NH)  14.7.19.242  Mount  Popa,  Upper  Burma  (Holotype).  (Fig.  3c) 
BM(NH)  14.7.19.240  Mandalay,  Burma. 

Pipistrellus  angulatus 

BM(NH)  67.2125  Schoolmaster's  House,  Nuhu,  Guadalcanal  I,  Solomon  Is  (ponceleti).  (Fig.  4d) 

Pipistrellus  collinus 

BM(NH)  50.983  Baiyanka,  Purari-Ramu  Divide,  SE  Bismarck  Range,  Papua  New  Guinea.  (Fig.  4b) 

Pipistrellus  coromandra 

BM(NH)  32. 1 1 . 1 .7  Nam  Tamai,  Upper  Burma. 

BM(NH)  50.478  Ningma,  Upper  Burma. 

BM(NH)  76.1263  Sumka  Uma,  Upper  Burma. 

HZM  1.7317,  HZM  2.73 18  Near  Mirzapur,  India.  (Fig.  7c,  HZM  2.7318) 

HZM  4.7320  Dalatpur,  near  Mirzapur,  India. 

BM(NH)  4.6.8.1  Annam,  Vietnam  (tramatus).  (Fig.  7b) 

BM(NH)  27.12.1.40  Bac-kan,  Tonkin,  Vietnam  (tramatus)  (Original  No.  444,  seen  by  Thomas,  1928a). 

Pipistrellus  mimus 

BM(NH)  98.5.5.20  Dangs,  Bombay,  India. 

HZM  1 .10456  Vikas  Vidyalaya,  near  Ranchi,  Bihar,  India.  (Fig.  7g) 

Pipistrellus  murrayi 

BM(NH)  99.8.6.34  Christmas  I,  Indian  Ocean  (Holotype).  (Fig.  4c) 

BM(NH)  9.1.16.7  Flying  Fish  Cove,  Christmas  I,  Indian  Ocean. 

Pipistrellus  papuanus 

BM(NH)  22.2.2.3  Fredrik  Hendrik  I,  Irian  Jaya.  (Fig.  2c) 

BM(NH)  34.1.14.8  Kokoda,  Papua  New  Guinea. 

Pipistrellus  tenuis 

BM(NH)  85.912  Coast  of  Sabah,  Borneo  (nitidus).  (Fig.  9d) 

Pipistrellus  ceylonicus 

BM(NH)  95.6.12.1  Pundibiya,  India. 

BM(NH)  2.4.2.8  Astoli,  Belgoum,  India.  (Fig.  7d) 

BM(NH)  1 1.4.5.5  Lanje,  Konkan,  India. 

BM(NH)  13.9.8.102  Gujerat,  India. 

BM(NH)  9.1.4.73  Mangalore,  Malabar  Coast,  India  (Holotype  indicus) 

BM(NH)  4.6.8.7-8  Tonkin,  Vietnam  (raptor).  (Fig.  3d,  BM(NH)  4.6.8.7  Holotype). 

Pipistrellus  crassulus 

BM(NH)  4.2.8.1  Efulen,  Cameroun  (Holotype).  (Fig.  7e) 

Pipistrellus  nanulus 

BM(NH)  4.2.8.8  Efulen,  Cameroun  (Holotype).  (Fig.  If) 

BM(NH)  79.508  South  Nimba,  Liberia. 

Pipistrellus  rueppellii 

BM(NH)  68.12.22.3  Zanzibar  (Holotype pulcher).  (Fig.  lOa) 

BM(NH)  99.9.9.20  Egypt. 

BM(NH)  -          -  Uganda.  (Fig.  lOb) 

HZM  3.3170  Kabompo  Boma,  Zambia. 

HZM  7. 12109  Suez,  Egypt. 

Pipistrellus  deserti 

BM(NH)  79.987  Hoggar  Plateau,  Algeria.  (Fig.  5c) 

NMW  27503  (?)  Upper  Egypt. 


VESPERTILIONINE  SYSTEMATICS  273 

Pipistrellus  kuhlii 

BM(NH)  92.9.9.25  Upper  Egypt. 

BM(NH)  Argostoli,  Cephaloni,  Greece.  (Fig.  5a) 

BM(NH)  63.335  Sangha,  Malya  Khola,  E  Nepal. 

HZM  5. 1 1607  Horefto,  near  Volos,  Greece. 

HZM  11.1016  Rapallo,  N  Italy. 

HZM  138.4563  Yal  bu  Hillal,  Batinah,  Oman. 

HZM  154.4619  Saham,  Batinah,  Oman. 

HZM  203.7232  Dig  Dagga,  Ras  al  Khaima,  United  Arab  Republic. 

HZM  218.7402  Benghazi,  Libya. 

HZM  227.91 10  Kapsowat,  Marakwat,  Kenya. 

Pipistrellus  maderensis 

BM(NH)  86.528  Madeira.  (Fig.  5b) 

Pipistrellus  rusticus 

BM(NH)  35.9.1.108  Okavango-Omatako  Junction,  Grootfontein  District,  Namibia. 

BM(NH)  79.1731  Oli  River,  Borgu  G.R.,  Nigeria.  (Fig.  6c) 

HZM  4.3285  Sentinel  Ranch,  River  Limpopo,  Zimbabwe.  (Fig.  5d) 

Pipistrellus  ( Vespadelus} 

Pipistrellus  pumilus 

BM(NH)  70.1093  E  Bonithon  Range,  C  Australia  23°42'S,  129°02'E,  1400  ft. 

BM(NH)  71.1497  Westwood,  near  Rockhampton,  Queensland,  Australia.  (Fig.  12k) 

Pipistrellus  (Perimyotis) 

Pipistrellus  subflavus 

HZM  1.2422  Big  Wyandotte  Cave,  Crawford  County,  Indiana,  USA.  (Fig.  2d) 

Pipistrellus  (Hypsugo) 

Pipistrellus  anchietae 

BM(NH)  69.1248  Ngoma,  Zambia. 

BM(NH)  70.2632  Balovale,  Zambia.  (Fig.  6e) 

BM(NH)  89.5.1.5  Caconda,  Angola  (Syntype  of  Vesperus  bicolor  Bocage,  1889).  (Fig.  9e) 

Pipistrellus  bodenheimeri 

HZM  3.3786  Jazirat  al  Abid,  Aden,  South  Yemen. 

HZM  5.8279  Bin  Gedi,  Israel.  (Fig.  90 

Pipistrellus  savii 

BM(NH)  31.11.11.13,  BM(NH)  66.4644  E  slope  of  Mount  Olympus,  Greece. 

BM(NH)  61.395  Ainab,  Lebanon.  (Fig.  6a) 

Pipistrellus  arabicus 

HZM  4.10060  Wadi  Sahtan,  Oman. 

HZM  5.1 1625  Wadi  Fidah,  Dank/Ibri,  Oman.  (Fig.  7a) 

Pipistrellus  helios 

BM(NH)  39.133  N  Guaso  Nyiro,  Kenya.  (Fig.  6d) 

BM(NH)  69.207  Kangatet,  S  Turkana,  Kenya. 

HZM  2.4086  Archer's  Post,  Northern  Frontier  District,  Kenya. 

Pipistrellus  nanus 

BM(NH)  49.484  Kontaur,  Gambia. 

HZM  3.2778  Sokoto,  N  Nigeria. 

HZM  3.4026,  HZM  4.4027  Near  Monrovia,  Liberia. 

HZM  83.4387  Haroni-Lusitu  Beacon  74,  Zimbabwe. 

HZM  107.3212  Kabompo  Boma,  Zambia. 

HZM  146.5161,  HZM  147.5162  Rondo,  Lindi,  Tanzania. 

HZM  165.5321  Liwale,  Tanzania. 

HZM  200.6581  Karonga,  Malawi. 

HZM  258.1 1469  Kunyale  Stream,  Mwinilunga  District,  Zambia. 


274  J.  E.  HILL  &  D.  L.  HARRISON 

HZM  260.12175  Lamto,  Ivory  Coast. 

HZM  26 1 . 1 2 1 76  Ivory  Coast. 

HZM  263.12451,  HZM  264.12452  Kamuani  Area,  Machakos  District,  Kenya.  (Fig.  6b,  HZM  263.12451) 

Pipistrellus  pulveratus 

BM(NH)  79.702  Near  Nicholson  Goat  Bungalows,  Hong  Kong  I. 

BM(NH)  79.903  Peace  Mansion,  Tai  Hang  Road,  New  Territories,  Hong  Kong  (Fig.  8c) 

Pipistrellus  Hesperus 

BM(NH)  98.3.1.8  Sierra  Laguna,  Baja  California,  Mexico. 

BM(NH)  29.1 1.7.10  Panamint  Mts,  California,  USA. 

HZM  4.1 1219  Sycamore  Well,  Hidalgo  County,  New  Mexico,  USA.  (Fig.  8d) 

Pipistrellus  eisentrauti 

BM(NH)  84.1684,  BM(NH)  84.1686  Mount  Cameroun,  Cameroun.  (Fig.  9g,  BM(NH)  84.1684) 

Pipistrellus  imbricatus 

BM(NH)  9.1.5.286  Buitenzorg,  Java.  (Fig.  9a) 

Pipistrellus  macrotis 

BM(NH)  23.1.2.12  Sabang,  NW  Sumatra.  (Fig.  9b) 

Pipistrellus  kitcheneri 

BM(NH)  10.4.5.47  Boentok,  Barito  River,  Kalimantan,  SC  Borneo.  (Fig.  8e) 

Pipistrellus  lophurus 

BM(NH)  14.12.1.6  Maliwun,  Victoria  Province,  Tenasserim,  Burma  (Holotype).  (Fig.  8f) 

Pipistrellus  stenopterus 

BM(NH)  60.1537  Institute  of  Medical  Research  Compound,  Kuala  Lumpur,  Malaya. 

BM(NH)  65.135  Pasir  Road,  Kuala  Lumpur,  Malaya.  (Fig.  7h) 

Pipistrellus  (Falsistrellus) 

Pipistrellus  affinis 

BM(NH)  83.3.3.2  Wynaard,  India.  (Fig.  8a) 

BM(NH)  72.4224  Argarawa,  Nevrawa  Elwa,  Central  Province,  Sri  Lanka. 

Pipistrellus  peter  si 

BM(NH)  23. 1 .2.3.  Buru  I,  Molucca  Is  (Fig.  8b) 

Pipistrellus  tasmaniensis 

HZM  1.8712  Barrington  Tops  National  Park,  New  South  Wales,  Australia.  (Fig.  8g) 

Pipistrellus  (Neoromicid) 

Pipistrellus  capensis 

BM(NH)  32.9.1.249  Broken  Hill,  Zambia. 

BM(NH)  54.859  Elizabethville,  Zaire. 

BM(NH)  61.1078  Doddieburn  Ranch,  West  Nicholson,  Zimbabwe,  2300  ft,  21°24'S,  29°21'E. 

BM(NH)  72.4383  E  of  Lake  Margharita,  Bulcha  Forest,  Ethiopia,  1800  m,  06°  1 1'N,  36°10'E. 

BM(NH)  72.4391  Didessa  River,  Wollega  Province,  Ethiopia,  1 190  m,  09°02'N,  36°09'E.  (Fig.  12g) 

BM(NH)  75.561  Mole  National  Park,  Ghana.  (Fig.  12b) 

BM(NH)  83.200  Mcheni  Gorge,  Chizarira  National  Park,  Binga  Province,  Zimbabwe,  17°40'S,  27°52'E. 

HZM  36.4514  40  m  NW  of  Serowe,  Botswana. 

BM(NH)  66.6057  Ambositra,  Madagascar  (matrokd).  (Fig.  12a) 

BM(NH)  77.2.19.6  Anzahameru,  Madagascar  ('minutus').  (Fig.  12i) 

Pipistrellus  guineensis 

BM(NH)  70.2224,  BM(NH)  70.2228,  BM(NH)  72.4373  Gambela,  Ethiopia,  8°15'N,  34°35'E  (BM(NH) 

72.4373  at  515  m)  (Fig.  12c,  BM(NH)  70.2224) 
BM(NH)  76.293  Shagamu,  Nigeria. 
BM(NH)  84.1019  Bontioli,  Bougouriba  River,  Burkina  Faso  (Upper  Volta). 

Pipistrellus  melckorum 

BM(NH)  83.216  Mcheni  Gorge,  Chizarira  National  Park,  Binga  Province,  Zimbabwe,  17°40'S,  27°52'E. 
(Fig.  12Q 


VESPERTILIONINE  SYSTEMATICS  275 

Pipistrellus  somalicus 

BM(NH)  70.484  Mouth  of  Fincha  River,  Blue  Nile  Gorge,  Ethiopia,  10°03'N,  37°20'E.  (Fig.  12h) 

BM(NH)  76.814  S  bank  of  Ganale  Doria,  Sidam-Bale  Bridge,  Sidamo  Province,  Ethiopia,  5°45'N,  39°37'E. 

BM(NH)  84.1016  Comoe  River,  Burkina  Faso  (Upper  Volta),  260  m,  9°57'N,  4°38'W. 

CMNH  MJS  2846  Snai  Sugar  Plantation,  l£  km  S,  \  km  E  of  Giohar,  Somalia,  2°46'N,  45°3 1  'E. 

Pipistrellus  zuluensis 

BM(NH)  83.212  Mchesu  River,  Chizarira  National  Park,  Binga  Province,  Zimbabwe,  17°47'S,  27°39'E. 

BM(NH)  83.215  Singama,  Sibuwa,  Binga  Province,  Zimbabwe,  17C36'S,  27°51'E.  (Fig.  12d) 

Pipistrellus  rendalli 

BM(NH)89.12.12.1  Bathurst,  Gambia. 

BM(NH)  7.12.17.1-2  Gondokoro,  White  Nile,  Sudan. 

BM(NH)  23.4.12.1-2  Bugala,  Sesse  Is,  Victoria  Nyanza,  Uganda.  (Fig.  12e,  BM(NH)  23.4.12.2) 

BM(NH)  48.702  N'ko,  Obubra  Division,  S  Nigeria  (Ibrunneus).  (Fig.  14b) 

Pipistrellus  tenuipinnis 

BM(NH)  47.350  Umuahia,  E  Nigeria. 

BM(NH)  54.917  Bonthe,  Sierra  Leone. 

BM(NH)  67.1734  Bota,  Victoria,  Cameroun,  4°00'N,  9°05'E.  (Fig.  12j) 

Pipistrellus  (Arielulus) 

Pipistrellus  circumdatus 

BM(NH)  73.618  Telecommunications  Tower,  Fraser's  Hill,  Pahang,  Malaya.  (Fig.  2e) 

Pipistrellus  cuprosus 

BM(NH)  83.351  Sepilok,  Sabah,  Borneo,  5°52'N,  1 17°56'E  (Holotype).  (Fig.  9h) 

Pipistrellus  societatis 

BM(NH)  67.1605  Base  Camp,  Gunong  Benom,  Pahang,  Malaya,  800  ft  (Holotype).  (Fig.  9c) 

Nyctalus  noctula 

BM(NH)  -  -  Locality  unknown. 

HZM  10.613  Bottisham,  Cambridgeshire,  England. 

HZM  33.8888  Winchelsea  Beach,  Sussex,  England.  (Fig.  100 

Laephotis  botswanae 

BM(NH)  -  -  Zomba,  Malawi  (original  No.  2269;  damaged). 

Laephotis  wintoni 

HZM  1.3020  Nyeri,  Mount  Kenya,  Kenya.  (Fig.  160 

Glischropus  tylopus 

BM(NH)  10.4.5.136  Upper  Barito  River,  Kalimantan,  SC  Borneo.  (Fig.  18a) 

Scotozous  dormeri 

BM(NH)  12.3.8.30  Furdapur,  Ajanta,  Khandesh,  India. 

BM(NH)  -  -  Kathiawar,  India  (Original  No.  BNHS  2007).  (Fig.  1 6d) 

Scoteanax  rueppellii 

BM(NH)  80.3.25.1  Richmond  River,  New  South  Wales,  Australia.  (Fig.  16i) 

Scotorepens  balstoni 

BM(NH)  10.6.21.9  Hermannsburg,  Northern  Territory,  Australia.  (Fig.  16g) 

Scotorepens  greyii 

BM(NH)  75.2261  Pine  Creek,  20  m  ESE  of  Candy's  Hill,  Northern  Territory,  Australia,  13°49'S,  131°49'E. 
(Fig.  16h) 

Nycticeinops  schlieffenii 

BM(NH)  14.7.3 1.14  Wei  Wei  River,  Kenya. 

BM(NH)  15.3.6.66  Kamisu,  Dinda  River,  Sudan. 

BM(NH)  71.675  Awash,  Filhoa,  Ethiopia,  09°00'N,  38°58'E. 

HZM  5.2120  Ikau,  Rukwa,  Tanzania.  (Fig.  16e) 

Scotoecus  albigula 

BM(NH)  63.1042  Calundo,  Lunda,  Angola.  (Fig.  20a) 


276  J.  E.  HILL  &  D.  L.  HARRISON 

Scotoecus  albofuscus 

BM(NH)  96. 1 2.3 1 . 1  Sierra  Leone.  (Fig.  20e) 

Scotoecus  hindei 

BM(NH)  14.7.31.13  30m  NW  of  Baringo,  Kenya.  (Fig.  20d) 

BM(NH)  66.1466  Jos,  Nigeria  (falabae).  (Fig.  20b) 

Scotoecus  hirundo 

BM(NH)  76.771  Mole  National  Park,  Ghana.  (Fig.  20c) 

Scotoecus  pallidus 

BM(NH)  86.531  Afghanistan  (damaged). 

Philetor  brachypterus 

BM(NH)  -  -  New  Guinea.  (Fig.  16b) 

Hesperoptenus  (Milithronycteris) 

Hesperoptenus  blanfordi 

BM(NH)  83.853  Sepilok,  Sabah,  Borneo,  5°52'N,  1 17°56'E.  (Fig.  21g) 

Hesperoptenus  tickelli 

BM(NH)  71.12.26.1  Sri  Lanka.  (Fig.  21b) 

Hesperoptenus  tomesi 

BM(NH)  7.1.1.428  Malacca,  Malaya  (Holotype).  (Fig.  2 la) 

Chalinolobus  gouldi 

BM(NH)  71.1504  Westwood,  near  Rockhampton,  Queensland,  Australia.  (Fig.  17b) 

Chalinolobus  morio 

BM(NH)  6.8.1.60  (King  River,  Western  Australia.  (Fig.  17a) 

Chalinolobus  nigrogriseus 

BM(NH)  44.6.13.2  Port  Essington,  Northern  Territory,  Australia  (rogersi). 

BM(NH)  75.2260  Pine  Creek,  20  m  ESE  of  Gandy's  Hill,  Northern  Territory,  Australia,  13°49'S,  131°49'E. 
(rogersi).  (Fig.  17c) 

Chalinolobus  picatus 

BM(NH)  9.3.7.2  Gunnamulla,  Queensland,  Australia.  (Fig.  17d) 

Chalinolobus  tuberculatus 

BM(NH)  89.10.27.1  Outlying  islands  near  Stewart  I,  New  Zealand.  (Fig.  17e) 

Nyctophilinae 

Nyctophilus  bifax 

BM(NH)  67.5.6.5  Cape  York,  Queensland,  Australia. 

BM(NH)  77.3.28.1  Islands  of  Torres  Straits,  Australia. 

BM(NH)  86.1 1.8.12  Somerset,  Cape  York,  Queensland,  Australia. 

BM(NH)  15.3.13.1  Cloncurry,  Queensland,  Australia. 

BM(NH)  15.3.13.3  Herberton  District,  Queensland,  Australia  (Holotype).  (Fig.  22a) 

Nyctophilus  daedalus 

BM(NH)  47.7.2 1 . 1 6,  BM(NH)  -  -  Port  Essington,  Northern  Territory,  Australia.  (Fig.  22g,  BM(NH) 

47.7.21.16) 
BM(NH)  97.4.12.5  Daly  River,  Northern  Territory,  Australia. 

Nyctophilus  gouldi 

BM(NH)  15.3.13.7  Ash  I,  Hunter  River,  New  South  Wales,  Australia  (damaged,  part  lost). 

BM(NH)  15.3.13.8  Sydney,  New  South  Wales,  Australia. 

BM(NH)  -  -  Botany,  Sydney,  New  South  Wales,  Australia  (Original  No.  164)  (Fig.  22d) 

HZM  1.12085  Werrikimbe,  Hastingsshire,  New  South  Wales,  Australia.  (Fig.  16c) 

BM(NH)  52. 1 . 1 5.30  Tasmania  (sherrini).  (Fig.  220 

Nyctophilus  geoffroyi 

BM(NH)  15.3.13.1 1  Kosciusko,  New  South  Wales,  Australia  (pacificus). 

BM(NH)  -  -  Tasmania  (pacificus)  (Original  No.  M.I 735). 


VESPERTILIONINE  SYSTEMATICS  277 

BM(NH)  -  -  Launceston,  Tasmania  (pacificus)  (Original  No.  M.  168)  (Fig.  22e) 

BM(NH)  7.1.4.3  Alexandria,  Northern  Territory,  Australia  (pallescens).  (Fig.  22b) 

Nyctophilus  microtis 

BM(NH)  88.4.18.1  Sogeri,  Papua  New  Guinea  (Holotype).  (Fig.  22c) 

Pharotis  imogene 

BM(NH)  97.8.7.21  Kamali,  Papua  New  Guinea.  (Fig.  22h) 


278 


J.  E.  HILL  &  D.  L.  HARRISON 


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Table  2     Usual  incisive  and  premolar  dental  formulae  in  the  Vespertilioninae  and  Nyctophilinae.  Total 
number  of  teeth  (including  four  canines  and  twelve  molars)  in  parentheses.  Dental  notation  of  Miller  ( 1 907). 


(38)     Myotis,  Pizonyx 

(36)     Lasionycteris,  Plecotus,  Idionycteris,  Eudiscopus 

(34)     Euderma,  Barbastella,  la,  Pipistrellus,  Glischropus,  Scotozous,  Nyctalus, 

Chalinolobus 
(32)     Eptesicus,  Vespertilio,  Histiotus,  Tylonycteris ,  Mimetillus,  Glauconycteris, 

Pipistrellus,  Laephotis,  Philetor,  Hesperoptenus 
(32)     Lasiurus 
(30)     Rhogeessa,  Baeodon,  Nycticeius,  Otonycteris,  Dasypterus,  Scotomanes, 

Scotophilus,      Scoteanax,      Scotorepens,      Nycticeinops,      Scotoecus, 

Nyctophilus,  Pharotis 
(28)     Antrozous,  Bauerus 


Table  3    Classifications  of  the  Vespertilioninae  and  Nyctophilinae.  That  of  Tate  (\942a)  is  concerned  primarily  with 
Oriental  and  Australasian  taxa,  those  of  Koopman  with  Australasian  (1973)  and  predominantly  African  (1975)  forms. 


Tate(1942a) 


Koopman  (1973,  1975) 


Hill  &  Harrison 


Pipistrellus 
abramus  group 
abramus 
akokomuli 
bancanus 
camortae 
irretitus 
paterculus 
pumiloides 
pipistrellus  group 

pipistrellus  (Including 

bactrianus) 
nathusii 

coromandra  group 
aladdin 
angulatus 
collinus 
coromandra 
imbricatus 
meyeni 
micropus 
murrayi 
ponceleti 
portensis 
regulus 
sturdeei 
subulidens 
tramatus 
tenuis  group 
mimus  (Including 

glaucillus) 
nitidus 
papuanus  (Including 

orientalis) 
principulus 
tenuis 
ceylonicus  group 

ceylonicus  (Including 

chrysothrix,  indicus, 

subcanus) 


Pipistrellus 

Amalgamates  pipistrellus,  abramus 
( =javanicus),  coromandra  and 
tenuis  groups  of  Tate  (1942a) 
pipistrellus  group 
imbricatus 
javanicus  (Including 

abramus) 
meyeni 
nanus  (Including 

(?)  Helios) 
permixtus 

tenuis  (Including  angulatus, 
collinus,  nitidus,  papuanus, 
ponceleti,  murrayi,  sewelanus, 
subulidens,  westralis  [Koopman, 
1984c]) 


ceylonicus  group 
ceylonicus 


Pipistrellus 

Pipistrellus  (Pipistrellus) 
pipistrellus  group 
pipistrellus  subgroup 
pipistrellus  (Including  aladdin, 

bactrianus,  lacteus, 

mediterraneus) 
nathusii 
permixtus 
javanicus  subgroup 
abramus  (Including 

akokomuli,  irretitus, 

pumiloides) 
babu 
endoi 
javanicus  (Including  bancanus, 

camortae,  meyeni, 

'tralatitius") 
paterculus 
peguensis 

coromandra  subgroup 
adamsi 

angulatus  (Including  ponceleti) 
collinus 
coromandra  (Including  afghanus, 

portensis,  tramatus) 
mimus  (Including 

glaucillus,  principulus) 
murrayi 
papuanus 
sturdeei 
tenuis  (Including  nitidus, 

sewelanus,  subulidens) 
wattsi 
westralis 

ceylonicus  subgroup 
ceylonicus  (Including  borneanus, 

chrysothrix,  indicus,  raptor, 

shanorum,  subcanus) 
(?)  minahassae 


VESPERTILIONINE  SYSTEMATICS 


281 


Table  3-cont. 


Tate(1942a) 


Koopman  (1973,  1975) 


Hill  &  Harrison 


minahassae  group 
minahassae 

rueppellii  group 
coxi 


kuhlii  group 
babu 
canus 
kuhlii  (Including 

ikhwanius,  lepidus) 
leucotis 
lobatus 


Eptesicus 
pumilus  group 
pumilus  (Including 
caurinus,  darlingtoni, 
vulturnus) 
pygmaeus 


Pipistrellus 
savii  group 
austenianus 
cadornae 
curtatus 
macrotis 
savii 
vordermanni 


minahassae  group 

minahassae 
rueppellii  group 

rueppellii  (Including  (?) 
fuscipes;  pulcher) 


kuhlii  group 
aero 

anchietae 
deserti 
inexspectatus 
kuhlii  (Including  (?) 

aegyptius;  fuscatus) 
rusticus  (Including  marrensis) 


savii  group 
ariel 
macrotis 
maderensis 


Hesperus  group 
hesperus 
musciculus 


joffrei  group 
anthonyi 
joffrei 
stenopterus 


joffrei  group 
stenopterus 


rueppellii  group 
crassulus 
nanulus 

rueppellii  (Including  coxi, 
fuscipes,  leucomelas,  pulcher, 
senegalensis ,  vernayi) 
kuhlii  group 
aero 
deserti 

inexspectatus 
kuhlii  (Including  (?) 
aegyptius;  fuscatus, 
ikhwanius) 
maderensis 

rusticus  (Including  marrensis) 
Pipistrellus  ( Vespadelus) 
douglasorum 
pumilus  (Including 

darlingtoni) 
regulus 
sagittula 
vulturnus 
Pipistrellus  (Perimyotis) 

subflavus 

Pipistrellus  (Hypsugo) 
savii  group 
savii  subgroup 
anchietae  ( =  'bicolor'7) 
ariel 

austenianus 
bodenheimeri 
savii  (Including  caucasicus, 

darwini,  maurus) 
nanus subgroup 
arabicus 
helios 
musciculus 
nanus  (Including  culex, 

stampflii) 
pulveratus  subgroup 

pulveratus 
hesperus  subgroup 

hesperus 
eisentrauti  subgroup 

eisentrauti 
imbricatus  subgroup 
curtatus 
imbricatus 
macrotis 
vordermanni 
lophurus  subgroup 
cadornae 
kitcheneri 
lophurus 

stenopterus  group 
anthonyi 
joffrei 
stenopterus 


282 

Table  3-c  ont. 


J.  E.  HILL  &  D.  L.  HARRISON 


Tate(1942fl) 


Koopman(  1973,  1975) 


Hill  &  Harrison 


affinis  group 

affinis  group 

Pipistrellus  (Falsistrellus) 

affinis 

kitcheneri 

affinis  group 

kitcheneri 

petersi 

affinis 

lophurus 

(?)  mordax 

petersi 

petersi 

pulveratus 

tasmaniensis  group 

tasmaniersis  group 

tasmaniensis  (Including 

mackenziei 

krefftii) 

tasmaniensis  (Including 

circumdatus  group 

circumdalus 

mordax 
Eptesicus 

Eptesicus  (Amblyotus) 

alaschanicus 

bobrinskoi 


matschiei  (Including 

pellucens) 
nilssonii  (Including 

caucasicus,  centrasiaticus, 
gobiensis,  kashgaricus, 
pallescens,  tamer lani,  velox) 
tauricus 

walli  [alaschanicus, 
caucasicus,  pallescens, 
tamerlani,  tauricus, 
velox  allocated  to 
Pipistrellus  savii  by 
Kuzyakin,  1950] 
Eptesicus  (Rhyneptesicus) 
nasutus  group 

nasutus 

Eptesicus  (Eptesicus) 
fuscus  group 
bottae 
hingstoni 
serolinus  (Including 

andersoni,  brachydigitus, 
mirza,  pachyomus,  pollens, 
shirazensis,  sinensis) 


Eptesicus 
capensis  group 
brunneus 
capensis  ( =  notius)  (Including 

garambae,  grandidieri) 
guineensis  (Including 

(?)  rectitragus) 
melckorum 
somalicus  (Including  ugandae, 

vansoni,  zuluensis) 

tenuipinnis  group 
flavescens  ( =  angolensis) 
rendalli  (Including 

faradjius,  phasma) 
tenuipinnis  (Including  ater) 
Pipistrellus 

circumdatus  group 
circumdatus 
mordax 
Eptesicus 


serotinus  group 

bottae  (Including  innesi) 
hottentotus  ( =  megalurus) 

(Including  smithi) 
loveni 
platyops 

serotinus  (Including 
isabellinus) 


krefftii) 

Pipistrellus  (Neoromicia) 
capensis  group 
brunneus 

capensis  (Including  garambae 
grandidieri,  notius,  matrokd) 
guineensis  (Including 

rectitragus) 
melckorum 
somalicus  (Including 

ugandae) 

zuluensis  (Including  vansoni) 
tenuipinnis  group 
flavescens  (Including  angolensis) 
rendalli  (Including 

faradjius,  phasma) 
tenuipinnis  (Including  ater) 
Pipistrellus  (Arielulus) 
circumdatus 
cuprosus 
societatis 
Eptesicus 

Eptesicus  (Eptesicus) 
nilssonii  group 
bobrinskoi 
gobiensis  (Including 

centrasiaticus,  kashgaricus) 
nilssonii  (Including  japonensis, 

(?)  parvus;  propinquus) 
nasutus  group 

nasutus  (Including  batinensis 
matschiei,  pellucens, 
walli) 


serotinus  group 
serotinus  subgroup 

bottae  (Including  anatolicus, 

hingstoni,  innesi,  ognevi, 

omanensis) 
brasiliensis  (Incluidng  andinus, 

argentinus,  chiriquinus, 

melanopterus) 
diminutus  (Including 

dorianus ,  fide  Us) 
furinalis  (Including  inca, 

montosus) 
fuscus  (Including  hispaniolae, 

peninsulae) 
guadeloupensis 
hottentotus  ( =  megalurus) 


VESPERTILIONINE  SYSTEMATICS 


283 


Table  3-cont. 


Tate(  19420) 


Koopman(1973,  1975) 


Hill  &  Harrison 


sodalis  (Including 
ognevi) 


demissus  group 

demissus 

Eptesicus  (Pareptesicus) 
pachyotis  group 
pachyotis 


flower  i  group 
floweri  ( =  lowei) 


(Including  smithi) 
innoxius  (Including 

punicus) 
loveni 
lynni 

serotinus  (Including  andersoni, 
brachydigitus ,  horikawai, 
inter medius,  isabellinus, 
mirza,  pachyomus,  pollens, 
pashtomus,  platyops, 
shirazensis,  sinensis, 
sodalis,  turcomanus) 
tatei 
demissus  subgroup 

demissus 
(?)  pachyotis  subgroup 

pachyotis 

Eptesicus  (Rhinopterus) 
floweri  (Including  lowei) 


284 


J.  E.  HILL  &  D.  L.  HARRISON 


base 


shaft 


1 I 


Fig.  1     Bacular  types  in  Pipistrellus  and  Eptesicus  (see  text).  Scale  a-c  =  0-5  mm;  d-g  =  1  mm. 


VESPERTILIONINE  SYSTEMATICS 


285 


Fig.  2    Baculum  of  a,  Pipistrellus  pipistrellus  (D,  LL,  reversed);  b,  P.  nathusii  (D,  RL,  RVL);  c,  P. 
papuanus  (D,  RL);  d,  P.  subflavus  (D,  RL);  e,  P.  drcumdatus  (D,  LL,  RVL).  Scale  =  0-5  mm. 


286 


J.  E.  HILL  &  D.  L.  HARRISON 


Fig.  3     Baculum  (D,  RL)  of  a,  Pipistrellus  abramus;  b,  P.  endoi;  c,  P.  paterculus;  d,  P.  ceylonicus  (raptor). 

Scale  =  2  mm. 


VESPERTILIONINE  SYSTEMATICS 


287 


mm*  ..„.„.„„..,,... .' 

»^V.MtV.V.Jm...->-^' 


Fig.  4     Baculum  (D,  RL)  of  a,  Pipistrellus  babu;  b,  P.  collinus;  c,  P.  murrayi;  d,  P.  angulatus  (ponceleti). 

Scale  =  1  mm. 


^.•......:n.<^v,,.:...;:...1......,.,. "w"r'^> 


Fig.  5     Baculum  (D,  RL)  of  a,  Pipistrellus  kuhlii;  b,  P.  maderensis;  c,  P.  deserti;  d,  P.  rusticus. 

Scale  =  0-5  mm. 


VESPERTILIONINE  SYSTEMATICS 


289 


Fig.  6     Baculum  of  a,  Pipistrellus  savii  (D,  RL);  b,  P.  HO/IMS  (D,  RL);  c,  P.  rusticus  (D,  RL);  d,  P.  helios 
(D,  RL);  e,  P.  anchietae  (D,  LVL,  reversed).  Scale  =  0-5  mm. 


S?>t^KW^^       .,•,,...-,.,.:;:.,.    ,  :  -..,..  wmc.r.l.UW 


Fig.  7  Baculum  of  a,  Pipistrellus  arabicus  (D,  RL,  RVL);  b,  P.  coromandra  (tramatus)  (D,  RL);  c,  P. 
coromandra  (D,  RL);  d,  P.  ceylonicus  (D,  RL);  e,  P.  crassulus  (D);  f,  P.  nanulus  (D,  RL);  g,  P.  mimus 
(D,  RL);  h,  P.  stenopterus  (D,  RL,  RVL).  Scale  =  1  mm. 


Fig.  8  Baculum  of  a,  Pipistrellus  affinis  (D,  RL);  b,  P.  peter  si  (D,  RL);  c,  P.  pulveratus  (D,  RL,  RVL);  d, 
P.  Hesperus  (D,  LL,  reversed,  LVL);  e,  P.  kitcheneri  (D,  RL,  RVL);  f,  P.  lophurus  (D,  RL);  g,  P. 
tasmaniensis  (D,  RL,  V).  Scale  =  1  mm. 


Fig.  9  Baculum  (D,  RL  except  where  stated)  of  a,  Pipistrellus  imbricatus;  b,  P.  macrotis;  c,  P.  societatis, 
d,  P.  tennis  (nitidus)  (D,  RL,  RVL);  e,  P.  anchietae  ('Vesperus'  bicolor);  f,  P.  bodenheimeri;  g,  P. 
eisentrauti;  h,  P.  cuprosus.  Scales  a-g=  1  mm;  h  =  0-5  mm. 


_l 


Fig.  10  Baculum  (D,  RL)  of  a,  Pipistrellus  rueppellii  (pulcher);  b,  P.  rueppellii;  c,  P.  adamsi;  d,  P. 
westralis;  e,  P.  javanicus;  f,  Nyctalus  noctula;  g,  P.  wattsi;  h,  P.  mackenziei  (c,  d,  g,  h  from  Kitchener 
etal,  1986).  Scales  =  a,  b,  e,  f=2mm;c,  d,  g,  h=  1  mm. 


294 


J.  E.  HILL  &  D.  L.  HARRISON 


Fig.  1 1  Baculum  (V,  RL)  of  a,  Pipistrellus  pumilus  pumilus;  b,  P.  pumilus  (caurinus);  c,  P.  vulturnus;  d,  P. 
douglasorum;  e,  P.  regulus;  f,  P.  sagittula  (a-c,  e,  f  from  McKean  et  al.,  1970;  d  from  Kitchener,  1976). 
Scale  =  2  mm. 


Fig.  12  Baculum  (D,  RL)  of  a,  Pipistrellus  capensis  (matrokd);  b,  P.  capensis;  c,  P.  guineensis;  d,  P. 
zuluensis;  e,  P.  rendalli  (with  anterior  view);  f,  P.  melckorum;  g,  P.  capensis;  h,  P.  somalicus',  i,  P. 
capensis  ('minutus');  j,  P.  tenuipinnis;  k,  P.pumilus.  Scale  =  1  mm. 


Fig.  13  Baculum  (D,  RL)  of  a,  Eptesicus  fuscus;  b,  E.  hottentotus  (megalurus);  c,  E.  furinalis;  d,  E. 
brasiliensis  (andinus);  e,  E.  bobrinskoi;  f,  E.  floweri;  g,  E.  serotinus;  h,  E.  serotinus  (isabellinus);  i,  E. 
fuscus  (hispaniolae);  j,  £.  bottae  (innesi);  k,  £.  brasiliensis:  1,  E.  flower i(lowei).  Scale  =  1  mm. 


VESPERTILIONINE  SYSTEMATICS 


297 


Fig.  14  Baculum  (D,  RL  except  where  stated)  of  a,  Eptesicus  bottae  (omanensis);  b,  Pipistrellus  rendalli 
(?  brunneus);  c,  Eptesicus  nasutus;  d,  Plecotus  teneriffae  (D)  (from  Ibanez  &  Fernandez,  1986). 
Scales  =  1  mm. 


298 


J.  E.  HILL  &  D.  L.  HARRISON 


Fig.  15  Baculum  (D,  RL  except  where  stated)  of  a,  Eptesicus  nilssonii  (D)  (from  Topal,  1958);  b, 
Baeodon  alleni  (from  Brown  ef  a/.,  1971);  c,  Pipistrellus peguensis  (from  Sinha,  1969);  d,  P.  camortae; 
e,  Idionycteris  phyllotis,  f,  Plecotus  townsendii  (pallescens),  g,  P.  raftnesquii  (e-g  from  Nader  & 
Hoffmeister,  1983;  h,  P.  rafinesquii  (macrotis)  (from  Hamilton,  1949);  i,  Bauerus  dubiaquercus  (from 
Pine  etal.,  1971).  Scales  a  =  0-5  mm;  b,  e-i  =  l  mm;c,  d  =  2mm. 


I       .      I  I      I      I 

Fig.  16  Baculum  (D,  RL  except  where  stated)  of  a,  Otonycteris  hemprichii;  b,  Philetor  brachypterus  (D, 
RVL);  c,  Nyctophilus  gouldi,  d,  Scotozous  dormeri;  e,  Nycticeinops  schlieffenii',  f,  Laephotis  wintoni;  g, 
Scotorepens  balstoni;  h,  S.  greyii;  i,  Scoteanax  rueppellii.  Scales  a-h  =  1  mm;  i  =  2  mm. 


Fig.  17  Baculum  of  a,  C//a//no/o^w5  wor/o  (D);  b,  C.  gouldi  (D);  c,  C.  nigrogriseus  (rogersi)  (D);  d,  C. 
picatus  (D);  e,  C.  tuberculatus  (D);  f,  Lasionycteris  noctivagans  (D,  RL);  g,  Scotophilus  nigrita  (gigas) 
(D,  RL);  h,  5.  heathii(D,  RL);  i,  S.  kuhlii(D,  RL);j,  5.  nigritellus  (D,  RL);  k,  Nycticeius  humeralis  (D, 
LL,  reversed).  Scale  =  1  mm. 


Fig.  18  Baculum  (D,  RL)  of  a,  Glischropus  tylopus;  b,  Antrozous  pallidus;  c,  Histiotus  velatus;  d,  H.  (?) 
macrotis;  e,  ^.  macrotis;  f,  Dasypterus  argentinus;  g,  Scotomanes  ornatus;  h,  Tylonycteris  pachypus;  i, 
r.  robustula;j,  Barbastella  barbastellus;  k,  Rhogeessa  tumida.  Scale  =  1  mm. 


I 

Fig.  19  Baculum  of  a,  Glauconycteris  poensis  (D);  b,  G.  variegata  (D);  c,  G.  beatrix  (D);  d,  G.  argentata 
(D);  e,  G.  humeralis  (D);  f,  G.  variegata  (papilio)  (D);  g,  Plecotus  auritus  (D);  h,  /*.  austriacus  (D);  i, 
Myotis  ridleyi  (D,  RL);  j,  M.  nattereri  (D,  RL);  k,  Pizonyx  vivesi  (D,  RVL);  1,  Lasiurus  cinereus  (D, 
RVL).  Scale  =  1mm. 


Fig.  20     Baculum  (D,  RL)  of  a,  Scotoecus  albigula;  b,  5.  hindei  (falabae)  c,  5.  hirundo;  d,  S.  hindei;  e,  S. 

albofuscus.  Scales  a-d  =  2  mm;  e  =  1  mm. 


d       %**^ 


I I 


Fig.  21  Baculum  (D,  RL  except  where  stated)  of  a,  Hesperoptenus  tomesi,  b,  H.  tickelli,  c,  H.  doriae  (a-c 
from  Hill,  1 976);  d,  la  io  (D)  (from  Topal,  1 970);  e,  Scotorepens  orion,  f,  S.  sanborni  (e,  f  from  Kitchener 
&  Caputi,  1985);  g,  Hesperoptenus  blanfordi  (from  Hill  &  Francis,  1984);  h,  Scotoecus  pallidus  (from 
Agrawal  &  Sinha,  1973);  i,  Vespertilio  murinus  (V,  RL)  (from  Topal,  1958);  j,  V.  orientalis  (from 
Wallin,  1969).  Scales  a-c  =  2  mm;  d-h,  j=  1  mm;  i  =  0-5  mm. 


_-   '•••    •- 


Fig.  22     Baculum  (D,  RL)  of  a,  Nyctophilus  bifax;  b,  AT.  geoffroyi  (pallescens);  c,  M  microtis;  d,  Af.  gouldi; 
e,  A1,  geoffroyi  (pacificus);  f,  A7^.  gouldi  (sherrini);  g,  A7,  daedalus;  h,  Pharotis  imogene.  Scale  =  2  mm. 


British  Museum  (Natural  History) 

The  birds  of  Mount  Nimba,  Liberia 

Peter  R.  Colston  &  Kai  Curry-Lindahl 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1 962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Trmg,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1986,  129pp.  Hardback.  0  565  00982  6  £17.50. 


Titles  to  be  published  in  Volume  52 


Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophora  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxella.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata). 

By  P.  J.  Chimonides 


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


Bulletin  of  the 

British  Museum  (Natural  History) 


Notes  on  some  species  of  the  genus  Amathia 
(Bryozoa,  Ctenostomata) 


P.  J.  Chimonides 


Zoology  series    Vol  52    No  8    27  August  1987 


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 
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World  List  abbreviation:  Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


©  Trustees  of  the  British  Museum  (Natural  History),  1987 


The  Zoology  Series  is  edited  in  the  Museum's  Department  of  Zoology 

Keeper  of  Zoology  :  MrJ.  F.  Peake 
Editor  of  Bulletin  :  Dr  C.  R.  Curds 
Assistant  Editor  :  Mr  C.  G.  Ogden 


ISBN  0565  05032  X 

ISSN  0007  1 498  Zoology  series 

Vol  52  No.  8  pp  307-358 
British  Museum  (Natural  History) 
Cromwell  Road 
London  SW7  5BD  Issued  27  August  1987 


Notes  on  some  species  of  the  genus  A  mat  hi; 
Ctenostomata) 

P.  J.  Chimonides 

Department  of  Zoology,  British  Museum  (Natural  History),  Cromwel 


Contents 

Synopsis ......... 

Introduction 

Definition  and  assessment  of  taxonomic  characters 

Materials  and  methods 

Key  to  species 

Systematic  section 

Discussion 

References 


Synopsis 

The  species  of  the  genus  Amathia  have  presented  difficulties  of  recognition  for  a  long  time.  Even  the  most 
recent  revisionary  accounts  have  failed  to  establish  the  separate  identity  of  many  of  the  species;  these  have, 
instead,  been  submerged  in  erroneous  synonymies.  Fifteen  species  are  dealt  with  in  full.  In  the  main,  species 
without  significant  spiralling  of  the  autozooid  groups  are  considered.  Three  new  species  are  introduced:  A. 
guernseii,  A.  intermedis  and  A.  tricornis.  A  fourth  species,  A.  populea  Busk  MS  in  d'Hondt,  is  recognised  as 
new.  A.  distans  var  aegyptana  is  raised  to  specific  rank.  A.  cornuta  Lamouroux  (preocc.)  is  renamed  A. 
lamourouxi.  A.  obliqua  and  A.plumosa  MacGillivray  are  redescribed.  A.  alternata  Lamx.,  A.  biseriata  Krauss, 
A.  brongniartii  Kirkpatrick,  A.  lendigera  Linn.,  A.  pruvoti  Calvet,  A.  pinnata  and  A.  wilsoni  Kirkpatrick,  A. 
woodsii  Goldstein,  are  redefined  with  type  material  selected.  Characters  for  use  in  taxonomic  and  systematic 
discrimination  are  introduced,  and  brief  comment  is  made  on  the  zoo-  and  palaeo-geography  of  the  species 
dealt  with. 


Introduction 

The  genus  Amathia  was  erected  in  1812  by  J.  V.  F.  Lamouroux,  consequential  to  the  study  of 
material  collected  from  Tasmania  and  the  south  coast  of  Australia  by  C.  A.  Lesueur  and  F.  Peron, 
during  the  years  1800-1804  (Tenison  Woods  1880,  d'Hondt  1979). 

However,  Sertularia  lendigera  of  Ellis  (1755)  from  European  waters  became  the  type  species  of 
the  genus,  by  virtue  of  being  Lamouroux's  sole  mentioned  species  at  the  introduction  of  the  genus 
(I.C.Z.N.  article  69(d)).  The  species  itself  was  validated  with  the  publication  of  the  10th  edition  of 
the  Sy sterna  Naturae  by  Linneaus  in  1758  (and  is  therefore  technically  that  of  Linneaus). 

Ryland  (1982)  gave  a  revised  perspective  classification  of  the  genus  but  there  are  differences 
between  his  definitions  of  higher  categories,  including  Amathia,  and  the  characters  of  the  genus 
presented  here.  At  the  Family  level,  Ryland  described  the  'zooids'  as  being  'radially  symmetrical, 
no  face  being  partly  membranous';  and  at  the  Superfamily  level,  he  described  'branching  being 
irregular'.  Both  descriptions  are  inaccurate  for  Amathia.  Similarly  d'Hondt  (1983)  for  the  Family 
level,  also  described  'External  autozoecial  symmetry'  as  'radiated',  while  at  the  Superfamily  level, 
there  was  some  ambiguity  in  the  definitions  of  the  characteristics  employed  e.g.  for  the  'Zoarium 
. . .  autozoecia  unconnected  to  their  neighbours'.  Clearly  there  is  need  for  a  review  of  the  characters 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  52  (8):  307-358 


Issued  27  August  1987 


307 


308  P.  J.  CHIMONIDES 

used  in  the  definition  of  the  higher  taxonomic  levels,  although  this  is  beyond  the  scope  of  this 
account. 

The  persistent  problem  has  been  how  to  differentiate  between  the  numerous  species.  The  brief 
descriptions  often  given  are  of  little  help,  and  in  fact  have  led  to  some  confusion.  Often,  widespread 
geographical  distributions  have  been  suggested.  Where  no  figures  or  specimens  are  available,  it  is 
unlikely  that  the  true  identity  of  some  species  will  ever  be  recognised.  Despite  the  efforts  of  d'Hondt 
(1979,  1983)  the  picture  still  remains  clouded. 

This  account  attempts  to  establish  criteria  for  species  differentiation  within  the  genus;  to  identify 
some  species  groupings  based  upon  these  criteria  and  in  the  process  to  discuss  and  correct  past 
misconceptions. 

In  general,  the  species  of  Amathia  may  be  assigned  to  either  of  two  groupings:  those  with 
autozooids  spirally  disposed  about  the  stolon;  those  with  autozooids  disposed  linearly  along  the 
stolon.  It  is  mainly  the  latter  group  which  is  discussed  here.  Where  spirally  disposed  species  are 
dealt  with,  this  is  mainly  to  obviate  possible  confusion  with  those  species  (i.e.  A.  alternata  and 
A.  pruvoti)  in  which  marked  twisting  of  the  autozooid  groups  occurs  along  the  stolon.  It  is  in  the 
context  of  comparison  with  A.  pruvoti  that  A.  distans  var  aegyptana  is  considered. 

Definition  and  Assessment  of  Taxonomic  Characters 

Waters  (1910)  in  his  brief  account  of  the  genus,  outlined  a  number  of  characters  which  may  serve  as 
a  foundation  on  which  to  build  an  understanding  of  both  the  genus  and  its  species.  These  charac- 
teristics may  be  added  to,  and  arranged  in  what  is  considered  here  to  be  an  order  of  decreasing 
reliability,  reflecting  an  increase  in  their  intraspecific  variability. 

List  of  species  discrimination  characteristics  in  order  of  reliability 

( 1 )  budding  pattern  of  stolons 

(2)  development  of  any  kenozooidal  processes  or  rhizoids  and  their  orientations 

(3)  arrangement  of  autozooids  about  the  stolons 

(4)  autozooidal  thickening 

(5)  profile  of  autozooids  and  stolons 

(6)  number  of  autozooids  and  proportion  of  stolon  occupied  by  autozooids 

(7)  dimensions  of  components 

The  potentially  informative  characters  of  larval  type,  larval  metamorphosis,  ancestrula 
formation  and  initial  colony  development  are  generally  not  known  and  hence  cannot  be  evaluated. 
Despite  the  explicit  account  given  by  Barrois  (1877)  for  A.  lendigera,  the  ancestrula  and  earliest 
astogenetic  stages,  for  example,  have  not  been  recognised  in  any  of  the  specimens  examined.  In 
some  specimens  it  was  clear  that  this  part  of  the  colony  was  absent;  in  others,  it  was  impossible  to 
see  because  of  heavy  overgrowth  by  the  colony's  own  rhizoids  or  by  spatial  competitors.  For  these 
same  reasons,  in  the  following  systematic  accounts,  no  information  is  given  on  the  non-erect  part 
of  the  colony  for  the  majority  of  species.  It  is  possible  that  some  colonies  are  the  result  of  associ- 
ation between  the  products  of  more  than  one  ancestrula,  (without  the  necessity  for  fusion  to 
have  occurred,  especially  in  the  non-arborescent  growth  forms).  Zimmer  and  Woollacott  (1977a) 
suggested  that  the  larval  type  of  all  stoloniferan  ctenostomes  is  the  same.  It  would  appear  however, 
that  their  conclusions  were  drawn  from  only  three  species:  'Amathia  lendigera',  Bowerbankia 
pustulosa  (Ellis  and  Solander)  and  Zoobotryon  verticillatwn  (Delle  Chiaje).  Furthermore,  Zimmer 
and  Woollacott  (19776)  pointed  out  that  past  accounts  of  metamorphosis  of  larvae  in  this  group 
were  inconsistent,  and  that  'additional  work  is  essential  to  clarify  the  pattern(s)  of  metamorphosis' 
of  the  larval  type.  Waters'  (1910)  intuitive  suggestion  of  the  'valuable  assistance'  which  the  primary 
zooecia  might  give  must,  therefore,  be  discounted  for  the  present. 

Extensive  examination  of  several  large  colonies  indicates  that  the  branching  pattern  of  stolons 
remains  remarkably  consistent  within  species.  Differences  in  branching  pattern  may  be  inferred  to 
have  been  microenvironmentally  induced  in  that  they  tend  to  be  sporadic,  involve  the  development 
of  new  stolens  from  astogenetically  early  regions  of  the  colony,  and  are  often  associated  with  the 


AMATHIA  309 

presence  of  epibionts.  The  budding  patterns,  together  with  kenozooidal  processes,  rhizoids,  auto- 
zooidal  thickenings  and  disposal  of  autozooids  about  the  stolons,  can  give  rise  to  characteristic 
colony  shapes  which,  with  familiarisation,  allow  identification  of  species  by  casual  inspection. 

The  growth  of  a  colony  relies  essentially  on  the  production  of  the  supporting  'stolonal'  keno- 
zooids.  Where  such  a  kenozooid  is  destined  to  bear  autozooids,  the  autozooids  are  usually  seen  to 
develop  at  about  the  same  time  as  the  kenozooid  lengthens  through  apical  growth.  Autozooid- 
bearing  kenozooids  are  here  termed  stolons.  The  position  of  the  autozooids  and  the  proportion  of 
stolon  occupied  by  them  is  highly  regular.  Stolonal  and  autozooidal  growth  is  considered  to  have 
ceased  with  the  production  of  septa  at  the  distal  end  of  the  stolon,  and  the  subsequent  appearance 
of  daughter  stolonal  buds.  The  kenozooidal  processes  of  character  No.  2  in  the  above  list  appear  to 
be  growth-terminating  features.  These  are  usually  distinguishable  from  potential  autozooid- 
bearing  stolonal  kenozooids  by  being  straighter,  narrower,  often  tapering  to  a  point,  and 
frequently  being  subdivided  by  septa. 

Where  rhizoids  are  to  be  produced  by  a  structure,  the  origin  of  each  rhizoid  is  marked  first  by  the 
appearance  of  an  oval  window  in  the  cuticle.  Rhizoids  then  develop  as  papilliform  outgrowths  of 
these  windows,  proximally  directed  along  the  colony,  growing  towards  the  colony  base.  The 
rhizoids  sometimes  fuse  with  each  other  en  route,  overgrowing  and  obscuring  underlying  stolons, 
and  forming  a  trunk-like  mass.  Autozooids  and  lateral  branches  of  overgrown  stolons  are  often 
shed.  When  rhizoids  are  produced,  the  resulting  colony  form  is  usually  arborescent. 

The  overall  cuticular  thickening  of  any  colony  appears  uniform,  except  at  the  regions  of  the 
growing  tips,  where  it  is  thinner.  This  level  of  cuticular  thickening  can  differ  between  colonies  of  the 
same  species.  As  this  thickening  is  contributory  to  colour,  it  follows  that  colour  is  also  variable. 
Within  all  species,  there  are  localised  areas  of  thickening,  which  tend  to  be  constant.  Thickening  in 
the  autozooids,  which  gives  their  groupings  a  characteristic  appearance,  may  be  used  to  discrimi- 
nate between  species.  Two  conditions  occur:  one  where  the  walls  between  autozooids  are  differen- 
tially thickened  (inner-wall  thickening);  the  other  where  the  outer  walls  are  differentially  thickened 
(outer-wall  thickening)  (see  Fig.  IB,  C). 

The  arrangement  of  autozooids  on  the  stolon  is  usually  described  as  being  paired,  or  as  a  biserial 
row.  Although  this  appears  correct,  in  all  specimens  examined,  displacement  of  autozooids  occurs, 
so  that  the  autozooids  of  one  row  interlock  with  the  recesses  between  the  autozooids  in  the  other 
row  (Dalyell,  1847  for  A.  lendigerd).  Very  frequently,  this  emphasises  a  single  proximal-most 
autozooid  in  each  group.  No  pairing  of  autozooids  may  be  confidently  assigned  throughout  a 
colony  in  any  species,  and  there  can  be  odd  or  even  numbers  of  autozooids  in  any  autozooid  group. 
In  some  cases,  notably  those  with  inner-wall  thickening,  the  proximal-most  autozooid  tends  to  be 
larger  in  cross  section  and  displaced  centrally,  such  that  it  can  be  very  difficult  to  assign  it  to  a  row 
of  origin  at  any  stage  in  its  ontogeny. 


Materials  and  methods 

Specimens  used  for  study  were  mainly  those  of  the  British  Museum  (Natural  History),  London, 
(BMNH)  and  The  Manchester  Museum,  (MM),  with  additional  material  referred  to  as  follows: 
Laboratoire  de  Biologic  des  Invertebres  Marins  et  Malacologie  of  the  Museum  National 
d'Histoire  naturelle,  Paris,  (LBIMM);  the  National  Museum  of  Victoria,  Melbourne,  (NMV); 
the  Rijksmuseum  van  Natuurlijke  Historic,  Leiden  (RM);  and  the  U.S.  National  Museum, 
Washington,  (USNM). 

Often,  the  material  for  study  had  been  preserved  dried,  with  resulting  distortions.  To  observe 
the  autozooidal  characteristics  preserved  in  the  cuticular  thickenings,  it  was  found  far  better  to 
rehydrate  the  specimens  although  it  was  still  possible  to  make  identifications  without  treatment. 
Rehydration  was  carried  out  using  tri-sodium  phosphate  in  7-10%  aqueous  solution,  with  subse- 
quent transfer  to  distilled  water  and  then  via  a  sucession  of  increasing  concentrations  of  alcohol,  to 
80%  concentration  for  storage.  From  this  process,  specimens  regained  the  turgidity  associated 
with  their  living  state.  It  was  from  specimens  in  this  turgid  state  that  measurements  were  taken. 


310  P.  J.  CHIMONIDES 

Some  care  was  needed,  as  rupture  of  specimens  was  possible  through  the  initial  high  osmotic 
differential  established  on  transfer  to  distilled  water.  In  some  cases,  specimens  failed  to  reflate 
because  of  existing  ruptures  in  their  cuticles. 

In  the  ensuing  descriptions,  anterior  is  used  to  denote  the  side  at  that  location  bearing  auto- 
zooids,  and  posterior,  the  side  opposite  (see  Fig.  ID,  E).  Dimensions  are  given  in  millimetres  and 
are  means  of  a  minimum  of  30  measurements.  Where  shape  negated  the  validity  of  a  single  sample 
measurement,  extreme  dimensions  are  given,  these  also  being  the  means  of  30  measurements  each. 
Measurements  were  made  in  ontogenetically  complete  components,  near  distal  regions,  avoiding 
where  possible,  astogenetically  earlier  (older)  regions  of  the  colony.  No  attempt  was  made  to 
determine  intra-colony  variations  quantitatively.  Where  these  were  noted,  they  were  assessed 
subjectively. 
The  following  abbreviations  are  used: 

SI.  length  of  stolon 

Sd.  diameter  of  stolon,  at  location  specified;  usually  midway  along  the  proximal  autozooid-free  end. 

Zh.  autozooid  height  to  the  highest  point  on  the  rim  of  thickening,  of  tallest  autozooids,  unless  otherwise 

specified. 

Zw.  autozooid  width,  measured  along  the  stolonal  axis. 

Z/S.  the  linear  proportion  of  stolon  occupied  by  autozooids 

Zn.  the  number  of  autozooids  per  autozooid  group  (and  apparent  number  of  'pairs') 

Tpl.  length  of  terminal  process 


Key  to  species 

(Identification  is  best  attempted  with  plentiful  material.) 

1  Rhizoids  developed,  colony  frequently  aborescent 

Rhizoids  not  developed,  colony  not  arboresecent,  no  terminal  processes  and  Z/S  ratio  <  50%     .       1 3 

2  Autozooidal  thickening  inner-wall brongniarttt 

Autozooidal  thickening  outer-wall 

3  Branching  nearly  always  bifurcate 4 

Branching  primarily  tri- and  tetrafurcate,  bifurcation  may  also  be  present 10 

4  Terminal  processes  developed 5 

Terminal  processes  not  developed 

5  A  pair  of  lanceolate,  single-kenozooidal  terminal  processes  developed  at  the  distal  end  of  each 

autozooid  group,  arising  in  the  same  direction  as  the  autozooids     .        .        .        lamourouxi 
Lanceolate  terminal  processes  of  compound  kenozooidal  construction,  each  filament  developed  in 
place  of  a  normal  stolon,  sometimes  branched 6 

6  Rhizoids  developed  postero-laterally,  terminal  processes  never  branched    .        .        .     populea 
Rhizoids  developed  anteriorly,  terminal  processes  often  forked woodsii 

1    Autozooids  re-orientated  by  approx.  180  deg.  from  stolon  to  stolon,  polyrhizoidy  (see  page  335) 

possible alternata 

Autozooid  orientation  from  stolon  to  stolon  maintained  within  30  deg.,  rhizoids  paired  at  most  .         8 

8  Rhizoids  developed  anteriorly,  autozooids  with  marked  distal  inclination,  autozooid  group 

profile  diminishing  distally guernseii 

Rhizoids  developed  postero-laterally,  autozooid  group  profile  horizontally  even  i.e.  level    .        .         9 

9  Stolons  curved  anteriorly,  curvature  increasing  distally,  autozooid  group  arranged  in  line  with 

stolonal  axis biseriata 

Stolons  straight,  autozooid  group  set  obliquely  to  stolonal  axis obliqua 

10  Autozooid-bearing  stolons  developed  laterally  from  a  central  axis  of  stolon-sized,  or  larger, 

kenozooids.  Autozooid-bearing  stolon  sequences  end  with  compound  terminal  processes,  these 

often  forked plumosa 

Autozooid-bearing  stolons  developed  laterally  from  a  central  axis  of  other  autozooid  bearing 

stolons  11 


AMATHIA  311 

1 1  Central  axis  stolons  undergo  trifurcation  only;  indistinguishable  from  lateral  stolons  .      pinnata 
Central   axis   stolons   usually   undergo   tetrafurcation,   a   fourth   autozooid-bearing   stolon 

developed  posteriorly.  Central  axis  stolons  morphologically  distinguishable  from  lateral 
stolons,  differences  may  be  slight 12 

12  Autozooid-bearing  stolon  sequences  end  with  lanceolate,  compound  kenozooidal,  terminal  pro- 

cesses, each  replacing  a  normal  stolon  and  thus  in  complements  of  three.  Pronounced  difference 

between  central  axis  and  lateral  stolons tricornls 

Autozooid-bearing  stolon  sequences  with  pinnate,  compound  terminal  processes;  each  assemb- 
lage replacing  stolons  in  other,  regular  positions,  giving  characteristic  arched  colony  sub  units. 
Difference  between  central  axis  and  lateral  stolons  slight wilsoni 

13  Autozooid  groups  regularly  twisted  along  stolon  length pruvoti 

Autozooid  groups  rarely  showing  any  twist 14 

14  Stolons  often  in  rectilinear  series,  straight,  sometimes  undergoing  trifurcation.  Autozooid  groups 

often  remote  from  subsequent  branching  point.  Autozooids  usually  erect        .        .  intermedis 

Stolons  of  variable  length,  usually  short,  sculptured  and  posteriorly  deflected.  Autozooid  groups 

overlie  subsequent  branching  point,  autozooids  inclined  distally,  the  lean  increasing  distally    . 

lendigera 


312  P.  J.  CHIMONIDES 

Systematic  Section 

Phylum  BRYOZOA  Ehrenberg,  1831 
Class  GYMNOLAEMATA  Allman,  1856 

Order  CTENOSTOMATA  Busk,  1852 
Genus  AM  A  THIA  Lamouroux,  1812:  p.  184 

Part  Sertularia  Linnaeus,  1758. 

Serialaria  Lamarck,  1816. 

Part  Valkeria  Dalyell,  1847. 

AmathellaGray,  1858. 

CharadellaGray,  1858. 

Serialia  Gray,  1858.  (errorum  pro  Serialaria  Lamarck,  1816). 

SpiraliaGray,  1858. 

CornaliaGray,  1858. 

Amathia:  Bobin  &  Prenant,  1956:  (incomplete  cum.  syn.,  NB.  Gray  1858  misquoted  as  1848);  Ryland,  1982; 

Winston,  1982;d'Hondt,  1979,  1983;Hayward,  1985. 

TYPE  SPECIES.  A.  lendigera  (Linnaeus  1758  sensu  Ellis  1755)  Lamouroux  1812:  p.  184. 
GENERIC  DESCRIPTION.  Colonies  mainly  erect  with  a  creeping  base,  this  sometimes  extensive. 
Autozooid  groups  displaced  towards  the  distal  portion  of  the  stolon.  Stolons  may  produce 
rhizoids,  proximally  disposed.  Distal,  mainly  growth-terminating  kenozooidal  processes  may  be 
developed  from  various  positions.  Autozooids  with  gizzards,  borne  on  kenozooidal  stolons, 
arising  from  rosette  plates,  in  groups,  connate  for  at  least  part  of  their  length,  appearing  biserially 
arranged  as  a  straight  or  spiral  series. 

REMARKS.  The  only  attempt  to  regroup  species  comprising  the  genus  Amathia  was  made  by 
Gray  (1858,  duplicated  1859).  Gray  introduced  several  indeterminate  subgeneric  or  generic 
groups,  the  type  species  of  which  were  insufficiently  described  and  not  illustrated.  The  great 
majority  of  the  limited  characteristics  employed  are  variable  within  species,  such  that  none  of  the 
divisions  Gray  introduced  exclusively  defines  any  species  group  identifiable  within  the  genus. 
Bobin  &  Prenant  (1956)  are  followed  here  in  assigning  all  species  described  to  the  genus  Amathia. 

Amathia  lendigera  (Linnaeus,  1758) 
(Figs  6A,  7A) 

?  Sertularia  lendigera  Ellis,  1755:  27,  pi.  15  (figs  24B,  24b). 

Sertularia  lendigera  Linnaeus,  1758:  812. 

Amathia  lendigera  Lamouroux,  1812:  184. 

Not  Amathia  lendigera:  MacGillivray,  1895:  135,  pi.  B  (fig.  1). 

Not  Amathia  lendigera:  O'Donoghue  &  de  Watteville,  1944:  430  (=  A.  populea). 

Part  Amathia  lendigera:  Bobin  &  Prenant,  1956:  280. 

Amathia  lendigera:  Hay  ward,  1985:  134,  fig.  45B. 

MATERIAL  EXAMINED 
Neotype  (selected  here):  BMNH;  1963.1.8.3,  Chichester  Harbour,  H.  G.  Stubbings  collected. 

OTHER  MATERIAL 

BMNH;  1827.11.18.8,  no  locality.  1852.3.16.62,  Weymouth.  1882.7.7.85,  no  locality.  1887.7.23.5,  Solent, 
I.O.Wight.  1891.8.7.18,  Portland,  Dorset.  1897.8.9.67,  Weymouth  Bay,  Portland,  10  fthms.  (18.29m). 
1899.5.1.21  l,?OffSaints  Bay,  Guernsey?  1900.10.30.10-11,  Weymouth.  1912.12.21.681,  Plymouth. 

MM;  7093-4,  Naples.  7095,  Roscoff,  France.  7096-9,  Swanage.  7105,  Naples.  7106,  St.  Raphael,  S.  France. 
7107,  Rapallo(  =  Rapolla,  Italy?). 

DESCRIPTION.  Colonies  tend  to  have  to  have  a  moderately  extensive  creeping  component  of 
stolonal  kenozooids.  These  are  adpressed  to  the  substratum  and  closely  follow  its  profile,  showing 
reduced  branching  in  some  places  and  multiple  branching  in  others.  These  stolonal  kenozooids  are 
usually  of  irregular  form  and  length,  and  only  rarely  bear  autozooids.  Bilateral  palmate  processes 
are  often  produced,  through  which  adhesion  to  the  substratum  is  effected.  Erect  components  may 


AMATHIA  313 

be  produced  at  any  branching  point,  with  or  without  continuation  of  the  creeping  component.  The 
erect  components  develop  as  the  characteristic  autozooid-bearing  stolons,  arranged  in  the  typical 
form  of  an  orbicular  mass,  cotton-wool  like  in  appearance,  utilizing  well  the  available  free  space 
near  to  the  substratum.  These  erect  components  appear  tangled,  but  are  rarely  so.  Any  erect 
component  may  resume  the  creeping  habit  on  contact  with  the  substratum.  Branching  in  the  erect 
part  of  the  colony  is  practically  always  bifurcate,  ranging  from  equally  dichotomous  to  almost 
rectilinear  with  side  branches,  these  appearing  on  alternate  sides.  Bifurcations  typically  form  an 
angle  of  90  deg.  Autozooid  group  orientation  about  the  stolon  is  not  usually  preserved  from  stolon 
to  stolon.  Daughter  stolons  often  arise  deflected  anteriorly  to  maternal  stolons.  Autozooid  groups, 
with  relatively  few  autozooids,  occurring  at  the  extreme  distal  end  of  stolons,  frequently  over- 
lapping the  subsequent  branching  point.  Stolons  are  often  deflected  posteriorly  at  the  proximal  end 
of  the  autozooid  group,  and  also  raised  slightly  on  the  anterior  surface  at  this  same  region.  Stolons 
may  be  of  variable  length.  Autozooids  are  outer-wall  thickened.  Autozooid  profile  diminishes 
distally,  due  in  part  to  decreasing  autozooid  height,  and  in  part  to  increasing  distal  inclination  of 
the  autozooids.  A  proximal-most  autozooid  is  usually  prominent  in  each  group  and  is  displaced 
centrally.  Where  not  truly  central,  this  autozooid  remains  on  the  same  side  of  its  stolon  as  the 
direction  in  which  that  stolon  was  budded.  The  arrangements  of  autozooids  on  sister  stolons  are 
therefore  mirror  images  of  each  other  (see  Fig.  5B).  Where  stolons  form  linear  sequences,  auto- 
zooid groups  borne  on  such  series  tend  to  show  an  alternate  sequence  of  autozooid  displacements 
on  successive  stolons.  Sometimes  sister  stolons  carry  identical  autozooid  displacements,  these 
being  opposite  to  that  on  their  maternal  stolon.  No  overall  pattern  is  discernible  within  the  colony 
in  the  occurrence  of  this  second  state  of  succession  (see  Fig.  5C).  Rhizoids  are  absent. 

SI.     1.25-2.75  Z/S.  25-50% 

Sd.  0.75-0.97  Zn.  8-1 7  (appearing  as  4^8 'pairs') 

Zh.  0.33-0.50 

Zw.  0.10-0.12 


REMARKS.  According  to  Harmer  (1931),  the  original  specimens  described  and  figured  by  Ellis 
(1755)  were  not  kept  (I.C.Z.N.  article  73(b)  (i),  recommendation  69B).  Harmer  stated  that  speci- 
mens of  A.  lendigera  were  sent  to  Linnaeus  by  Ellis,  but  some  12  years  after  the  publication  of  the 
nomenclaturally  significant  10th  edition  of  Linnaeus'  Sy sterna  Naturae  (1758).  Two  specimens, 
under  the  original  name  ofSertularia  lendigera,  are  still  in  the  collections  of  the  Linnean  Society  of 
London  (Nos.  1298.17  and  1298.18).  The  specimens  are  preserved  pressed  dry  on  paper,  and  both 
are  labelled  as  'lendigera'  in  Linnaeus'  handwriting.  From  examination  of  these  specimens,  some 
doubt  arises  that  Ellis  and  Linnaeus  were  sufficiently  rigid  in  their  interpretation  of  A.  lendigera. 
Two  species  are  present:  specimen  1298.17  is  identifiable  as  A.  semiconvoluta  (see  pages  335,  338); 
while  specimen  1298.18  is  probably  A.  lendigera.  Linnaeus  (1758)  has  trustingly  used  Ellis'  (1755) 
description  verbatim.  If  the  specimens  originated  from  Ellis,  Linnaeus  may  also  have  accepted 
their  identity  from  him.  It  is  possible  therefore,  that  the  mistaken  identity  of  1298.17  could  be 
attributed  to  Ellis;  neither  man  realising  the  presence  of  mixed  material. 

However,  there  is  some  evidence  in  support  of  Harmer' s  statement  that  the  Linnaean  specimens 
are  not  Ellis'  original  (1755)  material.  Linnaeus  is  reported  to  have  been  in  the  habit  of  upgrading 
his  botanical  collections,  with  the  replacement  of  older  specimens  by  new,  'some  of  them  not 
conspecific  by  modern  taxonomic  standards'  (Stearn,  1957),  a  practice  which  could  also  have  been 
applied  to  herbarium  preparations  of 'zoophytes'.  In  addition,  none  of  the  figures  of  Ellis  (1755) 
correspond  with  either  of  the  Linnean  Society  specimens,  in  particular  specimen  1298. 18.  Features 
of  importance  are:  the  arborescent  and  open  appearance  of  the  colony  shape  in  figure  '24b';  the 
number  of  autozooids  per  stolon  indicated  by  the  magnified  view  in  figure  '24B'.  Although  only  a 
single  line  of  autozooids  is  drawn  in  the  latter  figure,  this  may  be  interpreted  as  showing  either:  a 
single  proximal-most  autozooid  with  indications  of  the  outlines  of  subsequent  'paired'  autozooids; 
or  possibly  a  line  of  'all  paired'  autozooids.  The  condition  depicted  is  readily  seen  in  many  dry 
preserved  specimens,  where  only  the  thickened  outer  walls,  forming  the  periphery,  survive  well.  As 


314  P.  J.  CHIMONIDES 

such,  1 7  or  1 8  autozooids  would  be  represented  on  three  of  the  five  stolons;  1 5  autozooids  would  be 
represented  on  one  of  the  remaining  two;  and  there  is  an  inexplicable  absence  of  autozooids  on  the 
remaining  fifth  and  final  stolon.  Although  notionally  possible,  it  is  very  unusual  for  A.  lendigera  to 
show  as  many  autozooids  per  stolon  in  direct  succession  in  a  colony.  The  importance  of  this 
analysis  is  that  figure  '24B'  is  claimed  as  an  exact  microscope  drawing. 

Harmer  (1931)  suggested  that  figures  of  Ellis  be  regarded  as  the  lectotype  of  the  species. 
However,  the  figures  are  inadequate,  no  rhizoids  are  shown,  and  their  presence  or  absence  is  not 
indicated  in  the  description.  On  the  cumulative  evidence  (see  above),  figures  '24b'  and  '24B'  could 
thus  be  depictions  of  A.  intermedis  or  even  A.  guernseii. 

Selection  of  a  neotype  specimen  is  the  only  satisfactory  way  to  resolve  the  identity  of  A. 
lendigera;  particularly  important  as  the  species  is  the  type  of  the  genus.  There  is  no  indication  that 
the  Linnean  Society  specimen  1298.18  formed  any  basis  for  the  description  for  the  species.  In 
addition  to  the  uncertainties  surrounding  its  status,  1298.18  unfortunately  also  lacks  sufficient 
locality  data,  is  not  in  an  adequate  state  of  preservation,  and  so  should  not  be  considered.  Specimen 
BMNH  1963.1.8.3  is  therefore  selected  here  as  neotype.  It  is  preserved  in  alcohol,  growing  on 
Halidrys  siliquosa  as  is  the  Linnean  Society  specimen.  BMNH  1963.1.8.3  is  erroneously  listed  by 
d'Hondt  (1983)  as  A.  pruvoti,  a  very  different  species  (see  pages  336,  337). 

There  is  great  similarity  between  A.  lendigera  and  A.  intermedis  and  both  resemble  A.  guernseii 
(see  pages  316,  317).  The  morphologies  of  all  three  may  overlap  in  different  parts  of  the  colony.  A. 
lendigera  differs  from  A.  intermedis  in  that:  it  tends  to  have  fewer  autozooids  per  autozooid  group; 
the  autozooids  have  an  increased  distal  inclination;  the  autozooid  group  profile  diminishes  distally 
more  rapidly;  the  autozooid  groups  and  subsequent  bifurcation  sites  are  more  condensed  relative 
to  each  other;  it  has  a  more  compact  colony  form,  with  low  incidence  of  rectilinear  succession. 
Great  care  is  needed  to  distinguish  between  the  trifurcation  that  may  occur  in  the  erect  part  of  the 
colony  of  A.  intermedis,  and  the  multiple  branching,  including  trifurcation,  which  occurs  in  the 
immediate  vicinity  of  the  non-erect  part  of  A.  lendigera,  as  detachment  from  the  substratum  is 
frequent  in  preserved  specimens.  Non-erect  stolons  may  usually  be  identified  by  the  nearby 
presence  of  palmate  processes  (see  Fig.  8A),  and  the  irregular  morphology  associated  with  the 
creeping  mode. 

The  displacement  of  the  proximal-most  autozooids  in  maternal  and  daughter  stolons,  may 
reflect  the  timing  of  the  production  of  daughter  stolons  relative  to  each  other.  The  mirror  image 
arrangement  (see  Fig.  5B)  possibly  results  from  the  simultaneous  production  of  the  daughters. 

Most  of  the  published  records  for  A.  lendigera  are  listed  in  a  lengthy  synonymy  by  Bobin  and 
Prenant  (1956).  However,  many  of  these  records  are  unsupported  by  specimens  available  for 
examination  and  are  thus  equivocal.  In  addition,  the  account  these  authors  give  mentions  the 
occurrence  of  rhizoids,  and  thus  includes  another  species,  probably  A.  guernseii. 

Three  specimens  in  the  Waters  Collection  in  the  MM.  (7100,  7101,  7102)  from  Zanzibar,  are 
superficially  similar  to  A.  lendigera.  However,  notwithstanding  the  little  material  present,  it  is 
possible  to  see  that  the  autozooid  groups  lack  any  characteristic  distal  inclination.  In  addition,  the 
linearly  disposed  stolons  seem  to  be  arranged  in  true  rectilinear  fashion  and  lack  any  posterior 
deflection  associated  with  stolons  of  their  length  as  in  A.  lendigera.  Another  specimen  (7104)  from 
Menton  (southern  France)  labelled  'A.  lendigera',  shows  trifurcation  at  four  stolons  in  almost 
direct  succession,  but  conforms  in  most  other  characteristics.  These  stolons  are  all  in  proximity  to 
substratum  attachment  sites  and  are  probably  not  typical  of  the  whole  colony  budding  pattern. 
These  is  not  enough  material  to  be  certain  about  this  or  the  true  identity  of  the  specimen.  The 
locality  is,  however,  within  the  expected  distribution  area  of  A.  lendigera.  Some  of  MacGillivray's 
specimens  (NMV  65387-8)  marked  'British',  are  A.  lendigera.  Additional  material  (NMV 
65383-5)  labelled  'A.  lendigera'  and  from  Western  Port,  Australia,  is  a  different  species.  These 
specimens  bear  little  resemblance  to  the  'British'  material,  and  in  addition,  show  evidence  of 
rhizoids.  Where  the  rhizoids  are  not  obvious,  careful  illumination  is  required  to  observe  the  oval 
window  precursors.  The  specimens  are  probably  early  astogenetic  stages  of  A.  lamourouxi,  but 
there  is  not  enough  material  to  be  certain;  the  characteristic  terminal  processes  are  not  present,  and 
the  identity  is  inferred  from  the  branching  characteristics.  The  'Australian'  specimens  may  be  the 


AMATHIA  315 

material  described  as  A.  lendigera  by  MacGillivray  (1895,  pi.  B,  fig.  1),  although  the  actual 
specimen  figured  does  not  appear  to  have  been  recorded. 

DISTRIBUTION.  A.  lendigera  is  known  from  the  Thames  estuary,  and  along  the  south  and  west  coasts 
of  England.  The  species  also  occurs  off  the  north  coast  of  Africa,  off  Mediterranean  southern 
France,  and  Naples  and  'Rapallo'  in  Italy.  Substrata  recorded  are  rocks  and  the  alga  Halidrys 
siliquosa. 

Amathia  intermedia  sp.  nov. 
(Figs.  6C,  7C) 

?  Serialaria  lendigera:  Johnston,  1838:  fig.  40. 

?  Serialaria  lendigera:  Johnston,  1847:  fig.  68. 

?  Serialaria  lendigera:  Couch,  1844:  pi.  16. 

Valkeria  lendigera  Dalyell,  1847:  249,  pi.  52  (fig.  2). 

?  Part  Amathia  lendigera:  Bobin  &  Prenant,  1956:  fig.  124, 1,  IV. 

Holotype:  BMNH;  1887.5.2.18  part,  Hastings,  England. 

Paratypes:  BMNH;  1842.12.9.14,  Belfast  Bay.  1847.9.24.184,  North'd  (  =  Northumberland?)  Coast. 
1887.5.2. 18  part,  Hastings.  1963.2.10.1,  Scarborough.  1985.3.2.1a,  Ib,  Yarmouth.  1985.3.2.2,  Bournemouth. 
1985.3.2.3,  no  locality. 

ETYMOLOGY.  The  species  at  one  time  seemed  intermediate  in  character  between  A.  lendigera  and 
A .  guernseii. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  primarily  bifurcate,  ranging  from  equally 
dichotomous  to  rectilinear  series  with  side  branches.  There  is  a  disposition  to  the  latter  condition, 
where  at  a  bifurcation,  one  daughter  stolon  usually  remains  in  line  with  the  main  axis  of  the 
maternal  stolon,  while  the  other  daughter  stolon  appears  sequentially  on  alternate  sides.  These 
lateral  daughter  stolons  are  produced  at  the  same  distal  inclination  to  the  maternal  stolon  axis  as 
the  maternal  stolon  autozooids.  Their  lateral  angular  inclination  may  be  from  0-90  deg.  to  the 
orientation  of  the  maternal  autozooids,  but  usually  ranges  from  10-30  deg.  Occasionally  there  is  a 
trifurcation,  in  which,  of  the  three  daughter  stolons  produced,  the  middle  one  lies  in  the  rectilinear 
position.  The  other  two  are  produced  one  on  either  side,  separated  from  the  central  one  by 
approximately  45  deg.  The  autozooids  on  the  maternal  stolon  bisect  this  angle.  Autozooid  groups 
occur  towards  the  distal  end  of  stolons,  but  there  is  usually  a  further  autozooid-free  portion  of 
stolon,  distal  to  the  autozooid  group.  This  is  often  axially  well  divided  into  small  branches,  the 
subdivisions  orientated  in  the  same  direction  as,  and  supporting,  the  daughter  stolons.  There  is 
frequently  a  further  autozooid-free  length  between  the  end  of  the  autozooid  group  and  this  region 
of  division,  approximately  equal  to  the  diameter  of  one  autozooid.  Stolons  are  often  straight, 
showing  little  sign  of  accommodating  the  autozooids  borne.  The  autozooids  tend  to  be  erect,  and 
of  even  height  throughout  the  autozooid  group,  although  autozooid  group  profile  sometimes 
diminishes  at  the  distal  end.  This  is  due  in  part  to  an  increased  inclination  in  the  autozooids,  and  in 
part  due  to  decreasing  autozooid  height.  Autozooids  are  outer-wall  thickened,  but  thinly  so 
overall,  and  pale  yellow  brown  in  colour.  Viewed  anteriorly,  a  proximal-most  autozooid  is  usually 
evident  in  each  autozooid  group.  The  occurrence  of  this  autozooid,  the  pattern  of  autozooid 
displacements,  and  the  succession  states  of  autozooid  displacements  on  the  stolons,  are  identical  to 
that  found  in  A .  lendigera  (see  page  3 1 3).  The  orientation  of  the  autozooid  group  about  the  stolon  is 
not  always  preserved  from  maternal  to  daughter  stolons;  re-orientations  of  up  to  180  deg.  may 
occur.  No  rhizoids  are  produced,  and  the  colony  attains  a  diffuse  cotton-wool  like  appearance.  The 
non-erect  part  of  the  colony  does  not  appear  as  extensive  as  the  erect  part.  Stolonal  kenozooids  in 
the  non-erect  part  of  the  colony:  produce  branches  occasionally;  tend  not  to  bear  autozooids;  are 
not  of  the  same  appearance  as  those  of  the  erect  part,  in  being  elongated,  sometimes  twisted,  and 
generally  following  the  profile  of  the  substratum.  Erect  components  may  be  produced  at  any 
branching  point,  these  assuming  the  normal  erect  growth  pattern.  Attachment  to  the  substratum  is 
effected  through  lateral  palmate  processes,  often  developed  bilaterally  from  the  adnate  stolonal 
kenozooids. 


316  P.  J.  CHIMONIDES 

SI.    1.75-3.25          Z/S.  35-50% 

Sd.  0.80  Zn.  8-29  (appearing  as  4- 14 'pairs') 

Zh.  0.35-0.45 

Zw.  0.10 

REMARKS.  A.  intermedis  resembles  A.  lendigera  and  A.  guernseii,  the  closest  similarity  being  with 
the  former.  A.  intermedis  may  be  distinguished  from  A .  lendigera  in  having  the  following  character- 
istics: trifurcations  in  the  erect  part  of  the  colony;  a  tendency  towards  higher  numbers  of  auto- 
zooids  in  the  autozooid  groups,  and  longer  stolons;  a  staggered  occurrence  of  autozooid  groups 
and  branching  sites;  a  more  open  colony  form,  resulting  from  a  higher  occurrence  of  rectilinear 
succession  in  the  stolons.  A.  intermedis  may  be  distinguished  from  A.  guernseii  primarily  in  the  fact 
that  A.  guernseii  develops  rhizoids. 

As  with  A.  lendigera,  the  displacement  of  the  proximal-most  autozooids  in  maternal  and 
daughter  stolons  may  reflect  the  timing  of  the  production  of  the  daughter  stolons  relative  to  each 
other  (see  page  314). 

BMNH  1842.12.9.14,  1847.9.24.184,  from  Johnston's  collection,  are  A.  intermedis,  but  it  is  not 
known  if  any  of  this  is  his  figured  material  (1838,  fig.  40,  1847,  fig  68). 

DISTRIBUTION.  The  species  is  known  from  the  east  and  south-eastern  coasts  of  England,  and  also 
from  Belfast  Bay.  The  only  substratum  recorded  is  the  alga,  Halidrys  siliquosa. 

Amathia  guernseii  sp.  nov. 
(Fig  2A,  6B,  7B) 

Holotype:  BMNH;  1898.5.7.189,  Saints  Bay,  Guernsey. 

Paratypes:  BMNH;  1912.12.21.682,  Guernsey.  1967.8.10.2,  Scilly  Is.  1984.2.26.31,  Gulland  Rock, 
Padstow,  Cornwall. 

ETYMOLOGY.  The  species  was  first  recognised  in  material  from  Guernsey. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  primarily  bifurcate,  ranging  from  equally 
dichotomous,  to  almost  rectilinear  series  with  side  branches.  The  angle  between  sister  stolons 
remains  approximately  60  deg.  There  is  a  strong  disposition  towards  the  rectilinear  condition 
where  at  a  bifurcation  one  daughter  stolon  tends  to  remain  in  line  with  the  main  axis  of  the 
maternal  stolon;  the  other  daughter  stolon  appears  sequentially  on  alternate  sides,  produced  at 
approximately  the  same  distal  inclination  to  the  maternal  stolon  axis  as  the  maternal  stolon 
autozooids.  The  lateral  angular  inclination  of  this  daughter  stolon  is  about  30  deg.  to  the  orien- 
tation of  the  maternal  autozooid  group.  Autozooid  groups  occur  at  the  extreme  distal  ends  of 
stolons,  frequently  overlying  the  subsequent  branching  point.  Stolons  are  usually  shaped  in 
accommodating  the  autozooids,  being  deflected  posteriorly  at  the  proximal  end  of  the  autozooid 
group.  At  their  distal  ends,  stolons  often  broaden,  as  if  to  subdivide,  providing  bases  for  the 
subsequent  daughter  stolons,  and  usually  curving  anteriorly  around  the  distal  end  of  the  autozooid 
group.  Occasionally  a  trifurcation  occurs,  three  daughter  stolons  being  produced.  The  third  stolon 
arises  from  a  posterior  projection  at  the  broadened  distal  end  of  the  maternal  stolon;  viewed 
anteriorly,  this  region  retains  a  bilateral  symmetry.  At  the  proximal  end  of  the  autozooid  group, 
autozooids  are  inclined  distally  at  about  30  deg.  to  the  stolon  main  axis.  The  autozooid  group 
profile  tends  to  be  level  at  the  proximal  end  of  the  autozooid  group,  decreasing  at  the  distal  end;  this 
is  due  in  part  to  increasing  inclination  of  the  autozooids,  and  in  part  to  diminishing  autozooid 
height.  The  profile  of  the  rims  of  the  autozooids  usually  reflects  the  angle  of  inclination  in  having  a 
stepped  appearance.  Viewed  anteriorly,  a  proximal-most  autozooid  is  usually  evident  in  each 
autozooid  group.  The  occurrence  of  this  autozooid,  the  pattern  of  autozooid  displacements,  and 
the  succession  states  of  autozooid  displacements  on  the  stolons,  is  identical  to  that  found  in  A. 
lendigera  (see  page  313).  The  orientation  of  autozooids  about  the  stolonal  axis  is  not  rigidly 
preserved  from  stolon  to  stolon,  with  variations  up  to  90  deg.  being  possible.  Over  an  area,  the  sum 
total  of  such  variations  is  to  an  extent  self  cancelling,  so  that  autozooids,  overall,  face  in  approxi- 
mately the  same  direction  i.e.  in  towards  a  central  axis,  and  thus  a  relatively  sheltered  colony- 
bounded  space  (see  page  341).  Rhizoids  are  produced  from  the  anterior  face  of  stolons,  just 


AMATHIA  317 

proximal  to  the  autozooid  groups.  These  arise  singly,  or  as  a  pair,  one  on  either  side  of  the  stolon,  at 
about  30  deg.  to  the  orientation  of  the  autozooids. 

SI.     1.75-2.75  Z/S.50% 

Sd.   0.80  Zn.  8-23  (appearing  as  4- 11 'pairs') 

Zh.  0.38-0.50 

Zw.  0.10 

REMARKS.  There  is  much  overlap  in  the  characteristics  of  A.  lendigera,  A.  intermedis  and  A. 
guernseii,  and  it  can  be  very  difficult  to  distinguish  among  them  unless  there  is  an  adequate  amount 
of  material.  A.guernseiimay  be  distinguished  on  the  following  basis:  the  autozooids  of  A.  guernseii 
have  a  pronounced  distal  inclination  through  the  entire  autozooid  group,  whereas  they  tend  to 
remain  erect  in  A.  intermedis;  in  A.  lendigera,  the  condition  of  the  autozooids  is  intermediate. 
A.  guernseii  is  the  only  species  of  the  three  to  produce  rhizoids.  This  in  turn  affects  the  overall  form 
of  the  colonies;  A.  lendigera  and  A,  intermedis  being  diffuse,  (the  latter  also  tending  to  be  less 
compact),  whereas  A.  guernseii,  with  its  aggregating  rhizoid  system,  has  a  more  organised  and 
directional  appearance.  These  differences  would  appear  to  be  independent  of  the  type  of 
substratum.  The  description  of  A.  lendigera  given  by  Prenant  and  Bobin  (1956)  probably  includes 
A.  guernseii,  as  they  mention  the  presence  of  rhizoids.  In  all  three  species,  some  twist  of  the  stolons 
can  occur  and  this  is  reflected  in  the  autozooids,  but  it  is  never  consistent  throughout  the  colony,  as 
in  A.pruvoti  (see  pages  336,  337). 

As  with  A.  lendigera,  the  displacement  of  the  proximal-most  autozooids  in  maternal  and 
daughter  stolons  may  reflect  the  timing  of  the  production  of  the  daughter  stolons  relative  to  each 
other  (see  page  314). 

The  holotype  is  an  alcohol-preserved  specimen,  originally  a  single  colony,  now  divided  into  two 
fragments.  The  substratum  is  not  present  in  any  of  the  specimens  examined. 

DISTRIBUTION.  The  species  is  known  only  from  the  localities  of  the  type  material. 

Amathia  populea  Busk  MS  in  d'Hondt,  1983 
(Figs  2B,  6D,  7D) 

Amathia  lendigera:  O'Donoghue  &  de  Watteville,  1944: 430. 

Part  Amathia  populea  Busk  MS  in  d'Hondt,  1983:  97,  pi.  3  (4). 

Not  part  Amathia  populea  Busk  MS  in  d'Hondt,  1983:  65,  (  —  A.  woodsii). 

MATERIAL  EXAMINED 

Lectotype  (selected  here):  BMNH;  1899.7.1.526,  Natal,  S.A.,  Busk  Collection. 

Paralectotypes:  BMNH;  1822.8.22.1,  Port  Alfred,  Pondoland,  S.  Africa.  1851.3.12.36,  Port  Natal,  S. 
Africa  1899.7.1.1 12  C,  513,  540,  Algoa  Bay,  S.  Africa. 

OTHER  MATERIAL 

BMNH;  1886.7.2.9,  1985.3.4.1,  Algoa  Bay,  S.  Africa.  1942.8.6.15,  Isipingo  Beach,  Durban,  S.  Africa. 
1963.2.14.7,  Cape  of  Good  Hope. 

MM;  7061/2,  Grahamstown,  S.  Africa.  7062/2,  S.  Africa.  7076/2,  no  locality.  7077/2,  Cape  Agulhas,  S. 
Africa. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  bifurcate;  rarely,  a  trifurcation  occurs.  At 
a  bifurcation,  the  two  daughter  stolons  are  produced  laterally  to  anterolaterally,  at  approximately 
30  deg.  and  60  deg.  to  the  maternal  stolon  axis,  respectively.  The  two  angular  displacements  may 
vary,  but  occur  on  alternate  sides  at  successive  bifurcations.  Development  in  parts  of  the  colony 
may  be  directionally  biased  giving  rise  to  plumes  of  stolons.  Plumes  may  be  up  to  7  cm.  in  length, 
with  those  stolons  forming  the  central  axis  appearing  sympodially  arranged.  This  axis  is  in  fact  a 
simple  linear  series  of  stolons  with  lateral  branches  occurring  on  alternate  sides.  Side  branches 
within  a  plume  are  usually  limited  to  4  or  5  stolons  in  sequence.  As  a  result  of  daughter  components 
frequently  being  produced  in  a  slightly  anterior  direction,  plumes  are  arc-shaped  to  cylindrical  in 
cross-section.  All  sequences  end  with  the  production  of  paired  lanceolate  processes,  each  process 
made  up  of  2-3  sequential,  progressively  tapering  kenozooids.  Sometimes,  the  production  of  a 
stolon  in  a  side  branch  is  replaced  by  that  of  a  lanceolate  process.  Autozooid  groups  reach  to  the 


318  P.  J.  CHIMONIDES 

distal  ends  of  stolons,  frequently  overlying  the  subsequent  branching  point.  Stolons  are  usually 
shaped  in  accommodating  the  autozooids,  appearing  raised  at  the  proximal  end  of  the  autozooid 
group,  becoming  shallower  distally  and  usually  curving  anteriorly  to  the  region  of  bifurcation. 
Autozooids  are  outer-wall  thickened,  but  they  often  appear  cylindrical.  Autozooid  group  profile 
diminishes  distally,  in  part  due  to  stolon  shape,  in  part  due  to  decreasing  autozooid  height. 
Autozooids  incline  distally  at  about  30  deg.  this  being  displayed  at  the  autozooid  rims,  the  rims 
usually  having  a  stepped  appearance.  Viewed  anteriorly,  a  proximal-most  autozooid  is  usually 
evident  in  each  autozooid  group.  The  occurrence  of  this  autozooid,  the  pattern  of  autozooid 
displacements,  and  the  succession  states  of  autozooid  displacements  on  the  stolons,  are  identical  to 
those  found  in  A.  lendigera  (see  page  313).  As  one  daughter  stolon  tends  to  remain  in  line  with  its 
maternal  stolon,  the  alternating  sequence  of  autozooid  displacements  on  linear  series  of  stolons  is 
more  prominent.  The  orientation  of  autozooid  groups  is  generally  well  preserved  from  stolon  to 
stolon.  Rhizoids  may  be  produced  at  the  proximal  end  of  stolons,  most  frequently  from  those  in  the 
central  axis  regions  of  plumes.  Where  rhizoids  are  produced,  it  is  as  one  per  stolon,  each  arising 
usually  from  the  outer  faces  of  bifurcations,  orientated  at  between  90-180  deg.  to  the  autozooids 
on  the  same  stolon.  The  resulting  colony  form  is  usually  arborescent.  Secondary  development  may 
occur  in  the  erect  part  of  the  colony  where  stolons  in  the  common  bases  of  plumes  resume  normal 
budding  of  daughter  stolons.  The  angular  displacements  described  above  are  retained,  but  without 
maintaining  the  autozooid  orientations  about  the  stolons,  or  the  directional  organisation  evident 
elsewhere  in  the  colony.  The  ensuing  compact,  cotton-wool  like,  mass  may  engulf  the  plume  and 
trunk  regions. 

SI.     1.00-1.40  Z/S.  30-55% 

Sd.  0.13-0.18  Zn.  6-13  (appearing  as  3-6 'pairs') 

Zh.  0.38  Tpl.  1.75  (2.60  max.) 

Zw.  0.10-0.13 

REMARKS.  An  association  with  a  sandy  environment  is  inferred  from  the  sand  grains  sometimes 
found  accreted  to  rhizoids  and  attached  epizoic  worm  tubes.  In  plume  portions  of  the  colony,  the 
preserved  orientation  of  the  autozooid  groups,  and  the  cross-sectional  profile  of  the  regions,  results 
in  autozooids  facing  into  a  relatively  sheltered  colony-bounded  space  (see  page  341). 

The  plume  portions  of  A.populea  strongly  resemble  the  figures  of  A.  lemaniiin  the  unpublished 
plates  of  Lesueur.  However,  it  is  equally  possible  to  draw  a  similarity  between  these  figures  and  A. 
woodsii  (see  page  324)  or  possibly  portions  of  A.  tricornis. 

Understandably,  A.populea  has,  in  the  past,  been  confused  with  A.  woodsii  and  A.  tricornis  (e.g. 
d'Hondt,  1979,  1983).  It  has  also  been  confused  with  A.  lendigera  (e.g.  O'Donoghue  and  de 
Watteville,  1944,  BMNH  1942.8.6.15).  A.  populea  may  be  distinguished  from  A.  lendigera  (and 
similar  forms  A.  guernseii  and  A.  intermedis)  primarily  through  the  occurrence  and  location  of 
rhizoids.  These  do  not  occur  in  A.  lendigera  or  A.  intermedis.  In  A.  guernseii,  the  rhizoids  are 
produced  anteriorly,  just  proximal  to  the  autozooid  group;  whereas  in  A.  populea  they  are  pro- 
duced latero-posteriorly  and  proximally  distant  from  the  autozooid  group.  A.  tricornis  and  A. 
populea  differ  in  many  characteristics  (see  page  321). 

D'Hondt  (1979)  placed  A.populea  Busk  MS  (part,  without  qualification)  into  synonymy  with  A. 
cornuta  (sensu  d'Hondt,  1979)  along  with  a  number  of  other  species,  including  A.  australis. 
D'Hondt  (1983)  then  drew  some  distinction,  first  indicating  (p. 65)  that  A.  populea  Busk  MS  part 
from  Australia  is  synonymous  with  A.  cornuta  (sensu  d'Hondt,  1983  i.e.  A.  woodsii  see  pages  320, 
323  etseq.}.  Later,  d'Hondt  (1983:  p.97)  also  gave  a  brief  description  and  a  figure  (p.  103)  of  a  South 
African  specimen,  BMNH  1899.7.1.526  of  A.  populea  Busk  MS  part,  so  validating  Busk's  manu- 
script name,  and  making  the  name  A.populea  available  for  this  species.  D'Hondt  referred  to  the 
specimen  as  'A.sp.'  yet  appears  to  have  remained  equivocal  by  suggesting  that  this  is  also  possibly 
'a  form  of  A.  cornutaT  (sensu  d'Hondt,  1983)  i.e.  A.  woodsii  (see  pages  320,  323  et  seq.). 

A.  populea  and  A.  woodsii  may  be  distinguished  in  the  following:  the  form  of  the  lanceolate 
processes,  being  simple  in  A.  populea,  often  branched  in  A.  woodsii;  the  autozooid  to  stolon  ratio, 
being  higher  in  A.  woodsii;  the  orientation  of  the  rhizoid  origins,  being  latero-posterior  in  A. 
populea  and  anterior  in  A.  woodsii. 


AMATHIA  319 

Small  quantities  of  material  may  be  very  difficult  to  distinguish  and  identify  with  certainty,  such 
that  even  Busk  made  errors.  Some  of  Busk's  A.  populea,  BMNH  1899.7.1.528  from  Algoa  Bay, 
South  Africa  and  BMNH  1899.7.1.4383  from  Australia,  is  in  fact  A.  woodsii.  BMNH 
1899.7.1.4383  is  the  only  specimen  in  the  BMNH  collections  from  Australia  labelled  A.  populea, 
and  so  is  undoubtedly  the  material  that  d'Hondt  (1983)  refers  to  under  the  name  "A.  populea  Busk, 
unpublished  (pars:  Australia)'. 

All  specimens  labelled  by  Busk  as  A.  populea  and  considered  by  d'Hondt  (1983)  are  certain 
syntype  material.  D'Hondt's  figured  specimen,  (BMNH  1899.7.1.526)  is  here  chosen  as  lectotype, 
the  remaining  Busk  material,  except  for  the  two  misidentifications  indicated  above,  has 
paralectotype  status. 

DISTRIBUTION.  The  species  is  known  from  the  south-eastern  coast  of  South  Africa,  possibly  also 
occurring  off  southern  Australia. 

A  mat  Ida  woodsii  Goldstein,  1879 
(Figs  2C,  9B,  D) 

Amathia  woodsii  Goldstein,  1879:  20,  pi.  3  (fig.  5). 
Amathia  australis:  MacGillivray,  1889:  310,  pi.  185  (figs  5,  5a). 
Amathia  woodsii:  MacGillivray,  1895:  138,  pi.  B  (figs  5,  5a). 
Part  Amathia  cornuta:  d'Hondt,  1983:  65,  fig.  36  (C). 

MATERIAL  EXAMINED 
Neotype  (selected  here):  BMNH;  1883.1 1.29.27,  Port  Jackson. 

OTHER  MATERIAL 

BMNH;  1861.9.20.17,  Fremantle.  1897.5.1.1189,  no  locality.  1897.5.1.1196,  Port  Phillip  Heads. 
1899.7.1.528,  1985.3.6.1,  Algoa  Bay,  S.  Africa.  1899.7.1.4383,  Australia.  1909.8.4.10,  Western  Port, 
Australia.  1963.3.28.4,  Adelaide. 

MM;  7075/2,  Queensland. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  bifurcate.  Stolons  are  arranged  to  form 
rectilinear  series  with  side  branch  stolons.  Side  branch  stolons  are  produced  on  alternate  sides  at 
each  subsequent  bifurcation,  arising  with  the  same  distal  inclination  as  the  autozooids  of  their 
maternal  stolons.  The  lateral  angular  displacement  of  the  side  branch  stolons  can  be  0-90  deg.  to 
the  autozooid  orientation,  but  usually  ranges  from  10-30  deg.  In  parts,  growth  appears  favoured 
along  the  rectilinear  series,  with  side  branches  usually  restricted  to  4-5  stolon  units  either  side. 
These  parts  of  the  colony  have  a  plume  like  appearance.  Branches  end  with  a  pair  of  lanceolate 
terminal  processes,  usually  produced  in  the  same  orientations  as  stolons.  The  processes  are  made 
up  of  3-4  sequential,  progressively  tapering  kenozooids,  often  bifurcating  at  the  distal  end  of  the 
basal  segment  kenozooid.  The  lanceolate  processes  in  which  bifurcation  occurs  are  most  usually 
produced  in  the  non-rectilinear  position.  Frequently,  the  production  of  a  side  branch  is  replaced  by 
the  production  of  a  lanceolate  process,  emphasising  the  appearance  of  directional  growth.  Auto- 
zooid groups  occur  towards  the  distal  ends  of  stolons,  but  often  there  is  further  autozooid-free 
part,  coinciding  with  the  production  of  a  side  branch  component.  Stolons  may  show  a  gentle 
anterior  curvature,  and  sometimes  curve  around  the  distal  autozooids  of  a  group.  Autozooid 
group  profile  diminishes  distally,  mainly  due  to  decreasing  autozooid  height,  but  sometimes  due  in 
part  to  an  increase  in  their  distal  inclination.  Autozooids  are  outer-wall  thickened,  the  walls 
appearing  cylindrical.  Viewed  anteriorly,  a  proximal-most  autozooid  is  usually  evident  in  each 
autozooid  group.  The  occurrence  of  this  autozooid,  the  pattern  of  autozooid  displacements,  and 
the  succession  states  of  autozooid  displacements  on  the  stolons,  are  identical  to  those  found  in  A, 
populea  (see  page  318).  The  orientation  of  autozooid  groups  is  generally  well  preserved  from  stolon 
to  stolon.  Along  a  plume  therefore,  autozooids  on  the  rectilinear  sequence  all  face  in  the  same 
direction,  with  lateral  stolon  autozooids  generally  facing  across  these.  Rhizoids  may  be  produced, 
one  per  stolon,  arising  near  to  and  at  about  the  same  orientation  as  the  autozooids.  Colony 
arrangement  is  similar  to  A.  populea. 


320  P.  J.  CHIMONIDES 

Zh.  0.65  Z/S.  50-70% 

Zw.  0.10  Zn.  8-23  (appearing  as  4- 11 'pairs') 

SI.    2.58  Tpl.  2.50  (4.80  max.) 

Sd.  0.20 

REMARKS.  Goldstein's  account  and  figure  are  a  very  good  representation  of  the  species;  the  only 
omission  is  information  on  rhizoid  production.  D'Hondt  (1983)  placed  the  species  in  synonymy 
with  A .  cornuta  Lamarck  (1816),  but  there  is  some  doubt  as  to  the  identity  proposed  for  Lamarck's 
specimen  by  d'Hondt  (1983),  and  the  distinction  between  A.  cornuta  Lamarck  and  A.  woodsii  is 
here  maintained  (see  page  323  et  seq.}. 

With  limited  material,  confusion  could  arise  between  A.  woodsii  and  A.populea  or  A.  tricornis. 
D'Hondt  (1983)  has  referred  specimens  of  these  last  two  species  to  A.  cornuta  Lamarck  (sensu 
d'Hondt  1983),  i.e.  A.  woodsii.  The  species  may  be  distinguished  in  the  following:  the  presence  of 
the  characteristic  subdivided  lanceolate  process  in  A.  woodsii,  this  being  simple  in  the  other  two;  the 
production  of  rhizoids  being  near  to,  and  in  the  same  orientation  as  the  autozooid  group  in  A. 
woodsii,  these  being  distant,  and  of  different  orientation  in  the  other  two;  the  budding  pattern  in  A. 
woodsii  is  never  as  complex  as  in  A.  tricornis,  and  the  predisposition  to  rectilinear  development  is 
more  prominent  than  in  A.populea,  in  which  there  is  a  tendency  for  a  sympodial  appearance. 

According  to  Stach  (1936)  specimens  from  Goldstein's  collection  were  deposited  in  the  NMV. 
However,  his  material  for  A.  woodsii  is  not  there  (NMV  in  litt.  6.12.1984).  In  view  of  the  confusion 
which  has  arisen,  there  is  a  need  for  type  material.  The  description  and  measurements  given  here 
are  based  on  BMNH  1883. 1 1 .29.27  from  Port  Jackson,  an  alcohol  specimen,  rehydrated  from  the 
dry  state.  The  specimen  is  here  selected  as  neotype.  Goldstein  does  not  give  a  locality  for  his 
specimen,  only  mentioning  that  the  species  was  found  on  a  previous  occasion  at  Portland, 
presumably  Victoria  State. 

DISTRIBUTION.  The  species  is  known  from  the  south-eastern  coast  of  South  Africa,  and  from 
Australia,  with  records  from  Fremantle,  Adelaide,  Port  Phillip  Heads,  Port  Jackson  and 
'Queensland'. 


Amathia  tricornis  Busk  MS 
(Figs2D,  12C) 

Holotype:  BMNH;  1899.7.1.6600,  Australia,  Busk  Collection. 
Paratypes:  BMNH  1899.7.1.4393, 4394,  Australia,  Busk  Collection. 

ETYMOLOGY.  Busk's  MS  name,  probably  indicating  the  occurrence  of  three  terminal  lanceolate 
processes. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  mainly  trifurcate,  although  tetrafur- 
cation  occurs  in  certain  regions.  The  latter  condition  is  associated  with  astogenetically  early  parts 
of  the  colony,  which  form  the  base  and  central  main-axis  regions.  These  regions  tend  to  be 
composed  of  lengthy  series  of,  what  are  here  termed,  type  'a'  stolons.  Type  'a'  stolons  are  longer 
than  other  type  'b'  stolons  found  in  the  trifurcate  portions  of  the  colony,  and  often  bear  rhizoids. 
Where  rhizoids  are  produced,  these  arise  from  the  proximal  end  of  a  stolon,  usually  singly,  at 
between  45-90  deg.  to  the  autozooid  orientation  on  the  same  stolon.  Autozooids  borne  by  type  'a' 
stolons  show  no  difference  in  size  from  those  on  type  'b'  stolons,  although  the  autozooid  groups 
tend  to  be  shorter.  The  linear  proportion  of  stolon  occupied  by  autozooids,  therefore,  is  lower.  The 
stolon  budding  arrangement,  in  both  tri-  and  tetrafurcate  conditions,  always  results  in  one 
daughter  stolon  lying  in  rectilinear  succession  to  the  maternal  stolon.  Two  other  daughter  stolons 
are  produced  laterally,  one  on  each  side,  at  about  60  deg.  to  this  central  axis.  All  three  of  these 
daughter  stolons  bear  autozooids,  usually  orientated  in  the  same  direction  as  those  on  the  maternal 
stolon,  with  some  exceptions.  In  the  tetrafurcate  condition,  a  fourth  daughter  stolon  is  produced, 
also  at  about  60  deg.  to  the  central  axis,  but  posteriorly  to  the  maternal  stolon.  The  autozooid 
group  orientation  of  the  maternal  stolon  is  preserved  in  this  daughter  component;  the  autozooid 


AMATHIA  321 

group  thus  faces  distally  along  the  rectilinear  series  of  the  central  axis  (see  Fig.  2D).  In  the 
tetrafurcate  condition  alone,  the  orientation  of  the  autozooid  groups  along  the  rectilinear  series  is 
not  always  maintained.  A  repeat  rotation  of  90  deg.  may,  instead,  be  observed  at  each  axial 
junction.  The  relationship  of  sister  daughter  stolons  to  the  axial  daughter  stolon  remains  fixed, 
and,  thus,  the  entire  assemblage  follows  the  re-orientation.  The  original  orientations  are  recovered 
every  fourth  axial  stolon  unit  along  the  sequence.  In  the  trifurcate  condition,  lateral  growth 
appears  to  be  limited  to  one  or  two  stolons  in  sequence  each  side.  Development  in  these  parts  of  the 
colony  is  therefore  directionally  biased  and  these  parts  have  a  plume  like  appearance.  Branches  end 
with  the  production  of  three  lanceolate  terminal  processes,  each  made  up  of  two  or  three  sequen- 
tial, progressively  tapering  kenozooids.  These  arise  from,  and  lie  approximately  in  line  with,  their 
maternal  stolons.  Sometimes,  the  central  terminal  process  does  not  form,  being  replaced  by  a 
stolon  instead.  This  may  be  repeated  so  that  occasionally,  lateral  branches  may  be  several  stolons 
in  length.  On  both  type  'a'  and  type  'b'  stolons,  autozooid  groups  occur  at  the  extreme  distal  end  of 
stolons,  frequently  overlying  the  subsequent  branching  point.  Autozooid  group  profile  diminishes 
distally  in  all  cases,  mainly  due  to  increasing  distal  inclination  of  the  autozooids.  In  all  parts  of  the 
colony,  autozooids  are  outer-wall  thickened.  Along  rectilinear  sequences  of  stolons,  there  is  a 
predictable  repetition  in  the  arrangement  of  the  autozooids  borne.  The  sequence,  progressing 
distally,  is  as  follows:  if,  in  an  autozooid  group,  there  is  one  proximal-most  autozooid  prominent, 
this  is  associated  with  one  side  of  the  stolon;  in  the  next  stolon,  no  single  autozooid  is  prominent 
proximally,  the  proximal  autozooids  being  paired  equally;  on  the  third  stolon,  a  proximal-most 
autozooid  is  prominent  once  again,  but  on  the  opposite  side  to  that  of  the  first  stolon;  on  the  fourth 
stolon,  the  proximal  autozooids  are  paired  as  on  the  second  stolon;  the  fifth  stolon  repeats  the 
arrangement  on  the  first  stolon.  On  laterally  branched  stolons,  a  proximal-most  autozooid  is 
prominent,  and  is  associated  with  the  side  nearest  the  rectilinear  stolon  sequence.  Autozooids  on 
stolons  continuing  in  rectilinear  series  which  develop  from  lateral  branch  stolons  subsequently 
follow  the  predictable  pattern  of  repetition  given  above. 

SI.    (a)  2. 10  Z/S.  (a)  30-40% 

Sd.   (a)  0.25  Zn.  (a)  8-1 1  (appearing  as  4-5  'pairs') 

SI.    (b)1.45  Z/S.  (b)  60% 

Sd.   (b)0.25  Zn.  (b)  10-21  (appearing  as  5-9 'pairs') 

Zh.        0.35  (all  autozooids) 

Zw.       0.10  (all  autozooids) 

Tpl.       2.10 

REMARKS.  No  evidence  exists,  in  the  limited  material  available,  that  the  trifurcate  condition  ever 
gives  rise  to  the  tetrafurcate  condition.  The  colony  form  is  inferred  to  be  arborescent,  resulting 
from  the  production  of  rhizoids. 

None  of  the  material  held  at  the  BMNH  named  A.  tricornis  in  MS  by  Busk  is  misidentified. 
D'Hondt  (1979  &  1983)  erroneously  placed  this  species  in  synonymy  in  part,  with  parts  of  A. 
cornuta  (Lamarck)  sensu  d'Hondt  (i.e.  A.  woodsii,  see  page  323),  and  in  part,  initially  with  A. 
pinnata  (1979)  (see  page  330),  and  subsequently  with  A.  inarmata  (1983)  (i.e.  A.  biseriata,  see  page 
332).  D'Hondt  on  each  occasion  mentioned  A.  tricornis  in  synonymy  only,  thus  not  making  the 
name  available  at  any  time  (I.C.Z.N.  article  1  le). 

Among  the  species  which  may  be  confused  with  A.  tricornis  are:  A.  populea;  A.  woodsii;  A. 
pinnata;  A.  biseriata.  In  brief,  A.  tricornis  has  a  more  complex  colony  construction  and  differs  from 
these  species  in  many  features,  for  example:  the  bimorphic  autozooid-bearing  stolons;  the  normal 
occurrence  of  tetra-  and  trifurcation,  including  the  production  of  triplet  lanceolate  processes  and 
their  permutations  with  autozooid-bearing  stolons;  autozooid  and  stolon  re-orientations;  the 
productions  site  of  the  rhizoids.  There  are  also  differences  in  the  autozooid  to  stolon  ratios. 

DISTRIBUTION.  The  species  is  known  only  from  material  described  as  being  from  'Australia',  sent  to 
Busk  by  Miss  Gore. 


322  P.  J.  CHIMONIDES 

Amathia  lamourouxi  nom.  nov.  for 

Amathia  cornuta  auctorem 

(Figs  3A,  8C,  9A,  C) 

?  Not  Serialaria  cornuta  Lamarck,  1816:  131. 
Amathia  cornuta  Lamouroux,  1816:  159,  pi.  4  (fig.  la,  IB). 
?  Not  Serialaria  australis  Tenison  Woods,  1877:  83, 1st  fig. 
?  Not  Amathia  australis:  Tenison  Woods,  1880:  102. 
Amathia  cornuta:  Tenison  Woods,  1880: 99,  fig.  3. 
Not  Amathia  australis:  MacGillivray,  1889:  310,  pi.  185 

(figs  5,  5a),  (  =  A.  woodsii). 

Amathia  cornuta:  MacGillivray,  1895:  137,  pi.  D  (fig.  1,  la). 
?  Amathia  cornuta:  d'Hondt,  1979:  10,  16. 
Part  Amathia  australis:  d'Hondt,  1983:  65,  fig.  36(F). 
Not  Amathia  cornuta:  d'Hondt,  1983:  65,  fig.  36(C)  (  =  A.  woodsii). 

MATERIAL  EXAMINED 
Neotype  (selected  here):  BMNH;  1887.12.10.70,  Port  Phillip,  J.  B.  Wilson  collection. 

OTHER  MATERIAL 

BMNH;  1842. 11. 4.50,  Sydney.  1899.7. 1.3,  New  Zealand.  1899.7.1.4325,  Victoria.  1899.7.1.4327,4329-31, 
4333,  Australia.  1899.7.1.4328,  Bass  Strait.  1899.7.1.4334,  Australia  &  New  Zealand.  1985.3.10.1,  Flinders 
Is.,  Bass  Strait.  1985.3.24.1,  mid  channel,  Port  Phillip  Heads,  15m. 

MM;  7078/2,  Australia. 

LBIMM;  bry  2821  part,  Australie  Occidentale/Nouvelle  Hollande  (see  below). 

ETYMOLOGY.  Lamouroux's  name  is  used  for  his  species  of  A .  cornuta  ( 1 8 1 6),  a  name  preoccupied  by 
A.  cornuta  (Lamarck,  1816). 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  bifurcate.  At  any  branching  point, 
daughter  stolons  may  be  produced  in  positions  ranging  from  almost  rectilinear,  to  90  deg.  to  the 
maternal  stolon.  A  minimum  separation  of  90  deg.  occurs  between  daughter  stolons.  These  usually 
arise  from  the  posterior  side  of  their  maternal  stolon.  Stolons  are  narrowed  proximally  and  usually 
curved  anteriorly,  being  reminiscent  of  a  short,  simple,  cow  horn.  Autozooid  groups  occupy  the 
greater  part  of  stolons,  and  frequently  overlie  the  subsequent  branching  point.  Autozooid  group 
profile  usually  increases  proximodistally  within  each  group,  or  may  remain  level.  Autozooids  are 
outer-wall  thickened.  Viewed  anteriorly,  a  single  proximal-most  autozooid  is  evident  in  each 
autozooid  group.  This  autozooid  is  usually  placed  just  off  the  stolon  mid-line,  thus  associated  with 
either  one  or  other  side  of  the  autozoid  group.  No  pattern  is  evident  from  group  to  group,  in  the 
location  of  this  autozooid.  At  the  distal  end  of  each  autozooid  group  and  contiguous  with  the 
autozooids,  are  produced  a  pair  of  tapering,  single-kenozooid  lanceolate  processes.  At  the  distal 
extremities  of  the  colony,  both  daughter  stolons  tend  to  be  produced  at  90  deg.  to  their  maternal 
stolons,  i.e.  separated  from  each  other  in  equal  dichotomy  by  180  deg.  In  less  distal  parts  of  the 
colony,  daughter  stolons  may  be  separated  from  each  other  in  equal  dichotomy  by  lesser  angles. 
Daughter  stolons  produced  in  the  linear  position  are  less  common  and  are  associated  with  more 
central  and  proximal  (astogenetically  earlier)  regions  of  the  colony.  Autozooid  orientation  is 
generally  not  preserved  from  stolon  to  stolon.  Often  there  is  an  equal  rotation  by  up  to  90  deg.  of 
each  daughter  stolon  in  opposite  directions  i.e.  were  both  daughter  stolons  to  lie  in  the  linear 
position,  their  anterior  faces  would  be  away  from  each  other.  Successive  daughter  stolons  actually 
lying  in  the  linear  position  and  forming  a  sequence,  are  all  produced  from  the  same  side,  i.e.  in  such 
a  sequence,  viewed  anteriorly  each  time,  they  would  all  have  been  budded  from  e.g.  the  left  side. 
The  orientation  of  the  autozooid  groups,  in  such  a  linear  sequence,  is  rotated  by  90  deg.  in  the  same 
direction,  with  each  successive  stolon.  The  original  orientation  is  recovered  every  fourth  stolon 
unit.  Superficially,  branching  can  appear  as  'alternate'  along  a  linear  sequence.  Rhizoids  may  be 
produced,  usually  from  stolons  along  these  linear  sequences.  Rhizoids  are  produced  one  per 
stolon,  arising  from  a  position  level  with,  and  at  90  deg.  to  the  proximal-most  autozoid,  and  on  the 
same  side  of  the  stolon  as  the  direction  in  which  the  stolon  was  budded. 


AMATHIA  323 

Zh.  0.40-0.50          Z/S.75% 

Zw.  0. 1 3  Zn.   1 1-1 5  (appearing  as  5-7  'pairs') 

SI.     1.13  Tpl.  2.00-2.25 

Sw.  0.25  (at  the  widest  region) 

REMARKS.  No  pattern  has  been  discerned  in  the  autozooid  arrangement  from  stolon  to  stolon, 
other  than,  if  the  proximal-most  autozooid  of  a  group  is  associated  with  one  side  of  the  stolon,  then 
that  association  may  remain  in  both  daughter  stolons  over  a  number  of  successive  bifurcations. 

The  identity  of  A.  cornuta  auct.  is  inextricably  associated  with  the  collections  made  by  Peron  and 
Leseur,  between  the  years  1800-1804  (see  page  307),  on  which  both  Lamarck  and  Lamouroux 
worked,  both  of  them  describing  a  'cornuta'. 

Tenison  Woods  (1880)  drew  a  distinction  between  his  A.  australis  and  A.  cornuta  sensu 
Lamouroux  (i.e.  A.  lamourouxi)  based  on  the  understanding  that  Lamouroux's  (1816)  figure 
indicates  a  single  line  of  autozooids.  Lamouroux  himself,  referred  in  the  singular  to  'the  largest  cell 
of  each  group . . .  garnished  with  two  setaceous  appendages'.  The  misinterpretation  of  a  single  row 
for  a  double  row  of  autozooids,  might  be  made  as  a  result  of  a  preservation  artifact  where,  in  dried 
specimens,  the  thinner  central  walls  between  autozooids  collapse  from  view,  leaving  only  the  outer 
walls  visible.  On  this  basis  there  is  no  distinction  between  A.  australis  and  A.  lamourouxi.  The 
additional  difference  claimed  by  Tenison  Woods,  in  the  form  of  the  'setaceous  appendages'  (his 
figure  of  1 877  shows  these  as  being  broad  and  less  trim  than  those  in  Lamouroux's  figure),  might  be 
accounted  for  in  terms  of  the  variation  which  may  occur  within  A.  lamourouxi.  However,  Tenison 
Woods'  figure  (1877)  shows  clearly  that  his  specimen  had  undergone  trifurcation.  Two  possibilities 
may  account  for  this:  the  first,  that  under  certain  conditions,  A.  lamourouxi  can  undergo  such  a 
division;  the  second,  that  Tenison  Woods  did  in  fact  have  a  separate  species.  Although  the  former 
may  be  possible,  trifurcation  has  not  been  recognised  in  specimens  here  assigned  to  A.  lamourouxi. 
The  whereabouts  of  Tenison  Woods'  material  is  not  known. 

MacGillivray  (1889)  considered  A.  australis  to  be  A.  cornuta  sensu  Lamouroux,  but  in  his 
description  (p. 310)  and  figure  (pi.  185,  figs  5,  5a)  gave  an  acocunt  of  A.  woodsii.  MacGillivray 
(1895)  subsequently  recognised  the  error,  and  correctly  referred  to  his  account  of  1889  as  being 
descriptive  of  A.  woodsii.  At  the  same  time,  MacGillivray  distinguished  between  A.  woodsii  and  A. 
cornuta,  and  reaffirmed  his  opinion  that  A.  australis  was  synonymous  with  the  latter,  but  gave 
Lamouroux  as  the  author  and  placing  Lamarck  in  synonymy. 

D'Hondt  ( 1 979)  has  found  a  specimen,  LBIMM  bry  282 1 ,  which  is  claimed  to  be  the  holotype  of 
A.  cornuta  (Lamarck).  D'Hondt  (1979)  placed  A.  australis  in  synonymy  under  A.  cornuta 
(Lamarck),  but  without  mention  of  A.  woodsii.  D'Hondt  (1983)  then  placed  A.  woodsii  in 
synonymy  under  A.  cornuta  (Lamarck),  but  excluded  A.  australis,  thus  revoking  his  opinion  of 
1 979  and  indicating  that  A .  australis  is  different  (d'Hondt's  reference  to  'parts'  of  A.  australis  at  this 
point  are  enigmatic).  In  this  reorganisation  of  the  species,  d'Hondt  (1983)  gave  two  figures:  36(C) 
as  A.  cornuta  (Lamarck),  and  36(F)  as  A.  australis.  Figure  36(C)  is  in  fact  A.  woodsii,  and  36(F)  is  A. 
cornuta,  both  of  common  usage,  the  latter  corresponding  with  Lamouroux  (1816),  of  which  A. 
australis  is  usually  taken  to  be  a  junior  synonym. 

D'Hondt  (1983)  appears  to  have  determined  A.  cornuta  (Lamarck)  to  be  different  from  A. 
cornuta  Lamouroux.  A.  cornuta  (Lamarck)  predates  A.  cornuta  Lamouroux  (Tenison  Woods, 
1880,  d'Hondt,  1983).  D'Hondt  (1983)  thus  relegated  A.  woodsii  as  a  junior  synonym  of  A.  cornuta 
(Lamarck),  and  assigned  the  name  A.  australis,  as  the  next  available  name,  for  what  was  previously 
accepted  as  A.  cornuta  sensu  Lamouroux. 

Unless  it  was  the  only  specimen  involved,  LBIMM  bry  2821  can  only  be  taken  as  the  holotype  if 
so  designated  at  the  time  of  introduction  by  the  original  author.  Lamarck  (1816)  did  not  do  this, 
and  the  number  of  specimens  involved  is  not  certain. 

LBIMM  bry  2821  is  recorded  as  being  one  of  three  specimens  of  A.  cornuta  so  identified  by 
Lamarck  and  in  the  Paris  Museum  at  the  time  of  the  compilation  of  the  first  catalogue  of  Bryozoa 
in  1 867.  The  other  two  specimens  appear  to  have  been  A .  cornuta  sensu  Lamouroux,  only  'possibly' 
originating  from  Peron  and  Lesueur.  Their  locality  is  given  as  'Australasie'.  The  specimens  were 


324  P.  J.  CHIMONIDES 

numbered:  172a,b,c.  (photocopy  of  the  1867  catalogue).  LBIMM  bry  2821  is  the  only  one  of  the 
three  which  is  known  to  have  come  from  Peron  and  Lesueur  (d'Hondt  in  litt.  10.12.1984).  The 
locality  for  this  specimen  is  'Australie  Occidental'  (d'Hondt,  1979),  and  also  as  'Nouvelle- 
Holland'  (loan  form  24th  Jan.  1985). 

D'Hondt  (1979)  reported  that  LBIMM  bry  2821  carries  the  label  'Amathia  lemanii  Lesueur'. 
This  would  appear  to  be  in  the  hand  of  Pergens,  the  original  label  having  been  lost  or  destroyed. 
The  specimen  is  taken  to  be  the  same  one  that  Pergens  (1887)  correlated  with  a  figure  in  the 
unpublished  plates  of  Lesueur,  these  in  turn  related  to  a  manuscript  of  Desmarest  and  Lesueur, 
deposited  at  Paris  in  1 829  (with  another  slightly  different  version  at  le  Havre).  Pergens  ( 1 887,  p.  88) 
ascertained  that  plate  13,  figure  6,  in  the  unpublished  plates,  is  Amathia  lemanii,  and  (p.  90)  then 
gave  the  identification  he  was  able  to  make  of  the  species  in  terms  of  what  was,  to  him,  a  valid  and 
available  name  i.e.  A.  cornuta  (Lamarck).  Copies  of  the  plates  of  Lesueur  exist  at  the  BMNH. 
Plate  13,  figure  6,  consists  of  three  representations  of  the  species  intended,  all  three  at  different 
magnifications.  The  species  represented  could  be  A.  woodsii  or  A.  populea  (see  page  318);  both 
species  are  capable  of  assuming  the  characteristics  portrayed.  Missing  from  the  figure(s)  is  any 
information  on  rhizoids  and  on  any  occurrence  of  the  characteristic  subdivided  terminal  filaments, 
which  might  serve  to  distinguish  between  the  two  species.  Only  the  actual  specimen  used  by 
Lesueur  will  determine  the  true  species  (taken  to  be  LBIMM  bry  2821).  The  identity  of  this  species 
is  of  little  taxonomic  consequence  however,  as  neither  plates  nor  descriptions  have  ever  been 
published.  LBIMM  bry  2821  could  have  been  the  holotype  perhaps,  of  Desmarest  and  Lesueur's 
species,  but  there  is  insufficient  evidence  published  to  suggest  that  it  was  that  of  Larmarck's. 

In  addition,  Larmarck  (1816)  gave  no  figure,  specimen  number,  or  dimensions,  with  which  a 
specimen  may  be  correlated.  Furthermore,  the  locality  information  (see  above)  for  the  specimen, 
although  close,  does  not  match  exactly  with  that  of  Lamarck  (or  of  Lamouroux).  Lamarck  gives 
TOcean  asiatique'  (Lamouroux  gives  'Sur  les  Fucus  de  1'Australasie').  That  the  specimen  was  part 
of  Peron  and  Lesueur's  collections,  may  not  in  this  case  be  sufficient;  Lamarck  himself  is  not 
definite  as  to  the  origins  of  his  specimen,  only  'believing'  it  to  be  from  Peron  and  Lesueur.  Pergens 
(1887)  merely  expressed  his  opinion  that  LBIMM  bry  2821  is  the  same  as  A.  cornuta  (Lamarck); 
how  he  reached  that  conclusion  is  not  clear.  The  specimen  appears  to  be  only  one  remaining  of  a 
number,  others  having  gone  astray  since  the  days  of  Peron  and  Lesueur;  the  1867  catalogue  of  the 
Bryozoa  was  compiled  some  51  years  after  Lamarck's  publication. 

It  is  possible  that  LBIMM  bry  2821  may  be  eligible  for  selection  as  lectotype  of  A.  cornuta 
(Lamarck),  if  it  can  be  shown  to  have  been  part  of  Lamarck's  original  syntype  series,  and  formative 
of  his  opinion.  However,  Larmarck  did  also  identify  two  different  specimens  as  being  his  species, 
these  being  A .  cornuta  sensu  Lamouroux  (see  above) .  These  specimens  might  also  have  been  eligible 
for  selection,  but  are  no  longer  to  be  found  in  the  Paris  Museum  (d'Hondt  in  litt.  10.12.1984, 
24.01.1985). 

Further  challenge  to  the  identity  proposed  by  d'Hondt  for  A.  cornuta  lies  in  the  evidence 
that  preceeds  Pergen's  opinion  (1887).  Tenison  Woods  (1880)  gave  information  on  the  working 
relationship  between  Lamarck  and  Lamouroux  concerning  the  Amathia  specimens  collected  by 
Peron  and  Lesueur.  Much  of  the  information  appears  to  be  derived  from  Lamouroux's  (1816)  own 
preface  and  introduction.  Lamouroux  had  'the  fullest  access'  to  Lamarck's  collection,  and  named 
at  least  part  of  this,  if  not  all  of  it. 

Neither  Lamarck's  (1816)  nor  Lamouroux's  (1816)  account  of  a  'cornuta'  contradicts  the  other. 
However,  whereas  Lamouroux's  account  is  quite  explicit,  and  furnished  with  figures,  such  that  the 
species  he  described  may  still  be  recognised;  Lamarck's  account  is  open  to  interpretation.  The 
descriptions  may  be  translated  as  follows: 

Lamouroux,  p.  159:  No.  266. 

(from  the  French) — The  largest  cell  of  each  group  having  its  free  border,  garnished  with  two 

setaceous  appendages. 

(from  the  Latin) — two  setaceous  filaments  from  the  first  rank  cell 

Lamarck,  p.  131:  No.  2. 

(from  the  French) — I  believe  it  (to  be)  from  the  voyage  of  Messieurs  le  Sueur  and  Peron.  It  is  a 


AMATHIA  325 

little  more  stout  and  less  capillary  than  the  preceeding  [i.e.  A.  lendigera],  at  the  extremities  curved 
and  as  curls. 

(from  the  Latin) — very  branched,  articulated,  somewhat  curled;  branches  alternate;  curved 
secondary  little  branches;  cells  in  distinct  series;  two  setae  at  the  most  distant  extremity. 

In  Lamarck's  account,  no  orientation  is  given  for  the  setae,  and  the  description  could  apply  to  A. 
cornuta  of  Lamouroux,  A .  woodsii  or  A .  populea.  Whether  the  reference  to  the  secondary  branches 
is  an  indication  of  an  arborescent  growth  form  i.e.  axial  development  with  lateral  branch  system,  or 
a  reference  to  the  stolons  themselves,  is  not  clear.  In  either  case,  the  description  is  insufficiently 
distinctive.  Branching  is  alternate  in  A.  woodsii  and  A.  populea  and  may  appear  so  in  A.  cornuta 
sensu  Lamouroux.  Finally,  Lamarck  makes  no  mention  of  any  subdivided  terminal  processes 
(present  in  LBIMM  bry  2821  part)  to  be  expected  if  his  'cornuta'  was  the  equivalent  of  A.  woodsii. 

Although  it  is  not  possible  to  recognise  a  single  species  from  Lamarck's  description,  the  identity 
of  A.  cornuta  (Lamarck)  has  been  understood  through  the  later  accounts  of  the  two  authors:  in 
Lamouroux  (1824)  and  Lamarck  (1836)  respectively,  each  recognises  the  other's  A.  cornuta  as 
synonymous  with  his  own;  from  this  derives  the  A.  cornuta  of  common  usage.  It  is  this  concordance 
which  d'Hondt  (1983)  has  in  effect  repudiated. 

In  the  strictest  sense,  A.  cornuta  (Lamarck)  should  have  been  classed  as  a  nomen  dubium,  and  not 
used.  This  is  historically  implied  by  Tenison  Woods  (1880)  who  acknowledged  that  Lamarck 
probably  predated  Lamouroux,  and  so  accepted  A.  cornuta  (Lamarck),  'but  with  reference  to 
Lamouroux  only'.  MacGillivray  (1895)  appears  to  have  been  of  the  same  opinion  (see  above). 

In  the  light  of  such  contradictions,  Lamarck's  A.  cornuta  must  be  taken  as  a  nomen  dubium,  and 
the  name  should  no  longer  be  used  for  Lamouroux's  species.  A.  australis  of  Tenison  Woods  would 
be  the  next  valid  name  available,  if  certainty  could  be  attached  to  the  identity  of  his  species  (see 
above).  Under  these  circumstances,  it  is  wiser,  in  the  interest  of  long  term  stability,  to  select  a  new 
name  for  A.  cornuta  Lamouroux,  accepting  either  his  figures  as  lectotype,  or  perhaps  selecting  a 
neotype.  It  is  here  proposed  that  A.  cornuta  auct.  be  known  as  A.  lamourouxi,  with  specimen 
BMNH  1887.12.10.70  as  neotype. 

DISTRIBUTION.  The  species  is  recorded  from  New  Zealand  and  southern  Australia. 

Amathia  plumosa  MacGillivray,  1890 
(Figs3C,  12A,  B) 

Amathia  plumosa  MacGillivray,  1890:  110. 

Amathia  plumosa:  MacGillivray,  1895:  139,  pi.  C  (figs  2,  2a). 

Amathia  plumosa:  d'Hondt,  1983:  67,  fig.  36  (B). 

MATERIAL  EXAMINED 
Holotype:  NMV;  H494,  Port  Phillip  Heads,  J.  B.  Wilson  Collection. 

OTHER  MATERIAL 

BMNH;  1963.2.12.354,  358,  Western  Australia.  1985.3.8.1,  no  locality. 

DESCRIPTION.  The  branching  pattern  on  the  erect  part  of  the  colony  is  based  on  both  bi-  and 
trifurcation.  Trifurcation  is  associated  with  non  autozooid-bearing  stolonal  kenozooids,  forming 
angularly  undulating  'main-stems'.  At  the  distal  end  of  each  of  the  main-stem  constituent  stolonal 
kenozooids,  are  produced:  a  single  continuing  stolonal  kenozooid,  deflected  by  approximately  30 
deg.  towards  the  central  axis  of  the  main-stem;  two  (autozooid-bearing)  side  branch  stolons,  one 
each  side.  The  side  branch  stolons  are  produced  in  the  same  plane  as  their  maternal  stolonal 
kenozooid,  but  diverge  from  each  other  equally,  by  an  approximate  total  angle  of  60  deg.  The 
autozooids  borne  on  these  side  branch  stolons  face  the  main-stem,  and  the  stolons  themselves  are 
curved  anteriorly.  Subsequent  branching  from  these  side  branch  stolons  is  usually  bifurcate, 
although  new  main-stem  sequences  may  be  produced,  showing  the  associated  trifurcation. 
Development  along  side  branches  is  usually  limited;  2-3  stolons  in  a  sequence  is  usual,  but  up  to  8 
stolons  in  succession  may  occur.  The  orientation  of  autozooid  groups  along  any  such  sequence 
remains  the  same.  These  side  branches  end  with  the  production  of  a  pair  of  usually  dichotomously 
branched  lanceolate  processes.  These  are  made  up  of  sequential,  progressively  tapering 


326  P.  J.  CHIMONIDES 

kenozooids.  Where  the  processes  are  branched,  this  occurs  at  the  distal  end  of  the  base  segment 
kenozooid.  This  may  be  repeated  in  one  or  both  of  the  next  resultant  segments.  Rarely,  a  lanceolate 
process  may  trifurcate.  On  occasions,  the  production  of  a  stolon  in  a  side  branch  is  replaced  by  the 
production  of  a  lanceolate  process.  Development  in  such  cases,  therefore,  tends  to  be  directionally 
biased.  The  colony  is  composed  of  such  quasi-cylindrical  assemblages,  circular  in  cross-sectional 
profile,  and  somewhat  reminiscent  of 'feather  boas'.  These  may  be  supported  on  a  trunk-like  part 
of  the  colony,  resulting  from  the  production  of  rhizoids  (see  page  309).  Rhizoids  are  produced  in 
two  ways:  they  may  be  produced  from  the  proximal  end  of  main-stem  kenozooids,  sometimes 
singly,  although  more  often  as  an  adjacent  pair,  in  the  same  orientation  as  the  side  branch  stolons 
lying  immediately  proximal;  they  may  be  produced  from  autozooid-bearing  stolons,  at  approxi- 
mately 120  deg.  to  the  orientation  of  the  autozooid  group  on  the  same  stolons.  Where  autozooid 
groups  develop  in  side  branches,  these  occur  towards  the  distal  ends  of  stolons,  but  often  there  is  a 
further  autozooid-free  portion.  This  portion  is  about  the  same  length  as  the  diameter  of,  and 
coincident  with  the  production  of,  a  daughter  component.  Autozooid  group  profile  tends  to 
remain  level  and  autozooids  are  outer- wall  thickened.  Where  a  proximal-most  autozooid  of  a 
group  is  evident,  its  occurrence,  and  the  pattern  of  autozooid  displacements  from  stolon  to  stolon, 
are  similar  to  those  of  A.  lendigera  (see  page  313)  except  that  the  second  succession  state-does  not 
appear  to  occur. 

kSl.  1-45  (main-stem  kenozooidal  stolons) 

kSd.  0- 1 9-0-29          (main-stem  kenozooidal  stolons) 

SI.    1-03-1-61          Z/S.  55-65% 

Sd.   0-15-0-26          Zn.  7-17    (appearing  as  3-8 'pairs') 

Zh.  0-32 

Zw.  0-10 

REMARKS.  The  species  is  so  distinctive  that  it  does  not  appear  to  have  been  confused  with  any 
other.  The  slide  mounted  specimen  NMV  H494,  is  accepted  here  as  the  holotype  of  A.  plumosa 
MacGillivray  (1890),  and  as  that  figured  by  MacGillivray  (1895).  The  label  on  the  slide  carries  the 
information:  'H494  Amathia plumosa  McG  P.P.H.  fig  J.B.W.'.  This  in  agreement  with  the  original 
description  in  which  the  locality  is  given  as  'Port  Phillip  Heads',  from  the  collection  of  J.  B.  Wilson. 
The  description  given  here  is  based  on  the  above  specimen.  Some  supplementary  information  is 
derived  from  BMNH  1963.2.12.354  and  BMNH  1963.2.12.358,  these  agreeing  well  with  the 
holotype. 

There  is  some  indication  that  the  repeated  branching  in  the  lanceolate  processes  coincides  with 
the  development  of  a  lanceolate  process  in  substitution  for  an  expected  stolon,  although  there  is  no 
certainty  to  this.  Autozooid  groups  are  orientated  about  main-axis  stolons  to  face  into  relatively 
sheltered,  colony-bounded  space  (see  page  341). 

DISTRIBUTION.  The  species  is  known  only  from  Australia,  recorded  from  'western'  Australia  and 
the  type  locality  of  Port  Phillip  Heads  in  the  south-east. 

Amathia  obliqua  MacGillivray,  1985 
(Figs  3D,  8B) 

Amathia  obliqua  MacGillivray,  1895:  135,  pi.  B  (figs  2, 2a). 

MATERIAL  EXAMINED 

Syntypes:  NMV;  H493  (old  number  65391),  H493  (old  number  65392),  Port  Phillip  heads,  J.  B.  Wilson 
Collection. 

OTHER  MATERIAL 

MM;  7108/2W,  Port  Phillip. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  bifurcate  with  repetitive  gradual  varia- 
tion evident  in  the  branching  angle.  This  ranges  from  equal  dichotomy,  to  the  condition  where  the 
daughter  stolons  are  produced  at  angles  to  the  maternal  stolon  axis  of  30  deg.  and  60  deg. 
respectively.  This  variation  occurs  over  a  sequence  of  four  stolon  units,  i.e.  if  one  daughter  stolon  is 


AMATHIA  327 

angled  at  60  deg.  to  the  right  of  a  maternal  stolon,  the  same  angular  displacement  appears,  to  the 
left  of  a  maternal  stolon,  four  stolon  units  further  on  in  a  stolon  sequence.  The  original  angular 
displacements  are  recovered  after  a  further  sequence  of  four  stolon  units.  In  between  each  of  these 
stages,  there  is  an  intermediate,  equally  dichotomous  condition.  Over  the  entire  sequence,  a  sig- 
moidal  pattern  in  stolon  arrangement  may  be  observed.  Autozooid  groups  occur  towards  the  distal 
ends  of  stolons,  but  usually  there  is  a  further  autozooid-free  portion  beyond  the  group,  of  variable 
length.  Stolons  tend  to  be  straight,  but  sometimes  the  distal  autozooid-free  portion  may  be  twisted 
slightly  or  deflected  anteriorly,  or  both.  Autozooid  groups  are  set  obliquely  on  the  stolons.  The 
autozooid  group  points  in  the  same  direction  as  that,  in  which  the  bearing  stolon  itself  was  budded 
i.e.  viewed  anteriorly,  on  a  right  hand  daughter  stolon,  the  autozooid  group  starts  proximally  on 
the  left  of  the  stolon  and  finishes  distally  on  the  right,  and  vice  versa.  Autozooid  group  orientation 
is  generally  well  preserved  from  stolon  to  stolon,  though  variations  of  up  to  30  deg.  may  occur. 
Autozooids  are  outer-wall  thickened,  and  the  autozooids  of  any  one  group  tend  to  be  about  the 
same  height.  Autozooid  group  profile  therefore  tends  to  be  level.  A  single  proximal-most  auto- 
zooid is  usually  prominent  in  each  autozooid  group.  Its  occurrence,  and  the  pattern  of  autozooid 
displacements  from  stolon  to  stolon,  are  similar  to  those  of  A.  lendigera  (see  page  313)  except  that 
the  second  succession  state  does  not  appear  to  occur.  Rhizoids  may  be  produced,  one  per  stolon, 
from  mid-way  along  the  proximal  autozooid-free  end.  These  are  orientated  at  about  90  deg.  to  the 
autozooids,  on  the  outer  faces  of  stolons  at  a  bifurcation  i.e.  on  the  side  of  a  stolon  away  from  its 
sister  stolon. 

Zh.  0-39  Z/S.65% 

Zw.  0-11  Zn.  11-21     (appearing  as  5-10 'pairs'). 

SI.     1-16-2-13 

Sd.  0- 1 3-0-20  (immediately  proximal  to  the  autozooids) 

REMARKS.  Little  material  is  available  for  study,  therefore  little  is  known  of  the  colony  form,  other 
than  from  MacGillivray's  original  description.  It  is  inferred,  from  the  presence  of  rhizoids,  that  the 
colony  attains  an  arborescent  form.  MacGillivray's  (1895)  description  seems  to  bear  this  out,  the 
colony  being  'attached  by  the  bases  of  main  stems  by  radical  tubes,  the  branches  being  quite  free 
and  not  intertwining  or  climbing  over  other  objects'.  MacGillivray  made  no  mention  of  the  non- 
erect  part  of  the  colony.  Neither  of  the  two  slide  specimens  from  the  NMV,  H493  (65391,  65392) 
matches  the  figure  of  MacGillivray  (1895)  exactly.  There  is  however  a  very  close  resemblance  to 
specimen  H493  (65391).  Some  of  this  colony  fragment  has  broken  away  which  may  account  for  the 
lack  of  congruence  with  the  figure. 

As  with  many  other  species  of  Amathia,  A.  obliqua  has  been  confused  with  A.  lendigera  (by 
MacGillivray  1895).  The  presence  of  rhizoids  and  their  orientation,  the  development  pattern  of 
autozooid  groups  and  the  overall  colony  form,  serve  to  distinguish  this  species  from  A.  lendigera 
(and  also  from  A.  inter medis  and  A.  guernseii). 

DISTRIBUTION.  The  species  is  recorded  only  from  the  Port  Phillip  Bay  region  in  Australia. 

Amathia  wilsoni  Kirkpatrick  1888 
(Figs  4D,  IOC,  D) 

Amathia  wilsoni  Kirkpatrick,  1888:  18,  pi.  2  (figs  4, 4a). 
Amathia  wilsoni:  MacGillivray,  1895:  139,  pi.  D  (figs  2, 2a,  2b). 
Amathia  wilsoni:  d'Hondt,  1983:  67,  fig.  36  (A). 

MATERIAL  EXAMINED 
Syntype:  BMNH;  1888.5.17.7,  Port  Phillip,  J.  B.  Wilson  Collection. 

OTHER  MATERIAL 

BMNH;  1821.5.24.16,  1985.3.12.2,  Portland,  Australia.  1882.7.7.54,  Wilsons  Promontory.  1886.6.8.3, 
Port  Phillip  1910.10.17.31-32  part,  north  end  Victoria  Tasman  Cable,  <50fthms.  (91.44m). 
1963.2.12.361,  Australia  1963.2.12.366,  Holdfast  Bay  nr.  Adelaide.  1985.3. 12.1a,b,  Flinders  Is.  1985.3.18.3, 
Hobart,  Tasmania. 

MM;  7136/3W,  off  Shark  Is.,  Port  Jackson.  7137/3W,  Port  Phillip. 


328  P.  J.  CHIMONIDES 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  based  on  tri-,  tetra-  and  pentafurcation. 
The  colony  is  constructed  of  three  types  of  stolons,  here  termed  'a',  'b'  and  'c'  (see  below).  Often, 
the  region  of  branching,  of  a  maternal  stolon,  is  prominently  thickened.  Pentafurcation  appears  to 
be  associated  with  external  influence  such  as  injury  or  the  presence  of  an  epibiont.  Tetrafurcation  is 
associated  with  astogenetically  early  regions,  forming  the  base  and  main-axis  regions  of  the  colony. 
Main-axis  regions  are  composed  of  type  'a'  stolons.  Trifurcation  is  associated  with  side  branches 
which  develop  from  main-axis  regions.  It  is  not  possible  to  predict  with  certainty,  the  direction  in 
which  stolons  will  be  produced  at  pentafurcation.  In  both  the  tetra-  and  trifurcate  conditions 
however,  one  resultant  component  is  produced  in  linear  succession,  and  two  others  are  produced 
laterally,  one  on  each  side.  These  are  lateral  side  branches,  composed  of  type  'b'  stolons  and  are 
produced  at  an  angle  of  about  60  deg.  to  the  central  axis.  In  the  tetrafurcate  condition,  the  fourth 
component,  comprising  type  'c'  stolons,  is  produced  posteriorly  to  its  maternal  stolon  in  the  central 
axis,  also  at  an  angle  of  about  60  deg.  This  is  a  posterior  side  branch.  Development  along  the  side 
branches  is  limited  and  ends  with  the  production  of  pinnately  arranged,  tapering  kenozooids.  The 
component  kenozooids  of  such  pinnate  groupings  are  arranged  as:  three  in  linear  succession,  with 
an  opposed  lateral  pair  at  both  inter-kenozooidal  junctions.  In  the  lateral  side  branches,  the 
pinnate  kenozooids  are  usually  produced  after  a  'linear'  succession  of  three  stolons;  in  the  posterior 
side  branch,  after  only  one.  The  orientation  of  autozooids  about  the  stolon  along  a  main-axis 
sequence  remains  the  same;  this  same  orientation  is  preserved  in  the  posterior  side  branch.  In  the 
lateral  side  branches,  the  autozooid  group  orientation  is  also  preserved  from  stolon  to  stolon,  but 
the  autozooids  are  usually  re-orientated  to  face  distally  along  the  main-axis;  also  in  these  branches, 
only  one  stolon,  of  a  possible  three,  is  usually  produced  at  each  branching  point.  On  one  side  of  the 
main-axis,  viewed  anteriorly,  this  is  in  the  extreme  right  position;  on  the  other  side  of  the  main-axis, 
this  is  in  the  extreme  left.  In  each  case,  the  other  two  positions  are  replaced  by  a  pinnate  terminal 
kenozooid  group.  The  stolons  along  a  lateral  side  branch  are  thus  deflected  anteriorly  at  each 
junction,  in  relation  to  the  main-axis  stolons.  The  branches  therefore  form  inward  facing  arches 
across  the  anterior  surface  of  the  main-axis  stolons.  The  resulting  form  is  a  long  'cylindrical'  plume, 
reminiscent  of  snake  vertebrae  with  ribs.  The  colony  is  composed  of  a  number  of  these  plumes, 
arising  from  various  positions.  Autozooid  groups  occur  towards  the  distal  ends  of  stolons.  In  the 
main-axis  stolons,  there  is  a  further,  distal,  autozooid-free  portion  to  each  stolon,  usually  corre- 
sponding in  length  to  the  width  of  a  daughter  stolon.  In  the  side  branches,  the  autozooids  fre- 
quently overlap  the  subsequent  branching  point.  All  stolons  may  be  curved  anteriorly.  Autozooids 
are  outer-wall  thickened,  but  the  thickening  differential  is  usually  low.  Autozooid  group  profile 
tends  to  be  level.  The  arrangement  of  autozooids  in  groups  along  main-axis  stolons  is,  to  some 
extent,  predictable.  A  proximal-most  autozooid  may  be  evident  in  a  group,  and  is  associated  with 
one  side  of  the  stolon.  This  autozooid  loses  its  prominence  over  the  next  few  stolons,  the  proximal 
autozoids  of  the  groups  appearing  equally  paired.  Eventually,  a  proximal-most  autozooid 
becomes  prominent  once  more,  but  this  time  is  associated  with  the  opposite  side  of  its  stolon.  Such 
a  sequence  is  estimated  to  occur  over  5  stolon  units.  The  original  condition  is  regained  after  a 
sequence  of  10  stolon  units.  Side  branches,  where  produced,  have  autozooid  groups  each  with  a 
prominent  proximal-most  autozooid  associated  with  the  side  of  its  stolon  nearest  the  main-axis 
stolons.  This  arrangement  is  preserved  in  subsequent  autozooid  groups  along  a  side  branch,  unless 
a  main-axis  sequence  is  produced.  Rhizoids  may  develop,  one  per  stolon,  from  the  proximal  most 
end  of,  usually,  main-axis  stolons.  Each  rhizoid  is  produced  at  about  30  deg.  to  the  orientation  of 
the  autozooids  on  the  same  stolon. 

SI.    (a)  2-44  Z/S.(a)50% 

Zn.  (a)  (14-25    (appearing  as  7-12  'pairs') 

SI.    (b)l-60  Z/S.(b)80% 

Zn.  (b)  18-28  (appearing  as  9-14  'pairs') 

SI.    (c)M3  Z/S.(c)80% 

Zn.  (c)  18-28  (appearing  as  9- 14 'pairs'). 


AMATHIA  329 

Sd.  0-35  (all  stolons) 
Zh.  0-35  (all  stolons) 
Zw.  0-1 3  (all  stolons) 

REMARKS.  The  cuticle  in  some  specimens  is  seen  to  bear  numerous  cyst-like  bodies,  whose  structure 
and  function  have  yet  to  be  determined.  These  cysts  appear  to  be  associated  with  the  distal, 
astogenetically  later  (most  recently  budded)  parts  of  the  colony. 

The  branching  pattern  and  resulting  shapes  in  parts  of  the  colony  are  quite  distinct.  The  overall 
result  is  that  autozooids  face  into  a  relatively  sheltered  colony-bounded  space.  This  arrangement 
may  have  some  protective  advantage  (see  page  341). 

D'Hondt  (1983)  places  A.  verticillata  Waters  MS  and  A.  delicatissima  Busk  MS  in  synonymy 
with  A.  wilsoni.  Only  the  latter  assertion  is  completely  correct.  The  only  apparent  record  of  A. 
verticillata  MS  is  of  slide  MM  7137,  bearing  the  legend  'so  named  by  Kirkpatrick  . . .  KP.  after- 
wards called  it  Amathia  wilsoni  K'.  A.  verticillata  is,  thus,  merely  Kirkpatrick's  MS  name  for  what 
he  subsequently  described  as  A.  wilsoni.  The  slide  was  part  of  Water's  collection,  from  which  the 
confusion  probably  arises.  There  appears  to  be  no  other  record  of  A.  verticillata  Waters  MS. 

In  the  original  description  by  Kirkpatrick  (1888),  a  BMNH  specimen  from  Port  Jackson  is 
apparently  indicated.  No  such  specimen  has  been  found.  The  entry  in  the  account  is  somewhat 
anomalous,  in  that  the  account  deals  with  'Polyzoa  from  Port  Phillip'.  It  seems  likely  that 
Kirkpatrick  was  referring  to  an  additional  specimen,  then  held  in  the  collections  at  the  BMNH,  but 
whose  whereabouts  cannot  now  be  determined,  simply  of  the  same  identity  as  that  which  he 
described.  At  the  beginning  of  the  account,  Kirkpartrick  stated  that  he  was  describing  new  species 
from  a  collection  made  by  J.  B.  Wilson  from  Port  Phillip,  subsequently  sent  to  the  BMNH. 
Specimen  BMNH  1888.5.17.7  matches  this  description  in  being  part  of  such  a  collection,  and  is 
indicated  as  type  material  in  catalogue  and  registration  records,  in  Kirkpatrick's  own  hand.  This 
specimen  is  clearly  syntype  material. 

DISTRIBUTION.  The  species  is  known  from  the  south-eastern  region  of  Australia,  ranging  from 
Holdfast  Bay  near  Adelaide  to  Port  Jackson  near  Sydney  and  Hobart,  Tasmania.  The  record  from 
Flinders  Island  is  not  clear;  it  could  refer  to  the  island  off  Tasmania  or  that  in  the  Great  Australian 
Bight. 

Amathia  pinnata  Kirkpatrick,  1888 
(Figs3B,  10A,B) 

Amathia  pinnata  Kirkpatrick,  1888:  19,  pi.  2  (figs  5,  5a). 

Amathia  pinnata:  MacGillivray,  1895:  136,  pi.  C  (figs  1,  la). 

Part  Amathia  pinnata:  d'Hondt,  1979:  16. 

Part  Amathia  inarmata:  d'Hondt,  1983:  67,  fig.  36  (G). 

Not  Amathia  inarmata:  d'Hondt,  1983:  67,  pi.  2  (fig.  \)(  =  A.  biseriata). 

MATERIAL  EXAMINED 

Lectotype  (selected  here):  BMNH;  1888.5.17.8  A,  Port  Phillip,  J.  B.  Wilson  Collection. 
Paralectotypes:  BMNH;  1888.5.17.8  B,  C,  Port  Phillip,  J.  B.  Wilson  Collection. 

OTHER  MATERIAL 

BMNH;  1 847.6.23. 14,  Tasmania.  1 884. 1 1 . 14.5-1 2  B,  Port  Phillip.  1 886.6.8. 1 ,  Griffths  Point,  Port  Jackson. 
1963.2.12.363,  George  Town,  ?Tasmania?  1985.3.28.1,  Port  Phillip  Heads,  15  m.  1985.3.30.2,  Algoa  Bay,  S. 
Africa. 

MM;  7109/2W,  Port  Phillip. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  regular  and  almost  always  trifurcate. 
This  results  in  a  typically  compound  pinnate  arrangement.  At  any  branching  point,  one  daughter 
stolon  is  produced  in  rectilinear  succession,  and  two  others  are  produced  laterally  opposing,  at  an 
angle  of  about  60  deg.  to  the  centre.  Stolons  tend  to  be  straight.  Autozooids  are  distally  located, 
occupying  the  greater  part  of  stolons.  Autozooid  groups  develop  as  far  as  the  subsequent  branch- 
ing point,  but  do  not  overlie  it.  Autozooid  group  profile  tends  to  be  level,  and  autozooids  are 
outer-wall  thickened,  although  there  is  a  tendency  for  both  differential  and  overall  thickening,  not 


330  P.  J.  CHIMONIDES 

to  be  great.  Autozooid  group  orientation  from  stolon  to  stolon  is  generally  well  preserved.  The 
arrangement  of  autozooids  on  stolons  lying  in  rectilinear  succession  is  to  some  extent  predictable. 
Viewed  anteriorly,  a  proximal-most  autozooid  may  be  prominent  in  an  autozooid  group,  and  is 
associated  with  one  side  of  the  stolon.  Over  successive  stolon  units,  each  autozooid  group  shows 
rearrangement  so  that  this  autozooid  loses  prominence.  The  proximal  autozooids  thus  appear 
equally  paired,  until  a  proximal-most  autozooid  becomes  prominent  once  more,  this  time 
associated  with  the  opposite  side  of  the  stolon.  Such  a  series  appears  to  occur  over  a  sequence  of  4 
stolon  units.  The  original  condition  is  recovered  on  the  seventh  or  eighth  stolon  unit.  In  lateral 
daughter  stolons,  a  single  proximal-most  autozooid  is  prominent  in  the  autozooid  group,  this 
being  associated  with  the  side  of  the  stolon  nearest  the  rectilinear  series,  i.e.  for  a  right-branched 
stolon,  the  proximal-most  autozooid  is  nearest  the  left  side  of  its  stolon,  and  vice  versa. 
Autozooid  groups  on  stolons  subsequently  produced  from  a  lateral  daughter  stolon,  display  the 
same  organisation  along  resultant  rectilinear  series  and  lateral  components.  Terminal  lanceolate 
processes  may  occasionally  be  produced;  each  one  is  made  up  of  a  tapering  series  of  three 
kenozooidal  sub-units.  These  terminal  lanceolate  processes  are  usually  produced  simultaneously 
as  a  group  of  three,  each  process  replacing  a  normal  stolon.  Rhizoids  may  be  produced, 
approximately  mid-way  along  the  proximal  autozooid-free  part  of  the  stolon.  These  arise  singly 
or  as  a  pair,  one  on  either  side  of  the  stolon,  at  about  30  deg.  to  the  autozooid  orientation. 
Colonies  may  be  large  and  arborescent. 

Zh.  0-45  Z/S.80% 

Zw.  0-13  Zn.  18-49    (appearing  as  9-24 'pairs') 

SI.     1-50-3.25          Tpl.  1-50 

Sd.   0-35    (just  proximal  to  the  autozooid  group) 

REMARKS.  Measurements  of  this  species  given  by  Kirkpatrick  (1888)  appear  to  originate  from  the 
same  material  as  he  figured.  There  is,  however,  some  discrepancy  between  the  figures  and  the 
description,  as  it  is  possible  to  infer  18  autozooid  'pairs'  from  his  figure,  whereas  he  described 
the  range  as  being  from  '12-16'. 

Kirkpatrick's  figure  corresponds  to  a  specimen  which  is  obviously  a  fragment  from  a  larger 
colony.  However,  this  specimen  and  another  which  greatly  resembles  it,  are  obviously  not  from 
other  material  stored  in  the  same  container  and  bearing  the  same  registration  number.  All  these 
specimens  are  Kirkpatrick's  A.  pinnata,  as  is  borne  out  by  catalogue  and  registration  records  in 
Kirpatrick's  own  hand.  There  is  a  suggestion,  in  the  stolon  shape  and  rhizoid  production  site,  that 
the  registration  may  harbour  two  species.  A.  pinnata  sensu  stricto,  is  taken  as  the  morph  which 
corresponds  with  Kirkpatrick's  figure;  the  registration  of  these  components  receiving  the  suffixes  A 
and  B  (the  remaining  component  the  suffix  C).  Component  A  is  the  figured  specimen,  and  is  here 
designated  the  lectotype,  the  remaining  portions,  B  and  C,  being  paralectotypes.  Provisionally,  all 
three  components  are  accepted  as  being  A.  pinnata. 

This  species  is  one  of  a  number  that  were  considered  by  d'Hondt  (1979, 1983)  to  be  synonymous 
with  one  another,  the  grouping  also  including:  A.  biseriata;  A.  tricornis  (part);  A.  brongniartii;  A. 
cygnea  MS;  A.  'polycistica'  MS;  A.  desmarestii  MS  (see  page  331).  D'Hondt  (1979)  indicated  A. 
pinnata  Kirkpatrick  1888,  as  the  senior  synonym  of  this  group.  D'Hondt  (1983)  then  indicates  that 
A.  inarmata  MacGillivray  1887,  is  the  senior  synonym  of  the  same  compositional  group,  thus 
subordinating  A.  pinnata  as  a  junior  subjective  synonym.  A.  pinnata  is  in  fact  not  synonymous  with 
any  of  the  species  in  this  grouping,  being  a  separate  and  distinct  species  (see  pages  332,  333). 

DISTRIBUTION.  The  species  is  known  from  the  south-eastern  region  of  Australia,  ranging  from  Port 
Jackson,  to  Port  Phillip  Heads  and  Tasmania,  also  being  recorded  from  Algoa  Bay,  South  Africa. 

A  mat hia  biseriata  Krauss,  1837 
(Figs4B,  11C,D) 

Amathia  biseriata  Krauss,  1837:  23,  fig.  1  (a,  b,  c). 

Not  Amathia  biseriata:  Busk,  1852:  385. 

?  Amathella  biserialis  Gray  1858:  320  (?  errorum  pro  Amathia  biseriata  Krauss,  1837). 


AMATHIA  331 

Amathella  uniserialis  Gray,  1858:  320. 

Amathia  inarmata  MacGillivray,  1887:  183. 

Amathia  biseriata:  Kirkpatrick,  1888:  17. 

Amathia  inarmata:  MacGillivray  1889:  309,  pi.  183  (fig.  4). 

Amathia  biseriata:  MacGillivray,  1895:  137,  pi.  B  (fig.  4). 

part  Amathia  pinnata:  d'Hondt,  1979:  16. 

part  Amathia  inarmata:  d'Hondt,  1983:  67,  fig.  36  (G). 

MATERIAL  EXAMINED 
Neotype  (selected  here):  BMNH;  1887.12.10.90,  Port  Phillip,  J.  B.  Wilson  collection. 

OTHER  MATERIAL 

BMNH;  1899.7.1.4317,4318,  1963.2.12.357,  Australia.  1899.7.1.4319,  New  Zealand.  1965.8.12.19,  Jervis 
Bay,  Huskisson,  N.S.W.  1985.3.14.1,  no  locality. 

NMV;  H492  (1-9),  Port  Phillip  Heads. 

RM;  1808,  Port  Natal,  Africa. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  always  bifurcate.  Viewed  anteriorly,  at 
each  branching  point,  one  daughter  stolon  tends  to  lie  approximately  in  line  with  the  maternal 
stolon,  although  deviations  of  up  to  20  deg.  may  occur.  The  other  daughter  stolon  is  usually 
produced  laterally,  at  an  approximate  angle  of  45  deg.  to  the  main  axis  of  the  maternal  stolon.  The 
side  on  which  a  lateral  branch  appears,  alternates  at  each  successive  bifurcation.  Stolons  have 
a  slight  constriction  near  the  proximal  end  and  are  usually  curved  anteriorly,  the  curvature 
becoming  progressively  more  acute  distally,  to  bend  around  the  distal  end  of  the  autozooid  groups. 
The  stolon  tends  to  remain  in  contact  with  the  distal  side  of  the  autozooids.  Where  this  is  not  so, 
this  portion  of  the  stolon  remains  autozooid-free.  Daughter  stolons  produced  in  the  'linear' 
position,  arise  from  the  distal  end  of  the  maternal  stolon;  lateral  daughter  stolons  are  produced 
from  the  most  sharply  curved  region  of  the  maternal  stolon.  The  distal  region  of  the  maternal 
stolon  may  show  some  axial  subdivision  to  bear  the  daughter  stolons,  more  so  when  it  is  not  in 
contact  with  the  distal  face  of  the  autozooids.  Autozooid  groups  on  maternal  stolons  are  rarely 
developed  distal  to  the  origin  of  the  lateral  daughter  stolon,  or  where  the  stolon  shows  division. 
Autozooids  are  outer-wall  thickened,  the  thickening  sometimes  being  accentuated  at  the  rims. 
Autozooid  group  profile  appears  level,  sometimes  slightly  concave  centrally,  or  diminishing 
slightly  proximodistally  along  the  stolon.  A  single  proximal-most  autozooid  is  evident  in  each 
group,  slightly  broader  than  the  rest,  usually  placed  just  off  the  mid-line,  and  thus  associated  with 
one  or  other  side  of  the  stolon.  No  pattern  has  been  discerned  in  the  location  of  this  autozooid  from 
stolon  to  stolon.  Sometimes  both  daughter  stolons  show  the  same  autozooid  arrangement  as  on 
their  maternal  stolon;  sometimes  the  opposite;  sometimes  combinations  of  the  two.  Autozooid 
orientation  is  generally  well  preserved  from  stolon  to  stolon.  Rhizoids  may  be  produced,  one  per 
stolon,  arising  from  a  position  level  with,  or  just  proximal  to,  the  proximal-most  autozooid  of  the 
autozooid  group.  The  orientation  of  the  rhizoids  is  between  90-135  deg.  to  the  autozooid  group, 
occurring  on  the  same  side  of  the  stolon  as  the  direction  in  which  that  stolon  was  budded.  Stolons 
and  their  autozooid  groups  become  shorter  nearer  the  distal  (astogenetically  later  i.e.  most  recently 
budded)  regions  of  the  colony. 

SI.    2-05  Z/S.75% 

Sd.   0-32  Zn.  5-25  (appearing  as  4-1 2 'pairs') 

Zh.  0-35 

Zw.  0-11 

REMARKS.  The  autozooid  pattern  from  stolon  to  stolon  remains  elusive  in  this  species,  primarily 
due  to  the  difficulties  of  observation  over  the  number  of  stolon  sequences  necessary. 

This  species  is  one  of  a  number  that  were  considered  by  d'Hondt  to  be  synonymous  with  one 
another.  D'Hondt  (1979)  indicated  that  A.  pinnata  Kirkpatrick  1888,  was  the  senior  synonym  of 
this  group,  inclusive  of  A.  biseriata,  and  then  (1983)  indicated  that  A.  inarmata  MacGillivray  1887 
was  the  senior  synonym  of  the  same  compositional  group.  Both  assertions  are  erroneous.  The 
inclusion  of  the  name  A.  desmarestii  in  this  group  is  of  little  consequence  as  Lesueur  never 


332  P.  J.  CHIMONIDES 

published  his  work.  The  name  itself  is  not  valid  in  being  published  only  in  synonymy  (I.C.Z.N. 
article  1  le).  Of  the  other  species  in  this  group:  A.  tricornis  is  a  separate  and  distinct  species  (see 
page  321),  with  a  more  complex  colony  composition  than  the  rest;  A.  cygnea  Busk  MS,  and 
A.  polycistica  (sic)  Busk  MS,  are  here  considered  to  be  synonymous  with  one  another  as 
A.  brongniartii  (see  page  333);  A.  pinnata  is  also  a  separate  and  distinct  species,  in  which 
trifurcate  branching  predominates.  There  are  no  indications,  in  any  of  MacGillivray's  accounts 
of  A.  biseriata,  of  the  trifurcate  branching  pattern  shown  by  d'Hondt  (1983),  who  reproduced 
MacGillivray's  (1895)  figure  of  A.  pinnata.  In  fact,  MacGillivray  and  Krauss  both  stated  that  the 
branching  pattern  in  A.  biseriata  is  dichotomous.  Supplementary  features  which  may  be  used  to 
distinguish  between  A.  biseriata  and  A. pinnata  are:  the  site  of  rhizoid  production;  the  occurrence  of 
terminal  kenozooids  in  the  latter  species.  Were  A.  biseriata  and  A.  pinnata  synonymous,  then 
A.  biseriata  would  be  the  senior  synonym  (cf.  d'Hondt  1979).  A.  biseriata  however,  is  synonymous 
With  A.  inarmata,  but  again,  it  is  A.  biseriata  which  is  the  senior  synonym.  In  this  case,  Krauss' 
publication  predates  that  of  MacGillivray  by  50  years.  Furthermore,  MacGillivray  (1895) 
accepted  his  species  to  have  been  the  same  as  that  of  Krauss.  MacGillivray's  syntypes  are  held  in 
the  NMV  (H492  1-9),  and  all  9  specimens  are  A.  biseriata. 

MacGillivray  (1895)  appears  to  have  been  under  the  misconception  that  Krauss'  material  of 
A.  biseriata  originated  from  south  Africa,  when  in  fact  it  was  from  New  Holland  i.e.  western 
Australia.  It  is  probable  that  MacGillivray  (1895)  was  actually  referring  to  material  received  from 
'Pergens' (MacGillivray,  1889). 

Krauss'  (1837)  description  and  figures  are  here  considered  to  be  more  than  adequate  to  dis- 
tinguish his  species  from  any  other;  his  only  mistake  was  to  make  the  assumption  that  the  rhizoids 
produced  the  stolons  and  their  autozooids.  Krauss'  material  does  not  appear  to  have  survived.  In 
view  of  the  subsequent  confusion,  selection  of  a  neotype  is  necessary.  BMNH  1887.12.10.90 
(although  from  Port  Phillip)  is  selected  here. 

DISTRIBUTION.  The  species  is  known  from  south  Africa,  southern  Australia  and  New  Zealand. 

Amathia  brongniartii  Kirkpatrick,  1888 
(Figs4A,  11A,B) 

Amathia  brongniartii  Kirkpatrick,  1888:  18,  pi.  2  (figs  3,  3a). 

Amathia  brogniartii  (sic):  lapsus  calami  MacGillivray,  1 895:  1 36,  pi.  B  (figs  3,  3a). 

Part  Amathia  pinnata:  d'Hondt,  1979:  16. 

Part  Amathia  inarmata:  d'Hondt,  1983:  67. 

Not  Part  Amathia  inarmata:  d'Hondt,  1983:  67,  fig.  36G,  (  =  A.  pinnata). 

MATERIAL  EXAMINED 
Neotype  (selected  here):  BMNH;  1888.5.17.6,  Port  Phillip,  J.  B.  Wilson  Collection. 

OTHER  MATERIAL 

BMNH;  1838.2.26.13,  1847.6.23.4,  1899.7.1.4379,  4381,  Tasmania.  1887.4.27.19,  Port  Jackson,  N.S.W. 
1887.12.10.98A  (part),  Port  Phillip,  Viet.  1899.7.1.4419,  6601,  Swan  Is.  ?Bass  Strait?  1927.9.26.21,  Swan  Is., 
Banks  Strait.  1984.12.4.1,  Portsea  Pier,  Victoria,  2m.  1985.3.16.1,  Victoria.  1985.3. 16.1.2a,b,  Flinders  Is. 
TTasmania? 

NMV;  65397,  Port  Phillip  Heads,  Viet. 

MM;  7074,  Lane  Cove,  Port  Jackson,  N.S.W. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  always  bifurcate.  At  each  branching 
point,  one  daughter  stolon  tends  to  continue  in  line  with  the  maternal  stolon,  often  giving  rise  to 
rectilinear  series,  although  deviations  by  up  to  15  deg.  may  occur.  The  other  daughter  stolon  of 
each  birfurcation  arises  laterally,  often  anterolaterally,  appearing  on  alternate  sides  along  a  series, 
at  an  angle  of  between  20-50  deg.  to  the  axis  of  the  maternal  stolon.  Stolons  are  usually  straight  and 
tend  to  have  a  slight  constriction  near  the  proximal  end.  The  distal  end  of  a  stolon  does  not  usu- 
ally show  any  axial  subdivision  or  widening  to  bear  daughter  stolons;  more  often,  the  maternal 
stolon  shows  some  abbreviation  into  a  wedge  shape  to  accommodate  these.  The  autozooid  groups 
develop  as  far  as,  and  often  overlie  the  subsequent  branching  point.  Autozooids  are  markedly 
inner-wall  thickened,  with  a  thin  walled  exterior.  The  autozooids  are  usually,  large,  prismatic,  and 


AMATHIA  333 

pentagonal  in  section.  Viewed  anteriorly,  the  walls  between  the  autozooids,  being  much  thicker, 
give  a  characteristic  zig-zag  backbone  appearance  to  the  autozooid  groups.  Autozooid  group 
profile  ranges  from  gently  arched  upwards  to  level  with  the  autozooids  shorter  at  each  end  of  the 
group.  A  single  proximal-most  autozooid  is  evident  in  each  autozooid  group,  slightly  broader  than 
the  rest,  usually  just  off  the  mid  line  and  thus  associated  with  one  or  other  side  of  the  stolon.  Viewed 
anteriorly,  this  autozooid  is  always  on  the  side  nearest  the  sister  stolon.  Autozooid  orientation 
from  stolon  to  stolon  is  generally  well  preserved.  Rhizoids  may  be  produced,  one  per  stolon,  from  a 
position  level  with  or  just  proximal  to  the  proximal-most  autozooid  of  a  group.  These  arise  at 
about  1 10-160  deg.  to  the  autozooid  orientation  on  the  same  stolon,  and  on  the  same  side  of  the 
stolon  as  the  direction  in  which  the  stolon  was  budded.  Stolons  and  their  autozooid  groups  appear 
to  be  shorter  nearer  the  distal  (astogenetically  newer)  regions  of  the  colony. 

SI.    1 -75^-00          Z/S.80% 

Sd.  0-25  Zn.  10-39  (appearing  as  5-18 'pairs') 

Zh.  0-48 

Zw.  0-15 

REMARKS.  This  species  was  considered  by  d'Hondt  to  be  a  junior  synonym  of:  (1979)  A.  pinnata; 
then  (1983)  of  A.  inarmata.  A.  brongniartii  differs  from  A.  pinnata  in  many  features,  such  as:  the 
autozooidal  thickening;  the  sites  of  rhizoid  production;  the  basic  branching  pattern.  A.  inarmata  is 
itself  a  junior  synonym  of  A.  biseriata  (see  page  332).  Kirkpatrick  (1888)  and  MacGillivray  (1895) 
indicated  differences  between  A.  biseriata  and  A.  brongniartii  in  their  accounts.  The  two  species 
may  be  distinguished  quite  readily  by:  the  shape  of  the  stolons;  to  some  extent,  the  site  of  rhizoid 
production;  the  autozooidal  thickening,  this  last  being  the  most  prominent  difference. 

A.  brongniartii  appears  to  display  a  large  variation  in  stolon  length  and  attendant  number  of 
autozooids  borne.  Such  variation  may  be  seen  within  single  colonies.  However,  colonies  may  often 
show  good  uniformity  in  stolon  lengths,  whether  long,  short  or  intermediate.  Busk,  in  his  unpub- 
lished notes  and  figures  stored  at  the  BMNH,  considered  the  possibility  that  the  extremes  of  the  size 
range  might  be  discrete.  He  appears  to  have  called  colonies  with  short  stolons  and  lower  autozooid 
number  A.  cygnea  (up  to  20  autozooids,  equivalent  to  8-12  'pairs'),  with  more  diminutive  versions 
as  A.  cygnea  var.  nana.  Colonies  with  higher  numbers  of  autozooids  (24-36  units,  equivalent  to 
12-18  'pairs')  and  longer  stolons,  he  called  A.polycystica.  In  Busk's  material,  the  specimens  which 
might  be  A.  polycystica  tend  to  be  dark  coloured,  but  other  than  this  there  seems  to  be  nothing 
which  distinguishes  them  taxonomically.  (Busk's  notes  make  no  recognition  of  the  A.  brongniartii 
in  the  unpublished  plates  of  Lesueur). 

Kirkpatrick  (1888),  in  his  account  of  A.  brongniartii,  erroneously  credited  the  species  to 
Desmarest  and  Lesueur,  citing  Lesueur's  figures  and  Pergens'  (1887)  collations  as  his  reference  for 
the  identity  and  name.  As  Desmarest  and  Lesueur  never  published  their  work,  the  Pergens  men- 
tions the  name  only  in  synonymy,  Kirkpatrick  is  the  authority  for  the  species  (I.C.Z.N.  article  1  le). 
As  Kirkpartick  did  not  consider  himself  the  author  of  the  species,  he  did  not  choose  any  type 
specimens.  Kirkpatrick's  figures,  like  those  of  Lesueur,  are  of  insufficient  quality  to  be  utilised  as 
reliable  references.  Kirkpatrick's  figures  hardly  show  any  detail  at  all,  whilst  Lesueur's  figures 
(pi.  13  fig.  5)  show  curved  stolons  and  tube-like  autozooid  anteriors,  both  characteristics  of 
A.  biseriata.  At  the  same  time  however,  Lesueur  shows  approximately  20  'pairs'  of  autozooids,  a 
number  high  enough  to  be  associated  with  A.  brongniartii.  Understandably,  confusion  has  arisen 
and  so  it  would  seem  appropriate  that  a  neotype  be  designated.  BMNH  1888.5.17.6  is,  therefore, 
selected  as  neotype.-  This  specimen  is  the  only  one  labelled  as  A.  brongniartii  by  Kirkpatrick  in  the 
collection  made  by  J.  B.  Wilson  from  Port  Phillip.  This  collection  is  the  subject  of  Kirkpatrick's 
publication  of  1888. 

Specimen  NMV  65397  is  believed  to  have  been  before  MacGillivray  at  the  time  of  his  writing  his 
1895  account  of  Amathia  species  (in  litt.  NMV.  30th  May  1983.).  The  specimen  is  undoubtedly  A. 
brongniartii  Kirkpatrick,  thus  MacGillivray's  name  'A.  brogniartii'  is  simply  a  misspelling,  as  his 
synonymy  indicates. 

DISTRIBUTION.  The  species  is  known  from  south-eastern  Australia,  ranging  from  Tasmania  to  Port 
Jackson. 


334  P.  J.  CHIMONIDES 

Amathia  alternata  Lamouroux,  1816 
(Figs4C,  13A,B) 

Amathia  alternata  Lamouroux,  1816:  160. 

Amathia  alternata:  Lamouroux,  1821:  10,  pi.  65  (figs  18, 19). 

Amathia  alternata:  Lamouroux,  1824:  44. 

Not  Amathia  alternata:  Osburn,  1932: 444,  pi.  1  (fig.  4). 

Part  Amathia  convoluta:  Mature,  1957:  22,  fig.  11. 

Not  part  Amathia  convoluta:  Mature,  1957:  22,  fig.  10. 

Amathia  alternata:  Winston,  1982:  108,  fig.  8. 

MATERIAL  EXAMINED 
Neotype  (selected  here):  USNM;  6307  (part),  Albatross  Stn.  2619,  off  Cape  Fear,  North  Carolina. 

OTHER  MATERIAL 

BMNH;  1964.7.10.1  A,B,  New  River  Inlet,  North  Carolina. 
1964.7.10.2,  Alligator  Harbour,  North  Carolina. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  always  bifurcate.  At  each  branching 
point,  one  daughter  stolon  tends  to  continue  in  line  with  the  maternal  stolon,  forming  a  linear 
series.  The  other  daughter  stolon  is  produced  at  approximately  45  deg.  to  the  maternal  stolon  axis 
at  that  location,  at  between  45-90  deg.  to  the  orientation  of  the  distal  autozooids  there.  Branching 
may  appear  equally  dichotomous  at  times.  Daughter  stolons  are  produced  from  the  posterior  side 
of  the  maternal  stolon,  this  showing  some  abbreviation  into  a  wedge  shape  to  accommodate  them. 
Stolons  are:  narrowed  proximally,  additionally  having  a  proximal  constriction;  often  curved 
posteriorly,  also  undergoing  a  slight  twist  along  their  length.  The  linearly  disposed  stolons  may 
thus  appear  as  an  undulating  progression.  Any  twist  in  these  is  normally  reflected  in  the  autozooids 
borne,  changing  the  orientation  between  proximal  and  distal  autozooids  in  a  group  by  up  to  90  deg. 
At  times,  this  may  give  the  impression  that  autozooid  groups  are  simply  arranged  obliquely  on  the 
stolons.  Autozooid  groups  may,  however,  be  arranged  along  the  stolonal  axis  without  any  evi- 
dence of  twist  at  all  in  either  component.  Where  the  twist,  proximodistally  along  a  stolon,  is 
clockwise,  the  left  daughter  stolon  is  produced  in  the  'lateral'  position,  and  with  anticlockwise 
twist,  the  right.  The  direction  of  twist  is  generally  well  preserved  from  stolon  to  stolon  (although 
both  directions  may  be  found  in  the  same  colony).  Lateral  daughter  stolons  are  thus  produced 
from  the  same  side  of  stolons  along  any  linear  sequence.  The  spatial  orientation  of  these  lateral 
branches  is  determined  by  the  maternal  axis  and  autozooid  orientation  at  that  point.  Autozooid 
orientation  changes  by  100-1 80  deg.  from  stolon  to  stolon.  This  is  taken  from  the  distal  autozooids 
on  the  maternal  stolon  to  the  proximal  autozooids  on  each  of  the  two  daughters.  Autozooids 
frequently  overlie  the  subsequent  branching  point  and  are  outer- wall  thickened.  Autozooids  are 
inclined  at  about  60  deg.  to  the  stolonal  axis,  and  group  profile  is  level  or  gently  convex.  A 
proximal-most  autozooid  is  frequently  evident  in  each  group.  Viewed  anteriorly,  this  is  associated 
with  the  side  of  the  stolon  in  which  the  direction  of  twist  occurs  e.g.  the  right  side,  with  clockwise 
twist  proximodistally.  Rhizoids  may  be  produced  at  any  point  on  a  stolon  from  the  constriction  to 
beneath  the  proximal  autozooids.  A  polyrhizoid  condition  may  result,  where  any  number  of 
rhizoids,  up  to  a  maximum  of  5,  possibly  more,  may  be  produced  from  a  single  stolon,  at  any 
orientation.  Two  orientations  appear  more  frequently  occupied  by  rhizoids:  within  10  deg.  of  the 
proximal  autozooid  orientation  on  the  same  stolon;  approximately  180  deg.  to  the  proximal 
autozooid  orientation. 

Zh.  0-42  Z/S.85% 

Zw.  0-10  Zn.  24-57  (appearing  as  12-28  'pairs') 

SI.     1-45-3-87 

Sd.  0-29-0-35  (just  proximal  to  the  autozooid  group) 

REMARKS.  In  the  non-erect  part  of  the  colony,  secondary  thickening  of  stolons  may  occur.  This  has 
the  appearance  of  a  sleeve  developing  along  existing  stolons. 

A.  alternata  can  display  a  consistent,  if  only  slight,  spiral  nature,  and  the  species  has  been 
confused   with   A.   convoluta  sensu  Lamouroux,   and  possibly  also  with  A.  semiconvoluta 


AMATHIA  335 

Lamouroux.  As  no  type  material  exists,  it  may  also  be  possible  to  confuse  A.  alternata  with  other 
spiral-autozooid  group  species,  for  example  A.  tortuosa  Tennison  Woods  or  A.  connexa  Busk. 

Mature  (1957)  described  and  gave  representative  figures,  under  the  name  of  A.  convoluta  (sensu 
Lamouroux),  of  material  collected  at  Fort  Macon  on  Bogue  Banks,  USA.  This  material  was  noted 
to  have  'straight  autozooid  groups,  alternately  placed  from  one  internode  to  the  next',  and  to  be  the 
same  as  a  specimen  in  the  USNM  labelled  A.  spiralis,  from  Albatross  Stn.  2619,  off  Cape  Fear, 
North  Carolina.  The  latter,  USNM  6307  (part),  has  been  examined  here.  Portions  of  this  specimen 
have  autozooids  arranged  and  placed  in  the  way  Mature  describes  and  illustrates  for  the  Fort 
Macon  material.  This  specimen,  Maturo's  account,  and  the  description  given  here,  conform  with 
Lamouroux's  brief  description  of  A.  alternata  in  1816,  his  subsequent  account  of  1821,  in  which  he 
presents  figures  (pi.  65,  figs  18,  19),  and  his  final  account  in  1824. 

Lamouroux's  (1821)  figures  lack  important  information,  and,  in  not  being  published  at  the  same 
time  as  the  original  description,  are  clearly  not  eligible  for  any  type  status.  Lamouroux's  collection 
was  destroyed  during  the  Second  World  War  (d'Hondt  in  litt.  27.10.1982),  but  material  from  his 
collection  obtained  via  Busk,  and  labelled  A.  alternata  is  stored  at  the  BMNH  as  1897.7.1.6606. 
This  material  was  originally  stored  dried  and  pressed,  but  has  subsequently  been  rehydrated,  and  is 
at  present  stored  in  alcohol.  As  recorded  on  a  label  with  the  material,  examination  by  Dr  F.  Mature 
before  rehydration  revealed  only  two  bryozoan  species,  these  being  other  than  A.  alternata, 
possibly  A.  brasiliensis  Busk  and  Zoobotryon  verticillatum.  Since  rehydration,  the  absence  of  A. 
alternata  is  here  confirmed,  and  the  identity  of  the  two  other  species  established  as  A.  wilsoniand  A. 
semiconvoluta. 

The  packet  originally  enclosing  the  specimens  bears  the  names  'A.  alternata'  and  'Amerique'  in 
what  is  taken  to  be  Lamouroux's  handwriting.  The  locality  mentioned  does  not  disagree  with  that 
of  Lamouroux's  accounts  (1816,  1821,  1 824)  of  A .  alternata,  (the  most  specific  locality  given  being 
the  Sea  of  Antilles  in  1824).  The  two  species  found  enclosed  in  the  packet,  however,  are  not 
expected  from  this  region;  all  other  records  of  A.  wilsoniare  from  southern  Australia.  Similarly,  all 
other  records  for  A.  semiconvoluta  are  from  the  Mediterranean,  the  species  possibly  extending  as 
far  along  the  west  African  coast  as  Nigeria.  It  is  unlikely  that  A.  wilsoni  would  have  been  confused 
with  A.  alternata  by  Lamouroux,  as  the  species  has  many  distinguishing  features  and  lacks  alter- 
nate autozooid  group  placings.  It  may  be  possible  to  confuse  dried  A.  semiconvoluta  with  A. 
alternata,  but  this  is  considered  unlikely  of  Lamouroux,  as  he  is  the  author  of  both  species. 
Lamouroux  introduced  A .  alternata  in  1 8 1 6,  redescribed  the  species  in  1 82 1  and  again  in  1 824  when 
he  introduced  A.  semiconvoluta,  the  descriptions  for  the  two  species  appearing  on  the  same  page.  It 
is  to  be  assumed  that  the  author  was  capable  of  recognising  and  distinguishing  his  own  species.  The 
fate  of  any  A.  alternata  that  may  have  been  present  in  BMNH  1897.7. 1 .6606  is  open  to  speculation. 
The  circumstances  of  Busk's  acquisition  of  specimens  enveloped  in  paper  bearing  Lamouroux's 
writing  are  unknown. 

In  the  interests  of  nomenclatural  stability,  a  neotype  is  required.  Specimens  considered  eligible 
are:  those  in  the  BMNH  under  1964  registrations,  all  from  North  Carolina,  donated  and  identified 
by  Dr  F.  Mature  as  A.  alternata;  specimen  USNM  6307  (part);  Maturo's  Fort  Macon  material. 
The  whereabouts  of  the  Fort  Macon  material  (Mature  1957)  is  not  known.  The  neotype  selected, 
therefore,  is  specimen  USNM  6307  (part)  from  Cape  Fear,  off  North  Carolina,  at  1 5  fthms. 
(27.43  m)  this  being  the  earliest  recorded  specimen  surviving.  The  polyrhizoid  condition  is  not 
readily  apparent  in  this  specimen,  but  it  does  show  the  possible  variation  in  the  arrangement  of 
autozooid  groups. 

The  difference  between  A.  alternata  and  A.  semiconvoluta  are  as  follows:  the  degree  of 'spirality' 
that  may  occur  is  much  greater  in  A.  semiconvoluta  (180-270  deg.,  cf.  A.  alternata  0-90  deg.);  the 
orientations  of  the  distal  end  of  one  autozooid  group  and  the  proximal  end  of  the  next  are  within  10 
deg.  of  each  other  in  A.  semiconvoluta,  but  a  distinctive  100-180  deg.  in  A.  alternata;  a  lower  linear 
autozooid  to  stolon  ratio  of  50%  for  A.  semiconvoluta,  compared  to  about  85%  in  A.  alternata. 
Although  rhizoids  may  appear  at  similar  orientations  in  both  species,  only  A.  alternata  shows 
polyrhizoidy  with  rhizoids  in  proximity  to  the  autozooids  (and,  additionally,  a  slightly  wider 
bifurcation  angle  between  daughter  stolons).  In  A.  semiconvoluta  the  rhizoids  appear  at  the 


336  P.  J.  CHIMONIDES 

proximal-most  end  of  the  stolons,  at  about  the  same  orientation  at  which  the  preceeding  autozooid 
group  terminates,  or  displaced  by  1 80  deg.,  or  when  two  rhizoids  are  present  on  the  same  stolon,  at 
both  orientations.  A.  semiconvoluta  is  understood  from  the  following  specimens:  BMNH; 
1885.12.5.12,13,  Marseilles.  1888.11.9.4,  Naples.  1899.5.1.290,  1912.12.21.687,  Adriatic. 
1 899.7. 1.6606pt.,  locality? 

The  other  species  mentioned  above  i.e.  A.  convoluta,  A.  tortuosa  and  A.  connexa,  are  also 
distinguishable  from  A.  alternata  by  their  degree  of  spirality.  This  is  significantly  greater  than  A. 
alternata  in  all  cases.  Problems  might  arise,  however,  in  distinguishing  these  three  species  from 
each  other,  and  establishing  their  validity. 

A.  convoluta  is  understood  from  BMNH  1899.7.1.6607.  This  specimen  is  from  Lamouroux's 
collection,  obtained  via  Busk,  and  is  labelled  'Amathia  convoluta,  Australasia'  in  what  is 
accepted  to  be  Lamouroux's  handwriting.  There  is  nothing  to  contradict  its  identity  from  any  of 
Lamouroux's  descriptions.  The  specimen  also  conforms  with  MacGillivray's  (1 895)  account  of  the 
species,  corroborated  by  his  opinion  on  Busk  1884  (pi.  6.  fig.  2,  there  misidentified  as  A.  spiralis). 
However,  there  is  nothing  to  suggest  that  MacGillivray  ever  saw  BMNH  1899.7.1.6607  at  any 
time.  This  specimen  is  noted  as  'type'  in  the  catalogue  of  the  BMNH,  though  no  formal  declaration 
of  its  purported  status  has  ever  been  made.  It  is  possible  that  the  specimen  was  formative  of 
Lamouroux's  opinions  of  the  species  and  thus  a  'type'  but  there  can  be  no  certain  evidence  for  or 
against  this  notion.  However,  the  specimen  appears  to  be  the  only  extant  material  which  bears  an 
unchallengeable  identification,  attributable  to  the  original  author.  D'Hondt  (1983)  indicated  that 
Lamarck's  name  for  the  species  (Amathia  crispa),  as  the  senior  synonym,  should  instead  be  used. 

DISTRIBUTION. The  species  is  recorded  off  North  Carolina,  USA,  and,  from  Lamouroux's  (1824) 
record,  from  the  Caribbean. 

Amathia  pruvoti  Calvet,  1911 
(Fig.  13D) 

Amathia  pruvoti  Calvet,  191 1:  59,  fig.  2. 
Amathia  pruvoti:  Bobin  &  Prenant,  1956:  287,  fig.  128. 
Amathia  pruvoti:  d'Hondt,  1983:  67,  fig.  35F. 
Amathia  pruvoti:  Hayward,  1985:  136,  figs  46A,  B. 

MATERIAL  EXAMINED 
Type:  LBIMM;  Bry  8205,  Calvet  collection:  no  locality. 

OTHER  MATERIAL 

BMNH;  1882.7.7. 1-2,  Trieste.  1882.7.7.-,  Mediterranean. 

1885.12.5. 14,  Montpellier.  1889.7.27.48, 1890.7.22.8  part,  Studland  Bay,  Dorset.  1975.7. 1.1 5,  Emborios  Bay, 
Chios,  90  ft.  1984.2.26.102,  Dhiaporia  Rock,  Chios,  100ft. 

DESCRIPTION.  In  the  erect  part  of  the  colony,  branching  is  always  bifurcate.  Daughter  stolons 
appear  to  diverge  equally,  lying  at  approximately  60  deg.  to  each  other,  thus  giving  the  impression 
of  equal  dichotomy.  In  fact,  at  each  branching  point,  one  stolon  tends  to  be  budded  in  a  linear 
position  and  is  subsequently  deflected,  whilst  the  other  is  produced  laterally.  The  linearly  dis- 
posed daughter  stolon  may  be  wider  than  its  sister,  with  little  deflection,  at  times  giving  a  strong 
impression  of  rectilinear  progression.  In  all  stolons,  there  is  a  slight  constriction  near  the  proximal 
end.  The  distal  end  does  not  show  any  axial  subdivision  or  widening  to  bear  daughter  stolons; 
however,  it  often  shows  some  abbreviation  into  a  wedge  shape  to  accommodate  these.  Autozooid 
groups  sometimes  overlie  the  subsequent  branching  point.  However,  it  is  more  usual  for  the 
autozooid  group  to  only  develop  as  far  as  the  branching  point,  or,  alternatively,  'stop  short'  and  be 
followed  by  an  autozooid-free  portion  of  stolon,  about  the  width  of  an  autozooid  in  length.  Stolons 
tend  to  be  straight  proximally  although  often  slightly  curved  posteriorly  and  undergoing  an  axial 
twist  in  the  region  of  the  autozooids.  The  twist  in  the  stolon  is  usually  reflected  in  the  autozooids 
borne,  changing  the  plane  of  their  orientation,  between  the  proximal-most  and  distal-most  auto- 
zooids, by  approximately  90  deg.  The  plane  in  which  the  subsequent  bifurcation  occurs  is  also 


AMATHIA  337 

affected  to  the  same  degree.  The  twists  are  normally  predictable.  Viewed  anteriorly,  in  the  left- 
branched  daughter  stolon,  the  twist  is  usually  clockwise  in  a  proximodistal  direction,  and  anticlock- 
wise in  a  right-branched  daughter.  These  twists  generally  occur  irrespective  of  the  twist  which 
occurs  in  the  maternal  stolon.  However,  there  can  be  variations  to  this.  Occasionally,  both 
daughters  may  twist  in  the  same  direction,  this  being  opposite  to  that  of  their  maternal  stolon. 
Occasionally,  the  inverse  to  the  normal  condition  occurs,  where  a  left  daughter  twists  anticlockwise 
and  the  corresponding  right  daughter  twists  clockwise.  Autozooids  are  outer-wall  thickened. 
Autozooid  group  profile,  where  discernible,  is  level  proximally,  diminishing  distally,  resulting 
from  decreasing  height  and  increasing  distal  inclination  of  the  autozooids.  A  single  proximal-most 
autozooid  is  evident  in  each  autozooid  group,  usually  off  centre  to  the  axis  of  the  stolon.  Viewed 
anteriorly,  this  autozooid  is  always  associated  with  the  same  side  of  the  stolon,  as  the  direction 
in  which  the  autozooid  group  twists,  e.g.  the  right  side,  with  clockwise  twist  proximodistally. 
Autozooid  orientation  from  stolon  to  stolon,  changes  by  1 80  deg.  between  the  distal  autozooids  of 
the  maternal  stolon  and  the  proximal  autozooids  of  each  of  the  two  daughter  stolons.  No  rhizoids 
are  known,  and  the  erect  part  of  the  colony  appears  as  a  diffuse  cotton- wool  like  mass.  Sometimes, 
erect  components  of  the  colony  come  into  contact  with  the  substratum,  and  their  characteristic 
stolonal  shape  is  lost.  These  components  do  not  bear  autozooids;  as  stolonal  kenozopids  (see 
page  309),  they  become  elongated  and  twisted,  occasionally  branching  and  producing  clumps  of 
flattened  lateral  processes.  Further  erect  components  may  be  produced  at  any  time  and  these  may 
resume  the  normal  erect  growth  pattern. 

SI.    2.40-3.75  Z/S.60% 

Sd.   0-13-0-15  Zn.  21-31  (appearing  as  10-15 'pairs') 

Zh.  0-40 

Zw.  0-13 

REMARKS.  Apart  from  the  ancestrula,  little  is  known  of  the  non-erect  portion  of  the  colony.  It  is 
assumed  that  this  would  resemble  the  contact-modified  erect  stolons  and  their  growth  behaviour. 
No  occurrence  of  two  autozooid  groups  on  the  same  stolon  has  been  encountered  in  any  of  the 
material  examined  (cf.  Calvet,  1911).  Such  an  instance  would  be  contrary  to  the  present  concept  of 
the  genus. 

There  is  a  specimen  at  the  LBIMM,  bry  8205,  originating  from  the  Station  Zoologique  de  Cette, 
Universite  de  Montpellier.  This  is  latterly  documented  (e.g.  LBIMM  loan  form  26th  Oct  1983)  as 
'the  probable  type  of  A.  pruvoti:  Calvet  (Cette),  with  a  handwritten  label  of  the  author  carrying  the 
name  A.  semiconvolutcC .  The  justification  for  regarding  LBIMM  bry  8205  as  the  type  specimen  of 
A.  pruvoti  Calvet,  is  not  given.  The  specimen  is,  however,  well  preserved,  and  would  serve  as  an 
excellent  basis  on  which  to  recognise  the  species  in  future.  It  is  proposed  here  that  the  specimen  be 
accepted  as  the  type  specimen  of  the  species.  If  no  historical  justification  for  its  claimed  status  as  a 
'type'  is  available  (see  below),  it  is  here  selected  as  neotype,  obviating  the  confusion  that  has  arisen 
between  A.  pruvoti  and  A.  lendigera  (sensu  lato). 

It  should  be  noted  that  there  is  some  difference  between  Calvet's  description  (1911)  and 
specimens  subsequently  recognised  as  A.  pruvoti,  including  specimen  LBIMM  bry  8205.  Calvet 
described  stolons  as  lying  in  rectilinear  series.  This  condition  is  not  readily  apparent  in  the  majority 
of  specimens,  except  in  two  specimens  from  Chios,  BMNH  1 975.7. 1.15,1 984.2.26. 1 02,  and  in  these 
there  is  also  little  evidence  of  the  proximal  stolonal  constriction.  It  is  not  possible  to  be  certain  of 
what  Calvet  meant  when  he  described  the  'stature'  of  A.  pruvoti  as  'erect',  then  drawing  a  compara- 
tive difference  between  it  and  A.  lendigera,  when  the  colony  budding  patterns  of  the  two  species  are 
in  fact  very  similar.  It  is  possible  that  the  supposed  distinction  may  reflect  an  opinion  that  A. 
lendigera  has  a  higher  proportion  of  the  non-erect  colony  component,  or  that  the  erect  part  of 
A.  lendigera  tends  to  be  spatially  more  condensed.  In  both  species,  there  is  some  variation  in  the 
overall  length  of  stolons.  This  variation  appears  less  extensive  in  A.  pruvoti.  The  most  obvious 
difference  between  the  two  species,  however,  lies  in  the  disposition  of  autozooids  about  the  stolons. 
A  degree  of  twist  is  usually  present  in  A.  pruvoti,  and  an  autozooid-free  distal  portion  of  the  stolon 
often  occurs. 


338  P.  J.  CHIMONIDES 

Calvet  also  drew  a  comparison  with  A.  semiconvoluta.  The  differences  in  the  erect  part  of  the 
colony  between  this  species  and  A.  pruvotiare  that,  in  A.  semiconvoluta:  the  curvature  of  the  stolon 
beneath  the  autozooids  is  much  shallower,  if  present  at  all;  the  autozooid  height  tends  to  be  equal 
throughout  the  autozooid  group;  the  autozooid  group  is  more  spiralled,  undergoing  twists  of 
180-270  deg.;  the  orientations  of  the  distal  end  of  the  autozooid  group  on  the  preceeding  stolon, 
and  the  proximal  ends  of  the  next,  on  the  succeeding  stolons,  occur  within  10  deg.  to  each  other; 
the  direction  of  spiral  tends  to  be  preserved  from  maternal  to  daughter  stolons,  although  both 
directions  may  be  found  in  the  same  colony;  branching  is  always  bifurcate  (as  in  A. pruvoti)  but  one 
daughter  stolon  is  always  linearly  disposed,  giving  rise  to  definite  rectilinear  series,  with  the  other 
daughter  stolon  produced  anterolaterally  at  about  30  deg.  to  the  stolon  axis  and  distal  autozooid 
orientation;  when  the  autozooid  twist,  proximodistally,  is  clockwise,  the  right  hand  daughter 
stolon  is  in  the  rectilinear  position,  and  with  anticlockwise  twist,  the  left  hand  daughter  stolon 
acquires  the  rectilinear  position;  autozooid  groups  always  overlie  the  subsequent  branching 
point;  rhizoids  are  produced  from  the  proximal  end  of  stolons.  Further  characteristics  of  A. 
semiconvoluta  are  as  follows:  rhizoids  arise  singly,  either  in  the  same  orientation  as  the  proximal- 
most  autozooids,  or  at  180  deg.  to  this  (see  pages  335,  336,  Figs  5A,  13C);  when  two  rhizoids  per 
stolon  are  produced,  these  arise  as  one  from  each  orientation;  the  production  of  rhizoids  would 
enable  the  colony  to  attain  an  arborescent  form,  but  this  has  not  been  confirmed. 

There  is,  in  addition,  some  similarity  between  A.  pruvoti,  A.  distans  Busk,  A.  distans  var. 
aegyptana  d'Hondt  and  A.  brasiliensis,  each  of  which  is  a  distinct  entity.  The  distinction  between 
the  species  may  be  found  in  the  following  characteristics.  In  the  last  three,  the  autozooid  groups  are 
more  spiral,  usually  describing  a  360  deg.  rotation  about  the  stolon  in  A.  brasiliensis  and  A.  distans, 
slightly  less  (270-360  deg.)  in  A.  distans  var.  aegyptana.  Of  this  group,  A.  brasiliensis  is  the  only  one 
which  produces  rhizoids,  these  arising  at  the  proximal  end  of  stolons,  orientated  within  10  deg.  to 
the  proximal  autozooids  on  the  same  stolon.  A.  distans  var.  aegyptana  has  the  distinction  of 
producing  autozooid  groups  in  which  the  direction  of  spirality  remains  preserved  from  maternal  to 
daughter  stolons  i.e.  all  clockwise  or  all  anticlockwise,  whereas  one  of  two  other  patterns  prevail  in 
A.  pruvoti,  A.  distans  and  A.  brasiliensis.  Using  the  distal-most  autozooids  as  the  orientation 
reference,  and  viewing  anteriorly:  in  A.  pruvoti  and  A.  brasiliensis,  the  left  daughter  stolons  carry 
autozooids  arranged  clockwise  in  a  proximodistal  direction,  and  the  right  daughters,  anticlock- 
wise; in  A.  distans,  the  left  daughter  stolons  carry  autozooids  arranged  anticlockwise,  and  the  right 
daughters,  clockwise.  The  distinctions  are  made  with  reference  to  type  material: 

For  ,4.  distans:  BMNH  1887.12.9.926,  Bahia,  10-12  fthms.  (18.29-36.58  m.). 
For  A.  brasiliensis:  BMNH  1887.12.9.927,  Bahia,  10-20  fthms.  (18.29-36.58  m.). 
For  A.  distans  var.  aegyptana:  BMNH  1926.9.6.25,  Suez  Canal. 
For  A.  pruvoti:  LBIMM  bry  8205,  no  locality. 

In  conclusion,  A.  distans  var.  aegyptana  should  be  considered  as  a  species  in  its  own  right,  and  is 
here  raised  to  specific  rank  as  Amathia  aegyptana. 

Harmer  (1915)  drew  attention  to  the  similarities  between  A.  distans  and  other  species,  including 
A.  pruvoti.  However,  his  understanding  of  A.  distans,  particularly  in  the  degree  of  spirality  which 
may  occur,  is  here  considered  insufficiently  rigorous.  Unfortunately,  it  is  Harmer's  understanding 
which  is  followed  by  Bobin  and  Prenant  (1956)  and  d'Hondt  (1983). 

DISTRIBUTION.  The  species  is  known  mainly  from  the  Mediterranean,  with  some  material  from 
Studland  Bay  in  Dorset,  England. 


Discussion 

It  is  readily  apparent  that  there  is  a  considerable  degree  of  regularity  and  possible  colony  inte- 
gration within  species  of  the  genus  Amathia.  Some  of  this  is  reflected  in  the  consolidation  of  a 
colony  by  rhizoids.  These  grow  back,  sometimes  fusing  with  each  other,  and  ultimately  interact 


AMATHIA  339 

with  the  substratum  to  provide  support.  The  various  arborescent  growth  forms  that  result  can  only 
be  maintained  through  continued  sustenance  of  these  rhizoids,  and  of  any  underlying  stolons 
which  will  usually  have  lost  their  feeding  autozooids.  This  implies  nutrient  transfer  to  them,  and 
thus  a  potential  ability  for  self  repair. 

The  most  basic  and  obvious  level  of  intergration,  however,  is  the  clustering  of  autozooids  into 
groups  on  septa-bound  kenozooidal  stolons,  to  form  intercommunicating  functional  units.  These 
can  show  changes  of  characteristics  with  astogeny.  Changes  may  be  gradual,  as  in  stolon  lengths 
and  autozooid  numbers  in  A.  biseriata;  or  discontinuous,  as  in  the  autozooid  complement  per 
stolon  in  parts  of  A.  tricornis. 

In  the  majority  of  cases,  the  polypide  appears  capable  of  retracting  to  about  the  level  of  the 
highest  part  of  its  associated  thickened  walls.  This  suggests  that  a  degree  of  protection  may  be 
afforded  by  the  thickening,  and  has  some  analogy  to  the  situation  found  in  other,  calcified, 
bryozoans.  It  is  not  clear  whether  the  mineral  salts  reported  to  be  found  in  the  body  walls  of 
Amathia  (Ryland  1970)  are  associated  with  any  particular  feature,  such  as  this  autozooidal 
thickening. 

From  the  autozooidal  organisation  evident,  there  are  indications  that  some  further  analogy  may 
be  drawn  between  species  of  Amathia  and  other  bryozoans,  in  terms  of  colony  integration  and 
co-ordinated  behaviour.  Together  with  regular  budding  patterns  and  specific  orientations  of  auto- 
zooids, the  localised  autozooidal  thickening  carries  with  it  implications  for  the  achievement  of 
lophophore  eversion  (and  retraction).  The  thickened  areas  of  cuticle  might  resist  the  deformation 
required  by  the  autozooid  to  change  its  volume  and  effect  these  actions.  There  is  little  constraint  on 
independent  action  of  autozooids  in  those  groups  with  inner  wall  thickening;  the  outer  face  of  each 
autozooid  is  able  to  move  freely  in  response  to  the  volume  changes  necessary.  In  groups  with  outer 
wall  thickening,  the  implied  compliant  boundaries  for  each  autozooid  are  those  walls  contiguous 
with  other  autozooids.  Thus,  attempted  changes  in  the  colume  of  any  one  autozooid  might  impinge 
on  the  status  of  those  adjacent.  If  these  adjacent  autozooids  resist  a  change,  then  the  eversion  in  the 
original  autozooid  will  be  hindered.  It  may  be  inferred,  therefore,  that  in  some  species  with  outer 
wall  thickening,  feeding  may  be  a  group  activity.  Advantages  of  group  feeding  would  lie  in 
combined  feeding  currents,  enhanced  by  specific  autozooid  orientations  within  colony  bounded 
space,  (Winston  1979,  McKinney  1984).  Independent  autozooid  behaviour  is  more  likely  if:  the 
thickened  outer  wall  has  localised  weak  patches  acting  as  diaphragms;  the  wall  is  sufficiently  folded 
to  allow  concertina-like  accommodation  of  volume  change;  the  thickening  differential  is  low; 
there  are  co-ordinated  inverse  volume  changes  of  autozooid  pairs.  It  cannot  be  discounted,  how- 
ever, that  collective  feeding  may  occur  in  either  wall-type  grouping,  simply  by  co-operation  of 
autozooids.  Confirmation  of  possible  patterns  of  feeding  behaviour,  however,  requires  the 
observation  of  living  colonies. 

In  the  autozooid  groups,  no  pairing  of  autozooids  may  be  confidently  assigned  throughout  a 
colony  in  any  species  (see  page  309).  Although  the  concept  of  biserial  rows  loses  some  ground,  it 
cannot  be  discounted  completely.  There  is  thus  equal  possibility  that  the  arrangement  of  auto- 
zooids into  groups  may  have  evolved  in  any  of  three  ways:  by  unification  of  two  separate  single 
rows  of  autozooids  with  subsequent  modifications;  by  the  linear  organisation  of  randomly 
clumped  autozooids;  by  spatial  condensing,  with  alternate  displacement,  of  one  single  row  of 
autozooids.  All  three  hypothetical  initial  conditions  have  some  analogues  in  extant  ctenostomes; 
the  first  in  Zoobotryon,  the  last  two  in  species  of  Bowerbankia.  Tenuous  indications  for  origins  via 
the  third  category  may  be  inferred  from  the  order  of  autozooid  production  on  stolons.  Autozooids 
in  a  group  are  developed  in  distal  sequence,  often  making  their  appearance  laterally  displaced  on 
alternate  sides.  It  is  possible,  however,  that  this  simply  reflects  the  fact  that  growth  proceeds 
distally  through  a  sequence  of  interlocked  autozooids,  as  autozooids  may  also  be  seen  to  be 
produced  as  equal  pairs. 

As  with  many  colonial  organisms,  a  large  epifauna  is  frequently  associated  with  colonies  of 
Amathia,  presumably  deriving  benefit  from  the  microenvironment  of  the  colony  interiors  (see 
below).  The  colonies  serve  as  a  substratum  for  some  organisms  and  as  shelter  for  others.  Great 
numbers  of  other  bryozoans,  coelenterates,  crustaceans,  annelids,  algae,  foraminifera  and 


340  P.  J.  CHIMONIDES 

molluscs,  are  often  found.  In  this  context,  the  record  of  Amathia  body  walls  containing  calcium 
salts  (Ryland  1970)  needs  re-investigation  from  material  in  which  the  absence  of  any  encrusting 
calcareous  epibionts  is  ensured,  as  these  can  be  extremely  diaphanous.  It  is  not  known  if  any  of  the 
associations  are  species-specific,  or  what  other  levels  of  interdependence  may  occur.  The  ecological 
criteria  which  determines  distribution  and  survival  of  the  species  of  Amathia  are  known  in  only 
most  general  terms,  and  nothing  is  known  of  the  relative  ecological  requirements  which  epibionts 
and  'hosts'  may  have.  All  that  might  have  been  expected  is  that  numbers  of  epibionts  might  be 
related  to  some  simple  factor,  such  as  the  degree  of  shelter  a  colony  provides.  However,  Murray 
(1970)  reported  that  the  entire  life-cycle  of  the  gastropod  Marginella  minutissima  is  spent  with 
Australian  A.  biseriata.  In  this  case,  the  Amathia  colony  serves  both  as  food  substrate  as  well  as  the 
physical  substratum.  Murray's  concluding  suggestion  was  that  it  is  the  occurrence  of  the  bryozoan 
which  actually  determines  the  mollusc's  distribution. 

The  observable  specific  variation,  and  the  limited  numbers  of  recognisable  characters  perceived 
in  these  non-rigid  animals,  has  made  past  workers,  for  example  MacGillivray  (1895),  Hastings 
(1927),  d'Hondt  (1979,  1983),  variably  reluctant  to  accept  the  existence  of  certain  species.  As  a 
result  it  has  been  suggested  that  some  species,  for  example  A.  lendigera  and  A.  distorts,  are  almost 
ubiquitous.  Wide  geographic  distributions,  continuous  or  discontinuous,  are  not  unknown 
amongst  marine  animals  (Ekman  1967,  Cook  and  Lagaaij  1973),  and  the  genus  has  been  reported 
from  nearly  all  marine  regions  except  the  polar  and  subpolar  seas.  However,  there  is  no  evidence 
that  any  species  of  Amathia  has  ever  achieved  and  maintained  a  cosmopolitan  distribution.  Any 
indications  to  the  contary  seem  based  on  misidentifications.  The  problem  is  compounded  in  one 
instance;  for  two  specimens,  A.  wilsoni  and  A.  semiconvoluta  ex  Lamouroux  collection  (BMNH 
1899.7.1.6606  parts),  there  is  doubt  that  the  locality  data  and  specimens  actually  belong  together 
(see  page  335). 

Although  Rao  and  Ganapati  (1975)  reported  'Amathia  distans1  as  'an  important  fouling  species 
at  the  Visakhaptnam  Harbour',  species  of  Amathia  are  not  noted  as  fouling  the  hulls  of  sea  going 
vessels,  and  there  is  no  indication  that  shipping  has  any  effect  (cf.  Ryland  1970)  on  distribution. 
From  the  information  available  (albeit  that  this  reflects  the  situation  around  the  turn  of  the 
century,  when  many  of  the  specimens  studied  were  collected)  the  species  determined  appear  to  have 
distributions  which  reflect  modern  oceanic  current  flows  (see  below).  This  is  not  unexpected,  as 
Amathia  colonies  are  sessile,  and  the  geographic  distribution  of  species  would  be  greatly  dependent 
on  dispersal  of  colony  fragments  and  larvae  by  water  currents. 

Taken  simplistically,  the  maintenance  of  widespread  distributions  suggests  the  need  for 
adequate  gene  flow  to  help  preserve  the  biological  unity  of  each  species  (Sheppard  1975,  Speiss 
1977),  and  may  be  influenced  by  physical  criteria.  To  some  extent,  this  would  involve  the  effects  of 
sperm  dispersal.  Assuming  some  general  similarity  of  ctenostomes  with  other  Bryozoa,  the  free- 
swimming  life  of  the  lecithotrophic  larvae  (Barrois  1877,  Nielsen  1971,  Zimmer  and  Woollacott 
1977),  might  be  estimated  at  about  24  hours.  Records  of  lecithotrophic  larval  life  in  Cheilostomata 
range  from  20-75  minutes  as  in  Parmularia  (Cook  and  Chimonides  1985),  to  a  maximum  of  3-5 
days  as  in  Crassimarginatellafalcata  (Cook  1985).  Under  the  same  assumption  of  similarity,  sperm 
life  might  be  estimated  as  up  to  1  hour  (Marcus  1926  for  Electra  pilosa,  Silen  1966  for  Electro 
posidoniae).  Lecithotrophic  larval  life  in  Bryozoa  is  generally  held  to  be  short  and  dispersal  limited 
(Ryland  1976,  Farmer  1977,  Hay  ward  and  Cook  1983).  Similarly,  the  contribution  sperm  dispersal 
makes  towards  preventing  speciation  must  also  be  limited. 

It  is  difficult  to  assess  what  contribution  fragmentation  makes  towards  species  distribution;  for 
the  present,  it  is  possible  only  to  speculate  on  the  effects  of  the  factors  involved.  It  is  unlikely  that 
colonies  of  Amathia  would  be  susceptible  to  the  same  shear  forces  that  might  cause  rigid,  calcified 
colonies  to  fail  structurally  (Cheetham  and  Erikson  1983).  The  shape  of  Amathia  colonies  results 
partly  from  the  exoskeletal  function  of  locally  thickened  cuticles,  but  derives  mainly  from  turgor 
pressure  of  the  various  coelomic  fluids  acting  on  the  cuticles.  The  cuticles  are  flexible  but  non- 
elastic.  Such  an  essentially  hydrostatic  support  system  would  be  capable  of  a  great  deal  of  deform- 
ation with  subsequent  recovery.  Structural  failure  results  when  drag  forces  exceed  tensile  strength. 
Tensile  strength  of  alcohol  preserved  specimens  examined  appeared  subjectively  high.  The  failure, 


AMATHIA  341 

near  a  bifurcation,  of  single  stolons  taken  from  distal  tips  of  a  specimen  of  A.  brongniartii  from 
Victoria  Australia  (BMNH  1984.12.4.1),  was  recorded  at  80  grams. 

Additional  resistance  to  fragmentation  is  likely  in  colonies  with  dense  branching.  In  these,  water 
flow  effects  are  prevented  from  acting  directly  on  all  the  constituent  components,  and  the  effective 
drag  of  a  colony  is  less  than  expected  (Cheetham  and  Erikson  1983).  Under  this  condition,  much 
water  flow  would  be  redirected  around  the  colony,  and  this  would  place  some  emphasis  on  the 
external  hydrodynamic  profile  that  a  colony  presents.  A  possible  reaction  to  this  is  suggested  in  the 
fact  that  autozooids  are  often  arranged  to  face  into  the  relatively  sheltered  space  within  the  colony 
interior,  as  for  example  in  A.  wilsoni,  A.  woodsii,  A.populea  and  A.  guernseii. 

The  characteristics  of  flexibility,  reasonable  tensile  strength  and  hydrodynamic  reaction  are, 
however,  the  very  features  which  have  allowed  Amathia  species  to  spread  into  the  kind  of  high 
energy  environments,  for  example,  much  of  southern  Australia  (Thomas  and  Shepherd  1982,  King 
and  Shepherd  1982)  where,  if  only  under  severe  storm  conditions,  fragmentation  of  colonies 
themselves  must  occur.  In  less  extreme  circumstances,  for  some  species,  fragmentation  of  the 
possible  algal  substratum  might  occur,  setting  entire  colonies  adrift. 

The  longevity  of  adult  colony  pieces,  is  potentially  much  greater  than  that  of  the  larvae  and 
sperm.  Under  laboratory  conditions  at  the  BMNH,  specimens  of  Flustrellidra  hispida  survived  for 
over  6  months  without  their  original  algal  substratum,  which  had  rotted  away.  The  colonies 
adopted  a  highly  mishaped  globular  form,  approximately  1-5  cms.  maximum  dimension,  lying  free 
on  the  gravel  filter  bed  of  their  container.  These  colonies  could  be  bowled  around  by  very  mild 
water  movement,  while  the  great  majority  of  the  autozooids  forming  their  surfaces,  retained  the 
ability  to  feed. 

As  colony  fragments  of  Amathia  do  not  readily  float,  it  is  to  be  expected  that  they  will  be 
transported  well  only  whilst  they  are  kept  clear  of  the  sea  floor.  Transportation  and  being  kept 
clear  of  the  sea  floor  will  take  place  only  as  long  as  there  is  the  appropriate  energy  in  the  water 
currents.  More  distant  dispersal  is  possible  if  rafting  on  a  more  bouyant  substratum,  such  as  algae, 
occurs  (Cheetham  1966,  Cook  and  Lagaaij  1973).  The  success  of  any  dispersals  would  require  the 
eventual  deposition  of  species  in  some  suitable  environment.  Three  levels  of  failure  seem  possible: 
that  destination  environments  outside  the  recorded  distribution  are  unsuitable  (in  which  case, 
under  certain  circumstances,  it  is  not  impossible  that  remnants  of  at  least  some  of  these  failures 
might  be  found);  that  dispersals  do  not  reach  wider  transportation  currents;  that  dispersals  do 
reach  wider  transportation  currents  but  suffer  mortality  en  route,  through  loss  of  the  'raft'  as  the 
alga  dies  and  rots.  A.  lendigera  and  A.pruvotican  be  algal  epibionts.  These  species,  if  any,  would  be 
expected  to  have  achieved  very  wide  distributions,  but  this  does  not  appear  to  be  the  case.  Their 
distributions  instead  appear  similar  to  those  of  well  documented  Lusitanian  faunas  (Hardy  1959, 
Ekman  1967,  Tait  1986,  Currie  1983)  (cf.  A.  semiconvoluta  recorded  from  the  west  coast  of  Africa 
to  the  Mediterranean). 

Regardless  of  the  dispersal  method  of  fragmentation  products,  direct  survival  of  fragments 
would  mainly  be  favoured  by  a  low  energy  environment.  Higher  energy  environments  might  allow 
survival  only  through  subsequent  release  of  larvae  and  their  settlement.  No  colonies  have  been 
encountered  where  direct  re-establishment  of  fragments  is  recognised  to  have  occurred.  It  is  quite 
possible,  however,  that  colony  fragments  of  variable  size  may  re-attach  and  grow,  and  even  that 
arborescent  colonies  resume  their  posture  and  growth  form  with  the  aid  of  rhizoids,  in  a  process 
analogous  to  that  observed  in  Parmularia  (Cook  and  Chimonides  1985).  However,  whatever  the 
frequency  of  fragmentation  and  outcome  of  subsequent  events,  the  effects  on  distribution  appear, 
for  the  present,  to  be  of  little  significance. 

The  earliest  record  of  fossil  Amathia  is  from  the  Late  Cretaceous,  with  a  species  appearing  in  the 
Maastrichtian  of  The  Netherlands  (Voigt  1972,  Cheetham  and  Cook  1983).  The  genus  is  not 
associated  with  very  deep  water,  the  deepest  record  encountered  being  150  fathoms  (275  metres 
approx.)  for  specimens  collected  off  Bahia  during  the  Challenger  Expedition.  It  seems  likely, 
therefore,  that  the  genus  achieved  its  present  day  tropical  to  cold-temperate  distribution  via  shelf 
waters  through  Tethys  and  the  Tehuantepec  Channel,  and  to  have  traversed  these  regions  before 
their  closure  in  the  mid  Miocene  (Ekman  1967,  Cook  and  Lagaaij  1973,  Haq  1981).  It  is  obvious 


342  P.  J.  CHIMONIDES 

that  more  evidence  is  required  to  support  these  suggestions,  although  this  may  not  be  readily 
available,  as  non-boring  ctenostomes  have  a  poor  preservation  record  (Cheetham  and  Cook  1 983). 

It  is  interesting  to  note  that  some  of  the  species  recognised,  A.  pinnata,  A.  woodsii,  A.  biseriata, 
seem  to  have  been  recorded  exclusively  from  both  south  Africa  and  southern  Australia.  Parallels 
exist  for  other  bryozoan  species  (Hayward  and  Cook  1983).  This  distribution  is  almost  certainly 
the  resultant  of  palaeogeographic  factors  rather  than  of  modern  current  flows  (see  below),  and 
implies  that  the  genus  was  established  and  speciated  by  the  time  Africa  has  moved  into  relative 
isolation  from  its  Antarctic  association.  This  does  not  extend  the  theoretical  age  of  the  group  much 
beyond  the  Maastrichtian  however  (see  above). 

Although  the  imprecision  of  past  records  is  criticized,  the  interpretation  here  of  both  A. 
brongniartii  and  A.  pinnata  from  Australia,  as  two  single  species,  rather  than  as  species  complexes, 
is  prehaps  lenient  even  on  present  evidence.  Similarly,  the  specific  genetic  unity  implied  in  each  case 
for  A.  biseriata,  A.  woodsii  and  A.  pinnata  in  both  south  Africa  and  southern  Australia,  although 
accepted  here,  must  be  viewed  with  caution.  No  linking  distributions  are  recorded  and  gene  flow 
through  dispersal  of  sperm,  larvae  and  colony  fragments  is  not  favoured  over  such  distances  and 
locations,  and  would  not  prevent  divergence  from  occurring.  Additionally,  long  term  genetic 
stability  of  species  is  implied. 

In  general,  it  may  be  said  that  the  members  of  the  genus  have  had  time  to  become  distributed 
widely.  There  has  also  been  enough  time  for  the  effects  of  isolation  and  isolating  mechanisms  in 
demes  to  have  come  into  play  (Schopf  1977,  Speiss  1977).  Furthermore,  if  the  cryptic  speciation 
indicated  by  Thorpe  and  Ryland  (1979),  for  species  of  the  ctenostome  Alcyonidium,  has  any 
parallel  in  this  ctenostome  group,  further  subdivisions  within  many  of  the  groupings  proposed  here 
should  be  expected. 


Acknowledgements 

I  would  like  to  thank  the  following:  Dr  P.  E.  Bock  as  a  Research  Associate  of  the  National  Museum 
of  Victoria;  Dr  A.  H.  Cheetham  of  the  Smithsonian  Institution;  Dr  J.-L.  d'Hondt  of  the  LBIMM  at 
the  Paris  Museum  and  Dr  C.  Fransen  of  Rijksmuseum,  Leiden,  for  information  and  the  loan  of 
specimens.  I  am  grateful  to:  Dr  R.  V.  Melville  of  I.C.Z.N.  for  advice;  Dr  P.  J.  Hayward,  Swansea 
University;  Dr  J.  D.  Bishop  and  Dr  N.  J.  Evans  of  the  BMNH,  for  constructive  criticisms;  Miss 
B.  C.  Househam,  Mr  A.  Ritch  and  Mr  M.  Viney  formerly  of  the  BMNH  for  their  shouldering  of 
distractive  duties;  the  BMNH  Photo  Unit  for  photographs.  I  am  grateful  to  Miss  P.  L.  Cook  for  her 
support  and  encouragement  in  the  execution  of  this  work. 


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AMATHIA  345 

Winston,  J.  E.  1979.  Current-related  morphology  and  behaviour  in  some  Pacific  Coast  bryozoans.  247-267. 
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Manuscript  accepted  for  publication  19  September  1986 


346 


P.  J.  CHIMONIDES 


tp 


po 


Fig.  1  Schematic  representation  of  erect  colony  components  with  reference  orientations:  (A)  az 
autozooecia,  paz  proximal-most  autozooid,  rh  rhizoid,  s  stolon,  tp  terminal  process;  (B.)  interior  wall 
thickening;  (C.)  exterior  wall  thickening;  (D,  E.)  a  anterior,  d  distal,  /  left  lateral,  p  proximal,  po 
posterior,  r  right  lateral. 


AMATHIA 


347 


Fig.  2    Relative  orientations  of  autozooecia  and  rhizoids  about  the  stolon:  (A.)  A.  guernseii;  (B.) 

A.populea;  (C.)  A.  woodsii;  (D.)  A.  tricornis. 


348 


P.  J.  CHIMONIDES 


Fig.  3     Relative  orientations  of  autozooecia  and  rhizoids  about  the  stolon:  (A.)  A.  lamourouxi;  (B.) 

A.pinnata;  (C.)  A.  plumosa;  (D.)  A.  obliqua. 


AMATHIA 


349 


Fig.  4    Relative  orientations  of  autozooecia  and  rhizoids  about  the  stolon:  (A.)  A.  brongniartii;  (B.) 

A.  biseriata;  (C.)  A.  alternata;  (D.)  A.  wilsoni. 


350 


P.  J.  CHIMONIDES 


Fig.  5  (A.)  Relative  orientations  of  autozooecia  and  rhizoids  about  the  stolon  in  A.  semiconvoluta. 
Exemplified  by  A.  lendigera:  (B.)  normal  autozooid  arrangement  on  a  triad  of  maternal  and  daughter 
stolons  with  sister  stolons  carrying  autozooid  displacements  to  each  other;  (C.)  alternative  autozooid 
arrangement  on  a  triad  of  maternal  and  daughter  stolons,  the  daughter  stolons  carrying  identical 
displacements,  both  opposite  to  the  condition  on  the  maternal  stolon,  dl  left  daughter  stolon,  dr  right 
daughter  stolon,  m  maternal  stolon. 


AMATHIA 


351 


Fig.  6  (A.)  A.  lendigera  BMNH  1942.8.6. 1 5,  Neotype,  Chichester  Harbour,  UK.  x  1 8;  (B.)A.guernseii 
BMNH  1898.5.17.189,  Holotype,  Guernsey,  Guernsey,  UK  x  17;  (C.)  A.  intermedis  BMNH 
1887.5.2.18,  Holotype,  Hastings,  UK  x  27;  (D.)  A.  populea  BMNH  1899.7.1.526,  Lectotype,  Natal, 
South  Africa,  site  of  rhizoid  origin  arrowed  x  44. 


352 


P.  J.  CHIMONIDES 


Fig.  7  (A.)  /4.  lendigera  BMNH  1942.8.6.15,  Neotype,  Chichester  Harbour,  UK  x  8;  (B.)  A.  guernseii 
BMNH  1898.5.17.189,  Holotype,  Guernsey,  UK  x  8;  (C.)  A.  intermedis  BMNH  1842.12.9.14,  Belfast 
Bay,  N.  Ireland  x  6;  (D.)  A.populea  BMNH  1899.7.1.526,  Lectotype,  Natal,  South  Africa  x  8. 


AMATHIA 


353 


Fig.  8  (A.)  A.  lendigera  BMNH  1942.8.6.15,  Neotype,  Chichester  Harbour,  UK,  palmate  processes 
x  28-  (B.)  A.  obliqua  NMV  H493  (65391)  Syntype,  Port  Phillip  Heads,  Aus.  x  10;  (C.)  A.  lamourouxi 
BMNH  1887.12.10.70,  Neotype,  Port  Phillip,  Aus.  x  20;  (D.)  A.  obliquaNMV  H493  (65391)  Syntype, 
Port  Phillip  Heads,  Aus.,  site  of  rhizoid  origin  arrowed  x  57. 


354 


P.  J.  CHIMONIDES 


Fig.  9  (A.)  A.  lamourouxi  BMNH  1887.12.10.70,  Neotype,  Port  Phillip,  Aus.  x  8;  (B.)  A.  cornuta  sensu 
d'Hondt  (A.  woodsii)  LBIMM  2821  part,  TOcean  asiatique',  bifurcate  terminal  process  arrowed  x  8; 
(C.)  A.  lamourouxi  BMNH  1899.7.1.3,  New  Zealand,  apparent  alternate  branching  x7;  (D.)  A. 
woodsii  BMNH  1 883. 1 1 .29.27,  Neotype,  Port  Jackson,  Aus.,  rhizoid  origin  arrowed  x  1 3. 


AMATHIA 


355 


Fig.  10  (A/M./wmataBMNH  1888.5.17.8  A,  Lectotype,  Port  Phillip,  Aus.  x6;(B.)A.pinnataBMNH 
1888.5.17.8  C,  Port  Phillip  Aus.  x  8;  (C.)  A.  wilsoni  BMNH  1888.5.17.7,  Syntype,  Port  Phillip,  Aus. 
x  5;  (D.)  A.  wilsoni  BMNH  1888.5.17.7,  Syntype,  Port  Phillip,  Aus.,  rhizoid  origin  arrowed  x  1 1. 


356 


P.  J.  CHIMONIDES 


Fig.  11  (A.)  A.  brongniartii  BMNH  1888.5.17.6,  Neotype,  Port  Phillip,  Aus.,  rhizoid  origin  arrowed 
x20;  (B.)  A.  brongniartii  BMNH  1888.5.17.6,  Neotype,  Port  Phillip,  Aus.  x8;  (C.)  A.  biseriata 
BMNH  1887.12.10.90,  Neotype,  Port  Phillip,  Aus.  x8;  (D.)  A.  biseriata  BMNH  1887.12.10.90, 
Neotype,  Port  Phillip,  Aus.,  rhizoid  origin  arrowed  x  20. 


AMATHIA 


357 


Fig.  12  (A.)  A.  plumosa  NMV  H494,  Holotype,  Port  Phillip  Heads,  Aus.  x  10;  (B.)  A.  plumosa 
BMNH  1963.2.12.354,  Western  Australia,  rhizoid  origin  arrowed  x  18;  (C.)  A.  tricornis  BMNH 
1899.7.1.6600,  Holotype,  Australia,  rhizoid  origin  arrowed  x  14;  (D.)  A.  convoluta  (A.  crispd) 
BMNH  1899.7.1.6607,  Australasia  x  10. 


358 


P.  J.  CHIMONIDES 


Fig.  13  (A.)  A.  alternata  USNM  6307,  Neotype,  Cape  Fear,  N.C.  USA  x  4  (B.)  A.  alternata  BMNH 
1964.7. 10.1A,  New  River  Inlet,  N.C.,  USA,  showing  polyrhizoid  condition,  the  rhizoid  origins  indi- 
cated x  10;  (C.)A.semiconvoluta  BMNH  1912. 12.2 1.687,  Adriatic,  rhizoid  origin  arrowed  x6;(D.)A. 
pruvoti  LBIMM  Bry  8205,  Type  x  8. 


British  Museum  (Natural  History) 

The  birds  of  Mount  Nimba,  Liberia 

Peter  R.  Colston  &  Kai  Curry-Lindahl 


For  evolution  and  speciation  of  animals  Mount  Nimba  in  Liberia,  Guinea  and  the  Ivory  Coast  is 
a  key  area  in  Africa  representing  for  biologists  what  the  Abu  Simbel  site  in  Egypt  signified  for 
archaeologists.  No  less  than  about  200  species  of  animals  are  endemic  to  Mount  Nimba.  Yet,  this 
mountain  massif,  entirely  located  within  the  rain-forest  biome,  is  rapidly  being  destroyed  by 
human  exploitation. 

This  book  is  the  first  major  work  on  the  birds  of  Mount  Nimba  and  surrounding  lowland 
rain-forests.  During  20  years  (1962-1982)  of  research  at  the  Nimba  Research  Laboratory  in 
Grassfield  (Liberia),  located  at  the  foot  of  Mount  Nimba,  scientists  from  three  continents  have 
studied  the  birds.  In  this  way  Mount  Nimba  has  become  the  ornithologically  most  thoroughly 
explored  lowland  rain-forest  area  of  Africa. 

The  book  offers  a  comprehensive  synthesis  of  information  on  the  avifauna  of  Mount  Nimba 
and  its  ecological  setting.  During  the  20  years  period  of  biological  investigations  at  Nimba  this  in 
1 962  intact  area  was  gradually  opened  up  by  man  with  far-reaching  environmental  consequences 
for  the  rain-forest  habitats  and  profound  effects  on  the  birds.  Therefore,  the  book  provides  not 
only  a  source  of  reference  material  on  the  systematics,  physiology,  ecology  and  biology  of  the 
birds  of  Mount  Nimba  and  the  African  rain-forest,  but  also  data  on  biogeography  in  the  African 
context  as  well  as  conservation  problems.  Also  behaviour  and  migration  are  discussed.  At 
Nimba  a  number  of  migrants  from  Europe  and/or  Asia  meet  Afrotropical  migratory  and 
sedentary  birds. 

Professor  Kai  Curry-Lindahl  has  served  as  Chairman  of  the  Nimba  Research  Laboratory  and 
Committee  since  its  inception  in  1962.  Peter  Colston  is  from  the  Subdepartment  of  Ornithology, 
British  Museum  (Natural  History),  Tring,  and  Malcolm  Coe  is  from  the  Animal  Ecology 
Research  Group,  Department  of  Zoology,  Oxford. 

1986,  129pp.  Hardback.  0  565  00982  6  £17.50. 


Titles  to  be  published  in  Volume  52 


Miscellanea 

A  revision  of  the  Suctoria  (Ciliophora,  Kinetofragminophora)  5.  The  Paracineta 
and  Corynophora  problem.  By  Colin  R.  Curds 

Notes  on  spiders  of  the  family  Salticidae  1.  The  genera  Spartaeus,  Mintonia  and 
Taraxetta.  By  F.  R.  Wanless 

Mites  of  the  genus  Holoparasitus  Oudemans,  1936  (Mesostigmata:  Parasitidae)  in 
the  British  Isles.  By  K.  H.  Hyatt 

The  phylogenetic  position  of  the  Yugoslavian  cyprinid  fish  genus  Aulopyge  Heckel, 
1841,  with  an  appraisal  of  the  genus  Barbus  Cuvier  &  Cloquet,  1816  and  the 
subfamily  Cyprininae.  By  Gordon  J.  Howes 

Revision  of  the  genera  Acineria,  Trimyema,  and  Trochiliopsis  (Protozoa, 
Ciliophora).  By  H.  Augustin,  W.  Foissner  &  H.  Adam 

The  baculum  in  the  Vespertilioninae  (Chiroptera:  Vespertilionidae)  with  a 
systematic  review,  a  synopsis  of  Pipistrellus  and  Eptesicus,  and  the  descriptions  of 
a  new  genus  and  subgenus.  By  J.  E.  Hill  &  D.  L.  Harrison 

Notes  on  some  species  of  the  genus  Amathia  (Bryozoa,  Ctenostomata). 

By  P.  J.  Chimonides 


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


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