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THE  NAUTILUS 


Volume  106 
1992-1993 


AUTHOR  INDEX 


Askew,  T.  M 130 

Bailey,  J.  F 60 

RiELER,   R 15 

Ross,  K.  J 21 

Emerson,  \V.  K 39,  147 

eversole,  .\.  g 119 

Goodfriend,  G.  a 55 

CioODSELL,  J.  G.  119 

I  I.^HASEWYCH,  M.  G 1,  43,  130 

Healy,  J.  M 1 

Hunter,  R.  U 60 

KooL,  S.  P 21 


Marshall,  B  .^ 24 

McLean.  J.  H 39,115,125 

Oleinik,  a.  E 137 

Petit,  RE 43 

Petuch,  E.  J 68 

Potter,  E,  72 

QuiNN.  J.  E,  Jr 50.  77 

Rehder,  ha 123,  127 

Rex,  ma 72 

Tucker,  J   K  76 

Verhecken,  a 43 


NEW  TAXA  PROPOSED  IN  VOLUME  106  (1992-1993) 


n.  sp. 


GASTROPODA 

Pleurotoniariidae 

Pcrotrochtis  maureri  Harasew\cli  &  Askew,  1993,  n 
C'lypeosectidae 

Pseitdorimula  inidatlaniica  McLean,  1992 
Trocliidae 

Calliostomii  allfrninn  Ouiiin,  1992,  n.  sp. 
Calliostoma  argcnlinn  Ouiiin,  1992,  n.  sp. 
CallioHloma  atlanluidcs  Quinn,  1992,  n.  sp. 
C'.aUiostoina  aiilicurn  Quitin,  1992,  n.  sp. 
CUilliosliima  axclolsaoni  Quinn,  1992,  n.  n. 
C'allioslotrm  hcnmidcnsc  Qninn,  1992,  n.  sp- 
CaUiofitunia  brunncopictum  Quinn,  1992,  n.  sp 
Calliostoma  cnidophiluni  Quinn,  1992,  n.  sp. 
Calliostoma  coronatum  Quinn,  1992,  n.  sp. 
Calliostoma  citbensc  Quinn,  1992,  n.  sp 
Calliostoma  debilc  Quinn,  1992,  n.  sp 
Calliostoma  denlalum  Quinn,  1992,  n.  sp. 
Calliostoma  furosnm  Quinn,  1992,  n.  sp.    .  . 
Calliostoma  gncsti  Quinn,  1992,  n.  sp.       ... 
Calliostoma  hilarc  Quiiui,  1992,  n.  sp. 
Calliostoma  hirlum  Quiiui,  1992,  n.  sp. 
Calliostoma  moscaltetlii  Quinn,  1992,  n.  sp. 
Calliostoma  purpureiim  Quinn,  1992,  n.  sp. 
Calliostoma  rota  Quinn,  1992,  n.  sp. 
Calliostoma  rude  Quiiwi,  1992,  n.  sp 
Calliostoma  rngosum  Quinn,  1992,  n.  sp. 
CUilliostoma  sralrnum  Quiiui,  1992,  n.  sp. 
Calliostoma  snirra  Quinn.  1992,  n.  sp. 
Calliostoma  semisuavc  Quinn,  1992,  n   sp 
Calliostoma  serratulum  Quinn,  1992,  n.  sp. 
Calliostoma  tenebrosum  Quinn,  1992,  n.  sp. 
Calliostoma  vinosum  Quinn,  1992,  n.  sp 
Calliostoma  viscardii  Quinn,  1992^  n.  sp. 
Solariella  cristata  Quiiui,  1992,  n   sp. 
Solariella  quadricincta  Quinn,  1992, 
Solariella  slaminea  Quinn,  1992,  ii.  sp 


130 
115 

96 

103 

102 
97 

105 
86 
85 
83 

102 
93 
90 
87 
95 

106 
78 
97 
87 
95 

100 
83 

103 
92 

110 
85 
78 
87 
96 
85 
52 
50 
53 


Colunibellidae 

Cotonopsis  monfilsi  Emerson,  1993,  n.  sp 147 

Faseiolariidae 

Buccinofiisus  patuxcntensis  Petuch,  1993,  n.  sp.  165 

Meloiigenidae 

Bustjcotyptis  choptankensis  Petuch,  1993,  n.  sp.  166 

Tiirrifulgur  manjlandicus  Petuch,  1993,  n.  sp.  166 

Turrifulgur  prunicola  Petuch,  1993,  n.  sp.  167 

Thaidinae 

Ecphora  (Ecphora)  chesapeakensis  Petuch,  1992,  n.  sp.  68 

Ecphora  (Ecphora)  turneri  Petuch,  1992,  n.  sp.  70 

Ecphora  (Trisecphora)  scientisensis  Petuch,  1992,  n.  sp.  70 

Ecphorosycon  lindajoyceac  Petucli,  1993,  n.  sp.  164 

Muricidae 

Patuxentrophon  Petuch,  1993,  n   gen 165 

Volutidae 

Ftilgoraria  (Musashia)  novoilpinica  Oleinik,  1993,  n.  sp.  138 

Ftilgoraria  (Musashia)  genuata  Oleinik,  1993,  n   sp.  140 

Fnlgoraria  (Musashia)  cordata  Oleinik,  1993,  n.  sp.  140 

Fulgoraria  (Musashia)  lilitschikensis  Oleinik,  1993,  n.  sp.  140 

Harpidae 

Harpa  cabriti  Relider,  1992  n.  n 124 

Harpa  goodtvini  Rehder,  1993,  n,  sp 127 

Gancellariidae 

Trilonoharpa  leali  Harasew  \ch.  Petit  & 

\  erhecken,  1992,  n.  sp  45 
Cancellaria  petuchi  Harasewych,  Petit  &  Verhecken, 

1992,  ii..sp ' 47 

Turridae 

Buridrillia  dcrmjorum  Emerson  &  McLean,  1992,  n.  sp.  39 

Calverturris  Petuch.  1993,  n.  gen.  167 

Calverturris  schmidti  Petuch,  1993,  n.  sp 168 

Drillia  macleani  Tucker,  1992,  n.  n 76 

Transmariaturris  Petuch,  1993,  n.  gen 168 

Helminthoglyptidae 

Hemitrochus  bowdcncnsis  (Jncidfriend.  1992,  n.  sp.  55 


rHE  NAUTILUS 


Volume  106,  Number  1 
February  27,  1992 
ISSN  0028-1344 

A  quarterly  devoted 
to  malacology. 


Dry 


MR  6  my      f 

i 

^^'  Hole,  Mass.      J 


EDITOR-IN-CHIEF 
Dr.  M.  G.  Harasewych 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 

ASSOCIATE  EDITOR 
Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
P.O.  Box  2255 
Melbourne,  FL  32902 


CONSULTING  EDITORS 
Dr.  Rudiger  Bieler 
Department  of  Invertebrates 
Field  Museum  of 
Natural  History 
Chicago,  IL  60605 

Dr.  Robert  T.  Dillon,  Jr. 
Department  of  Biology 
College  of  Charleston 
Charleston,  SC  29424 

Dr.  William  K.  Emerson 

Department  of  Living  Invertebrates 

The  American  Museum  of  Natural 

History 

New  York,  NY  10024 

Mr.  Samuel  L.  H.  Fuller 
1053  Mapleton  Avenue 
Suffield,  CT  06078 

Dr.  Robert  Hershler 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 

Dr.  Richard  S.  Houbrick 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


Mr.  Richard  I.  Johnson 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Department  of  Malacology 
Los  Angeles  County  Museum  of 
Natural  History 
900  E.xposition  Boulevard 
Los  Angeles,  CA  90007 

Dr.  Arthur  S.  Merrill 
%  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Ms.  Paula  M.  Mikkelsen 
Harbor  Branch  Oceanographic 
Institution,  Inc. 
Ft.  Pierce,  FL  33450 

Dr.  Donald  R.  Moore 

Division  of  Marine  Geology 

and  Geophysics 

Rosenstiel  School  of  Marine  and 

Atmospheric  Science 

University  of  Miami 

4600  Rickenbacker  Causeway 

Miami,  FL  33149 

Dr.  Gustav  Paulay 
Marine  Laboratory 
University  of  Guam 
Mangilao,  Guam  96923 

Mr.  Richard  E.  Petit 

P.O.  Box  30 

North  Myrtle  Beach,  SC  29582 

Dr.  Edward  J.  Petuch 
Department  of  Geology 
Florida  Atlantic  University 
Boca  Raton,  FL  33431 


Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Geerat  J.  Vermeij 
Department  of  Geology 
University  of  California  at  Davis 
Davis,  CA  95616 


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THEf7NAUTILUS 


CONTENTS 


e  106,  Number  I 

Februani  27 ,  1992 

ISSN  0028-1344 


John  M.  Healy 
!V1.G.  Harasewvch 


Spermatogenesis  in  Perotrochus  quoyanus  (Fischer  6i 

Rernardi)  (Gastropoda:  Pleurotomariidae)    1 


Riidiger  Bieler 

Tcnagodiis  of  Siliquaria'-'  L'rira\elini;  taxonornic  confusion 
in  marine  "worm-snails    iCerithioidea:  Sili(iuariidae) 

15 

Silvard  P.  Kool 

Niirclla  Hodinn,  1798  ((Gastropoda:  Muricidae)  t\pe  species 

21 

Kenneth  J.  Boss 

Bruce  A.  .Marshall 

A  revision  ot  the  Recent  species  of  Eudoliuni  Uall,  1889 
(Gastropoda:  Toiinoidea)           

24 

VCilliam  K.  Emerson 
James  H.  McLean 

Biiridrillia  dcroijorurn.  new  species  from  the  Galapagos 

Islands,  a  li\inu  record  of  a  Neogene  Turrid  genus     

39 

THE  NAUTILUS  106(1):1-14,  1992 


Page  1 


Spermatogenesis  in  Perot rochus  quoyanus  (Fischer  &  Bernardi) 
(Gastropoda:  Pleurotomariidae) 


John  IVI.  Healy 

Departnu'iil  of  Zoology 
The  University  of  Queensland 
Si.  Lueia,  Brisbane 
Queensland,  ALSTRALIA  4()H7 


M.G.  Harasewych 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC:  20560  USA 


ABSTRACT 

The  male  reproducti\e  system,  ultrastructure  of  spermatozoa 
and  spermatogenesis  are  described  for  the  pleurotomariid  Pcr- 
otrochus  quotjanus  (Fischer  i  Bernardi)  CIross  morpholog\  of 
the  male  reproductive  system  of  P.  qtiotjanus  agrees  in  all 
essential  details  with  that  of  Mikadot rochus  beyrichii.  In  all 
features,  spermatozoa  of  Perotrochus  quoyanm  closely  resem- 
ble those  of  Perotrochus  westralis  Whitehead,  1987,  as  well  as 
spermatozoa  of  certain  members  of  the  Trochoidea  (Trochidae, 
Liotiidae)  Spermatozoa  of  P.  quoyanus  have  a  conical  acro- 
somal vesicle  with  a  finely  ridged  anterior  layer,  a  short,  rod- 
shaped  nucleus  with  numerous  lacunae,  a  midpiece  consisting 
of  five  (rareK  four)  mitochondria  surrounding  a  pair  of  cen- 
trioles,  a  rootlet  cotmecting  the  centrioles  and  axonenie  to  the 
nucleus,  and  a  flagelluni  (55-58  ^m  long)  that  is  continuous 
with  the  distal  centriole.  Investigated  species  of  Haliotidae  and 
Scissurellidae  {Sinezona  sp.)  differ  from  Perotrochus  in  acro- 
somal substructure,  and,  in  the  case  of  Sinczuna.  also  in  mid- 
piece  and  nuclear  morpholog\ 

Key  Words.  Spermatozoa,  Spermatogenesis,  Mollusca,  Gas- 
tropoda, Pleurotomariidae,  Perotrochus,  male  reproductive 
tract 


INTRODUCTION 

Living  species  of  the  Pleurotomariidae  have  a  host  of 
primitive  gastropod  features  including  a  prominent  la- 
bial shell  slit  as  well  as  paired  gills,  auricles,  osphradia, 
kidneys,  and  hypobranchial  glands  (Woodward,  1901; 
Bouvier  &  Fischer,  1902;  Fret'ter,  1966;  Hickman,  1984; 
Haszprunar,  1988).  The  Haliotidae  and  Scissurellidae, 
which  share  these  features  and  classically  have  been  as- 
signed to  the  Pleurotomarioidea,  are  now  considered  suf- 
ficiently different  from  the  Pleurotomariidae  to  warrant 
their  placement  into  separate  superfamilies,  while  the 
Pleurotomarioidea  is  considered  most  closely  related  to 
the  Trochoidea  based  on  the  shared  presence  of  a  glan- 
dular urinogenital  duct  in  females  (for  discussion  see 
Haszprunar,  1988,  1989;  McLean,  1989).  Basic  features 
of  pleurotomariid  anatomy,  including  radular  morphol- 
ogy, have  been  known  for  more  than  a  century  (Dall, 


1889;  Bouvier  &  Fischer,  1899,  1902;  Pelseneer,  1899; 
Woodward,  1901;  Fretter,  1964,  1966),  but  it  is  only  in 
recent  years  that  the  advent  of  deep-sea  submersible  craft 
has  allowed  the  biology  and  habitat  of  living  specimens 
to  be  studied  in  detail  and  in  situ  (Yonge,  1973;  Hara- 
sewych et  ai,  1988,  1992). 

The  field  of  comparative  spermatology  has,  over  the 
last  twenty  years,  contributed  greatly  to  the  resolution 
of  taxonomic  and  phylogenetic  problems  in  numerous 
phyla  (Baccetti  &  Afzelius,  1976;  Wirth,  1984;  Jamieson, 
1987),  including  the  Mollusca  (Nishiwaki,  1964;  Popham, 
1979;  Giusti,  1971;  Kohnert  &  Storch,  1984a,b;  Koike, 
1985;  Healy,  1983,  1986,  1988a;  Hodgson  et  ai.  1988). 
Among  the  Gastropoda,  studies  of  archaeogastropod  (,s-./. ) 
spermatozoa  and  spermiogenesis  (Kohnert  &  Storch,  1983; 
Azevedo  et  ai,  1985;  Koike,  1985;  Hodgson  &  Bernard, 
1988;  Healy,  1988b,  1989,  1990a,b)  are  becoming  in- 
creasingly important  since  it  is  from  this  broad  assem- 
blage that  origins  for  the  caenogastropod  and  euthyneu- 
ran  groups  are  sought  (Cox,  1960;  Ponder,  1973; 
Haszprunar,  1988).  The  recent  discovery  of  pronounced 
sperm  dimorphism  in  the  trochoidean  Zaiipais  laseroni 
Kershaw,  1955,  including  a  multi-tailed,  oligopyrene 
paraspermatozoon  (Healy,  1990b),  has  drawn  attention 
to  the  fact  that  comparatively  little  is  known  of  the  range 
of  sperm  morphologies  existing  in  the  Vetigastropoda. 
Healy  (1988b)  provided  the  first  ultrastructural  infor- 
mation on  spermatozoa  of  the  Pleurotomariidae  [Pero- 
trochus westralis  Whitehead,  1987,'  as  Plcurotomaria 
ajricana  (Tomlin,  1948)],  but,  because  of  limitations  im- 
posed by  the  state  of  preservation  of  the  testes,  was  unable 
to  trace  e\ents  of  spermatogenesis  or  give  substructural 
detail  of  certain  sperm  features.  L'sing  glutaraldehyde- 
fixed  testicular  material  of  Perotrochus  quoijanus,  we 
present  the  first  ultrastructural  study  of  sperm  devel- 
opment in  a  pleurotomariid  gastropod. 


'  For  a  discussion  of  the  nomenclature  of  this  species,  see 
Wagner  and  O>onians  (1990). 


Page  2 


THE  NAUTILUS,  Vol.  106,  No.  1 


MATERIAL  AND  METHODS 

Three  male  specimens  of  the  pleurotomariid  Perot rochus 
quoyanus  (Fischer  &  Bernardi,  1856)  were  collect ed  us- 
ing the  research  submersible  JOHNSON-SEA-LINK  II, 
1.03  nautical  miles  west  of  Ilets-a-Goyaves.  off  Basse 
Terre,  Guadeloupe,  West  Indies'  (16°10'33"N, 
6r49'00"W)  at  a  depth  of  350-360  m.  Specimens  were 
maintained  in  refrigerated  aquaria  for  six  days  prior  to 
cracking  the  shells  and  excising  the  testes.  For  scanning 
electron  microscopy  (SEM),  samples  were  prepared  by 
teasing  apart  sections  of  fresh  testes  in  filtered  seawater, 
transferring  droplets  of  sperm  suspension  to  coverslips, 
and  fixing  with  glutaraldehyde  vapor  (25%  glutaralde- 
hyde  in  a  covered  petri  dish).  The  coverslips  were  passed 
through  a  graded  acetone  series  (20-100%),  critical-point 
dried,  and  coated  with  gold-palladium.  The  sperm  were 
examined  using  a  Hitachi  S-570  SEM  at  an  accelerating 
voltage  of  10  kv.  Measurements  are  based  on  SEM  pho- 
tographs of  sperm  and  calibration  grids  of  standard  size 
(2160  lines/mm  at  15,000  X  for  acrosomes,  nuclei,  and 
mitochondria,  19.7  lines/mm  at  1,500  X  for  tails).  For 
transmission  electron  microscopy  (TEM),  l-2mm^  pieces 
of  testicular  tissue  were  fixed  with  5%  glutaraldehyde  in 
0.2  M  cacodylate  buffer  and  shipped  to  the  senior  author. 
Upon  arrival,  samples  were  further  fixed  in  cold  3%  0.2M 
cacodylate-buffered  glutaraldehyde  and  washed  thor- 
oughly in  cacodylate  buffer  before  being  placed  into  a 
1%  solution  of  osmium  tetroxide  (prepared  in  0.2M  cac- 
odylate buffer)  for  two  hours.  Tissues  were  again  rinsed 
in  buffer,  then  dehydrated  using  an  ascending  series  of 
ethanols  (20-100%).  Spurr  s  epoxy  resin  was  used  to  em- 
bed the  tissues  (Spurr,  1969).  Ultrathin  sections  were  cut 
with  an  LKB  IV  Ultratome,  collected  on  uncoated  200- 
mesh  copper  grids,  and  stained  using  either  the  double 
lead  stain  of  Daddow  (1983)  or  a  single  leatl  procedure 
(20  minutes  uranyl  acetate,  10  minutes  lead  citrate).  Sec- 
tions were  examined  using  a  Hitachi  300  transmission 
electron  microscope  operated  at  75  kV.  Remaining  soft 
tissues  were  fixed  in  10%  formaldehyde  in  seawater  and 
transferred  to  70%  ethanol  for  dissection.  Shell  fragments 
retained  as  voucher  specimens  are  housed  in  the  National 
Museum  of  Natural  Historv,  Smithsonian  Institution 
(USNM  878154). 


RESULTS 

Male  Reproductive  Sy.stem 

The  mustard-colored  testis  (fig.  1,  te)  lines  the  right  wall 
of  the  digestive  gland  (fig.  1,  dg)  ,  and  empties  into  a 
thin-walled  testicular  duct  (fig.  1,  td)  situated  ventral  to 
both  these  organs.  This  duct  becomes  tubular  along  the 
ventral  surface  of  the  stomach  (fig.  1,  sto)  and  continues 
anteriorly,  emptying  (fig.  1,  ga)  into  the  ureter  portion 
of  tlie  right  kidney  (fig.  1,  u)  anterior  and  to  the  right 
of  the  opening  (fig.  1,  o)  of  the  anterior  lobe  of  the  right 
kidne\  (fig.  1,  ark),  which  is  situated  in  the  cephalic 
hemocoel.  The  ureter/ urinogcnital  duct  (figs.  1.  2,  u) 
runs  anteriorly  along  the  roof  of  the  mantle  cavity  to  the 


Sto 


rav 


rko 


Fipure§  1-2.  Male  reproductive  tract  of  Perotrochus  qiioy- 
(titiis  (Fischer  &  liernardi),  I.  Diagrammatic  representation 
of  male  reproductive  system,  viewed  from  right  side.  Walls  of 
pericardium  and  right  kidney  removed  to  reveal  contents  2. 
Transverse  section  midway  along  paiiial  gonoduct,  viewed  from 
anterior,  ark,  anterior  lobe  of  right  kidney;  dg,  digestive  gland; 
ga,  genital  aperture;  Ik,  left  kidney;  o,  opening  of  anterior  lobe 
of  right  kidney;  pc,  pericardium;  r,  rectum;  rav.  right  afferent 
liranchial  vessel;  rk,  right  kidney;  rko,  right  kidney  opening; 
sto,  stomach;  td,  testicular  duct;  te.  testis;  u,  urinogenilal  duct. 


right  of  the  rectum  (figs.  1,  2,  r),  envelops  the  right 
afferent  branchial  vessel  (figs.  1,2,  rav),  and  drains  into 
the  mantle  cavity  through  a  transversely  oriented  right 
kidney  opening  (fig.  1,  rko),  approximately  1/4  of  the 
distance  from  the  rear  of  the  mantle  cavity  to  the  rear 
of  the  mantle  slit  The  urinogenital  ducts  of  all  three 
individuals  lacked  a  glandular  lining. 

The  testes  of  two  animals  were  full  of  mature  sper- 
matozoa, while  that  of  the  third  animal  were  almost 
entirely  spent.  .Although  onl\  scattered  groups  of  devel- 
oping cells  remained,  we  were  able  to  identif>-  basic 
features  of  spermatogonia,  spermatocytes  and  sperma- 
tids. 

Mature  Testic;ular  Spermatozoa  (SKM  observations) 

Spermatozoa  of  Perotrochus  quoyanus  consist  of  a  con- 
ical acrosomal  complex  (fig.  4,  a),  a  rod-shaped  nucleus 
(fig  4,  ii),  a  cluster  of  five,  equal-sized,  spherical  mito- 
chondria (fig.  4,  m,  fig.  5,  mp)  at  the  base  of  the  nucleus, 
and  a  single  55-58  ^m  long  Dagellum  (figs.  3-4,  f,  table 


J.  M.  Healy  and  M.  G.  Harasewych,  1992 


Page  3 


Figures  3-5.    Perotrochus  quoyanus-  Mature  testicular  sperm,  SEM.  3.    Two  spermatozoa  including  entire  Qagella  (f).  4,5. 
Acrosome  (a),  nucleus  (n)  and  mitochondria  (m)  of  two  spermatozoa.  Nuclear  lacuna  (nl)  and  detached  midpiece  (mp)  consisting 
of  five  mitochondria  visible  in  figure  5. 
Scale  bars:  3=10  ^m;  4,5  =  2  /um. 


1).  The  acrosomal  complex  (externalK,  the  acrosomal 
vesicle  proper)  is  approximately  1.15  ^m  long,  tapers 
slightly  at  contact  with  the  nucleus,  and  has  a  maximum 
diameter  of  1.18  //m  (figs.  4,5,  table  1).  The  nucleus 
measures  3.7  ^m  in  length,  is  broadest  posteriorly,  with 
a  maximum  diameter  of  1.4  nm.  Irregular  indentations 
on  the  nuclear  surface  (Bg.  5,  nl)  can  be  correlated  by 
TEM  with  nuclear  lacunae  (figs.  6,  7,  11,  nl)  occurring 
beneath  the  nuclear  and  plasma  membranes.  These  in- 
dentations are  not,  therefore,  nuclear  pores.  Spherical 
mitochondria  (diameter  0.8  /im)  obscure  the  attachment 
site  of  the  Dagellum.  The  flagellum  narrows  markedly 
towards  its  insertion  point  within  the  midpiece  (figs. 
13,15). 

Mature  Testicular  Spermatozoa  (TEM  observations) 

Acrosome:  The  acrosomal  vesicle  is  broadly  conical, 
with  a  rounded  anterior  surface  and  flattened  basal  sur- 
face (fig.  7,  av).  The  vesicle  has  a  length  of  0.90-0.93 
nm  and  the  maximum  diameter  of  1.28  ixm  at  its  base 
is  wider  than  the  apex  of  the  nucleus  (figs.  7,8,  n).  A 


deep,  narrow  invagination  extends  anteriorly  from  the 
base  of  the  vesicle  and  is  filled  with  a  diffuse,  faintly 
fibrous  material  (figs.  8,  10,  sm).  Some  sections  clearly 
indicate  an  eccentric,  slightly  angular  alignment  for  the 
invagination  relative  to  the  sperm  longitudinal  axis  (figs. 
8,  10).  Beneath  the  anterior  face  of  the  acrosomal  vesicle 
is  an  electron-lucent  layer  containing  fine  ridges  with  a 
periodicity  of  12-14nm  (figs.  7,9,  rl).  A  similarly  electron- 
lucent  layer,  lacking  discernible  ridged  substructure, 
forms  the  basal  rim  of  the  acrosomal  vesicle  (fig.  7,  br). 
A  loose,  fibrous  deposit  of  subacrosomal  material  occupies 
the  space  between  the  base  of  the  acrosomal  vesicle  and 
the  nuclear  apex  (fig.  8,  sm). 

Nucleus:  The  mature  nucleus  (fig.  6,  n)  is  short  (3.7  ^m) 
and  almost  cylindrical,  with  a  shallow  depression  ante- 
riorly (figs.  7,  8,  n)  and  five  (rarely  four)  shallow  de- 
pressions surrounding  a  centriolar  fossa  posteriorly  (figs. 
14,  15,  n).  The  anterior  depression  is  associated  with 
subacrosomal  material  (fig.  8,  sm),  while  the  posterior 
depressions  act  as  sockets  for  the  midpiece  mitochondria 
(fig.   15,  m).  Dense  material  linking  the  proximal  and 


Page  4 


THE  NAUTILUS,  Vol.  106,  No.  1 


Table  1.  Dimensions  of  mature  spermatozoa  from  SEM  ob- 
servations. Linear  measurements  in  fin\.  (n  =  .30,  10  sperm 
from  each  of  three  individuals.) 


Standard 

deviation 

Mean 

Range 

(<7) 

Acrosome 

Length 

Lie 

1.01-1.19 

0.07 

Width 

LOS 

1.01-1.18 

0.05 

Nucleus 

Length 

3.67 

3.52-3.78 

0.08 

Width  (anterior) 

0.98 

0.95-1.03 

0.02 

Width  (posterior) 

L19 

1.13-1.34 

0.08 

Mitochondria 

Diameter 

0.80 

0.68-0.93 

0.09 

Flagellum 

Length 

56,5 

.52.7-61.1 

3.26 

distal  centrioles  is  continuous  with  a  hollow  rootlet  (figs. 
14,  15,  r),  the  bulbous  end  of  which  fills  the  centriolar 
fossa.  Numerous  irregularly  shaped  lacunae  (figs.  6,  7, 
11,  nl)  occur  within  the  nucleus,  some  of  which  open 
underneath  the  nuclear  membranes,  though  not  to  the 
plasma  membrane  or  cell  surface.  Nuclear  contents  are 
highly  electron  dense  and  consist  of  tightly  packed  fibers 
(diameter  16  nm)  set  in  a  finely  granular  matrix. 

Midpiece:  Five  (rarely  four)  spherical  (diameter  0.6- 
0.8  ^m)  mitochondria  (figs.  12,  15,  m),  each  having 
curved,  plate-like  cristae,  surround  the  proximal  and  dis- 
tal centrioles  to  form  the  sperm  midpiece  (fig.  12).  The 
centrioles  (figs.  14,  15,  pc,  dc),  arranged  at  a  90°  angle 
to  each  other,  are  hollow,  cylindrical  structures  composed 
of  triplet  microtubules  and  emliedded  in  a  pericentriolar 
matrix  (triplets  often  obscured  by  matrix,  see  fig.  15 
inset).  Nine  satellite  fibers  (figs.  15,  16,  sf)  connect  the 
distal  centriole  to  an  annulus  (figs.  15,16,  an),  a  ring- 
shaped  deposit  of  material  lining  the  inner  surface  of  the 
plasma  membrane.  The  flagellar  axoneme,  therefore,  is 
anchored  to  the  midpiece  and  nucleus  via  the  centrioles 
and  rootlet  as  well  as  by  the  radial  set  of  satellite  fibers. 

Flagellum:  The  flagellum  measures  approximately  55- 
58  yuni  ill  length  and  consists  of  a  9  +  2  axoneme  enclosed 
by  the  plasma  membrane  (figs.  15,  f;  17).  Many  sper- 
matozoa were  observed  with  an  angularly  offset  flagellar- 
centriolar  apparatus  (fig.  13).  This  misalignment  could 
be  due  to  tight  packing  of  sperm  within  the  testis  or  even 
slight  immaturity,  since  our  .SEM  observations  on  free 


sperm  show  a  normal,  posteriorly  projecting  flagellum 
(figs.  3-5).  Occasionally,  a  dense  body  is  enclosed  with 
the  axoneme  by  the  plasma  membrane  (fig.  17,  db).  Its 
position  along  the  flagellum  could  not  be  determined. 
Further  study  is  required  to  determine  whether  this 
structure  is  a  true  sperm  feature  of  P.  quoyanus  or  an 
artifact  of  fixation.  In  the  distal  region  of  the  flagellum, 
the  9  +  2  substructure  of  the  axoneme  degenerates  into 
singlet  microtubules  (fig.  17,  arrow). 

Spermatogenesis 

Spermatogenic  cells  present  within  the  testis  consisted 
principally  of  isolated  clumps  of  spermatocytes  and  sper- 
matids (fig.  18,  spc,  spt).  Most  of  the  testis  space  in  ripe 
males  was  found  to  be  almost  totally  occupied  by  tightly 
packed  mature  spermatozoa.  To  some  extent  the  process 
of  reconstructing  events  of  spermatogenesis  was  ham- 
pered by  the  occurrence  of  many  abnormally  developing 
spermatocytes  and  spermatids.  The  morphology  and  pos- 
sible significance  of  these  cells  is  treated  in  the  final 
section  of  these  results. 

Spermatogonia:  Spermatogonia  (fig.  19)  were  only  rare- 
ly observed.  They  can  be  distinguished  from  spermato- 
cytes and  spermatids  by  their  oblong,  usualK  lobulate 
nucleus  (fig.  19,  n;  length  6.0  ^m,  breadth  4.0 /um),  prom- 
inent nucleolus  (fig.  19,  nc;  diameter  0.7  nm),  well-de- 
veloped nuclear  pores  (fig.  19,  arrows),  numerous  small 
mitochondria  (fig.  19,  m;  diameter  0.3-0.4  ^m),  and  more 
extensive  cytoplasm.  Endoplasmic  reticular  cisternae, 
where  visible,  are  scattered  and  poorly  developed.  The 
presence  of  centrioles  and  Golgi  complex  could  not  be 
confirmed  in  the  limited  number  of  observed  cells. 

Spermatocytes:  Spermatocytes  (fig.  20)  have  a  spherical 
to  ovoid  nucleus  (fig.  20,  n;  diameter  4.0-4.5  ^m)  that 
appears  to  lack  either  a  nucleolus  or  prominent  nuclear 
pores.  The  small  electron-dense  patches  visible  in  many 
cells  (fig.  20,  arrowheads)  ma\  be  sites  of  s\naptinemal 
complexes,  although  these  structures  are  more  easily  dis- 
cerned in  moribund  spermatocytes  that  have  partially 
lost  nuclear  contents  (fig.  38,  arrows).  Mitochondria  (fig. 
20,  m;  diameter  0.6-0.75  jum)  markedK  larger  than  those 
of  spermatogonia  are  pressed  slightK  into  the  surface  of 
the  nucleus.  Highly  electron-dense  proacrosomal  vesicles 
(fig.  20,  pav;  diameter  0.1-0.2  ^l^\)  of  Golgian  origin  are 
foimd  throughout  the  cytoplasm.  The  axoneme  (fig.  21, 
ax)  de\elops  intracellularly  from  one  of  a  pair  of  or- 
thogonally arranged  centrioles  (fig.  21,  pc,  dc)  positioned 
close  to  the  concave  face  of  the  Golgi  complex  (fig.  21, 
G).  Even  at  this  early  stage  in  axoneme  formation,  sat- 
ellite fibers  (fig.  21,  sf)  are  associated  with  the  future 


Figures  6-17.  Perotrochus  quoyanus.  Mature  testicular  sperm,  TEM  6.  Acrosome  (a),  nucleus  (n),  nuclear  lacunae  (nl),  and 
mitochondria  (m)  of  two  spermatozoa.  7,8.  Acrosomal  vesicle  (av)  showing  ridged  layer  (rl),  basal  rim  (br),  subacrosomal  material 
(sm),  and  apex  of  nucleus  (n)  with  nuclear  lacuna  (nl).  9.  Detail  of  ridged  laser  (rl)  in  acrosome  10.  Transverse  section  throng!) 
acrosomal  vesicle  showing  subacrosomal  material  (sin).  I  1.  Transverse  section  through  nucleus  showing  nuclear  lacuna  (nl).  12. 
Transverse  section  ihrougli  midpiece,  five  mitochondria  (m)  surround  the  distal  centriole  (dc)  1.3.  ,\ngularl\  offset  centriolar  (c) — 
flagellum  (f)  apparatus  of  a  spermatozoon.  14.    Detail  of  centriolar  fossa  and  attached  rootJcl  \r),  proximal  (pc)  and  di.stal  centrioles 


•^i      -^ 


15 


(dc),  and  mitochondria  (m),  15.    Base  of  nucleus  (n),  rootlet  (r),  proximal  (pc)  and  distal  (dc)  centrioles,  satellite  fibers  (sf ),  annulus 
(an),  flagelluni  (f),  and  mitochondria  (m).  Inset:  triplet  microtubules  of  proximal  centriole  (arrowheads).    16.     Oblique  section 
showing  distal  centriole  and  three  of  nine  satellite  fibers  (sf)  attached  to  annulus  (an).  17.    Transverse  section  through  flagella.  Note 
distal  region  (right)  and  dense  bod>  (db)  (left).  .Arrow  indicates  singlet  microtubules  in  distal  region  of  flagelluni. 
Scale  bars:  6  =  1  Mm;  7,8,10-17  =  0.25  mi";  9  =  0. 1  m'". 


Page  6 


THE  NAUTILUS,  Vol.  106,  No.  1 


distal  centriole.  Endoplasmic  reticular  cisternae  are  poor- 
ly developed. 

Spermatids  (Speriiiiogenesis):  Spermatids  can  be  di- 
vided into  three  categories  based  on  the  condensed  state 
of  the  nucleus:  early  cells,  middle-stage  cells,  and  ad- 
vanced spermatids. 

In  early  spermatids  the  nucleus  (fig.  22,  n)  is  spherical 
with  pale-staining,  fibrous  contents.  Middle-stage  sper- 
matids (figs.  23,24)  are  distinguished  from  earlier  cells 
liy  a  marked  increase  in  the  electron  densit\  of  the  nu- 
clear fibers,  and  by  a  tendency  of  the  mitochondria  and 
centrioles  to  move  toward  the  incipient  posterior  pole  of 
the  nucleus.  Although  multiple  proacrosomal  vesicles  are 
still  apparent  within  the  c\ toplasm  of  middle-stage  sper- 
matids (figs.  24,  25,  pav),  it  is  during  this  phase  of  sper- 
miogenesis  that  the  definitive  acrosomal  vesicle  is  formed 
by  fusion  of  proacrosomal  vesicles.  In  advanced  sper- 
matids, mitochondria  and  the  acrosomal  vesicle  come  to 
lie  in  shallow  depressions  of  the  nucleus,  while  the  nu- 
cleus itself  becomes  oblong  and  its  constituent  fibers  more 
condensed  (figs.  26,27,36,37).  In  addition,  the  acrosomal 
vesicle  undergoes  pronounced  changes  in  shape  and  sub- 
structure. Initially,  the  acrosomal  vesicle  is  round  and 
underlain  by  a  thin  disjointed  layer  of  subacrosomal  ma- 
terial (fig.  26,  sm).  As  seen  in  figure  26,  the  site  of  first 
contact  between  the  definitive  acrosomal  vesicle  and  nu- 
cleus may  occur  close  to  where  the  mitochondria  are 
situated.  Following  attachment  of  the  acrosomal  vesicle 
to  the  condensing  nucleus,  vesicle  contents  become  dif- 
ferentiated into  a  cluster  of  coarse  granules  (fig.  27,  g) 
and  a  more  extensive  homogeneous  portion  (fig.  27,  h). 
These  granules  become  partitioned  into  two  deposits  that 
occupy  anterior  and  posterior  depressions  in  the  homo- 
geneous portion  (fig.  28).  Subsequently,  an  invagination 
of  the  homogeneous  portion,  but  not  the  acrosomal  mem- 
brane, begins  to  form  anteriorly  (fig.  29,  arrowhead). 
The  anterior  cluster  of  granules  transforms  into  a  finely 
ridged  layer  (figs.  28-34,  rl).  As  this  layer  grows,  it  ex- 
tends into  a  deepening  invagination  of  the  homogeneous 
portion  (figs.  29-31,  arrowhead).  The  posterior  cluster  of 
granules  forms  the  electron-lucent  basal  rim  of  the  ac- 
rosomal vesicle.  A  thin  deposit  of  dense  material  defines 
the  basal  region  of  the  acrosomal  membrane  (figs.  28,29, 
dm).  Late  in  spermiogenesis,  the  basal  invagination  of 
the  acrosomal  vesicle  develops  and  is  filled  with  suba- 
crosomal material  (fig.  31,  sm).  The  anterior  invagination 
of  the  homogeneous  portion,  which  is  not  an  invagination 
of  the  vesicle  membrane,  and  the  basal  invagination  of 
the  vesicle  are  distinct  and  unconnected  structures.  The 
anterior  invagination  ultimately  disappears,  perhaps  by 
a  process  of  eversion,  leaving  the  ridged  electron-lucent 
layer  (figs.  31,34,35,  rl)  and  a  small  electron-lucent  i)late 
(figs.  31,34,35,  asterisk). 

Nuclear  lacunae,  so  clearly  apparent  in  mature  testic- 
ular spermatozoa,  only  become  evident  in  the  very  last 
stage  of  spermiogenesis.  These  spaces  are  not  in  contact 
with  the  exterior  of  the  spermatid.  The  ccntriolar  fo.ssa 
(fig.  36,  arrowhead)  forms  through  invagination  ol  the 
nuclear  extension  that  lies  between  the  posteriorly  po- 


sitioned mitochondria  Origins  of  the  pericentriolar  ma- 
trix and  centriolar  rootlet  were  not  determined.  Presum- 
ably the  centrioles  play  some  role  in  the  growth  of  these 
structures. 

Aberrant  spermiogenic  cells:  In  addition  to  spermat- 
ogonia, spermatocytes  and  spermatids,  the  testes  also  con- 
tained numerous  abnormally  developing  spermatocytes 
and  spermatids.  Some  of  these  spermatocytes  appear 
moribund  (fig.  38).  The  spermatids,  however,  are  clearly 
recognizable  by  their  angular  shape,  evidently  the  result 
of  cytoplasmic  pressure  from  adjacent  cells  (figs.  18,  spt; 
39).  Nuclear  condensation  and  proacrosomal  vesicle  pro- 
duction seem  to  proceed  as  in  normally  developing  cells. 
Gradually,  however,  the  nucleus  becomes  oblong  then 
angular  and  ultimately  irregular  in  shape  (figs.  39-42). 
Like  normal  spermatozoa,  mature  nuclei  of  the  abnor- 
mal, presumably  abortive,  lines  have  numerous  lacunae 
(fig.  42,  nl)  and  a  fibro-granulate  substructure  (figs.  40- 
42).  The  proacrosomal  vesicles,  rather  than  forming  a 
definitive  acrosomal  vesicle,  remain  as  a  clump  of  un- 
fused  entities  (Figure  40,  inset,  pav).  Mitochondria,  lo- 
cated in  depressions  of  the  nucleus,  and  axonemal  profiles 
are  often  observed  in  developing  and  mature  aberrant 
spermatozoa  (figs.  40,  m;  41,  ax).  The  position  and  num- 
ber of  centrioles  was  not  determined. 


DISCUSSION 

Reproductive  System:  The  morphology  of  the  male  re- 
productive system  of  Perotrochus  quoyanus  agrees  in  all 
major  features  with  that  of  Mikadotrochus  bcyrichii.  the 
only  other  species  of  pleurotomariid  for  which  the  male 
reproductive  system  has  been  documented  (Woodward, 
1901).  The  female  reproductive  system  of  pleurotoma- 
riids  differs  from  the  male  reproductive  s\stem  in  that 
the  pallial  portion  of  the  right  kidney,  the  urinogenital 
duct,  is  glandular.  To  date,  only  M.  beyrichii  (Wood- 
ward, 1901)  and  Perotrochus  midas  (Fretter,  1966)  are 
confirmed  to  have  glandular  female  urinogenital  ducts. 
The  duct  of  the  holot\  pe  of  Perotrochus  amabilis  (Bayer, 
1963),  an  "immature'  female  on  the  basis  of  gonadal 
sections,  lacked  glandular  elements,  prompting  Fretter 
(1964:179)  to  suggest  that  this  was  a  young  individual 
that  had  never  spawned,  and  that  glands  may  develop 
in  the  walls  of  this  duct  only  as  the  gonad  becomes 
mature.  Examination  of  the  shells  of  more  than  a  dozen 
specimens  collected  in  the  intervening  decades  reveals 
that  the  holotype  of  M.  arnahilis  is  among  the  larger 
specimens  known  of  this  species.  It  is  therefore  unlikely 
that  the  holotype  is  an  immature  individual.  Gonadal 
development  of  several  western  Atlantic  pleurotomariids 
varies  with  season  (Harasewych,  unpublished  observa- 
tions), suggestitig  that  the  glandular  lining  of  the  uri- 
nogenital duct  of  females  may  develop  and  diminish 
cyclically.  As  evidenced  by  the  three  specimens  used  in 
this  stud)  ,  the  urinogenital  ducts  of  male  pleurotomariids 
arc  not  glandular,  even  during  the  spawning  season.  Nev- 
ertheless, absence  of  a  glandular  urinogenital  duct  may 


18 


•*••  19 


•  /' 


s^^ 


'M 


Figure§  18-24.  I'crotrvchus  qiioyanus.  Spermatogenesis,  18.  Survey  section  of  testis  showing  spermatozoa  (spz),  developing 
spermatocytes  (spc),  and  advanced  spermatids  (spt).  Arrows  indicate  aberrant  spermatids.  19.  Spermatogonium,  Note  lobulate 
nucleus  (n),  large  nucleolus  (nc),  nuclear  pores  (arrows),  and  numerous  small  mitochondria  (m),  20.  Spermatocytes.  Note  mito- 
chondria (m),  nucleus  (n),  presence  of  proacrosomal  vesicles  (pav),  and  putative  synaptinemal  complexes  (arrowheads).  21.  Sper- 
matocyte Golgi  complex  (G)  close  to  proximal  and  distal  centrioles  (pc,  dc)  and  axoneme  (ax).  Note  satellite  fibres  (sf)  associated 
with  distal  centriole.  22.  Early  spermatids  (spt)  with  homogeneously  granular  nuclei  (n)  and  proacrosomal  vesicles  (pav).  23. 
Middle  stage  spermatids  with  very  electron  dense  fibrillar  nuclei  (n)  and  mitochondria  (m).  24.  Middle  stage  spermatid  showing 
pair  of  centrioles  (c),  proacrosomal  vesicles  (pav),  and  mitochondria  (m). 
Scale  bars:  18  =  10  ^m;  19,20.22-24  =  1  Mm;  21  =  0.5  Mm. 


Pages 


THE  NAUTILUS,  Vol.  106,  No.  1 


y.  a; 


FiguiT.,  2.>   .>.i (.;.  Ini:.  quuy.iiui.;    \k  icwiiiif  (Ifvelupmeiil.  25.    (iruup  of  proacTo.sonial  vesicles  (pav)  near  mitochondrion 

(m)  and  axoneme  (ax).  26.    Spermatid  with  acrosomal  vesicle  (av)  contacting  nucleus  (n)  near  mitochondrion  (m)   Subacrosomal 


J.  M.  Heal)  and  M.  G.  Harasewych.  1992 


Page  9 


not  be  a  sufficient  criterion  for  identifying  male  speci- 
mens. 

Spermatogenesis:  Despite  the  complicating  factor  of 
moribund  and  abnormally  developing  cells  within  the 
testis,  spermatogenic  stages  of  Perotrochus  quoyanus  re- 
semble those  reported  for  the  Trochoidea  (Kohnert  & 
Storch,  1983;  Azevedo  et  a/.,  19S5;  Koike,  1985;  Healy, 
1989).  Using  museum-preserved  tissues,  Healy  (1988b) 
was  able  to  determine  that  acrosomal  development  in 
Perotrochus  westralis  involved  the  production  of  mul- 
tiple proacrosomal  vesicles.  Fusion  of  proacrosomal  ves- 
icles into  a  definitive  acrosomal  vesicle  has  been  dem- 
onstrated in  many  bivalves  (Longo  &  Dornfeld,  1967; 
Kubo,  1977;  Bernard  &  Hodgson,  1985;  Hodgson  &  Ber- 
nard, 1986;  Eckelbarger  et  ai,  1990),  and,  outside  the 
Mollusca,  in  groups  as  disparate  as  the  Polychaeta  (Fran- 
zen,  1987)  and  Echinodermata  (Dan  &  Sirakami,  1971; 
Chia  &  Bickell,  1983).  In  contrast,  acrosome  develop- 
ment in  patelloidean  gastropods  centers  on  the  produc- 
tion of  a  single,  electron-lucent  vesicle  to  which  small 
vesicles  from  the  Golgi  cisternal  edges  fuse  and  contrib- 
ute (Hodgson  &  Bernard,  1988).  Our  study  has  discov- 
ered details  of  acrosome  development  previously  unde- 
scribed  in  the  Vetigastropoda,  including  the 
differentiation  of  anterior  and  posterior  extremities  of 
the  vesicle  and  formation  of  fine  ridges  in  the  anterior 
electron-lucent  layer.  There  are  reasons  for  believing  that 
these  events  also  occur  in  the  Trochoidea.  Mature  ac- 
rosomes  of  trochids  frequently  show  anterior  and  pos- 
terior electron-lucent  layers  (the  anterior  layer  with  ridg- 
es: HeaK  &  Daddow  unpublished).  In  spermatids  of 
Omphalius  pfeifferi  (Philippi,  1846),  the  definitive  ac- 
rosomal vesicle  (Koike,  1985:plate  3D)  closely  corre- 
sponds to  the  stage  illustrated  herein  for  Perotrochus 
quoyanus  (fig.  30).  The  origin  of  the  subacrosomal  ma- 
terial in  P.  quoyanus  and  in  Giblntla  umbilicalis  (da 
Costa,  1778)  (see  Azevedo  et  ai,  1985)  is  unknown.  A 
Golgian  source  seems  unlikely,  as  this  secretory  organelle 
has  migrated  posteriorly  by  the  time  the  definitive  ac- 
rosomal vesicle  has  formed  (the  stage  when  subacrosomal 
material  becomes  visible).  Possibly,  the  acrosomal  vesicle 
itself  is  capable  of  organizing  the  accretion  or  polymer- 
ization of  extravesicular  materials  within  the  cytoplasm. 
Takaichi  &  Dan  (1977)  proposed  a  similar  origin  for 
subacrosomal  material  in  the  pulmonate  Euhadra  hick- 
onis  (Kobelt,  1879).  An  interesting  feature  of  spermio- 
genesis  in  Perotrochus  quoyanus  is  the  often  distant  po- 
sitioning  of    the    nuclear-contacted    acrosomal    vesicle 


relative  to  this  vesicle's  final  position  at  the  nuclear  apex 
(see  fig.  26).  A  comparable  situation  occurs  in  the  trochid 
Calliotropis  glyptus  (Watson,  1879)  (see  Healy,  1989) 
and  evidently  in  the  turbinid  Lunella  granulata  (Gmelin, 
1791)  (see  micrographs  of  Koike,  1985).  In  Perotrochus, 
Calliotropis,  and  Lunella,  however,  the  mature  acro- 
.somal  vesicle  lies  at  the  nuclear  apex,  indicating  that  by 
some  means,  perhaps  via  nuclear  shape  change  late  in 
spermiogenesis  or  acrosomal  movement,  the  vesicle  at- 
tains its  final  position. 

The  pattern  of  nuclear  condensation  in  Perotrochus 
quoyanus  differs  from  that  occurring  in  the  Trochoidea 
in  two  respects:  (1)  the  heterochromatin  forms  a  ho- 
mogeneous network  of  dense  fibers,  whereas  the  hetero- 
chromatin forms  distinct  granules  in  Trochoidea;  and  (2) 
nuclear  lacunae  appear  only  at  the  last  stage  of  spermatid 
development,  whereas  the  lacunae  are  well  developed 
and  visible  at  earlier  stages  in  Trochoidea  (Kohnert  & 
Storch,  1983;  Azevedo  et  a!.,  1985;  Koike,  1985;  Healy, 
unpublished  data).  Unfortunately,  no  comparative  in- 
formation exists  on  nuclear  condensation  or,  in  fact,  on 
any  aspect  of  spermiogenesis,  in  the  Haliotidoidea,  Scis- 
surelloidea,  or  Fissurelloidea.  Initially  thecentriolar  fossa 
of  Calliotropis  glyptus  spermatids  resemble  the  mature 
fossa  of  Perotrochus  spp.,  but  late  in  spermiogenesis,  the 
solid  rootlet  and  attached  centrioles  of  C.  glyptus  become 
drawn  into  a  greatly  expanded  fossa  (Healy,  1989). 

Incorporation  of  the  future  flagellar  axoneme  within 
the  cytoplasm  of  spermatocytes  and  spermatids  in  Per- 
otrochus warrants  some  comment.  The  same  phenom- 
enon can  be  seen  in  published  micrographs  of  developing 
spermatids  in  the  trochid  Monodonta  turbinata  (Born, 
1778)  (see  Kohnert  &  Storch,  1983)  and  in  the  turbinid 
Lunella  granulata  (see  Koike,  1985).  Unfortunately,  nei- 
ther Kohnert  and  Storch  (1983)  nor  Koike  (1985)  offer  a 
discussion  of  this  positioning  of  the  axoneme.  In  sper- 
matids of  the  caudofoveate  Chaetoderma  sp.,  the  prox- 
imal and  distal  centrioles  each  give  rise  to  an  axoneme 
within  the  cytoplasm  (Buckland-Nicks  &  Chia,  1989). 
Of  these  two  axonemes,  only  that  associated  with  the 
future  distal  centriole  survives  into  the  mature  sperma- 
tozoon. A  similar  situation  has  been  reported  by  Eckel- 
barger et  al.  (1989)  in  paraspermatozoan  development 
of  the  abyssal  sea  urchin  Phrissocijstis  multispina,  with 
the  exception  that  both  axonemes  survive  in  the  mature 
cell.  Given  the  large  number  of  abnormally  developing 
spermatids  observed  in  the  ripe  testes  of  our  specimens 
of  Perotrochus  quoyanus,  it  cannot  be  ruled  out  that  the 
intracellular  axoneme  in  spermatids  of  this  species  may 


material  (sin)  is  thin.  Arrowheads  indicate  axonemal  profiles.  27.  Acrosomal  vesicle  showing  granule  cluster  (g)  and  homogeneous 
portion  (h)  of  vesicle  contents.  Note  also  subacrosomal  material  (sm ),  28.  Granules  (g)  distributed  in  anterior  and  posterior  depressions 
of  homogeneous  portion  (h).  Note  ridged  layer  (rl)  and  basal  rim  defined  by  dense  material  (dm).  29.  Beginning  of  invagination 
(arrowhead)  of  homogeneous  portion  (h).  30.  Penetration  of  ridged  layer  (rl)  into  deepening  invagination  (arrowhead)  of  homo- 
geneous portion.  3 1 .  Acrosome  of  late  spermatid  showing  developing  basal  invagination  of  acrosomal  vesicle  as  well  as  invagination 
of  homogeneous  portion  (arrowhead).  Asterisk  indicates  electron-lucent  plate  32-34.  Sequence  of  ridged  layer  (rl)  development 
shown  in  detail  Granule  cluster  (g).  .'Vsterisk  indicates  electron-lucent  plate.  35.  Nearly  mature  acrosome.  Electron-lucent  plate 
indicated  h\  asterisk.  Subacrosomal  material  (sm). 
Scale  bars:  25,26  =  0.5  nm.  27-35  =  0.25  ^m. 


Page  10 


THE  NAUTILUS,  Vol.  106,  No.  1 


be  an  aberrant  rather  tlian  normal  feature.  Examination 
of  testes  from  animals  collected  at  the  commencement 
of  the  reproductive  season  should  resolve  this  question. 

Aberrations  in  spermatogenesis:  Few  ultrastructural 
studies  have  dealt  w  itli  the  incidence  of  spermatogenic 
abnormalities  in  niollusks.  Takaichi  (1979)  detailed  ra- 
diation-induced malformations  of  the  mitochondrial 
sheath  and  nucleus  and  duplication  of  the  axoneme  in 
spermatids  of  the  pulmonale  Euhadra  hickonis.  Dorange 
and  Le  Peiinec  (1989)  noted  binuclear  spermatids  and 
angularly  dislocated  axonemes  in  late  spermatids  of  Pec- 
ten  maximus  (Linne,  1758)  and  regarded  these  features 
as  true  aberrancies.  O'Foighil  (1985)  suggested  that  an- 
gular dislocation  of  the  axoneme  in  testicular  sperm  of 
the  bivalve  Lasaea  suhviridU  Dall,  1899  could  be  due  to 
slight  immaturity.  In  Perotrochus  quoyanus  we  have 
observed  numerous  spermatocytes  and  spermatids  that 
were  undergoing  a  form  of  development  clearly  different 
from  normal  spermatogenesis.  Leaving  aside  the  phe- 
nomenon of  sperm  dimorphism  (a  well-documented  and 
'normal'  occurrence  in  many  Caenogastropoda — see 
Healy,  1988a  for  discussion),  the  irregular  shape  of  the 
condensed  nucleus  (pressed  into  shape  by  abutting  cells), 
and  the  apparent  inability  of  proacrosomal  vesicles  to 
fuse  into  a  single  acrosomal  vesicle,  strongly  suggest  that 
these  are  abnormal  cells.  Bearing  in  mind  that  a  certain 
background  level  of  spermatogenic  abnormality  proba- 
bly exists  in  many  if  not  most  animal  species  (Bryan  & 
Wolosewick,1973;  Baccetti  &  Afzelius,  1976),  we  believe 
the  appearance  of  aberrant  cells  in  P.  quoyanus  is  prob- 
ably a  normal  event  heralding  the  end  of  the  annual 
reproductive  phase  in  this  species.  We  base  this  view  on 
the  fact  that  all  three  males  examined  were  either  spent 
or  contained  principally  mature  spermatozoa  in  the  testis 
(with  isolated  pockets  of  developing  and  aljnormal  sper- 
matogenic stages). 

Spermatozoa:  Healy  (1988b)  has  previously  drawn  at- 
tention to  the  structural  similarities  between  spermatozoa 
of  Perotrochus  westralis  and  those  of  the  Trochoidea, 
particularly  Trochidae.  Our  observations  on  glutaral- 
dehyde-fixed  testis  sperm  of  P.  quoyanus  have  enabled 
us  not  only  to  confirm  these  similarities  but  also  to  expand 
on  details  of  the  Perotrochus  spermatozoon  as  recon- 
structed by  Healy  from  sea-water  formalin/ethanol-pre- 
served  material. 

The  electron-lucent  anterior  layer  of  the  Perotrochus 
acro-somal  vesicle  contains  regularly  spaced  ridges.  Sim- 
ilar ridges  have  elsewhere  been  observed  in  the  Trochi- 


dae [A!/.s7ro('Of/j/ra  roni^ric^a  (Lamarck,  1822),  Banhivia 
australis  (Menke,  1830);  Healy  &  Daddow  unpublished] 
and  in  the  liotiid  Liotina  peronii  (Kiener,  1839)  (Healy 
&  Ponder  unpublished).  It  is  interesting  to  note  that  the 
acrosomal  vesicle  of  other  pleurotomarioidean  (s./.)  fam- 
ilies (Haliotis  spp. — Haliotidae,  Lewis  et  al.,  1980;  Sakai 
et  al.,  1982;  Sinezona  sp. — Scissurellidae,  Healy,  1990a) 
lack  an  electron-lucent  anterior  layer,  whereas  in  the 
fissurellids  Scutiis  antipodes  Montfort,  1810  and  Mont- 
fortuhi  conoidea  Reeve,  1842,  a  layer  is  present  but 
exhibits  no  discernible  ridged  substructure  (Healy,  un- 
published). The  acrosomal  complex  in  Haliotis  and  in 
Sinezona  also  differs  from  that  of  Perotrochus  b\  having 
an  extensive  subacrosomal  deposit  similar  to  that  seen  in 
spermatozoa  of  some  fissurellids  (Scutus  antipodes, 
Montforttila  conoidea — see  Healy,  1990a  for  illustra- 
tions) and  many  bivalve  species  (see  references  in  Po- 
pham,  1979).  The  difference  in  appearance  of  subacro- 
somal material  between  Perotrochus  westralis  (rod-like) 
and  P.  quoyanus  (diffuse,  with  some  evidence  of  fibrous 
texture),  may  be  due  to  use  of  different  fixation  methods 
(P.  westralis — sea  water  formalin/ethanol;  P.  quoyan- 
us— glutaraldehyde  in  cacodylate  buffer).  Azevedo  et  al. 
(1985)  state  that  exposure  of  spermatozoa  of  Gibbula 
unihilicalis  to  sea  water  for  five  minutes  resulted  in  a 
clearly  defined  rod  (or  perforatorium),  deri\ed  from  a 
formerly  diffuse  subacrosomal  substance.  It  therefore 
seems  possible  that  the  subacrosomal  rod  of  P.  westralis 
may  also  be  an  end  product  of  prolonged  exposure  to 
sea  water.  The  dense  layer  of  material  visible  within  the 
subacrosomal  material  in  the  vicinity  of  the  nuclear  apex 
(see  figs.  7,  8)  may  also  be  involved  in  rod  formation. 
This  layer  was  observed  by  Healy  (1988b)  in  sea  water- 
formalin/ethanol  fixed  sperm  of  P.  westralis  and  inter- 
preted as  the  possible  remnants  of  nuclear  membranes. 
Our  observations,  based  on  glutaraldehyde-fixed  sperm 
ot  P.  quoyanus,  show  that  such  material  truly  lies  outside 
the  intact  nuclear  and  acrosomal  membranes,  and  there- 
fore constitutes  part  of  the  subacrosomal  material. 

The  close  resemblance  of  the  crypt-like  nuclear  fossa 
of  Perotrochus  spp.  (HeaK,  198Sb;  this  study)  to  the 
spermatid  fossa  of  Calliotropis  glyptus  (Healy,  1989)  has 
already  been  mentioned.  In  most  vetigastropods  and  the 
Patellogastropoda,  the  centrioles  are  only  superficially 
attached  to  a  shallow  nuclear  invagination.  In  Haliotis, 
the  proximal  centriole  itself  sometimes  occupies  the  shal- 
low fossa  (Lewis  et  al,  1980;  Sakai  et  al..  1982),  while 
in  Sinezona  (Scissurellidae)  and  Calliotropis  (Trochidae) 
the  centriole(s)  and  proximal  portion  of  the  a.xoneme  are 


Figures  36-42.  Perotrochus  quoyanus  '.ib.'.M.  IJcvt'lopiiig  iiucleu.s  (n),  iiiidpiece  niitoclidndria  (m).  axoneme  (ax),  centrioiar 
fossa  (arrowhead),  and  proxiiiiu!  (pel  and  (iLslai  (("iitrioies  ulc)  of  advanced  spermatids.  36  Inset.  Triplet  microtubules  of  centriole 
in  advanced  spermatid  38.  Morihund  spermatoc\te  sliowing  s\  iiaptinemal  complex  (arrows),  mitocliondria  (m).  proacrosomal 
vesicles  (pav),  and  axoneme  profiles  (arrowheads).  3'>.  .\herrant  spermatit!  Note  angular  shape  of  cell  and  its  condensing  nucleus 
(n),  as  well  as  the  presence  ot  proacrosomal  vesicles  and  milocliondria  (m)  M).  Mature'  aberrant  spermatozoon  wedged  between 
early,  prohabb  normal  spermatids.  Note  mitociiondria  (ni)  in  depressions  al  base  of  nucleus  (n)  Inset.  Detail  of  unfused  proacrosomal 
vesicles  (pav)  from  aberrant  spermatozoon.  41.  Fully  "condensed'  nucleus  (n)  of  aberrant  spcini  showing  irregular  shape  and 
multiple  axonemal  profiles  (ax).  42.  Nuclear  lacunae  (nl)  of  aberrant  spermatozoon 
Scale  bars:  36,37,40  Inset,4l,42  =  0.25  tim.  38-40  =  1  m"i 


J.  M.  Healy  and  M.  G.  Harasewych,  1992 


Page  11 


Page  12 


THE  NAUTILUS,  Vol    106,  No.  1 


actually  contained  within  the  fossa  (Healy,  1989,  1990a). 
The  ball-and-socket  fitting  of  rootlet  and  centriolar  fossa 
of  Perot rochtis  spp.  is  unusual  among  gastropods,  al- 
though a  similar  configuration  occurs  in  the  shipworm 
bivalve  Lyrodus  bipartita  (Jeffreys,  1860)  (see  Figure  4 
of  Popham,  1974).  Examination  of  other  genera  {Mi- 
kadotrochtts.  Entemnotrochus)  may  show  this  type  of 
nuclear  fossa  to  be  a  feature  of  all  Pleurotomariidae. 

Nuclear  lacunae  are  widely  reported  in  spermatozoa 
of  externally  fertilizing  mollusks,  polychaetes,  brachio- 
pods,  echinoderms,  as  well  as  of  some  internally  fertil- 
izing groups  (e.g.  some  teleosts,  Homo)  (Baccetti  &  Afzel- 
ius,  1976).  Their  occurrence  or  degree  of  development 
seems  to  be  more  closely  linked  with  the  mode  of  nuclear 
condensation  than  with  the  degree  of  modification  of 
nuclear  shape  occurring  during  spermiogenesis.  For  ex- 
ample, in  the  trochoid  Zaiipais  lascroni.  the  euspermatid 
nucleus  undergoes  marked  elongation  during  conden- 
sation (fibro-granular  pattern),  but  retains  lacunae  that 
ultimately  fuse  to  form  an  axial  tube  within  the  mature, 
filiform  nucleus  (Healy,  1990b).  Lacunae  are  usually  not 
observed  where  nuclear  condensation  proceeds  through 
either  or  both  longitudinal  fibrillar  and  lamellar  phases 
(see  Kaye,  1969;  Horstman,  1970,  Maxwell,  1983;  Koh- 
nert  &  Storch,  1984b;  Koike,  1985). 

The  midpiece  and  satellite  fiber/centriole  complex  of 
Perotrochus  spp.  are  essentially  as  observed  in  the  ma- 
jority of  Vetigastropoda  and  Patelloidea  (Koike,  1985; 
Hodgson  &  Bernard,  1988;  Healy,  1990a;  Healy  &  Dad- 
dow  unpublished),  the  Bivalvia  (for  references  see  Po- 
pham, 1979),  Scaphopoda  (Dufresne-Dube  et  al,  1983) 
and  Caudofoveata  (Buckland-Nicks  &  Chia,  1989).  The 
same  arrangement  of  these  organelles,  clearly  one  asso- 
ciated with  sperm  tail  attachment  and  stability,  also  oc- 
curs in  spermatozoa  of  many  other  externally  fertilizing 
animal  species  (for  major  references  see  Baccetti  &  Afzel- 
ius,  1976;  Wirth,  1984). 

The  flagellum  consists  of  an  axoneme  (9  +  2  microtu- 
bular  substructure)  sheathed  by  the  plasma  membrane. 
Our  scanning  electron  micrographs  reveal  that  the  fla- 
gellum is  narrower  in  diameter  close  to  the  nucleus.  TEM 
observations  suggest  that  this  is  probably  the  result  of  a 
more  closely  applied  plasma  membrane  in  this  region  of 
the  flagellum,  although  slight  narrowing  of  the  axoneme 
does  occur  near  the  distal  centriole  (see  figs.  4,  15).  At 
present  we  caruiot  clarify  the  origin  of  the  dense  body 
.sometimes  observed  within  the  flagellum  (see  fig.  17).  It 
was  not  observed  in  longitudinal  sections  through  the 
immediate  post-nuclear  region  of  the  flagellum  and  could 
yet  prove  to  be  an  artifact  of  fixation. 

Systematic  Considerations:  If  spermatozoa  of  Perotro- 
chus spp.  are  representative  of  the  Pleurotomariidae, 
then  a  closer  relationship  between  this  family  and  the 
Trochoidea  (particularly  Trochidae)  than  with  the  other 
pleurotomarioidean  {s.l.)  families  Haliotidae  and  Scis- 
surellidae  seems  evident.  This  conclusion  accords  both 
with  Ilas/.prunar's  (1988,  1989)  finding  that  no  synapo- 
morphies  exist  to  unite  the  Pleurotomarioidea  (s.l.),  and 
with  his  decision  to  place  the  Haliotidae  and  Scissurel- 


lidae  into  separate  superfamilies  within  the  Vetigastro- 
poda. The  question  as  to  whether  ancestral  vetigastropods 
were  more  like  scissurellids  than  pleurotomariids  (see 
Haszprunar,  1988,  1989  for  discussion)  cannot  yet  be 
resolved  using  sperm  data  alone  because  too  many  sig- 
nificant taxa  (including  the  new  hydrothermal  vent 
groups)  remain  unstudied.  Based  on  the  present  evi- 
dence, however,  we  suspect  that  spermatozoa  of  any  stem 
vetigastropod  would  have  resembled  more  closely  the 
unmodified  type  of  Perotrochus  (Healy,  1988b;  this  pa- 
per) than  the  modified  type  of  Sinezona  (Healy,  1990a). 
Vetigastropoda,  Patellogastropoda  and  Neritimorpha 
can  be  distinguished  on  the  basis  of  sperm  features  (es- 
pecially acrosomal  and  nuclear)  and  features  of  sper- 
miogenesis (dimorphic  in  the  case  of  the  Neritimorpha; 
rarely  so  in  the  Vetigastropoda)  (Koike,  1985;  Healy, 
1988a,  1990a, b).  It  will  be  interesting  to  determine 
whether  the  cocculinids — once  included  in  the  Vetigas- 
tropoda (Salvini-Plawen,  1980)  but  since  removed  to  a 
separate  archaeogastropod  suborder,  Cocculiniformia 
(Salvini-Plawen  &  Haszprunar,  1987) — also  show  char- 
acteristic sperm  and  spermiogenic  features. 

ACKNOWLEDGMENTS 

We  are  grateful  to  the  crews  of  the  Johnson-Sea- Link 
II  submersible  and  the  R/V  Seward  Johnson  for  their 
assistance  in  collecting  and  maintaining  the  specimens 
upon  which  this  study  is  based.  Thanks  are  are  extended 
to  Professor  G.  Grigg  of  the  Department  of  Zoology, 
University  of  Queensland  for  providing  access  to  TEM 
facilities  and  to  Mrs.  L.  Daddow  and  Mr.  T.  Gorringe 
(also  Department  of  Zoology)  for  assistance  with  TEM 
and  photography.  Financial  support  for  the  work  has 
been  provided  by  a  Queensland  Museum  Postdoctoral 
Research  Fellowship  (to  J.M.H.).  This  study  represents 
contribution  number  284  of  the  Smithsonian  Marine  Sta- 
tion at  Link  Port,  and  contribution  number  881  of  the 
Harbor  Branch  Oceanographic  Institution. 

LITERATURE  CITED 

Azevedo,  C,  A.  Lobo-Da-Cunha,  and  E.  Oliveira  198.5  Ul- 
trastructure  of  the  spermatozoon  in  Gibbula  umbilicalis 
(Gastropoda,  Prosobranchia),  with  special  reference  to  ac- 
rosomal formation.  Journal  of  Submicroscopic  Cytolology 
17;6()9-614. 

Baccetti.  B.  and  B.  .\,  Afzelius.  1976.  The  bioiog\  of  the 
sperm  ceif  Karger,  Basel   254  pp. 

Bernard,  R.  T.  F.  and  .\.  N  Hodgson  1985  The  fine  structure 
of  the  sperm  and  spermatid  differentiation  in  the  brown 
mussel  Perna  perna.  South  African  Journal  of  Zoology  20: 
5-9. 

Bouvier,  E.  L  and  H  Fischer,  1899.  Etude  monographique 
des  Pleurotomaires  actuels  Bulletin  of  the  Museum  of 
Coniparali\e  Zoolog),  Har\arcl  University  ;'52;193-249. 

Bouvier,  EL.  and  H.  Fischer.  1902.  L'organisation  et  les 
affinites  des  gasteropodes  primitifs  d  apres  I'etude  anato- 
mique  du  I'Icurotornaria  hcyriclii  Journal  de  Conchy- 
liologie  50(2):  117-272,  pis,  2-6. 

Buckland-Nicks,  J.  and  F.-S.  C:hia.     1989     Spermiogenesis  in 


J.  M.  Healy  and  M.  G.  Harasewych.  1992 


Page  13 


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THE  NAUTILUS  106(l):15-20,  1992 


Page  15 


Tenagodus  or  Siliqiiaria?  Unraveling  Taxonomic 
Confusion  in  Marine  "Worm-Snails" 
(Cerithioidea:  Siliquariidae) 


Riidiger  Bieler 

Department  of  Zoology 
Field  Museum  of  Natural  History 
Roosevelt  Road  at  Lake  Shore  Drive 
Chicago,  IL  60605,  L'SA 


ABSTRACT 

The  nomenclatural  history  and  availability  of  some  genus-  and 
family-group  names  in  the  marine  "worm-snair'  family  Sili- 
quariidae are  discussed:  (1)  Tenagodus  Guettard,  1770  (type 
species;  Serpula  anguina  Linne,  17.5iS)  has  priority  over  its 
objective  junior  synon\m  Siliqiiaria  Bruguiere,  1789.  Tena- 
godes.  Tenagoda  and  Silicaria  are  unjustified  emendations 
Siliqiiaria  Schumacher,  1817  (also  subsequently  emended  to 
Silicaria)  is  a  junior  homonym  described  in  the  Bivalvia.  (2) 
Siliquarius  Montfort.  1810  (type  species:  Siliqitarim  angiiiliis 
Montfort,  1810)  is  an  available  name.  (3)  Angiiinaria  Schu- 
macher, 1817,  is  preoccupied  by  Angninaria  Lamarck,  1816 
(Bryozoa).  (4)  Montfortia  Delia  Campana,  1890  (non  Mont- 
forlia  Recluz,  1843),  and  its  replacement  name  Hcmitcnago- 
diis  Rovereto,  1899,  are  based  on  the  type  species  Tenagodus 
bernardii  Morch,  1860.  (.5)  Agaihirscs  Montfort,  1808,  is  based, 
b\  original  designation,  on  (he  t\pe  species  .A^af/ilr,sc.s/iir('<'//i(.s 
Montfort,  1808  (a  senior  synon\ni  of  Siliqiiaria  spinosa  La- 
marck, 1818).  Agathyrsus  Herrmannsen,  1846,  is  an  unjustified 
emendation;  "Agathinus,"  "Agathirsis"  and  'Agathirsus  '  are 
incorrect  subsequent  spellings.  (6)  The  family  name  Siliquarii- 
dae Anton,  1838,  has  priority  over  its  objective  junior  synonym 
Tenagodidae  Gill,  1871 

Key  uords:  Ta\<iniini\ ,  Nomenclature.  Synonym) .  (laslrnpoda 


INTRODUCTION 

Marine  "worm-snails  '  are  now  understood  to  be  mem- 
bers of  three  different  families,  the  Vermetidae,  genus 
Vertincularia  in  the  Turritellidae,  and  Siliquariidae  (e.g., 
Morton,  1951,  1953;  Bieler,  1990)  The  siliqiiariids,  often 
(but  not  always)  recognizable  by  longitudinal  shell  slits 
or  series  of  holes,  are  an  enigmatic  group.  Little  is  known 
about  their  biology  and  geographical  distribution,  and 
the  published  literature  provides  seldom  more  than  de- 
scriptions of  empty  shells  or  shell  fragments.  Consider- 
able confusion  exists  about  the  synonymy  and  relation- 
ships of  nominal  taxa  such  as  Tenagodus  and  Siliqiiaria, 
or  Tenagodidae  and  Siliquariidae.  In  the  aftermath  of 
more  than  200  years  of  misspelling,  emending  and  re- 


placing certain  names,  today  s  worker  is  left  to  wonder 
about  how  many  "real  '  taxa  are  hidden  behind  Tena- 
godtis-Tenagodes-Tenagoda,  Siliqiiaria -Silicaria -Sili- 
quarius, or  Agathirses-Agathijrsiis-Agathinus.  This  is 
complicated  further  by  family  names  that  were  later 
based  on  some  of  these  names. 

This  paper  will  not  discuss  phylogenetic  relationships, 
a  topic  that  demands  detailed  anatomical  studies  that 
are  in  progress  (Bieler,  in  preparation).  No  interpretation 
of  subjective  synonymy  between  nominal  genera  in  the 
Siliquariidae  will  be  made  before  that  study  is  complete. 
However,  taxonomic  problems  often  can  be  solved  by  a 
careful  study  of  existing  literature  and  application  of 
ICZN  rules.  This  contribution  is  meant  as  a  taxonomic 
"house  cleaning,  '  addressing  problems  of  type  species, 
priority  and  availability  of  various  generic  and  family 
names  for  this  group.  A  main  goal  is  to  identify  objective 
synonymies  (and  consequently  reduce  the  number  of 
nominal  taxa  in  future  publications). 


TAXONOMY 

Genus-group  names 

Tenagodus  Guettard,  1770 

Type  species:    Serpula  anguina  Linne,  1758. 

The  name  Tenagodus  was  first  made  available  by 
Guettard  (1770:128).  Several  later  authors  {e.g.,  Herr- 
mannsen. 1849:540;  Delia  Campana,  1890:139;  Coss- 
mann,  1912:146)  refer  to  an  earlier  work  by  Guettard 
(usually  cited  as  "1760")  for  the  introduction  of  Tena- 
godus or  "Tenagoda."  However,  that  publication,  "An- 
nee  1760"  of  the  Histoire  de  T Academie  Royale  des 
Sciences  (1766a,  b),  contains  only  vernacular  names  in 
text  and  figure  captions.  Guettard  (1770)  did  not  indicate 
a  type  species.  The  first  subsequent  designation  of  a  type 
species  appears  to  be  that  of  Adams  and  Adams  (1854: 
360-361),  who  selected  "T[enagoda].  anguina,  Linnae- 
us" [Serpula  anguina  Linne,  175S]. 


Page  16 


THE  NAUTILUS,  Vol.  106,  No.  1 


Two  different  eiiu'iidatioiis  of  Tenagodus  (unjustified 
in  the  sense  of  ICZN  Art.  33)  were  introduced: 

Tenagodes  P.  Fischer,  1885  (p.  692);  Sacco,  1896:17;  Cossmann, 
1912:146. 

Tenagoda  Agassiz,  1848  (p.  1049);  Herrmannsen,  1849:540; 
Adams  &  Adams,  1854:360,  corrected  to  Tenagodus  in 
1858:656;  Chenu,  1859:321;  Paetel,  1869:9. 

Siliquaria  was  first  introduced  by  Bruguiere  (1789:xv). 
The  type  species  is  Serpula  angiiina  Linne,  1758,  by 
subsecjuent  monotypy  (Lamarck,  1799:79).  Bruguiere's 
Siliquaria  was  later  erroneously  credited  to  Lamarck  (e.g.. 
Schumacher,  1817:262;  Rang,  1829:187).  Silicaria,  an 
emendation  of  Siliquaria  Bruguiere  or  "Lamarck'  (un- 
justified in  the  sense  of  ICZN  Art.  33),  was  introduced 
and  used  by  some  authors  (e.g.,  Bosc,  1802:157;  Daudin, 
1800:32;  Audouin,  1829b). 

Siliquaria  was  also  described  as  a  genus  of  bivalves, 
by  Schumacher  (1817:43,  129).  Schumacher  knew  of  the 
conflict  with  the  earlier  name,  and  conserved  his  nominal 
taxon  by  dismissing  the  senior  homonym.  For  the  latter 
he  introduced  the  new  worm-snail  genus  Anguinaria  (see 
below):  "Je  n'ai  pu  conserver  le  nom  de  ce  genre  donne 
par  Mr.  Delamarck  savoir:  Silicaire  (Silicaria),  comme  il 
y  a  un  genre  parmi  les  bivalves  qui  porte  un  nom  presque 
semblable  "  (1817:262).  Siliquaria  Schumacher  was  also 
subsequently  emended  to  Silicaria  (e.g.,  Paetel,  1875: 
191). 


Discussion  of  the  type  species 

Linne's  (1758:787)  Serpula  anguina  was  originally 
based  on  more  than  one  species,  and  the  Linnean  col- 
lection contains  specimens  of  several  forms,  with  no  clear 
type  specimen  indicated  (Hanley,  1855:448).  Linne  had 
adopted  the  species  name  from  the  non-binominal  "Solen 
anguinus'  of  Rumphius  (1705:125)  and  referred  to  fig- 
ures in  that  work  (Rumphius,  1705:  pi.  41,  fig.  H)  and 
in  Argenville  (1742:  pi.  29,  fig.  H).  Rumphius'  figure 
shows  a  spiny  specimen  of  Tenagodus,  Argenville's  il- 
lustration is  that  of  a  member  of  the  Vermetidae  and 
was  later  used  by  Linne  (1767:1266)  as  reference  to  Ser- 
pula arenaria  Linne,  1758.  The  type  locality  for  S.  an- 
guina was  given  as  "India  "  (Linne,  1758:788).  Linne  also 
listed  several  references  to  a  variety  "/3  "  {e.g.,  Rumphius, 
1705:  pi.  41,  fig.  2;  Gualtieri,  1742:  pi.  10,  fig.  Z).  Most 
of  these  latter  illustrations  show  a  spineless  Mediterra- 
nean form  of  Tenagodus  later  described  as  Anguinaria 
ohtusa  Schumacher,  1817  (see  below). 

Born  (1780:440),  possibly  misled  by  Linne's  statement 
in  the  Museum  Ulricae  (1764:701)  that  the  variety  had 
a  spiny  shell,  then  confused  the  issue  by  applying  the 
name  Serpula  anguina  to  the  smooth  form  and  intro- 
ducing the  name  Serpula  rnuricata  for  the  true  anguina. 
Lamarck  (1801:98)  based  his  interpretation  of  S.  anguina 
on  yet  another  illustration  of  a  smooth  form,  by  Davila 
(1767:  pi.  4,  fig.  E),  later  described  ds  Siliquarius  anguilus 
Montfort,  1810  (see  below).  Manv  subsequent  autliors 
(e.g.,  Lamarck,   1818:337;   Philippi,   1836:173;  Chenu, 


1843:1;  Cossmann,  1912:146)  followed  and  sometimes 
defended  (Weinkauff,  1868:330)  Born's  misconception. 
Although  the  problem  was  discussed  at  several  occasions 
[Hanley,  1855:448;  M6rch,  1860b:403;  Tryon,  1886:189; 
Sowerby,  1876:(1)],  the  nsune  anguina  remained  mistak- 
enly in  use  also  for  the  smooth  Mediterranean  form, 
especially  in  paleontological  works  (e.g.,  Strausz,  1966: 
122). 

When  Lamarck  (1799:78)  and  Adams  &  Adams  (1854: 
361)  employed  the  name  anguina  in  fixing  the  type 
species  of  Siliquaria  and  Tenagodus,  respectively,  it  was 
in  this  misidentified  fashion.  Lamarck  ( 1799:78-79)  men- 
tioned a  shell  without  spines  for  Siliquaria,  and  his  later 
works  (e.g.,  1801,  1818)  clearly  identify  the  Mediterra- 
nean form.  Adams  and  Adams  (1854:361)  mentioned  that 
the  "typical  species  is  found  in  the  Mediterranean  '  and 
their  illustrations  in  the  same  work  (pi.  39,  fig.  5)  show 
smooth  Mediterranean  specimens. 

To  best  serve  stability  in  nomenclature,  and  in  agree- 
ment with  current  usage  (e.g..  Wenz,  1939:679;  Mala- 
testa,  1974:200;  Squires,  1990:286),  the  nominal  species 
named  in  fixation  are  here  accepted  as  type  species, 
regardless  of  misidentification.  A  referral  to  the  Inter- 
national Commission  on  Zoological  Nomenclature,  as 
specified  in  ICZN  Article  70(b),  seems  unnecessary  in 
this  case. 

The  synonym}  of  Tenagodus  and  Siliqiuiria  has  long 
been  known  (e.g..  Chenu,  1843: 1;  Herrmannsen,  1849: 
540;  Paetel,  1869:xiii,  56;  Keferstein,  1862-66:1056),  and 
the  priority  of  Tenagodus  was  stressed  by  some  authors 
(e.g..  Strausz,  1966:122).  Other  workers,  however,  re- 
jected Tenagodus  as  non-binominal  (e.g..  Daii,  1889:259; 
Gould,  1966:3;  Abbott,  1974:96),  and  both  names  have 
been  used  in  parallel  for  the  past  200  years. 

The  genus-group  name  Tenagodus  is  available  and 
has  priority  over  Siliquaria  (a  junior  objective  synonym), 
for  reasons  given  below: 

(1)  The  name  Tenagodus  was  used  as  a  scientific  name 
by  the  author  when  published  [ICZN  Art.  11(b)]. 
Guettard  s  description  (1770:128)  begins  "Genre  X. 
Tenagodus.  Tenagode  [the  French  vernacular].  Ca- 
ractere  generique.  ..." 

(2)  The  name  Tenagodus  was  introduced  in  a  work  pub- 
lished before  1931  and  containing  uninominal  genus- 
group  names  without  associated  nominal  species.  It 
can  be  accepted  "as  consistent  with  the  Principle  of 
Binominal  Nomenclature  in  the  absence  of  evidence 
to  the  contrary"  [ICZN  Art.  ll(c)(i)]. 

(3)  The  original  work  contains  a  description,  illustra- 
tions, and  bibliographic  references  to  additional  il- 
lustrations, thus  fulK  satisf\ing  ICZN  Art.  12  (a  new 
scientific  name  pui)lished  before  1931  must  have  been 
accompanied  by  a  description,  definition  or  indica- 
tion). 

(4)  The  work  has  not  been  placed  on  the  Official  Index 
of  Rejected  and  Invalid  Works  [ICZN  Art.  78(h)]. 

(5)  Other  genus-group  names  introtluced  in  the  same 
work  in  similar  fashion  are  in  current  use,  such  as 


R.  Bieler,  1992 


Page  17 


Kuphus,  Uperotiis  and  Brechitcs,  which  were  ac- 
cepted by  later  workers  (e.g..  Turner,  1966;  Smith, 
1971)  for  same  reasons.  A  suppression  of  Guettard  s 
work  by  plenary  power  of  ICZN  would  create  new 
taxonomic  problems. 

(6)  The  name  Tcnagodus  (in  various  spellings)  has  been 
used  frequenti\  {e.g.,  Morcii,  lS6()a,  1);  Fischer,  1885; 
Sacco,  1896;  Martin,  1899;  Simroth,  1906;  Clossmann, 
1912;  Thiele,  1929;  Wenz,  1939;  Strausz,  1966;  Mala- 
testa.  1974;  Boss,  1982).  It  does  not  qualify  as  an 
"unused  senior  synonym"  [ICZN  Art.  79(c)]. 

(7)  Siliqiiaria  Bruguiere,  1789,  and  Tcnagodus  Guet- 
tard, 1770,  have  (by  subsequent  designations)  the 
same  type  species;  they  are  objective  synonyms. 

Siliquarius  Monlfort,  1810 

Type  species:    Siliquarius  anguilus  Montfort,  1810. 

Siliquarius  was  introduced  by  Montfort  (1810:39).  The 
type  species,  by  original  designation,  was  given  as  Sili- 
quarius anguilus  (Montfort,  1810).  The  description  was 
accompanied  by  a  drawing  of  a  siliquariid  shell  without 
spines,  and  Montfort  made  reference  to  a  figure  by  Davi- 
la  (1767:  pi.  4,  fig.  E)  and  to  Siliquaria  anguina  sensu 
Lamarck,  1801  (who  had  based  his  concept  of  S.  anguina 
on  Davila's  figure). 

Siliquarius  was  used  by  some  later  authors  [e.g.,  Morch, 
1860b)  because  Siliquaria  was  considered  "preoccupied 
for  a  genus  of  plants  by  Forskal  (1775).  The  name  Sili- 
quarius Montfort  is  available  for  nomenclatural  purposes. 

Anguinaria  Schumacher,  1817 

[preoccupied] 

Schumacher  (1817:262)  introduced  Anguinaria  and  in- 
cluded two  new  species:  Anguinaria  obtusa.  referring  to 
"Serpula  anguinaria  [err.  pro  anguina]"  sensu  Born 
(1780:440,  pi.  18,  fig.  15),  and  Anguinaria  rubra,  referring 
to  Serpula  anguina,  sensu  Martini  (1769:50,  pi.  2,  figs. 
13-14).  No  subsequent  type  species  designation  was  lo- 
cated. 

The  name  Anguinaria  Schumacher,  1817,  is  not  avail- 
able for  nomenclatural  purposes,  because  it  is  preoccu- 
pied by  Anguinaria  Lamarck,  1816  (p.  142;  Bryozoa). 

Hemitenagodus  Rovereto,  1899 

Type  species:    Tenagodus  bernardii  Morch,  1860. 

Delia  Campana  (1890:139-140)  introduced  the  genus- 
group  name  Montfortia  as  a  subgenus  of  Tenagodus  (p. 
139)  or  new  genus  (p.  140).  The  name  Montfortia  was 
based,  by  original  designation,  on  Tenagodus  (Siliquari- 
us) bernardii  Mcirch,  1860  (1860a:368,  a  sponge-dwelling 
Recent  form  from  unknown  type  locality).  Later  workers 
(e.g.,  Wenz,  1939:680)  erroneously  considered  the  fossil 
species  Montfortia  ligustica  Delia  Campana,  1890,  as 
type  species  by  monotypy. 

Rovereto  (1899:108)  recognized  Montfortia  Delia 
Campana,  1890,  as  preoccupied  by  Montfortia  Recluz, 
1843,  and  introduced  the  replacement  name  Hemitena- 


godus. Thus,  Hemitenagodus  is  also  based  on  Tenagodus 
bernardii  Morch. 


Agathirses  Montfort,  1808 

Type  species:    Agathirses  furcellus  Montfort,  1808. 

Montfort  (1808:399)  introduced  Agathirses  with  A. 
furcellus  as  type  species  by  original  designation  ( 'Espece 
servant  de  type  au  genre")  for  an  Eocene  fossil.  The 
species  name  was  borrowed  from  "furcelle  (furcella),  ' 
listed  as  a  "singulier  tube  testacee  multioculaire  "  of  un- 
certain systematic  placement  by  Lamarck  (1801:104). 
Montfort  referred  to  tiie  'Siliquaire  de  Grignon "  as  il- 
lustrated by  Faujas-St.-Fond  (1803:  pi.  3).  Lamarck  (1818: 
338)  eventually  also  named  the  Grignon  fossil,  as  Sili- 
quaria spinosa.  Sherborn  (1930:6087)  erred  when  he 
credited  the  name  Siliquaria  spinosa  to  G.  Fischer  (1807: 
244),  who  had  only  used  the  French  vernacular  "Sili- 
quaire epineuse."  Later  authors  erroneously  cited  the 
junior  synonym,  Siliquaria  spinosa  Lamarck,  as  type  spe- 
cies of  Agathirses  (e.g.,  Herrmannsen,  1846:25;  Coss- 
mann,  1912:148;  Wenz,  1939:680). 

At  least  one  intentional  emendation  of  Agathirses  (un- 
justified in  the  sense  of  ICZN  Art.  33)  was  introduced: 

Agothijrsiis  Herrmannsen,  1846  (p.  25),  Agassiz  (1848:29);  Coss- 
mann,  1912:148, 

"Agathinus  Mft."  Paetel  (1888:499),  "Agathirsis  Mft." 
Paetel  (1875:5)  and  "Agathirsus  Cossmann,  1912"  Wenz 
(1939:680)  are  here  considered  incorrect  subsequent 
spellings  without  nomenclatural  bearing  [ICZN  Art.  33(c)]. 

Family-group  names 

Siliquariidae 

[based  on  Siliquaria  Bruguiere,  1789] 

Siliquariidae  was  first  introduced  by  Anton  (1838:xiii), 
as  "Siliquariacea."  The  name  was  frequently  credited  to 
later  authors,  such  as  Chenu,  1859;  it  was  also  indepen- 
dently proposed  by  several  later  workers  (e.g.,  Morton, 
1951:40)  unaware  of  its  earlier  introduction.  This  is  Sil- 
iquariadae  Schaufuss  in  Paetel  (1869:9)  and  "Siliquari- 
dae"  auct.  (e.g.,  Paetel,  1869:9). 

Tenagodidae 

[based  on  Tenagodus  Guettard,  1770] 

Tenagodidae  was  first  proposed  by  Gill  (1871:8).  It  was 
also  independently  introduced  by  later  authors  who  over- 
looked the  earlier  introduction  (e.g.,  Malatesta,  1974: 
200). 

Although  the  genus-group  name  Tenagodus  has  pri- 
ority over  Siliquaria,  the  family-group  name  Siliquari- 
idae has  priority  over  Tenagodidae: 

(1)  Both  generic  names  have  been  the  basis  of  family 
names. 

(2)  The  family-group  names  Siliquariidae  and  Tenago- 
didae are  objective  synonyms,  because  their  name- 


Page  18 


THE  NAUTILUS,  Vol.  106,  No.  1 


bearing  genera  are  objective  synonyms  [ICZN  Art. 
61(c)(ii)]. 
(3)  As  the  senior  s\  iionym,  Siliquariidae  can  continue  in 
use  altliough  Siliqiiaria  is  a  junior  synonym,  because 
the  Principle  of  Priority  also  applies  to  family-group 
names  concerned  [ICZN  Art.  40(a)(i)]. 


ACKNOWLEDGMENTS 

This  paper  required  access  to  some  rare  and  obscure 
works  and  would  not  have  been  possible  without  the 
excellent  library  facilities  at  the  Academy  of  Natural 
Sciences  of  Philadelphia  (ANSP),  the  Delaware  Museum 
of  Natural  History  (Wilmington),  the  Field  Museum  of 
Natural  History  (Chicago),  the  Museum  of  Comparative 
Zoology  (Harvard  University,  Cambridge  [MCZ]),  and 
the  National  Museum  of  Natural  History  (Smithsonian 
Institution,  Washington,  D.C.  [USNM]).  Special  thanks 
are  due  to  Dr.  Kenneth  J.  Boss,  Mrs.  Marion  Britz,  Dr. 
Silvard  Kool  (MCZ),  Dr.  R.  Tucker  Abbott  (Melbourne, 
Florida),  Dr.  Robert  Robertson  (ANSP),  Dr.  M.  G.  Har- 
asewych  (USNM),  and  Dr.  Alan  Kabat  (formerly  MCZ), 
who  helped  with  the  literature  search.  Dr.  James  F.  Quinn 
(Florida  Marine  Research  Institute,  St.  Petersburg)  ex- 
pertly commented  on  an  earlier  draft  of  the  Tenagodus- 
Siliquaria  discussion,  and  Mr.  Richard  E.  Petit  (North 
Myrtle  Beach)  kindly  reviewed  the  manuscript.  Two 
anonymous  reviewers  provided  valuable  critique  and 
suggestions. 


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THE  NAUTILUS  106(1  ):21-23,  1992 


Page  21 


Nucella  Roding,  1798  (Gastropoda:  Muricidae):  Type  Species 


Silvard  P.  Kool 
Kenneth  J.  Boss 

Moliusk  Ueparlmt-iit 
Museum  of  Comparative  Zoology- 
Harvard  I'liiversit) ,  Cambridge,  MA, 
0213.S 


ABSTRACT 

Bticciniim  filosum  Gmelin,  1791  [junior  synonym:  Nucella 
tlu'obronui  Roding,  1798;  senior  synonym  Biiccinurn  lapillus 
Linnaeus,  1758;  now  known  as  Succlla  lapillus  (Linnaeus)],  is 
the  type  species  of  Succlla  Roding  1798  In  subsequent  des- 
ignation of  Stewart  (1927:386,  footnote  #260), 

Key  words:  Nucella  lapillus.  type  species;  taxonomy;  Murici- 
dae, 


INTRODUCTION 

Roding  (1798:130-131)  introduced  the  genus  Nucella 
and  included  five  names:  iV,  reticulata,  N .  nioschatellina 
with  Buccinum  laeve  Gmelin,  1791  listed  as  a  synonym; 
N.  macina;  N.  lapillus  with  Buccinum  rusticurn  Gmelin, 
1791  as  a  synonym;  and  N.  theohroma  with  Buccinum 
filosum  Gmelin,  1791  as  a  synonym.  Roding  did  not 
specify  a  ty  pe  species.  Because  the  type  species  was  not 
fixed  in  the  original  publication,  only  the  subsequent 
designation  of  a  type  species  from  the  nomina  enumer- 
ated above  is  valid  in  accordance  with  Article  69  (Code 
of  the  International  Commission  on  Zoological  Nomen- 
clature [hereafter  I.C.Z.N.]  (1985)).  Several  authors  have 
either  incorrectly  designated  a  type  species  for  Nucella 
or  discussed  efforts  to  designate  a  type.  This  confusion 
was  partly  caused  by  the  fact  that  the  binomen  Nucella 
lapillus  R(")ding  is  a  secondary  homonym,  but  not  a  syn- 
onym, of  Buccinum  lapillus  Linnaeus,  1758. 

TAXONOMIC  HISTORY:  A  COMEDY  OF  ERRORS 

Dall  (1906;  1909),  Clench  (1947),  and,  in  more  detail, 
Rehder  (1962)  discussed  the  species  introduced  by  Rod- 
ing (1798)  in  Nucella.  The  first  species,  N.  reticulata,  is 
the  cancellariid  Cancellaria  reticulata  (Linnaeus,  1767); 
the  second,  N.  moschatellina  is  a  nomen  dubium;  the 
third,  N.  macina  is  a  nomen  nudum,  the  fourth,  N. 
lapillus,  not  to  be  confused  with  and  not  the  same  as 
Buccinum  lapillus  Linnaeus,  1758,  had  in  its  synonymy 
Buccinum  rusticum  Gmelin,  1791  and  is  identified  as 
Latirolagena  smaragdula  (Linnaeus,  1758);  the  fifth  spe- 
cies, Nucella  theobroma,  with  Buccinum  filosum  Gme- 


lin, 1791  listed  as  a  synonym,  is  a  subjective  junior  syn- 
onym of  Buccinum  lapillus  Linnaeus,  1758,  now  referred 
to  as  Nucella  lapillus  (Linnaeus,  1758)  (Kool,  1989). 

Dall  (1906),  after  prompting  acceptance  of  Roding's 
names,  acknowledged  that  tletermining  the  correct  type 
species  for  Nucella  would  "recjuire  special  study  to  un- 
ravel." In  a  later  work,  Dall  (1909)  provided  not  one, 
but  three  different  binomina  for  the  type  species  des- 
ignation, without  mentioning  the  word  "type.  " 

First,  in  his  list  of  subgenera  within  the  genus  Thais. 
he  (1909:46-47)  cited  "Purpura  filosa  Menke  =  P.  la- 
pillus L."  after  Nucella.  Wenz  (1941:1123)  mistakenly 
interpreted  this  as  a  species  designation  and  credited  Dall 
(1909)  with  the  designation  of  "N.  filosa  (Gmelin)  [Buc- 
cinum] =  lapillus  (Linne)  [Buccinun^]."  Whatever  the 
inappropriateness  of  this  invalid  designation,  Dall  left  no 
doubt  that  he  recognized  these  specific  nomina  as  syn- 
onyms by  placing  "P.  lapillus  L."  in  synonymy  with 
Purpura  filosa  "Menke  "  [=  Buccinum  filosum  Gmelin]. 
It  should  be  mentioned  that  Dall  s  attribution  of  filosa 
to  Menke  (1830:62)  was  an  erroneous  concession  to  the 
custom  of  the  nineteenth  century  where  some  authors 
credited  themselves  for  a  specific  name  after  having  al- 
tered the  generic  placement  and,  thus,  the  binominal 
combination,  in  this  case,  from  Buccinum  filosum  to 
Purpura  filosa. 

Secondly,  in  the  same  work,  Dall  (1909:48)  stated  ".  .  . 
thus  fixing  the  name  Nucella  on  tlie  well-known  Purpura 

lapillus by  which  action  he  obviously  meant  to 

restrict,  by  subsequent  designation,  the  concept  of  Nu- 
cella, but  he  again  did  not  mention  the  necessary  word 
"ty  pe  " 

Thirdly ,  in  another  listing  of  subgeneric  and  sectional 
taxa,  Dall  (1909:50)  designated  many  type  species,  and 
he  placed  "T.  lapillus  (Linn)  '  right  after  Nucella  s.s., 
probably  meaning  this  to  be  a  type  designation.  But 
although  Nucella  is  on  that  list,  he  again  failed  to  use 
the  word  "ty  pe.  Therefore,  v\  hatever  his  intentions,  Dall 
(1909)  provided  three  different,  ambiguous,  indefinite 
and  invalid  subsequent  designations. 

Later  works  that  treated  Nucella  include  Suter  (1913: 
425),  who  listed  "Purpura  lapillus  L.  '  as  type  species  of 
Nucella,  an  invalid  designation,  because  lapillus  Lin- 


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THE  NAUTILUS,  Vol.  106,  No.  1 


naeus  was  not  included  among  the  species  of  Nucella  in 
Rfiding  (1798),  and  i.s  therefore  not  available  (I. CI. Z.N. , 
Article  (59). 

Iredale  (1915:472),  without  realizing  that  Dall  had  not 
designated  a  valid  type  species  for  Nucella,  reprinted 
Dall's  (1909:50)  list  and  added  the  word  "Type,"  but 
misquoted  Dall  by  listing  "T  lapillus  Lam."  instead  of 
"T.  lapillus  (Linne)." 

Dall  (1915:557)  discussed  the  type  species  designation 
oi  Nucella  and  stated:  "The  type  of  the  subgenus  is  the 
solitary  North  Atlantic  species  of  N.  lapillus  of  Linnaeus, 
..."  However,  although  Dall  mentioned  the  word 
"type,  lapillus  Linnaeus  is  not  available  because  it  was 
not  a  name  included  under  Nucella  by  Roding  (1798). 

Stewart  (1927:386)  wrote  in  a  footnote  (#260):  "[Nu- 
cella:] Type  species  Buccinunt  filusuni  Gmelin  =  B.  la- 
pillus h."  Stewart  was  the  first  author  to  designate  a  valid 
type  species  for  Nucella,  without  realizing  this  himself; 
he  continued:  "It  is  probably  safe  to  say  that  Dall  fixed 
the  t\pe  in  1909.  The  species  Buccinuni  filosum,  orig- 
inally used  by  Roding  [as  a  synonym  ot  Nucella  theo- 
broma  Roding]  in  Nucella,  is  nomenclatorially  available 
for  the  type  designation.  Stewart  also  recognized  that  B. 
filosum  was  synonymous  with  B.  lapillus  Linnaeus. 

Treating  Nucella  as  a  separate  genus  with  four  sec- 
tions, Thiele  (1929:298)  mentioned  Nucella  (Nucella) 
lapillus  (Linne)  as  an  example;  he  did  not,  however,  use 
the  word  "type,"  and  none  of  the  available  nomina  was 
cited. 

Grant  and  (;ale  (1931:716),  following  Stewart  (1927), 
also  credited  Dall  (1909)  as  the  subsequent  designator 
and  gave  the  type  as  "Nucella  lapillus  Linnaeus."  How- 
ever, aside  from  not  being  available,  this  binomen  was 
not  one  of  the  three  combinations  cited  by  Dall  as  possible 
types. 

In  his  list  of  the  type  species  of  marine  mollusca  of 
the  British  Isles,  Winckworth  (1932:229)  listed  N.  theo- 
lyronia  Roding,  1798  as  the  "genotype"  of  Nucella,  how- 
ever, Stewart  s  (1927)  type  designation  preceded  this  one. 

Winckworth  (1945:141),  as  he  had  done  in  his  1932 
paper,  again  cited  N.  theobroma  Roding  as  type  species 
of  Nucella  and  correctly  noted  that  it  was  a  junior  syn- 
onym of  Buccinuiu  lapillus  Linnaeus.  However,  he  er- 
roneously attributed  the  designation  to  Dall  (1909),  who 
never  used  the  name  theubroma. 

Clench  (1947:86)  in  an  ambiguous  treatment  of  lapil- 
lus Linnaeus,  placed  the  species  in  the  geiuis  Thais  (Riid- 
ing,  1798)  and  used  the  subgeneric  name  Polytropa 
Swainson,  1840  [type  species  Buccinuni  lapillus  Linnaeus 
by  subsequent  designation,  j.E.  Gray,  1847],  rather  than 
Nucella.  (bench's  argument  for  not  using  Nucella  was 
that  Roding  never  intended  to  include  Buccinuni  lapillus 
Linnaeus  in  his  genus  Nucella.  Glench  arrived  at  this 
conclusion  after  comparing  the  hgures  in  Martini  (1777: 
pi.  121,  figs.  1 1 1 1-11  12)  tor  lapillus  Limiaeus  and  those 
ioT  filosum  Gmelin  (pi.  121,  figs.  1 113-1 1 14).  According 
to  Clench,  the  figures  for  Riiding's  Nucella  theubroma 
and  its  synonym  Buccinum  filosum  Gmelin,  are  unrec- 
ognizable, whereas  the  figures  of  i^(")ding's  Nassa  rudis 


and  its  associated  synonym,  "Buccinum  lapillus  Gme- 
lin," are  those  of  Buccinum  lapillus  Linnaeus  The  latter 
portion  ot  this  argument  is  true,  but  the  former,  as  Reh- 
der  (1962)  pointed  out,  is  not  tenable:  the  "unrecogniz- 
able" figures  oi  Nucella  theobroma  in  Martini  (1777:  pi. 
121,  figs.  1113-1114)  also  clearly  represent  Nucella  la- 
pillus Linnaeus,  in  one  of  its  many  variations.  Rehder 
(1962,  figs.  1-2)  illustrated  a  specimen  oi  lapillus  from 
France  that  closely  matched  Martinis  (1777:  pi.  121,  figs. 
1113-11 14)  illustrations.  It  has  long  been  known  that  the 
species  displays  a  high  degree  of  variation  in  shell  mor- 
phologv  and  color  (Martini,  1780:  pi.  122,  figs.  1124- 
1 125,  1128-1 129;  Forbes  &  Hanley,  1851;  Crothers,  1985). 
In  his  discussion  of  Nucella,  Clench  did  not  mention  the 
attempted  designations  of  Dall  (1915),  Suter  (1913), 
\Ninckworth  (1932),  and  the  valid  designation  of  Stewart 
(1927). 

Clench's  usage  of  Polytropa  instead  of  Nucella  is  not 
followed  herein.  Nucella  is  a  valid  taxon  to  be  used  for 
Linnaeus  lapillus.  Interestingly  enough.  Clench  cited 
"Purpura  filosa  Menke"  and  "Buccinun}  filosum  Gme- 
lin" in  his  synonymy  of  "Thais  {Polytropa)  lapillus 
Linne,"  which  contradicts  his  own  argument  that  Roding 
never  intended  to  include  Buccinum  lapillus  Linnaeus 
in  his  genus  Nucella. 

Finally,  in  an  attempt  to  resolve  some  of  the  irregu- 
larities and  inconsistencies  discussed  so  far,  Rehder  (1962: 
110)  stated  that  Dall  (1909:50)  intended  to  designate  a 
type  and  that  another  possible  attempt  at  a  designation 
(Dall,  1909:46)  is  invalid  because  Dall  cited  fi/o.sa  Menke, 
no\.  filosa  Gmelin.  Further,  Rehder  attributed  the  earliest 
type  designation  to  Winckworth  (1932:229)  who  listed 
Nucella  theobroma  and  pointed  out  that  Clench  (1947) 
had  overlooked  this  type  designation. 

SUMMARY 

In  conclusion,  the  correct  tv  pe  species  for  Nucella  Rod- 
ing, 1798  is  Buccinum  filosum  Gmelin,  1791,  a  subjective 
junior  synonym  of  Buccinum  lapillus  Linnaeus,  1758, 
and  an  objective  senior  synonym  of  Nucella  theobroma 
Roding,  1798,  by  subsequent  designation  of  Stewart  (1927: 
386,  footnote  #260). 

ACKNOWLEDGMENTS 

The  authors  thank  Mr.  Richartl  I  Johnson  for  a  critical 
re\  iew  of  an  earlier  draft  of  this  paper,  and  are  indebted 
to  two  anoiiN  nious  reviewers. 

LITERATURE  CITED 

clench,  W.J  1947.  Tlic  geiu'ra  Purpura  and  Thais  in  the 
western  Atlantic,  Joliii.soiiia  2(23):61-91,  pis.  32-40. 

(Mothers,  J.H.  1985  Dog-whelks:  an  iiitrodiictioii  to  the  bi- 
ology of  Nucella  lapillus.    Field  Studies  6:291-360 

D.ill,  VV.n.  1906.  Early  history  of  the  generic  name  Fiisus. 
Journal  of  Coiichology  1 1(  l'o):289-297. 

Dall.  W.H.  1909.  Contrihtitions  to  the  Tertiar\  paleontology 
(il  till-  Pacilic  coast.  1.  The  Miocene  of  .Astoria  and  Coos 


S.  p.  Kool  and  K.  J.  Boss,  1992 


Page  23 


Bay,  Oregon  United  States  Geological  Survey  Professional 
Paper  59,  278  pp.,  23  pis,  14  figs. 

Dall,  \V  H.  1915.  Notes  on  the  species  of  the  niollnscaii  sub- 
genus Mticclhi  inhabiting  the  northwest  coast  of  America 
and  adjacent  regions.  Proceedings  of  the  L'niteii  States 
National  Museum  49:557-572,  pis.  74-75. 

Forbes,  E.  and  S.  Hanle>.  1851.  A  histor\  of  British  Mollusca. 
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ternoster Row.  [3]  vi-\,  [1]  pp.  2-616. 

(wnelin,  J  F  1791  Caroli  a  Linne,  Svstema  Nat\irae  per  Reg- 
na  Tria  Naturae  .  Editio  Decima  Tertia,  Aucta,  Refor- 
mata.  Tomus  1,  Pars  6.  Beer,  Leipzig.,  pp.  3021-3910  [for 
a  full  citation  and  discussion  of  the  tv\'0  printings  of  this 
work  see  Kabat  and  Petit  (1988)]. 

Grant,  I'S.,  IV  and  H.R.  Gale.  1931.  Catalogue  of  the  marine 
Pliocene  and  Pleistocene  Mollusca  of  Galifornia  and  ad- 
jacent regions.  Memoirs  of  the  San  Diego  Society  of  Nat- 
ural History  1:1-1036,  15  figs.,  32  pis. 

Grav,  J.E.  1847  A  list  of  the  genera  of  Recent  Mollusca,  their 
sv  nonvma  and  tv  pes  Proceedings  of  the  Zoological  Societv 
of  London  15:129-219 

International  ("onnnission  on  Zoological  Nomenclature.  1985 
International  Code  of  Zoological  Nomenclature  Third 
Edition,  i-xx  -t-  338  pp.  International  Trust  for  Zoological 
Nomenclature/British  Museum  (Natural  Historv),  Lon- 
don; LIniversity  of  California  Press,  Berkclev  anil  Los  .An- 
geles. 

Iredale,  T  1915  Art  XLVTI — A  commentary  on  Suter  s 
"Manual  of  the  New  Zealand  Mollusca.  Transactions  and 
Proceedings  of  the  New  Zealand  Institute  for  the  year 
1914,  47:417-497. 

Kabat.  A.R.  and  RE.  Petit.  1988  The  two  printings  of  J  F. 
('•mvUu's.  Syslcma  Naturae,  13th  Edition  (1788-96).  The 
Nautilus  102(4)  164-166. 

Kool,  S.P.  1989.  Phylogenetic  analysis  of  the  subfamily  Thai- 
dinae  (Prosobranchia:  Neogastropoda;  Muricidae).  Ph.D. 
Dissertation,  The  George  Washington  L'niversity,  Wash- 
ington,!) C,  i-xiii  -I-  342  pp. 


Liimaeus,  C.    1758.    Caroli  Linnaei  .  .     Systema  Naturae  per 

Regna  tria  Naturae,  secundum  classes,  ordines,  genera, 

species,  cum  characteribus,  differentiis,  synonymis,  locis 
Editio   decima    reformata     Impensis   Direct     Tom 

l.Animalia.  Salvii:  Holmiae,  824  pp 
Martini,   F  H.W.      1777.     Neues  systematisches  Conchylien- 

('abinet.  Volume  3.  Raspe,  Niiremberg,  i-vi  +  434  pp  , 

pis.  Ixvi-cxxi. 
Martini,   F.H.W.     1780.     Neues  systematisches  Conchylieii- 

Cabinet.  Volume  4,  Raspe,  Nuremberg,  [24]  -I-  344  pp., 

pis.  cxxii-clix 
Menke,  K.T.     1830.    Synopsis  methodica  inolluscorum  gener- 

um  omnium  et  specierum  earum Uslar,  Pyrmonti, 

i-xvi  -I-  169  pp. 
Rehder,  H.  A.    1962.    The  status  of  Nucclla  Roeding.  The  Nau- 
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Roding,  P.F.     1798.    Museum  Boltenianum       .  :  pars  secunda 

continens  Conchylia     .     Hamburg  (Trapii),  i-viii  -I-   199 

pp 
Stewart,  R.B.     1927.    Gabb's  (California  fossil  type  gastropods 

Proceedings  of  the  Academy  of  Natural  Sciences  of  Phil- 
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teste  [Table  of]  Contents). 
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an  atlas  of  Quarto  Plates.  Mackay,  Government  Printer, 

Wellington,  N.  Z.,  i-xxiii  4-1120  pp. 
Swainson,  W.     1840.    A  treatise  on  malacologv,  or  shells  and 

shell-fish.  London.  419  pp. 
Thiele,  J      1929.     Handbuch  der  Systematischen  Weichtier- 

kunde.  Volume  1.  F'ischer,  Jena,  376  pp. 
Wenz.  W,    1941.    Prosoliranchia.  [in]  OH.  Sehindewolf  (ed). 

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5):  136-148. 


THE  NAITILUS  106(1  ):24-.38,  1992 


Page  24 


A  Revision  of  the  Recent  Species  of  Eudolium  Dall,  1889 
(Gastropoda:  Tonnoidea) 


Bruce  A.  Marshall 

National  Museum  of  New  Zealand 
P.O.  Box  4(i7,  Wellington,  New 
Zealand 


ABSTRACT 

The  tonnid  genus  Eudolium  contains  three  Recent  species:  E. 
crosseanum  Monterosato,  1869  (=  thornpsoni  McGinty,  1955), 
£.  hairdii  X'errill  &  Smith  (=  solidior  Dautzenberg  &  Fischer, 
1906  =  lineata  Schepnian,  1909  =  inflatum  Kuroda  &  Habe, 
1952  =  kuroharai  .Azuma,  1960)  and  £.  pyriformc  Sowerbv, 
1914.  Eudolium  crosseanum  and  E.  pyrijorme  respectively 
have  amphiatlantic-Mediterranean  and  Indo-Pacific  distribu- 
tions, while  £.  hairdii  occurs  widely  in  the  Atlantic,  Pacific  and 
Indian  Oceans.  Genus  and  species  group  synonymies  and  dis- 
tributions are  discussed,  and  shells,  radulae  and  male  external 
anatomies  are  illustrated. 

Key  words:  Mollusca;  prosobranch  gastropods;  Tonnidae;  Eu- 
dolium. systematics;  zoogeography 


INTRODUCTION 

The  taxonomy  of  the  species  of  the  prosobranch  family 
Tonnidae  is  poorly  imderstood.  Atlantic  species  were  re- 
vised by  Turner  (1948),  but  there  has  been  no  critical 
modern  revision  of  the  Indo-Pacific  species.  The  present 
contribution  is  the  resuh  of  an  attempt  to  determine  the 
identity  of  two  species  of  the  genus  Eudolium  Dall,  1889 
obtained  off  New  Zealand  in  recent  years. 

Since  Dail's  (1889a)  review,  Eudolium  bairdii  (Verrill 
&  Smith,  1881)  has  been  treated  as  a  synonym  of  E. 
crosseanum  (Monterosato,  1869)  by  the  majority  of  au- 
thors. From  examination  of  numerous  specimens,  how- 
ever, it  transpires  that  not  only  are  they  distinct  species, 
but  that  five  subsequently  introduced  taxa  are  synonyms. 
Moreover,  £.  crosseanum  is  more  closely  related  to  the 
Indo-Pacific  £.  pyriforrne  (Sowerby,  1914)  than  to  £. 
bairdii. 


ABBREVIATIONS  AND  TEXT  CONVENTIONS 

AMS — Australian  Mihseum,  Sydney. 
MCZ — Museum  of  (Comparative  Zoology,  Harvard. 
MNHN — Museum  National  d'Hi.stoire  Pslaturelle,  Pari.s. 
NMNZ — National  Museum  of  New  Zealand,  Wellington. 
NMP — Natal  Museum,  Pietermaritzburg. 
USNM — National  Museum  of  Natural  History,  Wash- 
ington, DC. 


In  captions  to  illustrations  shell  height  dimension  pre- 
cedes diameter. 


SYSTEMATICS 

Class  Gastropoda 

Superfamily  Tonnoidea  Suter,  1913 
Family  Tonnidae  Suter,  1913 
Genus  Eudolium  Dall,  1889 

Doliopsis  Monterosato,  1872(not  Vogt,  1852,  nor  Conrad,  1865): 
8.  Type  species  (by  monotypy):  Dolium  crosseanum  Mon- 
terosato, 1869;  Recent,  Mediterranean 

Eudolium  Dall,  1889a:232.  Substitute  name  for  Doliopsis  Mon- 
terosato (preoccupied). 

Galeodolium  Sacco,  1891:4.  Type  species  (by  subsequent  des- 
ignation of  Voices,  1986:178):  Cassidaria  mutica  Michelot- 
ti.  1861;  Oligocene,  Italy 

■'Tuberculodolium  Sacco,  1891:9.  Type  species  (by  subsequent 
designation  of  Vokes,  1986:178):  Eudolium  antiquum  Sac- 
co, 1890;  Oligocene,  Italy 

?Simplicodolium  Sacco,  1891:13.  Type  species  (here  designat- 
ed): Pyrula  fasciata  Borson,  1821;  Pliocene,  Italy. 

Remarks:  The  shells  of  Eudolium  species  differ  from 
those  of  Tonna  Brunich,  1771  in  having  more  narrowly 
tapered  bases,  consistently  narrow,  widely  spaced  spiral 
cords  with  narrow  secondary  spirals,  and  fine  axial  sculp- 
ture on  all  teleoconch  whorls.  The  two  groups  are  cer- 
tainly closely  related,  with  similar  shells,  radulae,  jaws 
and  external  anatomies.  An  operculum  is  lacking  in  both 
groups,  at  least  in  the  adults.  Eudolium  species  are  most 
commonly  encountered  at  200-600  m  depth,  while  few 
Tonna  species  range  much  deeper  than  50  m.  Although 
Eudolium  has  been  placed  as  a  subgenus  of  Tonna  by 
some  authors  {e.g..  Dall,  1889a, b;  Vredenburg,  1919; 
Thiele,  1929;  Kilias,  1962),  1  prefer  to  treat  it  at  generic 
level  because  of  the  distinctive  shell  facies  and  deeper 
center  of  bathymetric  distribution. 

As  here  interpreted  the  genus  Eudolium  contains  three 
Recent  species:  E.  crosseanum  (Monterosato,  1869),  £. 
bairdii  (Verrill  &  Smith,  1881),  and  £.  pyriforrne  (Sow- 
erby,  1914).  Of  other  species  that  have  been  referred 


B   A    Marshall.  1992 


Paae  25 


here,  Dolium  [Eudolium)  verrilli  Dall,  1889  has  been 
transferred  to  Hadroocorys  Quinn,  19S0  iQuinn.  1980). 
while  £.  aulacodes  Tomlin.  1927  belongs  in  Oocorys 
Fischer.  1883  (Kilbum.  1986).  Parvitonna  perselecta  Ire- 
dale,  1931  was  referred  to  Eudolium  by  Kilias  (1962). 
but  the  holotype  (.\MS  C. 57790)  is  clearK  a  species  of 
Tonna  (sensu  stncto).  Tonna  tessellatum  (Bruguiere. 
1789).  T.  fasciatum  (Bruguiere.  1789)  and  T.  zonatum 
(Green.  1830)  were  referred  to  Tonna  {Eudolium )  by 
Vredenburg  (1919).  but  the>  too  belong  in  Tonna  {sensu 
stricto). 

\okes  (1986'  interpreted  Cossmann's  1 190-'3:lo9!  state- 
ment "types  de  Galeodolium  et  de  Tuberculodolium:  E. 
antiquum  Sacco,  Cassid.  mutica  Michelotti.  d'apres  les 
types  communiques  par  M.  Sacco"  as  the  first  valid  sub- 
sequent designation  of  t\pe  species  for  Galeodolium  and 
Tuberculodolium.  Cossmann.  however.  inad\ertentK  re- 
versed the  order  of  the  ""t\p>e  sp>ecies  and  thus  selected 
species  that  were  not  originalK  included  (ICZN  .\rt.  69a). 
Although  N'okes  simpK  corrected  the  order.  Cossmann 
must  be  considered  to  have  associated  the  names  re- 
speciicely,  so  it  is  concluded  that  Cossmann's  selection 
is  invalid.  .AccordingK.  X'okes  vl986)  is  considered  to  be 
the  first  subsequent  designator  of  the  typ>e  species  of  these 
ta.xa. 

N'okes  (1986)  used  Galeodolium  as  a  subgenus  of  Eu- 
dolium for  species  with  strong  nodules  and  a  refle.xed. 
denticulate  outer  lip.  The  differences  between  the  t\pe 
si>eciesof  Eudolium  and  Galeodolium  Michelotti.  1S61: 
pi.  13.  fig.  16:  Sacco.  1S91:  pi.  1.  fig.  li.  however,  are 
simpK  matters  of  degree,  because  E.  bairdii  exhibits  a 
smooth  morphological  transition  bet^veen  the  extremes 
in  its  infraspecific  \ariabilit\  (see  below).  Placement  of 
Tuberculodolium  and  Simplicodolium  in  synon\mv  is 
tentative  pending  study  of  the  typ>e  material,  which  was 
not  available  to  me.  Judging  from  published  photo- 
graphs, the  undoubtedK  juvenile  holotype  of  Eudolium 
antiquum  Sacco.  1S90  .Ferrero  Mortara  et  al..  1984) 
seems  narrower  than  confirmed  species  of  Eudolium, 
while  the  lectotype  of  Pyrula  fasciata  Borson.  1821  (Pa- 
via,  1976)  may  be  a  juvenile  Tonna.  Because  Galeodoli- 
um and  Eudolium  are  regarded  as  synonyms,  and  as 
Eudolium  and  Tonna  are  undoubtedK  confamilial.  Ga- 


leodoliidae  Sacco,  1891  is  treated  as  a  synonym  of  Ton- 
nidae  Suter.  1913  (1825)  (Tonnidae  is  conserved  using 
ICZX  Art.  40b). 

Eudolium  sjjecies  occur  in  temp)erate  and  tropical  seas 
on  soft  substrata  on  continental  shelves  and  slopes  at  17- 
823  m  depth.  Undoubted  Eudolium  species  are  known 
from  the  EarK  Miocene  [e.g.,  E.  aoteanum  Beu.  1970 
and  another,  unnamed  species  from  New  Zealand;  £. 
biornatum  (Tate,  1894)  from  .Australia]  and  the  Oligo- 
cene  {e.g.,  Cassidaria  mutica  Michelotti,  1861).  .As  with 
Tonna  species  (Morton.  1991 1  the  diet  consists  of  holo- 
thurians:  the  gut  of  a  specimen  of  E.  bairdii  from  off 
Ma\or  Island.  New  Zealand,  contained  ossicles  of  the 
apodid  holothurian  Protankyra  rigida  Pawson,  1965  (det. 
D.  L.  Pawson)  Otherwise  nothing  is  known  of  Eudolium 
biolog). 

Eudolium  crosseanum  :  Monterosato.  1869) 

(figures  1-4.  9.  20.  23-26.  37 1 

Dolium  crosseanum  Monterosato,  1869:22S.  pL  12.  fig.  1:  Trvon. 

1885:263.  pi.  2.  fig.  11  (figs.  11  and  12  transptKed.  fig.  12 

=  D.  testardi  Montrouzier.  1863). 
Doliopsis  crosseana. — Monterosato.  1872:8;  Coen.   1930:147. 

figs.  1,  2  [in  part — figs.  6-9  =  Galeodea  echinophora  vLin- 

naeus,  1767),  P.  Bouchet.  personal  communication]. 
Dolium    Eudolium^  croaseanum. — Kobelt.  1908:155.  pi.  126. 

fig.  1  I  in  part — figs.  2.  3.  4  =  £.  bairdii'. 
Eudolium  crosseanum. — Wenz.  1941:1076.  fig.  3066:  Turner. 

1948:178  un  part— pi.  81.  figs.  1.  2.  te.xt  Hg.  5  =  £.  bairdii). 
Eudolium  testardi. — Osima,  1943:132,  pi.  4,  fig.  3  (not  Dolium 

testardi  Montrouzier.  1863). 
Eudolium  thompsoni  McGintv.  1955:80.  pi.  1.  figs.  5, 6;  .\bbott. 

1974:168,  fig.  178S:  Abbott  &  Dance.  1986:119.  text  fig 

(new  synonym). 
Tonna  {Eudolium)  cros.seana. — Kilias.  1962:14,  fig.  12  6:  Pi- 

ani.  1977:27.  38.  figs.  3.  6.  10  [in  part — fig.  5  =  Galeodea 

echinophora  ^ Linnaeus.  1767),  P.  Bouchet  personal  com- 
munication, fig.  11  =  £.  bairdii]. 
Tonna  ^Eudolium)  thompsoni. — Kilias.  1962:16,  fig.  12  5. 
NOT  Dolium  \EudoIium)  crosseanum. — Dall,  1889a.b;  Daut- 

zenberg  &  Fischer,  1906  (=  E.  bairdiiV 
NOT  Dolium  crosseanum. — Locard.  1S97  ^Talisman  stn.  63 

=  £.  bairdii,  and  stn.  139  =  a  turrid.  P.  Bouchet.  personal 

communication). 


Figure  I.  Label  in  Monterosato's  handwriting  gummed  to  dorsum  of  holot\-pe  of  Eutfo/ium  crosseanum.  Figure*  2-4.  Eudolium 
crosseanum.  2.  holot>pe.  off  SiciK.  Hebrew  Uni\ersit>.  Jerusalem  No.  21386  (79.8  x  58  mm).  3.  Oregon  station  3636.  off  Belize. 
USNM  751892  (32  8  x  22  m).  4.  Oregon  station  2021.  off  French  Guiana.  USNM  751885  (54  x  .39.5  mm).  Figures  5-8.  Eudolium 
pyriforme.  5.  7.  Bohol  Straits,  Philippines.  NMNZ  MF.  56381  (5.  51  x  34.5  mm;  7.  40.5  x  27.5  mm).  6.  David  Starr  Jordan 
station  TC  40  54.  off  Oahu.  Hawaii.  USNM  504464  i79  5  x  47  mm).  8.  Southwest  of  Taiwan.  NMNZ  MF  56380  S4  5  x  55  mm). 
.\\\  photographs  of  shells  are  to  scale 

Figure  9.  Eudolium  crosseanum.  Oregon  station  3636.  off  Belize.  USNM  751892.  detail  of  feleoconch  sculptm-e.  7  x.  Figiu-es 
10-19.  Eudolium  bairdii.  10.  detail  of  teleoconch  sculpture,  of  specimen  in  figure  16.  7  x.  H.  off  .\ldermen  Islands.  New 
Zealand.  D.  Gibbs  collection  (76  x  49.5  mm).  12.  13.  off  Tosa-Shimuzu.  Japan.  NMNZ  MF.563S3  ^figure  12.  51  x  37  mm;  figure 
13.  59  X  43  4  mm>  14.  Combat  station  C279.  Straits  of  Florida.  USNM  715002  (49.7  x  35.5  mml  15.  off  Scot  Reef.  Western 
Australia.  NMNZ  MF.57427  (58  x  38  mm).  16.  Oregon  11  station  11133.  off  Yucatan  Peninsula.  Mexico.  USNM  751930  v.>3  x 
22.7  mm).  17.  SMIB  4  station  D\V.55.  southwest  of  New  Caledonia.  MNHN  i45  x  28  5  mm'  18.  Oregon  station  5690.  off  Colombia. 
USNM  751869  (46  x  31.5  mm:.  19.  Albatross  station  5590,  Sebuku  Bay.  Borneo.  USNM  23911  ^46.5  x  31.7  mm).  .Ml  photographs 
of  whole  shells  are  to  scale. 


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THE  NAUTILUS,  Vol.  106,  No.  1 


B.  A.  Marshall,  1992 


Page  27 


Page  28 


THE  NAUTILUS,  Vol.  106,  No.  1 


Figures  20-22.  Riglit  lateral  views  of  head-foot  areas  of  male 
EudoliuTH  animals.  20.  E.  crosseanum,  Oregon  station  3636, 
off  Ik-lize,  USNM  751892  21.  E.  pyrijormc.  David  HI  an  Jor- 
dan station  T(;/40/5-l.  off  Oahu,  Hawaii,  USNM  804464  22. 
F..  hairdii.  M ascareigncs  III  station  24,  off  Madagascar,  MNHN, 
Scale  lines:  20  =  2.5  mm;  21,  22  =  5  mm.  ct,  cephalic  tentacle, 
pe,  penis;  pr,  proboscis;  pt,  penial  tentacle;  sg,  seminal  groove. 


NOT  Eudolium  crosseanuni. — Tomlin,  1927;  Barnard,  1963; 
Abbott,  1974;  Okutani,  1983;  Kilburn,  1986;  Horikoshi, 
1989;  Waren  &  Bouchet,  1990;  Poppe  &  Goto,  1991  (  = 
E  hairdii) 

Description:  Shell  up  to  81  mm  (est.)  high,  thin  to 
moderately  thick,  periostracum  thin,  straw-colored.  Pro- 
toconch  deep  yellowish  brown;  teleoconch  irregularly 
maculated  with  yellowish  brown  on  a  white  or  buff  white 
ground,  some  major  spiral  cords  with  small  yellowish 
brown  spots,  mature  outer  lip  typically  with  a  pinkish 
flush.  Protoconch  primarily  conchiolin,  conical,  of  about 
5  convex  worls,  3.46-4.44  mm  in  diameter,  sculptured 
with  3  rows  of  small  periostracal  spines.  First  whorl  of 
calcareous  mould  of  inner  surface  of  protoconch  (figure 
23 — exterior  removed  with  sodium  hypochlorite  solu- 
tion) 317  ^m  wide,  sculptured  with  very  fine  irregular 
network  of  crisp  threads  that  enclose  minute,  crowded, 
roughly  circular  pits.  Second  whorl  with  fine,  crisp  axial 
riblets  and  2  similar  crisp  spiral  threads,  one  at  shoulder 
angulation,  the  other  on  ramp  between  shoulder  spiral 
and  suture.  Shoulder  spiral  commencing  immediately, 
ramp  spiral  commencing  three  quarters  of  a  whorl  later. 
Axial  ribs  and  spiral  cords  becoming  obsolete  early  on 
2nd  whorl.  Subsequent  whorls  essentially  smooth,  round- 
ed. Teleoconch  of  up  to  4  evenly  convex  whorls,  sculp- 
tured with  numerous  rounded  spiral  cords  that  multiply 
by  intercalation;  and  fine,  crisp,  crowded  axial  riblets;  in 
rare  specimens  some  spiral  cords  becoming  very  weakly 
nodular  on  last  adult  whorl.  Thin  specimens  with  a  thin, 
flared,  weakly  dentate  outer  lip  and  smooth  columella 
and  parietal  area.  Thickened  specimens  with  strongly 
dentate  outer  lip,  spirally  plicate  columella,  and  with  1- 
3  small  spirally  elongate  denticles  on  parietal  area  close 
to  insertion. 

Animal:  (figure  20:  subadult  male,  shell  height  22.3  mm, 
Oregon  stn.  3636)  Everted  proboscis  large,  very  wide 
but  longer  than  broad,  thin-walled.  Cephalic  tentacles 
small,  slender,  rounded  in  cross  section,  gently  tapered, 
tips  roimded.  Eyes  small,  in  rounded  swellings  on  outer 
sides  of  tentacles  near  bases.  Penis  base  well  behind  base 
of  right  cephalic  tentacle,  penis  very  large,  spathulate, 
laterally  compressed,  tip  broadly  rounded,  deeply  grooved 
along  ventral  edge  to  tip,  minute  tentacle  protruding 
from  end  of  groove  at  distal  extremity.  Operculum  absent 
in  adults.  Jaw  plates  (figure  24)  ovate,  thin,  each  with 
strong,  thickened,  hooked  anterior  projection. 

Raditla:  (figures  25,  26)  with  the  formula  2.1.1.1.2,  teeth 
curved,  sharply  pointed,  very  stout,  strongly  interlocked. 
Central  tooth  broad;  cutting  area  acutely  angulate,  7-9 
small  sharp  cusps  on  each  side,  terminal  cusp  large,  long, 
narrow;  posterolateral  projections  on  face  strong,  sharp. 
Lateral  teeth  strongly  longitudinally  flanged  along  outer 


Figures  23-26.  Eudolitun  crosseanuni.  2.'{.  protoconch  with  periostracum  removed,  Oregon  station  5914,  I^esser  Antilles,  USNM 
751903.  24.  (jaw  plate),  2.^.  26.  (radnla)  from  shell  22  mm  high.  Oregon  station  3636.  off  Belize,  USNM  751892.  25.  width  of 
radula.  26.  detail  of  central,  lateral  ;ind  marginal  teeth  Figures  27-.'{0.  Eudoliunt  pyrijornie.  Hadiila  from  shell  41  mm  high, 
MUSORST(JM  6  station  I)W39i,  Loyalty  Islands.  27.  width  of  radula  28-30.  details  of  central  and  lateral  teeth.  Scale  bars:  27 
=  0.5  mm,  others  =  0  1  mm. 


B.  A.  Marshall,  1992 


Page  29 


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THE  NAUTILUS,  Vol.  106,  No.  1 


Figures  3 1 -.'ib.  Etidolium  hairdii.  3 1 -.'$3.  radula  from  slu'll  24  mm  higli,  Straits  of  Florida,  Florida  State  Museum  L'F  26666. 
31.  detail  of  central  and  lateral  teeth  from  anterior  end  of  radular  ribbon  representing  earliest  teeth  formed.  32.  detail  of  central, 
lateral  and  marginal  teeth  at  posterior  third  of  radular — note  size  and  morphological  differences  compared  with  teeth  formed 
earlier  (3 1 ).  3.3.  width  of  radula  at  posterior  third.  .34.  detail  of  central  and  lateral  radular  teeth  from  shell  .57  mm  high,  Mascareignes 
III  station  24,  oH'  Madagascar,  MNHN.  3,'>,  36.  radula  from  shell  76  mm  high,  off  Aldermen  Islands,  New  Zealand,  D.  Gibbs 
colleclion.  3.'j.  width  of  radula.  36.  detail  of  central  tooth   Scale  bars:  35  =  0.5  mm,  others  =  0.1  mm. 


edges  to  interlock  with  inner  marginals,  terminal  cusp 
very  large,  8-11  fine  cusps  heliiiKl  on  outer  edge.  Mar- 
ginal teeth  similar,  without  secondary  cusps. 

Type  data:  Dolium  crosseanum.  HOLOTYPE  Coen 
collection  (No.  2521),  Hebrew  I'niversity,  Jeru.salem 
21386,  off  Palermo,  Italy;  Eudoliurn  tlwinpsani.  HO- 
LOTYPE Florida  State  Museum,  Gainsville,  Florida  UF 


170426,  Triton  stn.  1206,  off  Sombrero  Ke\  Light,  Flor- 
ida Ke\s,  137  m 

Other  material  examined:    WESTERN  ATLANTIC— 

off  Cape  Ilatteras,  South  C^arolina,  south  to  off  French 
(iuiana,  35°06'N-7°18'N  (56  specimens  in  37  lots  USNM). 

Distribution:     (figure  37)  Mediterranean  and  Western 
Atlantic,  from  Cape  Hatteras  to  French  Guiana;  depth 


B.  A.  Marshall,  1992 


Page  31 


Figure  37.    Map  showing  ihstrihution  of  Eudolium  crosscanum  (•)  and  £.  pyrifornie  (A) 


range  of  material  examined  17-914  m,  maximum  known 
Hving  occurrence  457  m. 

Remarks:  As  discussed  by  Piani  (1977)  two  specimens 
have  been  interpreted  as  the  holotype  of  D.  crosseanum, 
one  in  the  Monterosato  collection,  Museo  Zoologico,  Roma, 
the  other  in  the  Coen  collection,  Hebrew  University  of 
Jerusalem.  Piani  correctly  concluded  that  the  Rome  spec- 
imen cannot  be  the  holotype  because  it  is  extensively 
covered  by  a  brown  periostracum  and  has  conspicuous 
nodules,  characters  not  mentioned  in  the  rather  detailed 
original  description,  and  because  it  lacks  the  maculations 
recorded  by  Monterosato.  From  comparison  of  dimen- 
sions of  the  Rome  and  Jerusalem  specimens  with  Mon- 
terosato's  original  published  measurements,  Piani  (1977: 
37)  concluded  that  the  Jerusalem  specimen  was  the  clos- 
est match  but  because  there  were  discrepancies  he  sug- 
gested that  it  was  perhaps  a  third  example  and  thus  by 
implication  not  the  holotype.  Piani  (1977:  figs.  6,  10) 
reproduced  Coen's  (1930:  figs.  1,  2  "tipo")  illustrations 
and  a  photograph  (provided  by  H.  K  Mienis)  of  the 
Jerusalem  specimen.  Upon  receipt  of  the  Jerusalem  spec- 
imen (figures  1,  2),  kindly  lent  to  me  by  H.  K.  Mienis, 
it  was  immediately  obvious  that  Piani's  illustrations  are 
of  one  and  the  same  specimen,  though  this  was  neither 
unequivocally  stated  by  Piani  nor  evident  from  the  il- 
lustrations. 

It  transpires  that  the  dimensions  of  the  Jerusalem  spec- 
imen were  wrongly  quoted  by  Piani  (1977:37),  as  they 
in  fact  closely  accord  with  Monterosato's  (1869)  original 
measurements  (  "long  81;  diam.  maj.  56  mill.;  apert.  62 
mill,  longa,  28  lata  "):  height  79.8  mm  (lacks  proto- 
conch — estimated  height  when  intact  81  mm),  diameter 
58  mm,  height  of  aperture  62  mm,  width  of  aperture  28 


mm.  The  general  shell  facies,  color  and  color  pattern  of 
this  specimen  agree  closely  with  the  original  description, 
while  the  number  of  denticles  on  the  outer  lip  (32)  are 
closely  accordant:  Monterosato  stated  that  there  were  33 
denticles,  but  it  is  possible  that  a  low  swelling  at  the 
abapical  extremity  of  the  lip  was  also  counted.  The  only 
discrepancy  is  the  thickness  of  the  outer  lip,  which  ranges 
from  3.5  to  4.0  mm  over  the  denticulate  area,  and  not 
3  mm  as  stated  by  Monterosato.  The  smooth  zone  at  the 
adapical  extremity,  however,  is  3  mm  thick,  suggesting 
that  Monterosato  may  have  measured  it  there  to  avoid 
including  the  denticles.  Diagnostic  features  of  this  par- 
ticular specimen  that  were  realised  on  the  original  draw- 
ing include  the  distinctive  pattern  of  chipping  at  the  rim 
of  the  anterior  siphonal  canal,  and  the  growth  scar  in 
front  of  the  columella.  Incontrovertible  proof  that  this 
specimen  belonged  to  Monterosato  is  the  label  in  Mon- 
terosato's handwriting  gummed  to  the  dorsum  (figure  1) 
"Genus  Doliopsis  mihi  inedito' . 

The  specific  identity  of  the  Rome  specimen  is  uncer- 
tain, but  judging  from  Piani's  (1977:  fig.  11)  illustration 
it  is  much  more  strongly  nodular  than  any  specimen  of 
E.  crosseanum  seen  during  the  present  study  and  seems 
more  likely  to  be  E.  hairdii  (P.  Bouchet  concurs).  Other 
published  records  of  E.  crosseanum  from  the  Mediter- 
ranean apart  from  references  to  the  holotype  are  based 
on  misidentification  (see  synonymy). 

By  direct  comparison  of  the  holotypes,  the  Western 
.Atlantic  E.  thompsoni  McGinty,  1955  is  specifically  in- 
distinguishable from  E.  crosseanum.  Judging  from  the 
fact  that  the  species  has  only  been  obtained  once  in  the 
Mediterranean  yet  it  is  relatively  common  in  the  western 
Atlantic,  it  is  likely  that  the  Mediterranean  specimen  is 
the  result  of  a  chance  introduction  of  a  teleplanic  larva 


Page  32 


THE  NAUTILUS,  Vol.  106,  No.  1 


Figure  38.     N4ap  showing  distribution  of  Eudolium  bairdii.  Eastern  Atlantic  distributions  provided  by  P.  Bouchet  (personal 
comnuinication). 


transported  by  the  Gulf  Stream  from  Western  Atlantic 
populations  (see  Discussion). 

Eudolium  crosseanum  is  closely  similar  to  the  Western 
Pacific  and  Hawaiian  species  E.  pyriforme  (see  below). 

Eudolium  crosseanum  was  the  first  species  described 
by  Monterosato  in  an  illustrious  malacological  career,  and 
the  holotype  must  have  been  one  of  his  most  prized 
possessions.  He  would  have  been  impressed  to  know  that 
his  specimen  would  still  be  the  only  one  known  from  the 
Mediterranean  122  years  after  it  was  described,  and  that 
it  would  remain  the  largest  known  example  of  the  species. 


Eudolium  pyriforme  (Sowerby,  1914) 
(figures  5-8,  21,  27-30,  37) 

Doliuni  pyriforme  Sowerby,  1914:37,  pi.  2,  fig.  14. 

Eudolium  piiriformc. — Osinia,  1943:131,  pi.  5,  fig.  4;  May- 
bloni,  1951:282;  Kira,  1959:55,  pi,  22,  fig,  5;  Garrard,  1961: 
17;  Kira,  1962:59,  pi.  23,  fig.  5;  Okutani,  1964:400,  pi.  1, 
fig.  23,  te.xt  fig.  2;  Habe  &  Kosuge,  1967:66,  pi,  25,  fig.  12; 
Kuroda,  Habe  &  Oyama,  1971:135,  pi.  37,  fig.  4;  Wolfe, 
1974:3,  text  fig.;  Powell,  1974:201;  Cernohorsky,  1976:1, 
fig.  1;  Cernohorsky,  1978:61,  pi.  16,  fig.  7;  Powell,  1979: 
163;  Kosuge,  1985:59,  pi.  22,  fig.  6;  Abbott  &  Dance,  1986: 
116,  text  fig.;  Kilburn,  1986:4,  fig.  11;  Okutani,  1988:78, 
fig  46. 

Description:  Shell  up  to  85.5  (est.)  mm  high,  thin;  peri- 
ostracurn  thin,  translucent,  straw-colored.  Protoconch 
yellowish  brown;  teleoconch  sparsely  to  densely  irregu- 
larly maculated  with  pale  to  deep  yellowish  brown,  major 
spiral  cords  alternately  spotted  yellowish  brown  and  white 
on  a  buff  white  ground,  outer  lip  of  many  specimens 


with  a  pinkish  flush.  Protoconch  primarily  conchiolin,  of 
about  5  convex  whorls,  2.95-3.45  mm  in  diameter,  es- 
sentially smooth  (worn?).  Internal  calcareous  mould  not 
seen.  Teleoconch  of  up  to  4.6  evenly  convex  whorls, 
sculptured  with  numerous  rounded  spiral  cords  that  mul- 
tiply by  intercalation,  and  fine  axial  riblets,  axials  crisp 
on  first  3  whorls,  less  crisply  defined  on  subsequent  whorls, 
no  nodules.  Outer  lip  of  most  specimens  weakly  thick- 
ened in  adults,  but  a  few  specimens  are  moderately  thick- 
ened and  dentate;  columella  and  parietal  area  simple. 

Animal:  (figure  21 — adult  male  minus  radula  that  had 
been  extracted  when  received  from  one  of  7  shells  64- 
79.5  mm  in  height,  from  off  Oahu,  Hawaii — USNM 
804464)  externally  similar  to  that  of  E.  crosseanum  (see 
above)  but  differing  in  that  the  penis  arises  nearer  outer 
base  of  right  cephalic  tentacle.  Jaw  similar  to  that  in  £. 
crosseanum. 

Radula:  (figures  27-30 — juvenile  female  22  mm  shell 
height,  Oregon  stn.  3636,  off  Belize  City,  USNM  751892) 
similar  to  that  of  £.  crosseanum  except  that  central  and 
lateral  teeth  have  smaller  cusps,  and  sides  of  central  tooth 
subparallel  instead  of  markedly  tapered. 

Type  data:  Dolium  pyriforme:  HOLOTYPE:  The  Nat- 
ural History  Museum,  London  1915.1.6.148,  off  Kii,  Ja- 
pan. 

Other  material  examined  (73  specimens):  INDIAN 
OC>EAN — Trawled  off  Umgababa,  Natal,  South  Africa 
(1  NMP);  trawled  ofi^  Durban,  South  Africa,  421  m  (1 
NMP);  trawled  off  Bazaruto  I.,  Mozambique  (1  NMP); 
12°39.8'S,  48°16.2'E,  N  Madagascar,  375-385  m  (1 
MNHN);  I2''42.4'S,  48°14.3'E,  N  Madagascar,  285-295 


B.  A.  Marshall,  1992 


Page  33 


m  (1  MNHN);  12°42.9'S,  48°12.1'E,  N  Madagascar,  445- 
455  m  (1  MNHN);  12°43'S,  48°15'E,  300-340  m  (1 
MNHN);  20°58'S,  55°15'E,  off  Reunion,  450-580  m  (1 
MNHN);  22°14.7'S,  43°04.5'E,  S  Madagascar,  470-475 
m  (1  MNHN);  Crevettiere  1986  stn.  80,  22°17'S,  43°04'E, 
SW  Madagascar,  530  m  (1  MNHN);  Crevettiere  1986 
stn.  76,  22°22'S,  43°03'E,  SW  Madagascar,  530  m  (2 
MNHN);  Crevettiere  1986  stn.  56,  22°26'S,  43°05'E,  SW 
Madagascar,  435  m  (2  MNHN);  Crevettiere  1986  stn. 
57,  22°26'S,  43°06'E,  SW  Madagascar,  460  m  (1  MNHN); 
Crevettiere  1986  stn.  58,  22°26'S,  43°06'E,  SW  Mada- 
gascar, 440  m  (1  MNHN);  SoWa  stn.  01/84/74,  14°16.5'S, 
122°54.4'E,  NW  of  Collier  Bay,  Western  Australia,  302 
m  (1  WAM);  Soela  stn.  01/84/54,  15°51.2'S,  120°44.3'E, 
WNW  of  Lacepede  Archipelago,  Western  Australia,  350- 
348  m  (6  WAM)  JAPAN— off  Tosa  Shimuzu,  ca.  350- 
400  m  (1  NMNZ);  off  Ashizuri-misaki  (2  MCZ);  oft  Tosa, 
ca.  183  m  (1  MCZ);  off  Kii  (1  MCZ).  TAIWAN— SW  of 
Taiwan,  ca.  128-183  m  (5  NMNZ).  PHILIPPINES— 
Albatross  stn.  5289,  Batangas  Bav,  Luzon,  315  in  (2 
USNM);  Coriolis  MUSORSTOM  3  stn.  CP103,  of  Min- 
doro,  193-200  m  (1  MNHN);  Coriolis  MUSORSTOM  2 
stn.  CP83,  13°55'N,  120°30'E,  off  Mindoro,  318-320  m 
(1  MNHN);  Bohol  Straits,  ca.  200  m  (5  NMNZ);  Albatross 
stn.  5519,  off  Pt  Tagalo,  N.  Mindanao,  333  m  (2  USNM); 
Albatross  stn.  5518,  off  Pt  Tagalo,  Mindanao,  366  m  (2 
USNM);  9°43'S,  130°00'E,  Timor  Sea,  ca.  250  m  (2 
NMNZ).  HAW  Ml— David  Starr  Jordan  stn.  TC-40-54, 
21°01.6'N,  156°43.00'W,  off  Oahu,  223  m  (7  USNM). 
QUEENSLAND  and  NEW  SOUTH  WALES— Off  Cape 
Moreton,  165  m  (1  AMS);  Kapata  stn.  71-08,  ca.  32°46'S, 
152°16'E,  off  Newcastle,  280-549  m  (1  AMS);  Kapala 
stn.  K75-05-03,  33°02'S,  152°31'E,  off  Newcastle,  475  m 
(1  AMS);  between  Newcastle  and  Sydney,  182-549  m  (1 
AMS);  off  Broken  Bay,  137  m  (1  AMS);  between  Sydney 
and  Norah  Head,  412-457  m  (1  AMS);  Kapala  stn'  K76- 
07-01,  33°33'S.  151°59'E,  off  Broken  Bay,  384  m  (1  AMS); 
Kapala  stn.  K76-24-05,  33°33'S,  151°59'E,  off  Broken 
Bay,  373-366  m  (1  AMS);  off  Svdnev,  420-440  m  (1 
AMS);  35°50'S,  150°34'E,  off  Batemans  Bay,  366  m  (1 
AMS).  CHESTERFIELD  REEFS— Corio/'is  MUSOR- 
STOM 5  stn.  376,  19°51'S,  158°30'E,  280  m  (1  MNHN). 
NEW  CALEDONIA— Vanfcan  MUSORSTOM  4  stn. 
CP193,  18°56'S,  163°23'E,  off  d'Entrecasteau  Reefs,  415 
m  (1  MNHN);  Vaiiban  SMIB  3  stn.  DW28,  22°47'S, 
167°12'E,  394  m  (1  MNHN);  Vaiiban  SMIB  1  stn.  DW2, 
22°52'S,  167°13'E,  415  m  (1  MNHN);  Vaiiban  SMIB  2 
stn.  DW15,  22°53'S,  167°11'E,  375-402  m  (1  MNHN); 
Vauban  SMIB  2  stn.  DW8,  22°54'S,  167°13'E,  435-447 
m  (1  MNHN);  Vauban  SMIB  2  stn.  DW17,  22°55'S, 
167°15'E,  428-448  m  (1  MNHN).  LOYALTY  IS- 
LANDS—A/is  MUSORSTOM  6  stn.  DW391,  20°47'S, 
167°06'E,  off  Ouvea,  390  m  (1  MNHN);  Alis  MUSOR- 
STOM 6  stn.  CP  465,  21°04'S,  167°32'E,  off  Lifou,  480 
m  (1  MNHN).  NEW  ZEALAND— off  Aldermen  I.,  ca. 
366  m  (1  Gardner  coll'n,  Auckland);  BS843  (0.589). 
37°14.6'S,  176°51.0'E,  Rangatira  Knoll,  NW  of  White  I., 
407-162  m  (fragment  NMNZ);  off  Tokomaru  Reef,  Gis- 
borne,  ca.  220  m  (1  Auckland  Institute  and  Museum). 


Distribution  (figure  37):  Southern  Africa,  Madagascar, 
Reunion,  Japan,  Taiwan,  Philippine  Islands,  Indonesia, 
Hawaii,  Australia,  New  Caledonia,  northern  New  Zea- 
land; depth  range  of  material  examined  137-580  m, 
deepest  known  living  occurrence  390  m. 

Remarks:  Eudolium  pyriforme  is  extremely  similar  to 
E.  crosseanum  in  shell  color,  color  pattern,  protoconch 
morphology,  shell  thickness,  size  relative  to  the  number 
of  whorls,  and  in  size  attained.  Although  they  are  also 
similar  in  sculpture,  E.  pyriforme  differs  in  having  axial 
riblets  that  are  consistently  lower,  broader  and  less  sharp- 
ly defined  than  those  of  E.  crosseanum,  especially  where 
traversing  the  spiral  cords.  As  described  above,  a  single 
adult  male  specimen  of  E.  pyriforme  differs  from  that 
of  an  immature  E.  crosseanum  in  having  the  penis  base 
close  beside  the  right  cephalic  tentacle  instead  of  well 
behind  it  (figures  20,  21).  The  external  anatomies  of 
males  and  females  are  otherwise  similar.  Although  the 
position  of  the  penis  seems  likely  to  be  a  major  distin- 
guishing feature,  with  only  two  different-sized  animals 
for  comparison  it  is  impossible  to  ascertain  whether  or 
not  its  position  is  infraspecifically  stable  or  changes  dur- 
ing ontogeny. 

As  described  and  illustrated  herein  (figures  25-30),  the 
radula  of  £.  pyriforme  differs  from  that  of  E.  crosseanum 
in  the  shape  of  the  central  tooth  and  in  the  size  of  the 
secondary  cusps.  With  only  a  single  radula  from  each 
species  from  dissimilar-sized  specimens,  however,  it  is 
impossible  to  ascertain  either  the  degree  of  infraspecific 
variability  at  equivalent  size,  or  the  degree  of  morpho- 
logical change  during  ontogenetic  development  (see  E. 
bairdii  below). 

Specimens  of  £.  pyriforme  tend  to  be  a  little  narrower 
than  those  of  E.  crosseanum.  although  there  is  a  broad 
overlap  in  shape.  Shell  height/diameter  ratio  in  E.  pyr- 
iforme ranges  from  1.41  to  1.72  (mean  1.53,  SD  0.095, 
n  =  30)  as  against  1.31-1.66  (mean  1.44,  SD  0.085,  n  = 
18)  in  £.  crosseanum.  Specimens  from  off  Hawaii  are 
uniformly  narrowly  ovate  (USNM  804464,  height/di- 
ameter ratio  1.59-1.72,  mean  1.67,  SD  0.047,  n  =  7, 
figure  6).  Some  specimens  from  Japan  (MCZ  293903), 
the  Timor  Sea  (NMNZ  MF.57428),  New  South  Wales 
(AMS  C.68870)  and  Mozambique  (NMP  H.7800)  are  as 
narrowly  ovate  as  specimens  from  Hawaii,  however,  and 
there  is  complete  integration  between  narrow  and  broad 
forms  in  material  from  west  of  Hawaii  (height/diameter 
ratio  1.41-1.62,  mean  1.48,  SD  0.052,  n  =  22). 

Eudolium  bairdii  (Verrill  &  Smith,  1881) 
(figures  10-19,  22,  31-36.  38) 

Doliuni  bairdii  Verrill  &  Smitii  in  Verrill,  1881:299;  Verrill, 
1882:.51.5;  Verrill,  1884253.  pi.  29,  figs.  2a,b. 

Dolium  hayardi  (sic,).~Paetel,  1888:221. 

Dolium  (Eudolium)  crosseanum. — Dull,  1889a:232  (in  part), 
pi.  15.  fig,  5. 

Dolium  {Eudolium)  crosseanum. — Dall,  1889b:134,  pi.  15,  fig. 
5,  pi.  44,  fig.  2,  pi.  62,  figs.  83,  83a;  Dautzenberg  &  Fischer, 
1906:38  (not  D.  crosseanum  Montern,sato,  1869). 


Page  34 


THE  NAUTILUS,  Vol,  106,  No.  1 


Dohum  crosseamim. — Locard.  1897293  (in  part  not  D.  cros- 
seanurn  Monterosato,  1869 — Talisman  stn.  139  record  is 
based  on  a  turrid:  P.  Bouchet,  personal  communication). 

Dolium  (Eudolium)  cro.sseanum  var.  solidior  Dautzenherg  & 
Fischer,  1906:38,  pi.  3,  fig.  1.  {new  synonym). 

Dolium  (Eudolium)  crosseanum. — Kobelt.  1908:1.5.5,  pi  126, 
figs.  2,  3,  -1  (in  part — fig.  1  =  £.  crosseanum). 

Morio  lineata  Schepman,  1909:124,  pi,  10,  fig,  5  (new  syn- 
onym). 

Eudolium  crosseanum. — Tomlin,  1927:82,  fig,  4b;  Barnard, 
1963:8;  .\bbott,  1974:168,  pi,  6,  fig,  1787;  Okutani,  1983: 
264,  text  fig,;  Kilburn,  1986:4,  fig,  12;  Horikoshi,  1989:63, 
pi,  15,  fig,  11;  Waren  &  Bouchet,  1990:89,  figs,  11,  78,  79; 
Poppe  &  Goto,  1991:128,  pi,  22.  figs,  1,  2  (not  D.  cros- 
seanum Monterosato,  1869), 

Eudolium  lincatum. — Osima,  1943:133,  pi,  5,  fig,  1, 

Oocorys  lineata.— Turner .  1948:178,  190;  .Abbott  &  Dance, 
1986:116,  te,\t  fig, 

Eudolium  inflatum  Kuroda  &  Habe,  1952:56  (replacement 
name  for  Eudolium  lincatum  Osima  not  Schepman);  Ku- 
roda &  Habe,  19.57:28,  figs,  2,  4,  5;  Kira,  1962:.59,  pi,  23, 
fig,  4;  Okutani,  1988:77,  fig.  45;  Bieler  &  Petit,  1990:137 
(new  synonym). 

Eudolium  lincatum  inflatum — Kira,  195444,  pi.  22,  fig,  4; 
Kira,  19.59:.55,  pi,  22. 

Eudolium  kuroharai  Azunia,  1960:98,  pi,  1,  fig,  8,  text  fig,  1 
(new  synonym). 

Tonna  (Eudolium)  crosseana. — Kilias,  1962:14  (in  part), 

Dolium  (Eudolium)  crosseanum  solida  (sic), — Settepassi,  1971, 
appendix  vii,  text  figs, 

Tonna  (Eudolium)  crosseana. — Piani,  1977,  fig.  11, 

Oocorys  solidior. — Piani,  1977:38,  figs.  1,  2.  7, 

Description:  Shell  up  to  75,5  mm  liigli,  thin  to  rather 
thick,  periostracum  thin,  straw-colored.  Protoconch  deep 
yellowish  to  reddish  brown;  teleoconch  white  or  buff 
white,  major  spiral  cords  yellowish  to  reddish  brown, 
outer  lip  white.  Protoconch  primarily  conchiolin,  conical, 
of  about  5  convex  whorls,  3.00-4.75  mm  in  diameter, 
sculptured  with  3  narrow  spiral  threads  with  small  spines 
(in  well  preserved  specimens).  Calcareous  mould  of  inner 
surface  of  protoconch  (Waren  &  Bouchet,  1990:  fig  79) 
similar  to  that  of  E.  crosseanum  but  with  3  instead  of  2 
spiral  threads.  Teleoconch  of  up  to  4.4  evenly  convex 
whorls,  sculptured  with  numerous  rounded  spiral  cords 
that  iTiultipK  by  intercalation,  and  fine  axial  riblets,  axials 
less  crisply  defined  after  third  whorl,  last  adult  whorl 
with  or  without  weak  to  strong  nodules  on  major  spiral 
cords.  Shell  ranging  through  intermediates  from  thin  to 
rather  thick.  Thin  specimens  with  thin,  flared,  weakly 
dentate  outer  lip  and  simple  columella  and  parietal  area. 
Most  thickened  specimens  with  thick,  strongly  dentate 
outer  lip,  thick  spirally  plicate  callus  at  base  of  columella, 
and  1-3  spirally  elongate  denticles  on  parietal  area  below 
insertion. 

Animal:  (figure  22,  adult  male,  shell  height  57.5  mm, 
Mascareignes  III  stn.  24,  off  Madagascar)  externally  sim- 
ilar to  that  of  E.  pyrijorme. 

Radula:  (figures  31-36)  (from  shells  25,  57  and  76  mm 
in  height)  similar  to  those  of  £.  crosseanum  and  E.  pyr- 
ijorme. With  increasing  size  central  and  lateral  teeth 


enlarging  while  size  of  secondary  cusps  and  area  they 
occupy  remain  rather  static.  Secondary  cusps  ultimately 
becoming  obsolete  on  laterals  in  large  adults. 

Type  data:  Dolium  bairdii:  HOLOTYPE  USNM  51385 
(51  X  25  mm).  Fish  Hawk  station  945,  39°58'N,  71°13'W, 
off  Barnegat,  New  jersev,  379  m;  Dolium  (Eudolium) 
crosseanum  var.  solidior.  HOLOTYPE  Musee  Oceano- 
graphique,  Monaco,  38°52.50'N,  27°23.05'W,  off  the 
Azores,  599  m;  Morio  lineata:  HOLOTYPE  Zoological 
Museum,  Amsterdam  3.09.008,  Sihoga  stn  173,  3°27'S, 
131°0.5'E,  Ceram  Sea,  567  m;  Eudolium  inflatum:  TYPE 
MATERIAL  (Osima,  1943,  pi.  5,  fig.  1)  possibly  in  Oshi- 
ma  collection,  which  was  destroyed  during  World  War 
II  (A.  Matsukuma,  personal  communication),  off  Wa- 
kayama  Prefecture,  Japan;  Eudolium  kuroharai:  HO- 
LOTYPE in  Mr.  M.  Azuma's  private  collection,  Takara- 
zuka  City,  Japan  (not  seen),  off  Tosa,  Japan. 

Other  materia!  examined  (161  specimens):  WEST- 
ERN ATLANTIC  OCEAN— Barnegat,  New  Jersey,  south 
to  off  French  Guiana,  39°58'N-7°37'N  (12  specimens  in 
9  lots  MCZ,  91  specimens  in  43  lots  USNM).  WESTERN 
INDIAN  OCEAN— 12°01'S,  49°26'E,  off  Madagascar,  450 
m,  A.  Crosnier  (1  MNHN);  12°42'S,  48°14'E,  off  N.  Mad- 
agascar, 380-375  m,  A.  Crosnier  (1  MNHN);  12°43'S, 
48°12'E,  off  N.  Madagascar,  445-455  m,  A.  Crosnier  (1 
MNHN);  Marion-Dujresne  crusise  32.  stn.  CP  144, 
20°50'S,  55°35'E,  off  Reunion,  605-620  m  (1  MNHN); 
Mascareignes  III  stn.  37,  22°18'S,  43°05'E,  off  S  Mada- 
gascar (1  MNHN);  Crevettiere  1986  stn.  80,  22°17'S, 
43°04'E,  SW  Madagascar,  530  m  (1  MNHN);  Crevettiere 
1986  stn.  81,  22''23'S,  43°03'E,  SW  Madagascar,  525  m 
(2  MNHN);  Mascareignes  III  stn.  24,  22°31'S,  43°07'E, 
off  S  Madagascar,  430-460  m  (1  MNHN);  trawled  off 
Umgababa,  Natal,  South  Africa  (1  NMP);  trawled  off 
Tongaat,  Natal,  South  Africa  (1  NMP);  off  Cape  St.  Fran- 
cis, South  Africa  (1  NMP).  WESTERN  AUSTRALIA— 
off  Augustus  I.  south  to  off  Rowlev  Shoals  (2  specimens 
in  1  lot  NMNZ,  23  specimens  in  19  lots  W.AM,  1  specimen 
USNM).  JAPAN— SW  of  Ashizuri  misaki,  Kochi  Prefec- 
ture, trawled  (1  MCZ);  off  Tosa  Shimuzu,  Kochi  Prefec- 
ture, 350-400  m  (2  NMNZ).  PHILIPPINES— Corio/is 
MUSORSTOM  3  stn.  CP106,  13°47'N,  120°30'E  off  Min- 
doro,  640-668  m  (1  MNHN).  MALA\Sl\— Albatross 
stn.  .5592,  Sebuku  Bay,  Borneo,  558  m  (1  USNM);  Al- 
batross stn.  5590,  Sebuku  Bav,  Borneo,  567  m  (1  USNM). 
NEW  CALEDONIA— Vai/fcan  MUSORSTOM  4  stn. 
DW156,  18°54'S,  163°19'E,  Grand  Passage,  525  m  (1 
MNHN);  Vauban  MUSORSTOM  4  stn.  DWT97,  18°51'S, 
163°21'E,  Grand  Passage,  550  m  (1  MNHN);  Vauban 
MUSORSTOM  4  stn.  CP  171,  18°58'S,  163°14'E,  Grand 
Passage,  425  m  (1  MNHN);  Jcan-Charcot  BIOCAL  stn. 
DW36,  23°09'S,  167°11'E,  650-680  m  (1  MNHN);  Alis 
SMIB  4  stn.  DW55,  23°21'S,  168''05'E,  215-260  m  (1 
MNHN).  LOYALTY  ISLANDS— A/is  Ml'SORSTOM  6 
stn.  CP  467,  2r05'S,  167°32'E,  off  Lifou,  575  m  (1 
MNHN).  NEW  SOUTH  WALES— off  Port  Stephens 
south  to  off  Brush  I.  (5  specimens  in  5  lots  AMS).  NEW 
ZEALAND— N  of  Mavor  I.,  Bav  of  Plenty,  380-420  m, 


B.  A.  Marsliall,  1992 


Page  35 


coll.  M.  Huaki  (1  NMNZ);  off  Aldermen  I.,  New  Zealand, 
400  m  (1  D.  Gibbs  coll'n,  Auckland). 

Distribution  (figure  38):  Atlantic,  Mediterranean  and 
Indo- Western  Pacific,  17-823  m,  deepest  known  occur- 
rence of  living  specimen  560  m. 

Remarks:  Eiidoliiim  bairdii  is  rendered  high!}  distinc- 
tive by  the  reddish  to  \ellowish  brown  spiral  bands  on 
the  major  spiral  cords,  a  pattern  in  marked  contrast  to 
the  irregular  mottling  of  E.  crosseanum  and  E.  pyri- 
forme.  Endolitim  bairdii  is  indistinguishable  from  the 
Pacific  form  that  has  usualK  been  known  as  E.  lineatum 
(see  below).  Compared  with  E.  crosseanum  and  £.  pyr- 
iforme  at  the  same  stage  of  growth,  the  primary  spiral 
cords  in  E.  bairdii  tend  to  be  more  prominent,  while  the 
secondary  spirals  generally  enlarge  more  slowly  relative 
to  the  primaries.  E.  bairdii  differs  further  in  having  con- 
siderably stronger  a.xial  riblets,  especially  where  travers- 
ing the  spiral  cords.  The  axial  riblets  in  £.  bairdii  tend 
also  to  be  less  crowded,  numbering  3-7  per  millimeter 
at  the  end  of  the  second  teleoconch  whorl  (mean  5.40, 
SD  1.04,  n  =  20),  compared  with  6-10  (mean  S  33,  SD 
1.07,  n  =  12)  in  £.  crosseanum,  and  6-11  (mean  8.69, 
SD  1.55,  n  =  13)  in  E.  pyriforme.  The  calcareous  internal 
mould  of  the  protoconch  of  E.  bairdii  ( Waren  &  Bouchet, 
1990:  fig.  79 — misidentified  as  E.  crosseanum)  differs 
from  that  of  E.  crosseanum  (figure  23)  in  having  3  instead 
of  2  spiral  threads.  With  only  a  single  protoconch  from 
each  species  for  comparison,  however,  the  degree  of  in- 
fraspecific  variability  in  protoconch  morphology  is  un- 
certain. 

Of  the  three  known  living  Eudolium  species,  E.  bairdii 
exhibits  the  greatest  variation  in  shell  morphology,  un- 
doubted adults  ranging  from  32.5  mm  (est.)  to  75.5  mm 
in  height,  with  lightly  built  or  strongly  thickened  shells, 
and  with  or  without  nodules  on  the  last  adult  whorl. 
There  is  smooth  intergradation  between  all  of  the  ex- 
tremes both  within  and  between  samples  from  the  At- 
lantic and  the  Indo-Pacific.  B\  direct  comparison  of  ho- 
loty  pes  and  topotypes,  I  am  unable  to  detect  any  constant 
differences  between  D.  bairdii,  D.  crosseanum  var.  so- 
lidior.  M.  lineatum,  E.  inflatum  and  E.  kuroharai,  and 
so  it  is  concluded  that  they  are  all  conspecific.  It  is  sig- 
nificant, however,  that  thick-shelled  forms  with  heavily 
thickened  outer  lips  and  strongly  dentate  apertures  (ex- 
treme solidior  form)  (figures  14,  16)  have  no  strict  par- 
allel (figure  17)  outside  the  Atlantic,  while  specimens 
from  off  eastern  Australia  are  thick-shelled  and  often 
particularly  heavily  sculptured,  yet  they  do  not  exhibit 
the  apertural  features  of  the  solidior  form.  These  differ- 
ences suggest  that  there  may  have  been  some  genetic 
drift  between  isolated,  probably  largely  self-recruiting 
populations,  but  these  may  also  be  non-genetic  responses 
to  local  environmental  conditions.  The  latter  interpre- 
tation is  suggested  by  the  fact  that  a  solidior-Vike  form 
is  also  rarely  exhibited  by  the  Atlantic  E.  crosseanum 
(figure  4)  yet  not  by  E.  pyriforme,  its  Indo-Pacific  sister 
species.  Heavy  shelled  forms  (estimated  height  34.5-61.0 
mm)  are  clearly  adults  and  perhaps  senescent,  and  it  is 


likely  that  many  lightly  built  forms  (32.5-75.5  mm)  are 
also  mature.  Although  type  specimens  of  D.  bairdii  and 
the  synonymized  taxa  are  not  illustrated  here,  illustra- 
tions of  strongly  similar  specimens  are  provided  for  ori- 
entation: D.  bairdii,  E.  inflatum  and  E.  kuroharai  (figures 
12,  13,  15),  D.  crosseanum  var.  solidior  (figures  14,  16), 
Morio  lineata  (figures  18,  19). 

E.  bairdii  and  E.  crosseanum  have  fully  overlapping 
geographic  and  bathymetric  ranges  in  the  Atlantic  (fig- 
ures 37,  38),  and  the  two  species  have  been  taken  together 
at  three  stations  in  the  Western  Atlantic  {Oregon  stn. 
2391,  USNM  878126  and  751929  respectivelv;  Oregon 
stn.  4911,  USNM  878128  and  751872;  Oregon  II  stn. 
11253,  USNM  766104  and  878129).  E.  bairdii  and  £. 
pyriforme  also  have  overlapping,  geographic  and  bathy- 
metric ranges  and  are  thus  probably  locally  sympatric. 

DISCUSSION 

Although  larvae  of  Eudolium  species  have  not  been  ob- 
served in  the  field  or  reared  in  the  laboratory,  the  pro- 
toconch morphologv  and  species  distributions  are  char- 
acteristic of  tonnoideans  with  teleplanic  larvae 
(Scheltema,  1966;  Laursen,  1981).  Teleplanic  larvae  re- 
main planktonic  for  many  months,  in  some  species  for 
a  year  or  more,  and  can  be  transported  great  distances 
in  ocean  currents  (Scheltema,  1966,  1971;  Pechenik  et 
ai,  1984).  Species  with  teleplanic  larvae  often  have  ex- 
tremely wide  amphioceanic,  Indo-Pacific,  or  Indo-Pa- 
cific-Atlantic  distributions  in  their  benthic  stage,  es- 
pecially those  with  wide  tolerances  (Scheltema,  1986  and 
references  therein).  Although  some  localities  for  individ- 
uals in  the  benthic  stage  do  not  necessarily  support  self- 
recruiting  populations,  it  is  clear  that  widely  separated 
self-recruiting  populations  are  able  to  maintain  their  spe- 
cific integrity  through  periodic  influx  of  larvae  from  up- 
current  populations. 

Whether  or  not  Eastern  Atlantic  and  Mediterranean 
specimens  respectively  of  £.  bairdii  and  £.  crosseanum 
arose  from  local  self-recruiting  populations  is  unknown, 
but  judging  from  the  fact  that  they  are  indistinguishable 
from  Western  Atlantic  specimens  and  that  both  species 
are  evidently  far  more  common  in  the  Western  Atlantic 
than  in  the  Mediterranean,  it  is  probable  that  some  if 
not  all  originated  from  Western  Atlantic  larvae  that  were 
transported  by  the  Gulf  Stream  (Scheltema,  1986).  While 
E.  bairdii  may  well  be  at  least  partly  self-recruiting  in 
the  Eastern  Atlantic,  this  is  unlikely  to  be  true  for  £. 
crosseanum  in  the  Mediterranean  with  a  single  con- 
firmed specimen.  Specific  integrity  of  the  strongly  iso- 
lated Hawaiian  population  of  £.  pyriforme  is  probably 
maintained  through  intermittent  recruitment  of  larvae 
from  the  Western  Pacific  via  the  Kuroshio  Current  (Zins- 
meister  &  Emerson,  1979). 

It  is  likely  that  the  two  most  closely  related  species, 
£.  crosseanum  and  £.  pyriforme  originated  from  dis- 
junct populations  of  an  ancestor  that  formerly  had  a 
continuous  distribution  in  the  Tethyian  Atlantic-Indo- 
Pacific.  Since  Eudolium  species  are  unknown  living  ei- 


Page  36 


THE  NAUTILUS,  Vol.  106,  No.  1 


ther  in  the  Eastern  Pacific  or  from  the  Western  Atlantic 
south  of  Columbia,  it  seems  probable  that  distribution 
was  continuous  via  Africa  rather  than  America.  Unfor- 
tunately, the  southwestern  coast  of  Africa  has  been  rel- 
atively poorly  sampled  at  depths  most  likely  to  yield 
Eudolium  species  (300-600  m)  and  it  is  unknown  wheth- 
er or  not  they  occur  there.  The  apparent  absence  of  £. 
crosseanum  off  South  Africa  suggests  that  larvae  are 
incapable  of  reaching  there  from  northeastern  South 
America  in  the  great  counterclockwise  South  Atlantic 
gyral.  Similarly,  larvae  of  E.  pyrifornie  in  the  warm 
Aguihas  Current  are  perhaps  unal)le  to  survive  injection 
into  the  Atlantic  Ocean  via  the  cold  Bengueia  CJurrent. 
.\tlantic  and  Indian  Ocean  populations  of  £.  bairdii  may 
be  tulK  isoialed  1)\  the  same  means.  Unless  hydroiogicai 
conditions  ofl  southwest  .Africa  were  formerly  more  fa- 
vorable, perhaps  during  Pleistocene  interglacials,  £. 
bairdii  and  the  ancestor  of  E.  crosseanum  and  £.  pyri- 
fornie may  have  ranged  through  the  circumtropical  Te- 
thys  Ocean,  in  which  case  isolation  of  eastern  and  western 
populations  would  date  from  the  late  Miocene  following 
establishment  of  the  Middle  Eastern  Landbridge  (Por, 
1986  and  references  therein).  The  second  alternative  is 
favored  here. 

ACKNOWLEDGMENTS 

For  the  loan  of  material  I  am  grateful  to  the  following: 
A.  G.  Beu  (Geology  and  Geophysics  Division,  Lower 
Hutt);  P.  Bouchet  (Museum  National  d'Histoire  Natu- 
relle,  Paris);  D.  Gibbs  (Auckland);  C.  Carpine  (Musee 
Oceanographique,  Monaco);  M.  G.  Harasewych,  R.  S. 
Houbrick  and  S.  Greenhouse  (National  Museum  of  Nat- 
ural History,  Washington,  D.C.);  D.  G.  Herbert  and  R. 
N.  Kilburn  (Natal  Museum,  Pietermaritzburg);  S.  Kool 
and  R.  D.  Turner  (Museum  of  Comparative  Zoology, 
Harvard);  A.  Matsukuma  (National  Science  Museum,  To- 
kyo); W.  F.  Ponder  and  I.  Loch  (Australian  Museum, 
Sydney);  T.  Schiotte  (Zoological  Museum,  Copenhagen); 
Y.  Shibata  and  R.  Yamanishi  (Osaka  Museum  of  Natural 
History);  F.  G.  Thompson  (Florida  State  Museum,  Gains- 
ville);  K.  M.  Way  (The  Natural  History  Museum,  Lon- 
don); F.  E.  Wells  (Western  Australian  Museum,  Perth). 
For  constructive  comments  on  the  manuscript  I  am  grate- 
ful to  A.  G.  Beu,  P.  Bouchet  and  R.  N.  Kilburn.  Special 
thanks  to  R.  Thomson  (Victoria  University,  Wellington) 
for  access  to  the  scanning  electron  microscope;  and  to 
my  staff  colleagues  J.  Lord  and  J.  Nauta  for  photography, 
A.  Marchant  for  printing,  K.  Ryan  for  word  processing, 
and  R.  Webber  for  the  excellent  illustrations  of  Eudolium 
animals. 

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THE  NAUTILUS  106(1  ):39-42.  1992 


Page  39 


Buridrillia  deroyortim.  New  Species  from  the  Galapagos  Islands, 
a  Living  Record  of  a  Neogene  Turrid  Genus 


William  K.  Emerson 

American  Museiiiii  ol  Natural  Histor\ 
Central  Park  West  at  79tl)  Street 
New  York,  NY  1002-1,  USA 


James  H.  McLean 

Natural  Histors  Museum  of  Los 
Angeles  C^ountv 
900  Exposition  Boulevard 
Los  Angeles,  C.\  90007,  USA 


ABSTRACT 

Buridrillia  (U'riiyiiruni  new  species  is  described  from  3 10  meters 
oft  Isla  Floreana  (l>pe  localit>  I  and  365  meters  off  Isla  Duncan, 
Galapagos  Islanils  It  is  the  first  li\ing  species  of  a  turrid  genus 
otherwise  known  from  Neogene  fossils  from  the  New  World 
tropics.  The  genus  is  assigned  to  Crassispirinae  on  shell  and 
radular  characters.  It  has  a  columellar  plication,  evidentK  con- 
vergent with  that  of  Borsoniinae, 

Key  M'ords:  Prosobranch  gastropod;  Turridae;  Clrassisiiirinae; 
Buridrillia  new  species;  Galapagos  Islands,  Ecuador. 


INTRODUCTION 

Andre  and  Jacqueline  DeRoy  were  for  many  years  avid 
collectors  of  Galapagan  marine  mollusks  and  are  knowl- 
edgeable students  of  the  molluscan  fauna  of  the  archi- 
pelago. Mr.  DeRoy  was  a  commercial  fisherman  who 
undertook  numerous  dredging  operations,  and  Mrs. 
DeRoy  spent  much  of  her  time  collecting  in  shallow 
waters.  For  the  past  30  years,  they  have  contributed 
specimens  to  the  American  Museum  of  Natural  History 
and  the  Natural  History  Museum  of  Los  Angeles  County 
for  study  and  report.  Among  these  specimens  is  a  new 
species  of  turrid  gastropod  collected  in  the  1970s.  We 
take  great  pleasure  in  describing  this  new  turrid  in  honor 
of  the  DeRoys. 

The  species  described  here  is  of  more  than  usual  in- 
terest for  two  reasons:  it  represents  the  first  living  record 
of  a  genus  previously  believed  to  have  become  extinct 
in  the  late  Neogene,  and  study  of  its  radular  characters 
results  in  its  assignment  to  a  subfamiK  other  than  that 
which  comes  to  mind  upon  first  examination  of  shell 
morphology. 

Abbreviations  for  institutions  used  in  the  text:  AMNH — 
American  Museimi  of  Natural  History,  New  York; 
LACM — Los  Angeles  County  Museum  of  Natural  His- 
tory, Los  Angeles. 


SYSTEMATICS 

Familv  TURRIDAE  Swainson,  1840 


Subfamily  CRASSISPIRINAE  Morrison,  1966 
Genus  Buridrillia  Olsson,  1942 

T\pe  species  (original  designation):  Clathrodrilla  (Bur- 
idrillia) panarica  Olsson,  1942:51.  Pliocene  of  the  Burica 
Peninsula,  Panama  and  Costa  Rica. 

Olsson  (1964:98)  extended  the  distribution  of  the  type 
species  to  the  Esmeraldas  formation  of  Ecuador,  from 
which  sediments  he  also  described  five  additional  species 
of  Buridrillia.  He  allocated  the  genus  to  "Drillinae"  as 
then  understood,  which  predated  the  proposal  of  Cras- 
sispirinae. 

In  his  revision  of  turrid  classification,  Powell  (1966: 
62)  assigned  Buridrillia  to  the  Borsoniinae,  evidently  based 
on  the  prominent  columellar  plication  of  the  type  species. 
However,  the  radular  tooth  of  the  new  species  B.  deroij- 
orum  is  clearly  of  the  duplex  (terminology  of  Morrison, 
1966)  or  modified  wishbone  type,  comparable  to  that 
illustrated  by  Kantor  and  Taylor  (1991,  fig.  2B)  for  Cli- 
onclla  sinuata  (Born,  1778).  This  tooth  type  is  charac- 
teristic of  (but  not  limited  to)  the  Crassispirinae  (McLean, 
1971;  McLean  in  Keen,  1971).  It  is  substantially  different 
from  the  long  hollow  tooth  of  the  Borsoniinae. 

Shell  characters  other  than  the  columellar  plication  of 
Buridrillia  are  remiiiLscent  of  the  crassispirine  genera 
Ciassispira  (Crassispira)  Swainson  (1840:152,  313),  and 
Hindsiclava  Hertlein  &  Strong  (1955:227),  in  which  there 
is  a  subsutural  cord  on  the  smooth  shoulder,  a  deep  sinus 
on  the  concave  shoulder,  and  axial  and  spiral  sculpture 
on  the  body  whorl.  The  columellar  plication  of  Buri- 
drillia is  evidently  convergent  with  that  of  Borsoniinae. 
Assignment  of  the  genus  to  Crassispirinae  is  supported 
both  on  radular  characters  and  the  balance  of  shell  char- 
acters. 

Buridrillia  deroijonim  new  species 
(figures  1-14) 

Description:  Shell  sturdy,  fusiform,  attaining  57-1-  mm 
in  length.  Spire  acuminate,  whorls  shouldered,  shoulder 
concave,  with  prominent  subsutural  cords;  axial  sculpture 
of  nodose  ribs;  spiral  sculpture  of  depressed  lirae  that 
decussate  the  axial  ribs  to  form  three  rows  of  nodules; 


Page  40 


THE  NAUTILUS,  Vol.  106,  No.  1 


Figures  1-14.  BuridriUia  dcroijorum  new  .specie.s.  1-3.  Holotype,  .AMNH  2321ti3;  4-6.  I'aratype  #8,  LACM  2461;  7,8.  Paratype 
#5,  AMNH  232165  (Kigure.s  l-«,  slightly  reduced,  see  Table  1  for  measurements).  9,  10.  Paratype  #8,  LACM  2461,  spire  enlarged 
about  X  2.5.  1 1,  12.  Paratype  #10,  Operculum,  1 1.  View  of  interior  side,  12.  View  of  exterior  side  (Figures  11,  12,  about  X  2.5). 
13,  ll-.  Paratype  #12,  Raciuiar  (icnlitinn  (Scale  bar  for  1:3  =  lOO^m,  for  14  =  .W^jin). 


body  whorl  with  numerous  spiral  lirae  that  interrupt  the 
axial  sculpture  on  the  anterior  half.  Protoconch  lost  on 
14  of  the  15  specimens,  protoconch  poorly  preserved  on 
paratype  #8  (Table  1),  of  2  1/2  apparently  smooth  w  horls 


(Figures  9,  10);  bod\  whorl  sculptured  with  12  to  13  axial 
ribs.  Aperture  large,  outer  lip  thin,  inflated,  explaned 
with  a  deep,  L'-shaped  anterior  notch,  posterior  notch 
deep,  w  idely  open;  columella  with  a  prominent  plication 


W    K.  Emerson  and  J.  H    McLean.  1992 


Page  41 


Table  1.  Buridrillia  dcroijorum  new  species.  Shell  measure- 
ments in  mm.  Spires  not  preser\'ed,  except  for  parat\pe  #8. 
Number  of  specimens  examined  =  15 


# 

Length 

Widtli 

Whorls 

AMNH  232163 

Holotype 

50.1 

17.8 

7',/2 

AMNH  232164 

Paratvpe  #1 

34.4 

13.7 

5'/2 

Paratvpe  #2 

34.2 

13.0 

5'/2 

Paratvpe  #3 

25.7 

10.5 

5'/2 

AMNH  232165 

Paratvpe  #4 

52.4 

18.7 

7'/2 

Paratvpe  #5 

45.5 

17.1 

6'/2 

Paratvpe  #6 

31,0 

16.1 

61/2 

Parat\'pe  #7 

24,3 

10,7 

6 

LA(.:M  2461 

ParatN'pe  #8 

57.3 

194 

9'/2 

Paratvpe  #9 

56.1 

18.8 

7'/2 

♦Paratvpe  #10 

(operculum) 

48.9 

17.3 

7'/2 

Parat\'pe  #1 1 

35  8 

13  1 

IVi 

*Paratvpe  #12 

(radula) 

22.4 

9.8 

6'/2 

Paratvpe  #13 

15  2 

6  4 

5 ',''2 

Paratxpe  #14 

12,3 

5.1 

5 

Range 

12  3-57  3 

5.1-19.4 

Mean 

:>  1  1 

13  7 

■  Specimens  providmg  radula  and  operculinn  tor  photograpin 


on  the  proximal  part  of  the  pillar.  Periostracum  dense, 
flak\,  greenish-brown.  Shell  color  buff  with  light  tan 
bands  in  the  subsutural  area  and  across  base;  aperture 
glossy  white,  tinged  with  tan.  Operculum,  small,  thin, 
light  brown,  long  (H=7.9  mm)  and  narrow  (W=2.6  mm) 
with  a  marginal  ridge  and  a  terminal  nucleus  (Figures 
II,  12).  Radula  of  both  rows  of  marginal  teeth  only,  of 
the  duplex  or  modified  wishbone  type  (Figures  13,  14). 

Type  locality:  North  of  Isla  Floreana  [also  known  as  Santa 
Maria  Island  or  Charles  Island]  I°I4'S,  90°26'VV,  Gala- 
pagos Islands,  Ecuador  in  310  m,  dredged  bv  the  DeRoys, 
April  15,  1979. 

Specimens  examined:  (All  dredged  by  A.  and  J.  DeRoy, 
in  the  Galapagos  Islands,  see  Table  I  for  measurements): 
Holotype  AMNH  232163  (Figures  1-3)  and  paratypes 
1-3,  AMNH  232164,  from  the  type  locality;  paratypes 
4-7,  AMNH  232165,  from  the  type  locality,  May  15, 
1978,  in  310  m;  paratypes  8-14;  LACM  2461,  off  Isla 
Duncan  [0°25'S,  90°43'W]  in  365  m,  March.  1979. 

Distribution:  Known  only  from  the  Galapagos  Islands 
from  the  t\pe  localitv  and  off  Isla  Duncan,  in  310  and 
365  m. 

Remarks:  Of  the  five  known  Neogene  species  of  Bur- 
idrillia from  the  Esmeraldas  formation  described  bv  Ols- 


son  (1964),  B.  deroyonim  most  resembles  the  type  spe- 
cies, B.  panarica.  from  the  Pliocene  of  Pacific  Panama 
(type  locality)  and  Costa  Rica  (Charco  Azul  formation) 
as  well  as  from  the  Pliocene  of  Ecuador  (Esmeraldas 
formation).  The  two  species  are  of  similar  size  and  have 
a  similar,  well-developed  columellar  plication.  From  B. 
panarica  it  differs  in  having  a  raised  subsutural  cord 
rather  than  a  constricted  subsutural  band,  and  in  having 
much  more  strongK  nodose  axial  ribs. 

The  columellar  plication  of  B.  deroyorum  is  present 
on  all  fifteen  specimens  regardless  of  size.  However,  the 
strength  of  its  continuation  on  the  internal  pillar  may 
differ.  A  strong  plication  is  visible  through  a  hole  on  the 
back  side  of  paratspe  10  made  to  extract  the  radula. 
However,  the  pillar  in  Paratype  8  (Figures  5,  6),  which 
is  visible  through  a  naticid  drill  hole,  has  a  very  taint 
plication.  Similar  variation  has  been  noted  for  fossil  spe- 
cies of  Buridrillia.  Olssoii  (1964:98)  pointed  out  that  some 
specimens  of  the  type  species  have  a  well-marked  col- 
umellar fold,  but  that  others  lack  it  or  have  slight  swell- 
ings or  raised  lines.  Other  species  described  by  Olsson 
have  poorly  marked  folds  or  lack  folds 

The  present  distribution  of  Buridrillia  is  that  of  a  relict 
genus,  a  survivor  in  the  eastern  Pacific  at  the  Galapagos 
refugium  of  a  turrid  group  that  was  once  more  widely 
distributed  in  the  New  World  tropics  during  the  Neo- 
gene. 


ACKNOWLEDGMENTS 

We  are  grateful  to  Andre  and  Jacqueline  DeRoy  of  Isla 
Santa  Cruz,  Galapagos  Islands,  for  donating  the  type 
specimens  to  our  respective  institutions.  We  gratefully 
acknow  ledge  the  assistance  of  our  colleagues:  Walter  E. 
Sage,  III,  Stephanie  Crooms  and  Andrew  S.  Modell  of 
the  AMNH  provided  technical  assistance,  word-process- 
ing and  photographic  services,  respectively;  C,  Clifton 
Coney  of  the  LACM  operated  the  electron  microscope 
at  the  I'niversit)  of  Southern  California.  We  thank  Yuri 
I.  Kantor  of  the  A.  N.  Severtzov  Institute  of  Animal 
Evolutionary  Morphology  and  Ecology,  Moscow  for 
commenting  on  a  draft  ol  the  manuscript. 

LITERATURE  CITED 

Born.  1  1778  Index  Rerum  Naturalium  Musei  Caesarei  V'in- 
dobonensis.  Part  1;  Testacea.  Vienna,  i-xlii  +  1-458  p 

Hertlein,  L.  G.  and  A.  M.  Strong.  1955.  Marine  mollusks 
collected  during  the  Askoy  Expedition  to  Panama,  Colom- 
bia, and  Ecuador  in  1941.  Bulletin  of  the  American  Mu- 
seum of  Natural  History  107(2):159-318. 

Kantor,  Y.  I.  and  J.  D.  Taylor.  1991.  Evolution  of  the  toxo- 
glossan  feeding  mechanism:  new  information  on  the  use 
of  the  radula.  Journal  of  Molluscan  Studies  57(11:129-134. 

Keen,  .\.  M  1971.  Sea  Shells  of  Tropical  West  .\merica  ed 
2,  Stanford  University  Press,  xiv  -I-  1064  p. 

McLean,  J.  H.  1971.  A  revised  classification  of  the  family 
Turridae,  with  the  proposal  of  new  subfamilies,  genera, 
and  subgenera  from  the  eastern  Pacific.  The  V'eliger  14(1): 
114-  130. 


Page  42 


THE  NAUTILUS,  Vol.  106,  No.  1 


Morrison,  J,  P  E.    1966,    On  the  hiMiilii-sol  Turridae,  American 

Malacological  L'nioii,  Annual  Reports,  tor  1965: 1-2, 
Olsson,  .'\.  .\.    1942,    Tertiars  atitl  (Quaternary  fossils  from  the 

Buriea  Peninsula  of  Panama  and  Costa  Hiea   Bulletins  oi 

Amerieau  Paleoiitolog)  27(  106):  1-106, 
Olsson,  A.   ,\,     1964,     Neogene  moiiusks  Irom   northwestern 

Ecuador,  Paleoiitological  Research  Institution,  Ithaca.  New 

York,  256  p,,  38  pis. 


Powell,  A,  W  B  1966,  The  moliuscan  families  Speightiidae 
and  Turridae  Bulletin  of  the  .Aucklanil  Institute  and  Mu- 
seum. 5: 184  p.,  23  pis, 

Swainsou,  W,  1840,  .\  Treatise  on  Malacology  or  the  Natural 
C;lassification  of  Shells  and  Shell-fish,  London,  419  p. 


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rHE  NAUTILUS 


Volume  106,  Number  2 
June  5,  1992 
ISSN  0028-1344 

A  quarterly  devoted 
to  malacology. 


Marine  Biological  Laboratory 
LIBRARY  I 

JUN  1  5  1992 


EDITOR-IN-CHIEF 
Dr.  M.  G.  Harase-wyeh 
Division  oi  Mollusks 
National  Museum  of 
Natural  Histor\ 
Smithsonian  Institution 
Washington,  DC  20560 


ASSOCIATE  EDITOR 
Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
P  O.  Ron  2255 
Melbourne,  FL  32902 


CONSULTING  EDITORS 

Dr   Riidiger  Bieler 
Department  of  Invertebrates 
Field  Museum  of 
Natural  History 
C:hicago,  IL  60605 


Dr.  Robert  T.  Dillon,  Jr. 
Department  of  Biology 
College  of  (;harleston 
Charleston,  SC:  29424 


Dr.  William  K.  Emer.son 

Department  of  Living  Invertebrates 

The  American  Museum  of  Natural 

History 

New  York,  NY  10024 


Dr.  Robert  Hershler 
Division  of  Mollusks 
National  Museum  of 
Natural  Histors' 
Smithsonian  Institution 
Washington,  DC  20560 

Dr.  Richard  S    Houbrick 
Division  ot  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC:  20560 


Mr.  Richard  I.  Johnson 
Department  of  Mollusks 
Museum  of  ("omparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Department  of  Malacology 
Los  Angeles  County  Museum  of 
Natural  History 
900  Exposition  Boulevard 
Los  Angeles,  CA  90007 

Dr.  Arthur  S.  Merrill 
%  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Ms.  Paula  M.  Mikkelsen 
Harbor  Branch  Oceanographic 
Institution,  Inc. 
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Dr.  Donald  R.  Moore 

Division  of  Marine  Geology 

and  Geophysics 

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Atmospheric  Science 

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Marine  Laboratory 
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Dr.  Ruth  D.  Turner 
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Museum  of  Comparative  Zoology 
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Davis,  CA  95616 


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THEt^NAUTILUS 


CONTENTS 


Volume  106,  Number  2 
June  5,  1992 

ISSN  0028-1344 

Marine  Biological  lahnr^tf;;^'] 
LIBRARY 

Two  new  species  of  Cancellariidae  (Gastropocfc:            JUN  1  S  IQQ? 
Neogastropoda)  from  Brazil   1 43 

New  species  of  Solariella  (Gastropoda:  Trochidae)  from  the 

western  Atlantic  Ocean         50 

A  new  fossil  land  snail  of  the  genus  Hemitrochus  from 

Bowden,  Jamaica  55 


M.  G.  Harasewych 
Richard  E.  Pelil 
Andre  V  erhecken 


James  F.  Quinn,  Jr. 


Glenn  A.  Goodfriend 


R.  Douglas  Hunter 
John  F.  Bailey 

Dreissena  pohjmorpha  (Zebra  mussel):  Colonization  of  soft 
substrata  and  some  effects  on  unionid  bivalves 

60 

Edward  J.  Petueh 

New  Ecphoras  (Gastropoda:  Thaididae:  Ecphorinae)  from 

the  Calvert  Formation  of  Maryland  (Langhian  Miocene)       

68 

Elizabeth  Potter 
Michael  A.  Rex 

Parallel  development-depth  trends  in  deep-sea  torrid  snails 

from  the  eastern  and  western  north  Atlantic    

72 

John  K.  Tucker 

Drillia  maclcani,  new  name  for  Drillia  sintiosa  McLean 

and  Poorman,  1971  (Gastropoda:  Turridae)     

76 

Notices 


58th    ANNUAL    MEETING    OF    THE    AMERICAN 
MALACOLOGICAL  UNION 

The  58th  annual  meeting  of  the  American  Malacolog- 
ical  Union  will  be  held  August  2-7,  1992,  at  the  Hyatt 
Sarasota,  Sarasota,  Florida.  Local  assistance  is  being  pro- 
vided by  the  Sarasota  Shell  Club  and  the  Mote  Marine 
Laboratory.  The  meeting  will  feature  contributed  pa- 
pers, bourse  and  exhibits,  a  collectors'  evening,  auction, 
banquet,  and  fieldtrips  to  marine,  freshwater,  and  ter- 
restrial habitats  and  a  nearby  fossil  pit.  Two  international 
symposia  are  planned:  Biology  of  Caribbean  Mollusks, 
organized  by  Dr.  Riidiger  Bieler  [Department  of  Zool- 
ogy, Field  Museum  of  Natural  History,  Roosevelt  Road 
at  Lake  Shore  Drive,  Chicago,  IL  60605,  Tel.  (312)  922- 
9410,  ext.  270,  FAX  (312)  663-5397],  and  Advances  in 
Gastropod  Phylogeny,  organized  by  Dr.  Terrence  M. 
Gosliner  [Department  of  Invertebrate  Biology  and  Pa- 
leontology, California  Academy  of  Sciences,  Golden  Gate 
Park,  San  Francisco,  CA  94118,  Tel.  (415)  750-7277,  FAX 
(415)  750-7090].  Information,  registration  forms,  and  a 
call  for  papers  will  be  included  in  the  Spring  1992  issue 
of  the  AMU  Newsletter.  Additional  information  about 
the  meeting  and  membership  applications  can  be  ob- 
tained by  contacting: 

Dr.  Robert  C.  Bullock,  President 
American  Malacological  Union 
Department  of  Zoology 
Biological  Sciences  Building 
University  of  Rhode  Island 
Kingston,  RI  02881 
Tel.  (401)  792-2372 
FAX  (401)792-4256 


WESTERN  SOCIETY  OF  MALACOLOGISTS 

The  Twenty-fifth  Annual  Meeting  of  the  Western  So- 
ciety of  Malacologists  will  be  held  at  the  Asilomar  Con- 
ference Grounds  on  Monterey  Bay,  Pacific  Grove,  Cal- 
ifornia, from  June  30  to  July  3,  1992.  In  addition  to 
contributed  papers  on  marine,  freshwater  and  terrestrial 
mollusks,  both  living  and  fossil,  the  agenda  includes  two 
special  symposia,  on  Opisthobranchs,  and  on  Cocos  Is- 
land, Costa  Rica.  A  shell  auction,  reprint  sale,  and  ban- 
quet round  out  the  program.  For  further  information 
contact  WSM  President  David  K.  Mulliner  (5283  Vickie 
Drive,  San  Diego,  California  92109;  (619)  488-2701)  or 
WSM  Treasurer  Henry  W.  Chane\  (Museum  of  Natural 
History,  2559  Puesta  del  Sol  Road,  Santa  Barbara,  Cal- 
ifornia 93105;  (805)  682-4711,  ext.  334;  fax  (805)  569- 
3170).  Deadline  for  receipt  of  abstracts  is  May  15,  1992. 


October  11-14,  1992 — Symposium  on  Freshwater  Mus- 
sel Conservation  and  Management.  Embassy  Suites  Ho- 
tel, St.  Louis,  MO.  Sessions  devoted  to  regulations,  com- 
mercial harvest,  conservation,  sampling  methodology, 
data  management,  and  environmental  awareness.  For 
info:  Kurt  Welkc,  Wisconsin  Dept.  of  Natural  Resources, 
11 1  W.  Dunn  St.,  Prairie  du  Chien,  Wl  53821,  608/326- 
0233. 


THE  NAUTILUS  106(2):43-49,  1992 


Page  43 


Two  New  Species  of  Cancellariidae 
(Gastropoda:  Neogastropoda)  from  Brazil 


M.  G.  Harasewych 
Richard  E.  Petit 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC  20560  USA 


Andre  Verhecken 

Scientific  Associate 
Malacology  Section 
Koninklijk  Belgisch  Instituut 
voor  Natuurwetenschappen 
B- 1040  Brussels,  BELGIUM 


ABSTRACT 

Two  new  species  of  cancellariids  are  described  from  shallow 
waters  of  the  Brazilian  Province.  Each  has  a  sister  species  in 
the  Caribbean  Province.  Tritonoharpa  leali,  n.  sp.  differs  from 
T.  lanceolala  in  having  a  more  inflated  and  thinner  shell,  in 
which  the  axial  costae  are  more  pronounced  than  the  spiral 
cords.  Cancellaria  petuchi  ii.  sp  differs  from  C.  reticulata  in 
having  a  thick,  w  hite  callus  on  the  columellar  side  of  aperture, 
a  broad,  rounded  shoulder,  and  a  proportionalK  longer  aperture 
and  larger  protoconch  than  C.  reticulata.  CaJicclUiria  petuclii 
lacks  strongU  cancellated  sculpture  on  the  bod\  whorl  and  the 
bifid  posteriormost  columellar  fold  of  C.  reticulata 

Key  words:  Cancellariidae;  Tritonoharpa;  Cancellaria.  new 
species;  Brazil. 


INTRODUCTION 

The  nioiluscan  fauna  (if  Brazil  has  been  regarded  bv 
mo.st  authors  (e.g.,  Abbott,  1974;  Rios,  1970,  1975,  1985) 
as  comprising  the  southernmost  component  of  the  Ca- 
ribbean Province.  Several  workers,  among  them  Ekman 
(1953),  Work  (1969)  and  Rios  (1970),  had  commented 
on  an  endemic  component  of  the  Brazilian  molluscan 
fauna,  while  others  (Briggs,  1974;  Coelho  &  Koening, 
1972;  Coelho  &  Ramos,  1972)  had  suggested  the  existence 
of  a  Brazilian  Subprovince  or  Province.  More  recentK , 
Petuch  (1988:166)  delimited  the  region  extending  from 
the  mouth  of  the  Amazon  River  southward  into  the  Mar 
de  Plata  as  the  Brazilian  Province  and  listed  a  number 
of  endemic  molluscan  indicator  species.  Other  authors 
(e.g.,  Yokes,  1990)  have  begun  to  distinguish  between 
Brazilian  species  and  their  often  closely  related  Carib- 
bean counterparts. 

The  cancellariid  fauna  of  Brazil  is  poorK  known.  Lange 
de  Morretes  (1949,  1953)  did  not  include  any  member 
of  this  family  in  his  catalogs  of  Brazilian  mollusks,  while 
Abbott  (1974:246)  and  Rios  (1970:111,  1975:127,  1985: 
126)  listed  only  Cancellaria  reticulata  Linne,  1767  and 
Tritonoharpa  lanceolata  (Menke,  1828)  (as  Cohibraria 
lanceolata)  as  occurring  in  the  Brazilian  fauna.  Ver- 
hecken (1991)  underscored  the  paucity  of  records  for  this 


family  in  the  fauna  of  the  subequatorial  western  Atlantic, 
and  described  two  bathyal  cancellariids  from  off  south- 
eastern Brazil. 

A  large  series  of  specimens  of  an  undescribed  Can- 
cellaria as  well  as  two  specimens  of  a  new  species  of 
Tritonoharpa  were  recently  made  available  to  us  from 
commercial  and  institutional  sources.  We  take  pleasure 
in  naming  these  new  species  after  the  persons  who  first 
brought  them  to  our  attention. 


ABBREVIATIONS  USED  IN  TEXT 

KBIN — Koninklijk  Belgisch  Instituut  voor  Natuurweten- 
schappen,  Brussels 

MNHN — Museum  national  d'Histoire  naturelle,  Paris 

MORG — Museu  Oceanografico  da  Funda^ao  Universi- 
dade  do  Rio  Grande,  Rio  Grande,  Brazil. 

UFJF — Universidade  Federal  de  Juiz  de  Fora,  Minas 
Gerais 

USNM — National  Museum  of  Natural  History,  Smith- 
sonian Institution,  Washington,  DC 

SYSTEMATICS 

Family  Cancellariidae  Forbes  &  Hanley,  1851 
Subfamily  Plesiotritoninae  Beu  &  Maxwell,  1987 

Genus  Tritonoharpa  Dall,  1908 

Beu  and  Maxwell  (1987)  reviewed  the  genus  Tritono- 
harpa, enumerating  the  following  diagnostic  characters: 
elongate-ovate  to  biconic  shells  with  spire  half  or  more 
the  shell  length;  weakly  to  moderately  convex  teleoconch 
whorls;  prominent  non-collabral  varices  on  early  as  well 
as  later  whorls;  well  developed  columellar  collar;  colu- 
mellar plait  either  single  and  weak  or  absent;  radula 
absent. 

Recent  western  Atlantic  species  of  Tritonoharpa  in- 
clude the  Carolinian  and  Caribbean  T.  lanceolata 
(Menke,  1828);  T.  cubapatriae  (Sarasua,  1975),  based  on 
a  unique  holotype  from  off  Havana,  Cuba;  T.  baijeri 
(Petuch,  1987)  from  the  Caribbean  coast  of  Colombia; 


Page  44 


THE  NAUTILUS,  Vol.  106,  No.  2 


M.  G.  Harasewvch  et  al.,  1992 


Page  45 


Figures  5-6.  Tritonoharpa  leali  new  species.  5.  Protoconch  of  holotype,  scale  bar  =  500  ^m.  6.  Surface  sculpture  of  holotype, 
scale  bar  =  200  ^m.  Figures  7-8.  Tritonoharpa  lanceolata  (Menke,  1828).  7.  Protoconch  of  specimen  in  figure  3,  scale  bar  = 
500  ^m.  8.  Surface  sculpture  of  specimen  in  figure  3,  scale  bar  =  200  nm. 


and  the  new  species  described  below.  Minihraria  mon- 
roei  (McGinty,  1962)  has  been  referred  to  the  Muricidae 
on  the  basis  of  the  morphology  of  its  varices  and  pro- 
toconch (Beu  &  Maxwell,  1987:56). 

Tritonoharpa  leali  new  species 
(figures  1,  2,  5,  6) 

Description:  Shell  (figures  1-2)  to  18.4  mm,  thin,  tall, 
narrow.  Spire  high  (spire  angle  31-35°),  stepped,  com- 
prising about  %  of  total  shell  length.  Protoconch  paucis- 


piral,  of  2y4  inflated,  weakly  shouldered  whorls  (figure 
5).  First  l'/2  whorls  nearly  planispiral,  descending  rapidly 
thereafter.  Protoconch  coaxial  with,  or  deviated  from, 
teleoconch  axis  by  up  to  5°  Transition  to  teleoconch 
abrupt,  marked  by  flared  protoconch  lip  and  onset  of 
pronounced  spiral  sculpture.  Teleoconch  with  up  to  5% 
slightly  inflated,  weakly  shouldered  whorls.  Suture  deep- 
ly impressed,  obscured  by  axial  costae  of  succeeding 
whorl.  Spiral  sculpture  of  strong,  low,  equally  spaced 
cords,  20-26  on  bod\'  whorl,  12-15  on  penultimate  whorl; 
cords,  0-3,  weak  or  absent  on  siphonal  canal.  Axial  sculp- 


Figures  1-2.  Tritonoharpa  leali  new  species.  1.  Holotype,  MORG  286.59,  Davis  Bank,  southeastern  Brazil,  20°40'S,34°41'W,  in 
60  m,  Marion-Dufresne  MD55  sta  DC40,  7.5  x.  2.  Paratype  1,  USNM  860521,  off  Itaparica  Island.  Bahia,  Brazil,  trawled  in  20 
m,  5.0x.  Figure  3.  Tritonoharpa  lanceolata  (Menke,  1828).  USNM  798073.  Off  St.  Thomas,  Virgin  Islands,  5.0x.  Figure  4. 
Tritonoharpa  haijeri  (Petuch.  1987),  Holotype,  USNM  859853,  Cabo  La  Vela.  Goajira  Peninsula,  Colombia,  in  35  m,  5.0 x. 


Page  46 


THE  NAUTILUS,  Vol.  106,  No.  2 


Figures  9-10.  Camcllaria  pcluchi  new  species.  *>.  Ilolotvpe,  MORG  286(i0,  2.0 x  10.  Paralvpe  4.  USNM  859409,  Off  Vitoria, 
Espirito  Santo,  Brazil,  trawled  on  sand  bottom  in  40  ni,  2  Ox  Figure  11.  Caiicellaria  adclac  Pilsbrs,  1940,  USNM  508663,  Little 
Duck  Key,  Florida,  1  5x.  Figure  12.  CanccUaria  rclirulata  (Linne,  1767),  LISNM  811479,  30  mi.  ENE  of  Eau  Gallie,  Florida 
[28''17'N,'80°01'W],  in  34-41  fms  (62-75  m),  K/V  Silver  Bav  Sta   2010,  2.0x. 


lure  of  pronounced  varices  and  narrow,  opistliocline  cos- 
tae.  Varices  high,  non-collabral,  narrow,  rounded  aba- 
pcrturally,  undercut  adaperturally,  180-200°  apart  on 
early  whorls,  up  to  238°  apart  on  later  whorls.  Costae  (7- 
22  between  varices,  15-33  per  whorl)  offset  from  varices 
by  5-10°,  producing  cancellate  sculpture  ar  their  inter- 
section with  spiral  cords.  Aperture  narrow,  elliptical,  with 


well  defined  posterior  notch.  Inner  lip  smooth,  \\ith  col- 
uiuellar  collar  reflected  over,  but  not  adherent  to,  fascicle 
and  p.seudouinbilicus.  Outer  lip  with  up  to  8  pairs  of 
denticles  confined  to  ilaretl  region  beyond  varix.  Siphonal 
canal  short,  dorsally  reflected,  partially  covered  abaxi- 
ally.  Rase  color  khaki  to  yellowish  tan,  with  darker  brown 
maculations  anterior  to  suture  and  along  varices. 


M.  G.  Harasewych  et  ai,  1992 


Page  47 


Table  1.  Shell  measurements  of  Canccllaria  reticulata  and  C.  petuchi.  All  measurements  in  mm  (N  =  10).  For  C.  reticulata,  the 
10  specimens  were  selected  from  throughout  the  species  range,  while  for  C.  petuchi.  data  is  based  on  specimens  from  a  single  lot 
(paratypes  4-13). 


Shell  length  (SL) 

Aperture  length  (AL) 

(AL/SL) 

No.  whorls,  teleoconch 

No.  whorls,  protoconch 

Diameter,  protoconch 

Diameter  no.  whorls,  protoconch 


c 

re  tic 

ulat 

a 

C. 

petuchi 

X 

(J 

X 

a 

39.67 

6.21 

33.68 

1-98 

25.64 

3.99 

23.64 

1.33 

0.65 

0.03 

0.70 

0.01 

6.34 

1.76 

5.51 

0.20 

2.67 

0.10 

2.74 

0.06 

1.23 

0.06 

1.60 

0.05 

0.46 

0.02 

0.59 

0  02 

Type  locality:  Davis  Bank,  southeastern  Brazil, 
26°40'S,34°41'W,  in  60  m,  Marion-Dufresne  MD.55  sta. 
DC40.  May  1987. 

Material  examined:  Holotype,  MORG  286.59,  9.22  mm; 
Paratype  1,  USN'M  860521,  off  Itaparica  Island,  Bahia, 
Brazil,  trawled  in  20  m,  18.44  mm. 

Distribution:  This  species  is  known  from  two  records, 
both  off  the  state  of  Bahia,  Brazil,  in  depths  of  from  20 
to  60  m. 

Etymology:  This  species  honors  Dr.  Jose  H.  Leal,  in 
recognition  of  his  contributions  to  the  study  of  Brazilian 
mollusks. 

Comparative  remarks:  Tritonoharpa  leali  most  closely 
resembles  T.  lanceolata,  from  which  it  differs  in  having 
a  somewhat  more  inflated  and  thinner  shell,  in  which 
the  axial  costae  are  more  pronounced  than  the  spiral 
cords  (compare  figures  5  and  7).  Tritonoharpa  haijeri  is 
even  broader  and  thinner  than  T.  leali,  and  is  the  only 
western  Atlantic  Tritonoharpa  in  which  the  aperture 
length  exceeds  half  the  shell  length. 


Subfamily  Canceilariinae  Forbes  &  Hanley, 
Genus  Cancellaria  Lamarck,  1799 


1851 


The  genus  Cancellaria  is  characterized  by  the  presence 
of  a  thick,  ovate  to  biconic  shell  with  strongly  cancellate 
sculpture,  prosocline  outer  lip,  a  weak  stromboid  notch, 
short  but  distinct  siphonal  canal,  prominent  siphonal  fas- 
cicle, lirate  outer  lip,  and  columella  with  two  to  three 
strong  folds,  the  posteriormost  strongest  and  usually  bifid. 
Jung  and  Petit  (1990:100)  reduce  Pijniclia  Olsson,  19.32 
to  subgeneric  status  within  Cancellaria,  and  distinguish 
it  from  the  nominotypical  subgenus  on  the  basis  of  its 
pyriform  shape  with  reduced  or  absent  sculpture  on  body 
whorl  and  a  columella  with  two  strong,  non-bifid  folds. 

Cancellaria  petuchi  new  species 
(figures  9,  10,  14-16) 

Description:  Shell  (figures  9-10)  to  37  mm,  heavy  bi- 
conic, pseudoumbilicate.  Spire  high  (spire  angle  62-70°), 
conical,  comprising  about  %  of  total  shell  length.  Pro- 
toconch paucispiral,  of  2'/2  low,  inflated,  glassy  whorls 


(figure  14).  Protoconch  may  be  co-axial  with,  or  deviated 
from  teleoconch  axis  by  up  to  15°.  Teleoconch  with  up 
to  6%  strongly  convex  whorls.  Suture  deeply  impressed. 
Spiral  sculpture  of  18-24  major  cords  on  body  whorl,  6- 
10  on  siphonal  canal,  6-8  on  penultimate  whorl.  Axial 
sculpture  of  20-38  prosocline  ribs  that  form  strongly 
cancellate  sculpture  at  intersection  with  spiral  cords,  es- 
pecially on  first  four  teleoconch  whorls.  Thereafter,  axial 
ribs  become  broader,  less  pronounced,  may  disappear 
entirely,  especialK'  below  the  shoulder.  Aperture  large, 
broad,  hemi-elliptical,  deflected  from  coiling  axis  by  12- 
18°  Outer  lip  with  shallow  indentation  posterior  to  junc- 
tion with  siphonal  canal  and  10-12  strong  lirae,  slightly 
recessed,  either  diminishing  Vi  whorl  into  aperture,  or 
more  usually,  disappearing  and  reappearing  V4  whorl 
within  the  aperture.  Inner  lip  with  2  columellar  and  1 
siphonal  folds.  Posteriormost  columellar  fold  largest, 
overlying  siphonal  fasciole.  All  folds  with  single,  sharp 
keel.  Siphonal  canal  short,  deflected  dorsally  and  abax- 
ially.  Base  color  white  to  light  tan,  with  spiral  bands 
ginger  to  dark  brown  interrupted  by  narrow  axial  bands 
of  base  color,  especially  on  early  whorls.  Some  specimens 
pure  white.  Aperture  with  white  overglaze  forming  thick 
callus  along  columella. 

Type  locality:  N.E.  of  Vitoria,  Espirito  Santo,  Brazil, 
19°35'S,  39°42'W,  in  15  m.  South  Eastern  Brazil  N.O. 
Marion-Dufresne  MD55  sta.  DC87,  May  1987. 

Material  examined:  Holotype,  MORG  28660,  28. 16  mm; 
Paratypes  1-3,  MNHN,  N.E.  of  Vitoria,  Espirito  Santo, 
Brazil,  19°34'S,39°34'W,  in  34  m,  N.O.  Marion-Dufresne 
MD55  sta.  CB90,  May  1987,  32.52  mm,  11.01  mm,  8.06 
mm;  Paratypes  4-23,  USNM  859409,  Paratypes  24-36, 
Petit  collection,  Paratypes  37-38,  KBIN,  Paratypes  39- 
52,  Verhecken  collection,  all  from  Off  Vitoria,  Espirito 
Santo,  Brazil,  trawled  on  sand  bottom  in  40  m,  29.26- 
36.56  mm.  Paratypes  53-.55,  Petit  collection,  Mangue  da 
Olaria,  Guarapari,  Espirito  Santo,  Brazil,  in  sand,  minus 
tide.  Paratypes  56-57,  UFJF,  Niteroi,  Rio  de  Janeiro, 
Brazil.  Paratype  58,  MORG  19472,  off  Salin6polis,  Pard, 
Brazil,  in  36  m.  Paratypes  59-62,  MORG  23307,  Coroa 
Vermeiha,  Abrolhos  Islands,  Bahia,  Brazil,  in  1  m. 

Distribution:  This  species  ranges  from  off  Salinopolis, 
Para,  to  off  Niteroi,  Rio  de  Janeiro,  Brazil.  It  occurs  from 
intertidal  depths  to  40  m. 


Page  48 


THE  NAUTILUS,  Vol.  106,  No.  2 


Figure  13.  Cancellaria  reticulata  (Linne,  1767).  Protoconch,  scale  bar  =  500  ^m.  Figure  14-16.  Cancellaria  petuchi  new- 
species.  14.  Protoconch  of  paratype,  same  data  as  fig.  10.  Scale  bar  =  500  iim.  15.  Frontal  and  16.  Lateral  view  of  distal  ends  of 
rachidian  teeth.  Scale  bars  =  5.0  nm  (fig.  15),  3.0  ^m  (fig.  16). 


Etymology:  This  species  honors  Dr.  Edward  J.  Petuch, 
as  a  tribute  to  his  contributions  to  our  knowledge  of  the 
inoUuscan  fauna  of  Brazil,  and  the  Neogene  zoogeog- 
raphy of  the  western  Atlantic  faunas. 

Comparative  remarks:  Cancellaria  pettichi  differs  from 
its  geographically  pro.xiniate  congener  C  reticulata  in 
having  reduced  or  absent  cancellate  sculpture  on  the 
body  whorl  and  lacking  a  bifid  posteriormost  columellar 
fold.  Cancellaria  petuchi  has  a  thick,  white  callus  on  the 
columellar  side  of  the  a|)erture  and  a  broad,  rounded 
shoulder,  features  lacking  in  C.  reticulata.  Cancellaria 
petuchi  also  has  a  proportionally  longer  aperture  and 
larger  protoconch  than  C.  reticulata  (table  1).  Radular 
morphology  of  Cancellaria  petuchi  (figures  15,  16)  agrees 
in  all  essential  features  with  that  of  C.  reticulata  (Har- 
asewych  &  Petit,  1982:figures  14,  15 — these  figures  show 
the  ventral  surfaces  of  the  distal  tips  of  two  radular  teeth). 
Of  the  western  Atlantic  species  of  Cancellaria,  C.  pe- 
tuchi most  strongly  resembles  C.  adelae  Pilsbry,   1940 


(figure  11),  a  species  endemic  to  the  Florida  Keys,  with 
which  it  shares  its  predominantly  smooth  body  whorl 
sculpture  and  thick  parietal  overglaze.  Cancellaria  ade- 
lae however,  has  a  strongK  bifid  columellar  fold,  and 
indeed,  may  have  an  additional  fold  along  the  posterior 
portion  of  columella. 

ACKNOWLEDGMENTS 

We  thank  Dr.  Philippe  Bouchet  (MNHN),  Dr.  Edmund 
Gittenberger,  (Nationaal  Natuurhistorisch  Museum,  Lei- 
den), Robert  Moolenbeek  (Instituut  voor  Taxonomische 
Zoologie,  Amsterdam),  Prof.  Maury  Pinto  de  Oliveira 
(UFJF),  Prof.  Elesar  C.  Rios  (MORG),  and  Dr.  Edward 
J.  Petuch,  (Florida  .Atlantic  I'niversity,  USA)  for  making 
specimens  available  for  study.  Drs.  Edward  J.  Petuch 
and  Jose  H.  Leal  critically  read  drafts  of  the  manuscript 
and  provided  some  of  the  references  on  the  biogeography 
of  the  Brazilian  fauna.  Ms.  Susanne  Braden  provided  the 
SEM  micrographs. 


M.  G.  Harasewych  et  a/.,  1992 


Page  49 


LITERATURE  CITED 

Abbott,  R  T.  1974,  American  seashells,  second  edition.  Van 
Nostrand  Reinhold,  Co.,  New  York.  663  p. 

Beu,  A.  G.  and  P  A  Ma.wveli.  1987  A  revision  of  tossij  and 
living  Gastr<ipods  related  to  Plesiotritun  Fisciier,  1884 
(Family  Cancellariidae,  SubfamiK  Plesiotritoninae  n. 
subfam.)  with  an  appendix:  Genera  of  Buccinidaee  Pisa- 
niinae  related  to  Colubraria  Schumacher,  1817.  New  Zea- 
land Geological  Survey  Paleontological  Bulletin  54.1-140, 
pis.  1-30. 

Briggs,  J.  C.  1974.  Marine  Zoogeography.  McGrav\-Hill,  New 
York.  475  p. 

Coelho,  P.  A.  and  M  L  Koening.  1972.  .\  distribui9ao  dos 
crustaceos  pertencentes  as  ordens  Stoniatopoda,  Tanai- 
dacea  e  Isopoda  no  Norte  e  Nordeste  do  Brasil,  Trabalhos 
Oceanograficos  da  Universidade  Federal  de  Pernambiico 
13:245-260. 

Coelho,  P.  A.  and  M  A.  Ramos.  1972.  A  constitu9ao  e  dis- 
tribui(jao  da  fauna  de  decapodos  crustaceos  do  litoral  leste 
da  America  do  Sul  entre  as  latitudes  de  5°N  e  39°S.  Tra- 
balhos Oceanograficos  da  Universidade  Federal  de  Per- 
nambuco  13:133-236, 

Ekman,  S.  1953.  Zoogeography  of  the  Sea.  Sidgwick  and 
Jackson,  London  .\iv  -I-  417  p. 

Harasewych,  M.  G.  and  R.  E.  Petit.  1982  Notes  on  the  mor- 
phology of  Cancellaria  reticulata  (Gastropoda:  Cancel- 
lariidae). The  Nautilus  96(3):104-113. 

Jung,  P.  and  R.  E.  Petit.  1990.  Neogene  Paleontology  in  the 
northern  Dominican  Republic,  10  The  family  Cancellar- 
iidae (Mollusca:  Gastropoda),  Bulletins  of  American  Pa- 
leontology 98(334):87-144, 


Lange  de  Morretes,  F.    1949.    Ensaio  de  catalogo  dos  moluscos 

do  Brasil   Arquivos  do  museu  Paranaense  7(1):5-216. 
Lange  de  Morretes,  F,     1953.    Adenda  e  corrigenda  au  ensaio 

de  catalogo  dos  moluscos  do  Brasil,  .Arquivos  do  Museu 

Paranaense  10(l):37-76, 
McGinty,  T,  L,    1962.    Caribbean  marine  shells.  The  Nautilus 

76(2):39-44,  pl.3. 
Petuch,  E.  J.    1987.    New  Caribbean  Molluscan  Faunas.  The 

Coastal  Education   &   Research  Foundation,  Charlottes- 
ville. 1-154,  A1-A4. 
Petuch,  E.  J      1988.     Neogene  history  of  tropical  American 

mollusks.  The  Coastal  Education  &  Research  Foundation, 

Charlottesville.  217  p. 
Pilsbrv,  H.  A.    1940.    A  new  race  of  Cancellaria  from  Florida. 

The  Nautilus  54(2):54. 
Rios,  E.  C.    1970.    Coastal  Brazilian  Seashells.  Museu  Ocean- 

ografico,  Rio  Grande.  255  p.,  60  pis. 
Rios,  E.  C.     1975.     Brazilian  Marine  Mollusks  Iconography. 

Museu  Oceanografico,  Rio  Grande.  331  p.,  91  pis 
Rios,  E.  C.     1985.    Seashells  of  Brazil.  Museu  Oceanografico, 

Rio  Grande.  328  p.,  102  pis. 
X'erhecken,  A     1991.    Description  of  two  new  speciesof  bathy- 

al  Cancellariidae  (Mollusca,  Gastropoda)  from  off  Brazil. 

Bulletin  Museum  national  d  Histoire  naturelle,  Paris.  Se- 
ries 4,  12,  Section  A  (3-4):547-553. 
\'okes,  E.  H.    1990.    Two  new  species  of  Chicoreus  subgenus 

Stratus  (Gastropoda:  Muricidae)  from  northeastern  Brazil. 

The  Nautilus  103(4):124-130. 
Work,  R.  C,     1969,    Systematics,  ecology,  and  distribution  of 

the  mollusks  of  Los  Roques,  Venezuela.  Bulletin  of  Marine 

Science  19(3):614-711. 


THE  NAUTILUS  106(2):50-54,  1992 


Page  50 


New  Species  of  Solariella  (Gastropoda:  Trochidae)  from  the 
Western  Atlantic  Ocean 


James  F.  Quinn,  Jr. 

Florida  Marine  Research  Institute 
Department  of  Natural  Uesiiurces 
100  Eighth  Avenue,  SE 
St.  Petersburg,  FL  33701-5095,  USA 


ABSTRACT 

Three  new  species  of  the  genus  Sulariclhi  Wood,  1842,  are 
described.  Solariella  qtiadricincta  is  known  only  from  the  con- 
tinental shelf  off  northeastern  Venezuela  in  deptlis  of  26-8(i  m, 
and  S.  staminea  is  known  only  from  the  Davis  Seamount  off 
southeastern  Brazil  in  60  m;  shells  of  both  species  are  very 
similar  to  those  of  S.  carvalhoi  Lopes  and  Sa  Cardoso,  1958. 
Solariella  cristata  is  known  from  the  upper  continental  slope 
of  Isla  Cancun,  Yucatan,  Mexico,  and  off  Key  Largo,  Florida 
Keys,  in  depths  of  155-256  m,  and  off  St.  Vincent,  Lesser 
Antilles  in  165-201  m. 

Key  words:  Trochidae;  Solariellinae;  Solariella;  systematics; 
new  species. 


INTRODUCTION 

Two  recent  monographs  of  western  Atlantic  Trochidae 
present  accounts  of  the  faunas  of  restricted  geographical 
areas:  the  Straits  of  Florida  (Quinn,  1979)  and  tlie  Gulf 
of  Mexico  (Quinn,  in  press).  In  the  process  of  examining 
specimens  for  those  reports,  several  new  species  from 
other  areas  of  the  western  Atlantic  Ocean  were  discov- 
ered. This  paper  presents  descriptions  of  three  new  spe- 
cies of  Solariella  Wood,  1842.  These  species  are  included 
in  Solariella  because  of  the  similarities  of  their  shells  to 
tho.se  of  species  such  as  S.  lacunella  (Dall,  1881);  how- 
ever, because  radular  characters  are  important  (Herbert, 
1987)  and  animals  of  the  three  species  were  unavailable 
for  study,  such  a.ssignment  of  these  species  is  tentative 
at  present. 

Institutional  abbreviations  used  in  this  paper  are  as 
follows:  MNIIN  (Museum  National  d'llistoire  Naturelle, 
Paris,  France);  MORG  (Museu  Oceanographico  da  Fun- 
da(,'ao  Universidade  do  Rio  Grande,  Rio  Grande,  RS, 
Brazil);  UMML  (Rosenstiel  School  of  Marine  and  At- 
mospheric Science,  University  of  Miami,  Miami,  Flori- 
da); USNM  (National  Museum  of  Natural  History,  Smith- 
sonian Institution,  Washington,  DC). 


SYSTEMATICS 

Genus  Solariella  Wood,  1842 

Type  species  (monotvpv):  Solariella  maculata  Wood, 
1842. 

Solariella  quadricincta  new  species 
(figures  1-4) 

Material  examined:  1  fragment,  UMML  30.6695  (para- 
type);  JOHN  ELLIOTT  >ILLSBURY  Station  P-727, 
10°20'N,  65°02' W,  64  m;  10-ft  otter  trawl;  21  July  1968.— 
1  specimen,  L'MML  uncatalogued  [examined  and  pho- 
tographed in  1975  (figs.  3,  4),  but  a  recent  attempt  to 
relocate  this  specimen  was  unsuccessful,  and  it  is  pre- 
sumed lost];  JOHN  ELLIOTT  PILLSBURY  Station  P- 
721,  ir06.5'N,  64''22.5'W,  26-27  m;  10-ft  otter  trawl; 
21  Julv  1968.  — 1  specimen,  USNM  859437  (holotype);  1 
specimen,  UMML  30.6528  (paratype);  JOHN  ELLIOTT 
PILLSBURY  Station  P-718,  1 1°22.5'N,  64°08.6'W,  60  m; 
10-ft  otter  trawl;  20  Julv  1968.— 1  specimen,  UMML 
30.6376  (paratype);  JOHN  ELLIOTT  PILLSBURY  Sta- 
tion P-705,  10°45'N,  62°00'W,  77-86  m;  10-ft  otter  trawl; 
18  July  1968. 

Description:  Shell  of  moderate  size  for  genus,  attaining 
6.55  mm  height,  7.15  mm  width,  umbilicate,  ivory  with 
light  orange-brown  spots  and  flammules,  nacreous  under 
thin  outer  porcelaneous  layer.  Protoconch  300-325  /um 
maximum  diameter,  of  about  1  whorl.  Teleoconch  whorls 
5.5,  tubular,  shouldered;  first  3  whorls  with  3  strong, 
subequal  spiral  cords  forming  whorl  periphery,  adapical 
cord  forming  whorl  shoulder;  suKsequent  whorls  with 
subsutural  spiral  cord  forming  narrow  channel  with  su- 
ture; last  whorl  with  fourth  primary  spiral  cord,  subequal 
to  other  peripheral  spiral  cords  and  coincident  with  su- 
ture on  previous  whorls;  1-2  weak  spiral  cords  sometimes 
occurring  between  primary  spiral  cords;  fine  spiral  threads 
overK  ing  interspaces  and  primary  spiral  cords  on  last  3 
whorls.  Axial  sculpture  consisting  of  rather  strong  riblets 
on  whorls  2  and  3,  fading  in  strength  to  numerous  col- 
labral  threads  in  interspaces  of  spiral  cords  on  subsequent 


J.  F.  Quinn,  Jr.,  1992 


Page  51 


r- 


K 


p^ 


^ 


r/ 


V, 


^ 


%^^^JI 


Figures  1-t.  SolaricUa  quadricincta  new  species.  1,  2.  .^pertural  and  basal  views  of  holotype,  USNM  8.594.37,  height  6.40  mm, 
widtli  6.40  mm,  from  oft  Isia  cle  Margarita,  Venezuela,  11°22..5'N,  64°08.6'W,  60  m.  3,  4.  Apertural  and  basal  views  of  specimen 
(lost;  measurements  imknowii),  UMML  .30.66.'59,  from  off  Isla  de  Margarita,  Venezuela,  11°06..5'N,  64°22..5'W,  26-27  m.  Figures 
5,  6.  SolaricUa  carvalhoi  L<ipes  and  Sa  C^ardoso,  19.58.  FSBC  1  39514,  height  7.9  mm,  width  8.2  mm,  from  Cagarras  Island,  Rio 
de  Janeiro,  Brazil,  32  m. 


whorls;  axial  threads  form  lamelliform  beads  on  subsu- 
tiiral  spiral  cord,  w  eakK  beading  shoulder  spiral  cord  on 
whorls  3-5.  Subsutural  shelf  rather  wide,  flat,  sloping 
abapically  from  subsutural  spiral  cord  to  shoulder  spiral 
cord;  fine  spiral  threads  appearing  on  third  whorl  near 
shoulder,  progressively  covering  shelf  surface  on  subse- 
quent whorls;  1-2  stronger  spiral  threads  appearing  on 
last  whorl.  Base  convex,  with  6-7  strong  spiral  cords; 
cords  and  interspaces  with  fine  spiral  threads;  interspaces 
with  fine  collabral  threads.  Umbilicus  wide,  about  40- 
45%  maximum  shell  widtli,  funnel-shaped;  v\  alls  convex, 
with  6  strongly  beaded  spiral  cords.  Shell  ground  color 
ivory,  with  regularly  spaced,  spirally  elongate  light  or- 
ange-brown spots  on  major  spiral  cords,  with  occasional 
flammules  of  same  color  on  subsutural  shelf.  Operculum 
thin,  corneous,  multispiral 

Type  locality:  Off  Isla  de  Margarita,  Venezuela, 
11°22.5'N,  64°08.6'W,  60  m. 

Remarks:  Shells  ot  SolaricUa  quadricincta  closely  re- 
semble those  of  S.  staminea.  new  species,  and  S.  car- 
valhoi Lopes  and  Sa  Cardoso,  1958,  from  Brazil  (figs.  5, 
6).  Shells  of  all  three  species  have  four  strong,  subequal 


primary  spiral  cords  on  the  last  whorl,  of  which  the 
shoulder  spiral  cord  is  the  most  distinctly  beaded;  have 
fine  spiral  threads  on  suprabasal  whorl  surface;  and  have 
flat  subsutural  shelves.  Shells  of  S.  quadricincta  differ 
from  those  of  S.  starnirica  by  being  slightly  narrower 
(height;width  ratios  =  0.92-1.01  and  0.79-0.90,  respec- 
tively); by  having  a  channeled  suture;  by  having  crisp 
rather  than  flattened  axial  threads;  by  having  fewer,  nar- 
rower, rounded  basal  spiral  cords;  by  having  a  narrow 
and  more  strongly  beaded  circumumbilical  cord;  by  hav- 
ing more  numerous,  more  strongly  beaded  intra-mnbil- 
ical  spiral  cords;  and  by  having  a  strong,  distinct  color 
pattern.  Shells  of  S.  quadricincta  differ  from  those  of  S. 
carvalhoi  by  being  smaller  at  similar  whorl  number;  by 
ha\  ing  canaliculate  sutures;  by  having  more  steeply  slop- 
ing subsutural  shelves;  by  having  finer,  crisper,  more 
closely  spaced  axial  threads  that  are  present  in  the  in- 
terspaces of  all  spiral  cords;  by  having  a  more  finely 
beaded  shoulder  cord;  by  having  wider  interspaces  be- 
tween the  basal  spiral  cords;  and  by  having  more  strongly 
beaded  intraumbilical  spiral  cords.  SolaricUa  quadri- 
cincta is  only  known  from  the  continental  shelf  off  north- 
eastern Venezuela  in  depths  of  26-86  m. 


Page  52 


THE  NAUTILUS,  Vol.  106,  No.  2 


"\ 


t 


10 


Figures  7-10.  Solariclla  cristata  new  species.  7,  8.  Holotype,  IISNM  859421,  height  9  1  mm.  width  9  4.5  mm,  from  SE  of  St. 
Vincent,  Lesser  Antilles,  13°11.2'N,  6r05.1'W,  16.5-201  m,  9,  10.  Paratype,  UMML  30.5669,  height  SI  mm,  width  8.0  mm,  from 
off  Isla  Mujeres,  Yucatan,  Mexico,  21°07'N,  86°21'W,  155-205  m. 


Solariella  cristata  new  species 
(figures  7-10) 

Material  examined:  1  specimen,  UMML  30.5669  (para- 
type); JOHN  ELLIOTT  PILLSBURY  Station  P-598, 
21°07'N,  86°21.0'W,  155-205  m;  10-ft  otter  trawl;  15 
March  1968.— 1  specimen,  UMML  30.7433  (paratype); 
GERDA  Station  G-6I,  25°14'N.  80°02'W,  256  m;'  6-ft 
otter  trawl;  29  August  1962.— 1  specimen,  L'SNM  859421 
(holotype);  JOHN  ELLIOTT  PILLSBURY  Station  P- 
874,  13''11.2'N,  61°05.1'W,  165-201  ni;  5-ft  Blake  trawl; 
6  July  1969. 

Description:  Shell  rather  large  for  genus,  attaining  9.1 
mm  height,  9.45  mm  width,  umbilicate,  ivory  vvith  light 
brown  spots  and  llannnulcs,  nacreous  under  thin  outer 
porcelaneous  layer.  Protoconch  300-320  ^lm  maximum 
diameter,  of  about  1  whorl.  Teleoconch  whorls  6.1,  tu- 
bular, shouldered;  first  3  whorls  with  4-5  spiral  cords, 
increasing  to  14  on  body  whorl;  abapical  cord  on  whorls 
3-5  strongest,  forming  peripheral  carina;  cords  on  last 
whorl  becoming  subequal  to  peripheral  cord,  giving  whorl 


more  evenly  rounded  appearance.  Axial  sculpture  ap- 
pearing on  whorl  2,  consisting  of  rather  strong,  low,  flat- 
tened folds  on  adapical  half  to  two-thirds  of  whorls  2- 
4,  becoming  narrower,  roimded,  more  crowded,  and  ex- 
tending to  level  of  suture  on  subsequent  whorls;  axial 
folds  forming  strong,  rounded  beads  on  spiral  cords,  beads 
strongest  on  adapical  part  ol  whorl.  Subsutural  shelf  nar- 
row, sloping  adapically  from  suture  to  shoulder  spiral 
cord;  single  strong  spiral  cord  bisecting  shelf.  Base  weakly 
convex,  with  7-10  strong,  smooth  spiral  cords.  Umbilicus 
wide,  about  30%  maximum  shell  width,  funnel-shaped; 
walls  convex,  with  4-6  strong,  strongly  beaded  spiral 
cords.  Shell  ground  color  ivory,  with  scattered  light  brown 
spots  appearing  on  fourth  whorl,  becoming  more  nu- 
merous and  forming  irregular  axial  flammules  on  last 
whorl. 

Type  locality:  SE  of  St.  Vincent,  Lesser  Antilles, 
I3°11.2'N,  6r°05.1'W,  165-201  m. 

Remarks:  The  peripherally  carinate,  rather  strongly  ax- 
ially  sculpted  shells  of  Solariella  cristata  most  closely 


J.  F.  Quinn,  Jr.,  1992 


Page  53 


Figure  1  1-12.    Soldriclla  staminea  new  species   Apertural  and  basal  views  of  holotype,  MORG  26530,  height  3.8  mm,  width  4  8 
mm,  from  Davis  Seamount,  Brazil,  20°40'S,  34°41'W,  60  m. 


resemble  shells  of  S.  cincta  (Pliilippi,  1836)  (see  Fretter 
&  Graham,  1977:46-48,  figs.  31,  32)  from  the  north- 
eastern Atlantic  Ocean.  However,  shells  of  S.  cristata  are 
larger,  have  a  narrower  subsutural  shelf,  and  have  a 
distinct  color  pattern,  whereas  those  of  S  cincta  are 
uniformly  ivor\ .  The  only  other  western  Atlantic  species 
having  shells  with  a  strongly  carinate  periphery  is  S. 
patriae  Carcelles,  1953,  but  shells  of  that  species  lack 
other  spiral  cords  except  the  shoulder  and  circumbasal 
spiral  cords,  and  have  only  three  basal  cords  (see  Rios, 
1985:23,  pi.  10,  fig.  97).  The  three  widely  separated  lo- 
calities from  which  S.  cristata  has  been  collected  (the 
Florida  Keys,  Yucatan,  and  the  Lesser  Antilles)  suggest 
that  the  species  is  widespread,  but  rare,  in  the  Caribbean 
Sea. 

Solariella  staminea  new  species 
(figures  11,  12) 

Material  examined:  1  specimen,  MORG  26530  (holo- 
type); 3  specimens,  MNHN  uncatalogued  (paratypes); 
MARION-DUFRESNE  Cruise  MD-55,  Station  DC-40, 
20°40'S,  34°41'W,  60  m;  dredge;  May  1987;  P.  Bouchet, 
J.  H.  Leal,  and  B.  Metivier  collectors. 

Description:  Shell  of  moderate  size  for  genus,  attaining 
6.7  mm  height,  7.45  mm  width,  umbilicate,  white  with 
few  light  yellow  to  orange-brown  spots  and  streaks,  na- 
creous under  thin  outer  porcelaneous  layer.  Protoconch 
about  300  ^m  ma.ximum  diameter,  of  about  one  whorl. 
Teleoconch  whorls  5.6,  tubular,  shouldered;  spire  whorls 
with  3  strong  spiral  cords,  adapical  one  forming  whorl 
shoulder;  last  whorl  with  fourth  strong  spiral  cord,  sub- 
equal  to  other  spiral  cords  and  coincident  with  suture  on 
previous  whorls;  1-2  additional,  weaker,  intercallary  spi- 
ral threads  sometimes  present;  fine  spiral  threads  over- 
ly ing  interspaces  and  lower  2-3  spiral  cords  on  last  3 
whorls.  Axial  sculpture  of  rather  strong  riblets  on  whorl 
2,  fading  in  strength  to  numerous,  crowded,  rather  flat- 
tened collabral  threads  in  interspaces  of  spiral  cords  on 
subsequent  whorls;  axial  threads  forming  weak,  rounded 


beads  on  shoulder  spiral  cord,  finely  beading  other  spiral 
cords.  Subsutural  shelf  moderately  wide,  flat,  sloping 
abapically  from  suture  to  shoulder  spiral  cord;  fine  spiral 
threads  appearing  on  third  whorl;  one  spiral  cord  ap- 
pearing near  beginning  of  first  whorl,  but  fading  to  ob- 
scure angulation  or  disappearing  on  second  whorl;  1-2 
spiral  cords  appearing  on  third  whorl,  one  near  middle 
of  shelf  strongest.  Base  weakly  convex,  with  9-11  strong, 
flattened  spiral  cords;  innermost  cord  strap-like  and 
weakly  beaded;  interspaces  with  fine  collabral  threads, 
forming  weak  rugae  on  spiral  cords  on  adaxial  half  of 
base.  Umbilicus  wide,  about  35-45%  maximum  shell 
width,  funnel-shaped;  walls  convex,  with  4-5  beaded 
spiral  cords.  Shell  ground  color  white,  with  few,  widely 
spaced  light  yellow  to  orange-brown  spots,  principally 
on  shoulder  cord,  but  occasionally  extending  abaxially 
as  short  streaks  or  flammules. 

Type  locality:  Davis  Seaniount,  off  southeastern  Brazil, 
20°40'S,  34°41'W,  60  m. 

Remarks:  Shells  of  Solariella  staminea  are  very  similar 
to  those  of  S.  quadricincta  and  S.  carvalhoi.  Differences 
that  distinguish  shells  of  S.  staminea  from  those  of  S. 
quadricincta  are  discussed  in  the  Remarks  section  of  the 
latter  species.  Shells  of  S.  staminea  differ  from  those  of 
S.  carvalhoi  by  being  broader  (height:width  ratios  = 
0.79-0.90  and'  0.93-0.98,  respectively);  by  having  a 
broader  subsutural  shelf  with  stronger  spiral  cords  and 
axial  threads;  by  having  a  shoulder  spiral  cord  bearing 
rounded  rather  than  spirally  elongate  beads;  by  having 
more  numerous,  flatter  basal  spiral  cords;  by  having  a 
circumumbilical  spiral  cord  that  is  broad,  flat,  and  weak- 
ly beaded  rather  than  narrow,  sharp,  and  strongly  bead- 
ed; by  having  stronger,  more  weakly  beaded  intraum- 
bilical  cords;  and  by  having  a  very  weak  color  pattern. 

ACKNOWLEDGMENTS 

I  thank  the  late  Gilbert  L.  Voss  (Rosenstiel  School  of 
Marine  and  Atmospheric  Science,  University  of  Miami) 


Page  54 


THE  NAUTILUS,  Vol.  106,  No.  2 


for  access  to  the  collections  under  liis  care  and  for  per- 
mission to  d(>.scribe  the  new  species  in  that  material.  The 
late  Joseph  Ilo.sewater  and  Richard  S.  Houbrick  (both 
USNM)  kindly  allowed  me  to  examine  types  and  other 
material.  Jose  H.  Leal  brought  the  specimens  of  Solariella 
staminea  to  my  attention  and  kindK  provided  the  SEM 
micrographs  of  that  species.  Eliezer  de  C.  Rios  (MORG) 
generous!)  provided  comparative  material  of  Solariella 
carvalhoi.  Marjorie  Myers,  Llyn  French,  and  Judy  Leiby 
provided  valuable  word-processing  and  editorial  assis- 
tance. Thomas  H.  Perkins  and  William  G.  Lyons  (both 
Department  of  Natural  Resources,  Florida  Marine  Re- 
search Institute),  and  two  anonymous  reviewers  provided 
valuable  critiques  of  previous  drafts  of  this  paper. 

LITERATURE  CITED 

C^arcelies,  A.  R.  19.53.  Nuevasespeciesdegastropoclosmarinos 
de  las  republicas  oriental  del  I'ruguay  y  Argentina  Co- 
municaciones  Zooiogicas  del  Museo  de  Historia  Natural 
de  Montevideo  4(70):  1-16. 

Dall,  W.  H.  1881.  Reports  on  the  results  of  dredging,  under 
the  supervision  of  Alexander  Agassiz,  in  the  Gulf  of  Mex- 
ico, and  in  the  Caribbean  Sea,  1877-79,  by  the  United 
States  Coast  Survey  steamer  "Blake",  Lieutenant-Com- 
mander C.  D.  Sigsbee,  U.S.N.,  and  Commander  J  l\.  Bart- 


lett,  U.S.N.,  commanding.  W.  Preliminary  report  on  the 
Mollusca.  Bulletin  of  the  Museum  of  Comparative  Zool- 
ogy, Har\ard  L'niversity  9(2)::3-3-144, 

PVetter.  \'  and  .\.  (Jraham  1977.  The  prosobrancli  molluscs 
of  Britain  and  Denmark.  Part  2 — Trochacea  Journal  of 
Molluscan  Studies,  Supplement  3:39-100. 

Herbert,  D.  G.  1987.  Revision  of  the  Solariellinae  (Mollusca: 
Prosobranchia:  Trochidae)  in  southern  Africa.  Annals  of 
the  Natal  Museum  28(2):283-382. 

Lopes,  H  de  S.  and  P  de  Sa  Cardoso.  1958.  Sobre  un  novo 
gastropodo  brasileiro  do  genero  "Solariella"  Wood,  1842 
(Trochidae).  Revista  Brasileira  de  Biologia  18(l):59-64. 

Philippi,  R  .\  18.3(1  Eniuneratio  molluscorum  Siciliae  cum 
viventium  tuni  tellure  fossilium,  quae  in  itinere  suo  ob- 
servavit.  Vol.  1.  Berolini,  Sinionis  Schroppii  et  Soc,  xiv  -I- 
267  pp. 

Quinn,  J.  F.,  Jr.  1979.  Biological  results  of  the  University  of 
Miami  Deep- Sea  Expeditions.  130.  The  systematics  and 
zoogeography  of  the  gastropod  family  Trochidae  collected 
in  the  Straits  of  Florida  and  its  approaches.  Malacologia 
19(l):l-62. 

Quinn,  J.  F.,Jr.  In  press.  The  Trochidae  t)f  the  Ckilf  of  Mexico 
(Prosobranchia:  Archaeogastropoda).  Memoirs  of  the 
Hourglass  Cruises. 

Rios,  E.  de  C.  1985.  Seashells  of  Brazil.  Funda^ao  Cidade  do 
Rio  Grande,  Rio  Grande,  RS,  Brazil,  328  -t-  [103]  p. 

Wood,  S.  V.  1842.  A  catalogue  of  shells  from  the  Crag,  .\nnals 
and  Magazine  of  Natural  History,  Series  1,  9:527-544. 


THE  NAUTILUS  106(2):55-59,  1992 


Page  55 


A  New  Fossil  Land  Snail  of  the  Genus 
Hemitrochus  from  Bowden,  Jamaica^ 


Glenn  A.  Goodfriend 

Geophysical  Laboratory 
Carnegie  Institution  of  Washington 
5251  Broad  Branch  Road,  NW 
Washington,  DC  20015  LISA,  and 
Department  of  Environmental 

Sciences  and  Energy  Research 
Weizmann  Institute  of  Science 
76100  Rehovot,  ISRAEL 


ABSTRACT 

A  new  species  of  Hemitrochus,  H.  bowdenensis,  is  described 
from  a  collection  made  in  the  Bowden  Beds  at  Bowden,  Ja- 
maica. The  species  differs  from  its  nearest  relative  H.  gramin- 
icola  (the  only  modern  Hemitrochus  found  in  Jamaica)  on  the 
basis  of  shell  sculpture,  the  siiape  of  the  whorls,  and  umbilical 
width.  Analysis  of  amino  acid  D/L  ratios  from  one  specimen 
indicates  a  late  Holocene  age  for  the  sample  and  that  therefore 
the  material  does  not  belong  stratigraphically  to  the  Bowden 
Beds  (of  early  Pliocene  age).  The  species  is  apparently  extinct, 
probably  as  a  result  of  either  habitat  destruction  following 
European  settlement  or  late  Holocene  climatic  changes. 

Key  words:  land  snail,  fossil,  new  species,  Jamaica,  Hemitro- 
chus. Bowden. 


INTRODUCTION 

The  Bowden  Beds,  located  near  Bowden,  St.  Thomas 
Parish,  in  southeastern  Jamaica,  have  yielded  an  ex- 
tremely rich  marine  mollusk  fauna  (Woodring,  1925, 
1928)  as  well  as  a  number  of  land  snails,  including  a 
species  of  helicinid  of  the  genus  Lticidella  (Simpson, 
1895),  three  species  of  poteriids  (Simpson,  1895;  Bartsch, 
1942;  Morrison,  1955),  and  two  species  of  camaenids  of 
the  genus  Pleurodonte  (Simpson,  1895;  Kimball,  1947). 
Various  other  land  snail  species  have  been  mentioned  as 
having  been  collected  from  the  Bowden  Beds  ("Thijsa- 
nophora",  Opeas  striata.  Succinea  latior,  Simpson,  1895; 
"Stenogyra",  "Melaniella",  "Truncatella",  Woodring, 
1928),  but  because  of  the  fresh  appearance  of  some  of 
these  shells,  it  has  been  suggested  (Woodring,  1928)  that 
these  mav  be  recent  material  washed  into  the  Bowden 


'  Contribution  No.  24,  Department  of  Environmental  Sci- 
ences and  Energy  Research,  Weizmann  Institute  of  Science 


sediments.  The  Bowden  Beds  are  considered  to  be  of 
early  Pliocene  age  based  on  analysis  of  the  foraminiferal 
fauna  (assigned  to  the  Globorotalia  margaritae  zone  by 
Bolli  and  Bermudez  (1965),  which  was  placed  in  the  early 
Pliocene  by  Bolli  and  Premoli  Silva  (1973)).  Recent  work 
on  marine  mollusks  from  Bowden  has  accepted  this  age 
assignment  (Jung,  1989). 

E.xamination  of  the  collections  at  the  Academy  of  Nat- 
ural Sciences  of  Philadelphia  (ANSP)  has  turned  up  a 
new  species  of  land  snail  collected  from  the  Bowden  Beds 
and  belonging  to  the  family  Helminthoglyptidae  (=  Xan- 
thonycidae  sf  nst/  Baker  (1943)  and  Nordsieck  (1987);  = 
Fruticicolidae  sensu  Turner  (1958)).  This  new  species  is 
described  below  and  its  relationships  are  discussed. 

Hemitrochus  boivdenensis  new  species 

Description:  Shell  of  average  size  for  the  genus  {ca.  11- 
12  mm  diameter),  low-trochoidal  (height  %  of  diameter), 
moderately  thin;  spire  weakly  convex;  shell  periphery 
subangular,  becoming  nearly  rounded  at  the  lip;  suture 
deeply  impressed;  base  weakly  convex,  with  the  apex  of 
the  convexity  shifting  from  a  position  in  the  middle  of 
the  base  in  the  younger  part  of  the  shell  to  a  position 
nearer  to  the  umbilicus  (than  to  the  periphery)  as  the 
adult  lip  is  approached;  descent  of  the  suture  behind  the 
lip  unknown,  since  the  upper  part  of  the  lip  is  not  pre- 
served in  the  specimens;  lip  unreDected  at  periphery  but 
gradually  becomes  reflected  on  the  base  in  the  direction 
of  the  umbilicus;  bordering  the  umbilicus,  the  lip  is 
strongly  reflected,  but  only  for  a  short  distance,  whereas 
in  the  middle  of  the  base  of  the  shell,  the  lip  reflection 
starts  earlier  but  is  weaker;  the  basal  insertion  of  the  lip 
is  drawn  out,  forming  a  rim  on  one  side  of  the  umbilicus 
which  is  reflected  over  a  small  part  of  the  umbilicus; 
aperture  round  to  weakly  elliptical,  with  the  width  usu- 
ally slightly  exceeding  the  height;  protoconch  1.6-1.7 
whorls,  smooth;  the  sculpture  of  later  whorls  consists  of 


Page  56 


THE  NAUTILUS,  Vol.  106,  No.  2 


Table  1. 

Measurements  of  the 

type  material 

of  Hemitrochus 

boudenensis. 

Specimen 

Life  history 
stage 

Diameter 

(mm) 

Height 

(mm) 

Height/ 
diameter 

Whorl 
number 

Aperture 
width 

(mm) 

Aperture 
height 

(mm) 

.Aperture 

height/ 

width 

Holotype 
Paratype 
Paratype 

adult 

subadult 

?juveniie 

12.4' 

10.5' 

8.6 

8.3 
6.9 
5.5 

0.67 
0.66 
0.64 

4.5 
4.4 
4.0 

5.2- 
4.6 
3.9 

49^ 
4.6 

3.8 

0.942 
1  00 
0.97 

'  Diameter  is  the  appro.ximate  adult  diameter;  a  precise  measurement  is  not  possible  since  the  lip  is  broken. 
^  Measured  0.1  whorl  behind  aperture,  due  to  broken  lip. 


low,  broad  ribs  {ca.  12-14  on  the  penultimate  V4  whorl) 
that  are  parallel  to  the  growth  lines  and  irregular,  with 
each  rib  varying  in  both  height  and  width  along  its  length; 
the  length  of  the  ribs  also  varies,  with  some  disappearing 
on  the  dorsum  of  the  shell,  and  others  continuing  across 
the  periphery  and  terminating  just  below  the  periphery; 
on  the  rest  of  base,  sculpture  consists  only  of  irregular 
growth  lines,  which  get  stronger  nearer  the  lip;  color 
opaque  white.  Measurements  of  specimens  are  presented 
in  Table  1. 

Material  examined:  The  holotype  (ANSP  75798)  and 
two  paratypes  (ANSP  75799)  are  the  only  known  spec- 
imens of  this  species.  The  holotype  (figures  1-3)  is  an 
adult  shell  missing  the  upper  lip  and  the  end  of  the  lip 
where  it  becomes  the  umbilical  wall.  The  first  paratype 
(figures  4,  5)  is  apparently  a  subadult  specimen.  The  lip 
shows  the  reflection  which  characterizes  the  adult  form 
but  the  lip  is  a  little  thinner  than  that  of  the  holotype, 
suggesting  that  the  shell  is  not  fully  mature.  The  dimen- 
sions of  the  shell  should  be  representative  of  the  adult 
dimensions,  since  further  growth  would  result  only  in 
thickening  of  the  shell.  The  upper  lip  of  this  specimen 
is  broken  as  in  the  holotype  but  the  umbilical  end  of  the 
lip  is  intact.  The  second  paratype  has  the  last  ca.  1  whorl 
missing,  the  remains  of  which  can  be  seen  only  as  a  rim 
around  the  umbilicus.  An  additional  ca.  'A  of  the  upper 
part  of  the  last  whorl  was  originally  present  but  was 
removed  for  amino  acid  analysis. 

Type  locality,  Mratigraphy,  and  collection  information: 

The  locality  data  provided  with  the  material  indicates 
that  it  was  collected  "Among  oligocene  fossils,  Bowden, 
Jamaica",  by  Uselma  C.  Smith  and  S.  L.  Schumo.  Bowden 
is  a  small  village  located  on  the  east  side  of  Port  Morant 
Bay  in  southeastern  St.  Thomas  Parish,  Jamaica.  No 
stratigraphic  information  is  provided  with  the  material. 
However,  concerning  the  well-known  Bowden  fossil  beds 
(later  considered  to  be  Miocene  in  age;  Woodring,  1928), 
Woodring  (1925:7)  stated  that  "At  the  type  locality  [i.e., 
at  Bowden]  a  bed  of  imj)erfectly  consolidated  gravel 
consisting  of  small  pebbles  and  grains  of  igneous  rocks, 
limestone,  and  other  sorts  of  rock  in  a  marly  matrix  crops 
out  in  the  road-cut  for  a  distance  of  several  hundred  feet 
along  the  bay.  This  bed,  which  is  not  more  than  2  or  3 
feet  thick,  contains  perfectly  preserved  fossils,  and,  so 
far  as  known,  has  furnished  all  the  fossils  collected  at 
Bowden."  Chubb  (1958:28)  indicates  that  at  Bowden 


there  is  "one  bed,  some  2-3  feet  thick  lying  a  few  feet 
above  road-level,  of  unconsolidated  gravel  consisting  of 
small  pebbles  and  grains  of  igneous  rocks  in  a  marly 
matri.x.  The  bed  is  lenticular  and  runs  for  only  about  a 
couple  of  hundred  yards,  and  it  is  extremely  rich  in  well- 
preserved  fossils.  ..."  The  land  snail  material  was  among 
an  extensive  collection  of  marine  mollusks  made  b\  Smith 
and  Schumo  and  it  seems  almost  certain  that  all  this 
material  was  collected  from  the  locality  described  by 
Woodring  and  by  Chubb.  In  a  visit  to  Bowden  by  the 
author  in  April,  1991,  it  was  found  that  the  beds  con- 
taining marine  mollusks  run  along  the  roadcut  on  the 
east  side  of  the  Bowden  road,  northward  from  the  junc- 
tion of  the  road  climbing  up  the  hill  from  Bowden.  This 
should  be  taken  as  the  type  locality  for  Hemitrochus 
bowdenensis. 

No  date  of  collection  is  associated  with  the  Schumo 
and  Smith  collection  from  Bowden,  but  the  catalogue  at 
the  Academy  of  Natural  Sciences  of  Philadelphia  indi- 
cates that  this  material  was  presented  to  the  museum  on 
January  7,  1902  (G.  Rosenberg,  personal  communica- 
tion), so  it  was  collected  some  time  before  this  date. 

Comparative  remarks:  The  only  other  helminthoglyp- 
tid  genus  found  in  Jamaica  is  Dialeuca.  The  shells  in  this 
genus  differ  from  Hemitrochus  shells  in  several  respects. 
The  umbilicus  (covered  in  adults)  is  very  narrow  and  the 
basal  lip  inserts  at  a  high  angle  to  the  columella  (in 
Hemitrochus  this  insertion  is  at  a  low,  near  vertical  angle); 
these  features,  together  with  a  higher  expansion  rate  of 
the  last  whorl,  result  in  the  aperture  being  considerably 
wider  than  high.  The  basal  lip  expands  gradually  and 
evenly  across  the  base  (this  also  occurs  in  some  Bahamian 
Hemitrochus  species)  and  the  dorsal  lip  is  reflected.  The 
sculpture  consists  only  of  weak  growth  lines  (this  occurs 
also  in  some  Bahamian  Hemitrochus  species).  The  ribbed 
sculpture  characteristic  of  most  species  of  Hemitrochus 
occurs  also  in  Plagioptycha,  a  genus  found  in  Hispaniola, 
the  Bahamas,  and  the  Lesser  Antilles.  But  Plagioptijcha 
differs  from  Hemitrochus  in  having  a  high-angle  inser- 
tion of  the  basal  lip  onto  the  columella  (as  in  Dialeuca) 
associated  with  a  relatively  wide  aperture;  the  shell  is 
more  depressed  and  a  ridge  is  usually  present  on  the 
inside  of  the  lower  lip  on  the  left  (umbilical)  side.  Thus 
the  new  species  clearly  belongs  in  Hemitrochus  rather 
than  in  a  related  helminthogU  ptid  genus. 

Hemitrochus  bowdenensis  most  closeH  resembles  the 
only  other  Hemitrochus  species  inhabiting  Jamaica,  H. 


G.  A.  Goodfriend,  1992 


Page  57 


Figures  1-3.  Heniitrochus  huwdencnsis  new  species,  dorsal,  apertural,  and  ventral  views  of  holotype.  4,  5.  Hernitrochus  bow- 
denensis  new  species,  apertural  and  ventral  viev\s  of  paratype.  6-8.  Hernitrochus graminicola  (from  Happy  News,  SE of  Alexandria, 
St.  Ann,  Jamaica),  dorsal,  apertural,  and  ventral  views.  Scale  line  (in  mm)  in  figure  2  refers  to  figures  1-5  and  scale  line  in  figure 
7  refers  to  figures  6-8. 


graminicola  (C.  B.  Adams).  H.  graminicola  (figures  6- 
8)  differs  in  having  a  more  weakly  developed  sculpture 
consisting  of  fine,  regular  ribs  {ca.  30  on  the  penultimate 
'/4  whorl)  which  cross  the  base  of  the  shell,  disappearing 
only  near  the  umbilicus.  It  also  differs  in  having  a  round- 
ed periphery  on  the  whole  of  the  last  whorl,  a  more 
inflated  base,  less  impressed  sutures,  and  a  generally  larg- 
er size  (12-16  mm  diameter).  However,  in  the  form  of 
the  lip  reflection  and  in  the  shift  of  the  apex  of  the  basal 


convexity  toward  the  umbilicus  in  the  direction  of  the 
lip,  H.  boivdenensis  resembles  H.  graminicola  precisely. 
H.  pseudogtjra  (Torre)  from  Cuba  closely  resembles  H. 
graminicola  but  is  less  close  to  H.  boivdenensis  in  that 
its  sculpture  is  both  weaker  and  finer  than  that  of  H. 
graminicola.  It  should  be  emphasized  that,  although  these 
are  the  species  nearest  phenotypically  to  H.  boivdenensis, 
they  are  not  very  closely  related  to  it — the  character  of 
the  sculpture  of  H.  boivdenensis  is  altogether  different 


Page  58 


THE  NAUTILUS,  Vol.  106,  No.  2 


Table  2.     D/L  amino  acid   ratios  in   paratype  specimen  of 
Hemitrochus  howdcncnsis 


Amino  acid 


D/L 


Alanine 

Alloisoleucine,  isoleucine 

Proline 

Aspartic  acid 

Methionine 

Glutamic  acid 

Phen\ialanine 


0.12 

0.053 

0.33 

0.27 

0.24 

0.062 

014 


from  that  of  any  modern  species.  H.  bowdenensis  is  not 
banded  as  other  Hemitrochus  species  are.  The  lack  of 
banding  on  the  shells  could  possibly  be  the  result  of 
fading,  but  this  seems  unlikely  for  such  young  material. 
Hemitrochus  has  been  considered  a  subgenus  of  Ce- 
polis  (Pilsbry,  1939;  Baker,  1943),  but  Turner  (1958) 
raised  it  to  generic  status  on  the  basis  of  anatomical 
characteristics.  Altliough  the  earlier  inclusion  of  Hemi- 
trochus within  Cepolis  was  retained  by  Nordsieck  (1987) 
and  Vaught  (1989),  the  taxonomy  of  Turner  (1958)  is 
followed  here. 


AGE  OF  THE  SAMPLE 

Although  the  Bowden  Beds  are  considered  to  be  of  early 
Pliocene  age,  the  possibility  of  inclusion  of  some  modern 
material  in  the  collections  still  exists,  for  example  if  ma- 
terial were  collected  from  slumped  sediments  which  could 
have  incorporated  modern  terrestrial  shells  or  from  ex- 
posed fissure  infills.  Woodring  (1928)  considered  that 
some  of  the  land  snail  material  that  had  been  collected 
from  the  Bowden  Beds  may  have  been  modern,  stating 
that  some  of  the  material  may  represent  "the  remains 
of  living  snails  that  fell  into  openings  in  the  ground  and 
thus  were  collected  with  the  fossil  material"  (p.  109). 
Because  of  this  possibility,  amino  acid  enantiomer/epi- 
mer  analyses  were  carried  out  on  a  fragment  of  one  of 
the  H.  bowdenensis  shells  as  a  check  on  its  age  (see 
Goodfriend,  1991,  for  analytical  methods).  Material  of 
early  Pliocene  age  would  be  expected  to  give  D/L  amino 
acid  ratios  near  equilibrium  (1.3  for  D-alloisoleucine/L- 
isoleucine  and  1.0  for  other  D/L  amino  acid  ratios)  and 
may  be  highly  depleted  in  amino  acid  content. 

The  analytical  results  (Table  2)  indicate  that  the  Hemi- 
trochus shell  is  relatively  young.  As  would  be  expected, 
faster-raccmizing  amino  acids  such  as  aspartic  acid,  pro- 
line, methionine,  and  phenylalanine  (Goodfriend,  1991) 
show  higher  D/L  ratios  than  the  slower  racemizing/ 
epimcrizing  amino  acids  glutamic  acid  and  isoleucine. 
A  calibration  of  the  rate  of  isoleucine  epimerization  in 
Pleurodonte  during  the  Holocene  at  a  site  on  the  north 
coast  of  Jamaica  (Goodfriend  and  Mittercr,  1988)  leads 
to  an  age  estimate  based  on  the  alloisoleucine/isoleucine 
(A/I)  ratio  of  the  Hemitrochus  of  1700  year  B.P.  (as- 
suming an  initial  A/I  value  of  0.013,  as  is  typical  of 
modern  land  snail  shells).  This  estimate  is  very  approx- 


imate since  there  are  a  number  of  errors  involved  which 
compound  to  produce  the  total  error:  the  error  of  mea- 
surement of  the  A/I  ratio  of  the  Hemitrochus  sample 
(5-10%),  possible  differences  in  the  epimerization  rate 
between  different  genera  (usually  on  the  order  of  10%), 
and,  most  importantly,  the  uncertainty  of  the  rate  dif- 
ference between  Bowden  and  the  north  coast  calibration 
site.  For  example,  a  2°  difference  between  the  sites  would 
lead  to  a  40%  difference  in  the  epimerization  rates  (equa- 
tion 3  in  Goodfriend  and  Mitterer,  1988).  Allowing  a 
total  uncertainty  of  ±50%  would  indicate  a  probable  age 
between  800-2400  year  B.P.  Thus  it  is  clear  that  the 
sample  is  of  late  Holocene  age,  and  therefore  represents 
material  which  does  not  belong  to  the  Bowden  Beds.  A 
visit  to  the  Bowden  Bed  type  locality  revealed  that  the 
moUusk-containing  unit  was  in  many  cases  covered  by 
colluvial  material,  slumped  down  from  the  steep  slope 
above.  It  seems  likely  that  the  H.  bowdenensis  material 
was  collected  from  such  a  slumped  deposit,  perhaps  at 
the  same  level  as  the  mollusk  beds. 

DISCUSSION 

This  new  species  of  Hemitrochus  is  not  represented  in 
modern  collections  from  Jamaica  and  is  therefore  pre- 
sumably extinct.  Whether  this  presumed  extinction  was 
the  result  of  forest  clearance  subsequent  to  European 
settlement  of  Jamaica  or  occurred  before  this  time,  as  a 
result  of  natural  processes  such  as  climatic  change,  cannot 
be  ascertained  without  additional  dated  records  of  this 
species.  Evidence  of  human  induced  local  extirpations 
of  land  snail  species  exists  for  the  north  coast  of  Jamaica 
(Goodfriend  and  Mitterer,  1988).  But  climatic  changes 
in  the  late  Holocene  have  also  been  documented  in  Ja- 
maica (Goodfriend,  1987). 

The  occurrence  of  this  apparently  extinct  species  in 
the  recent  fossil  record  of  Jamaica  is  somewhat  surprising 
in  view  of  the  fact  that  other  Holocene  and  late  Pleis- 
tocene deposits  on  the  island  contain  only  extant  species 
(Goodfriend  and  Mitterer,  1988;  Goodfriend,  1989).  Re- 
cent extirpations  of  species  in  north-central  Jamaica 
(Goodfriend,  1987)  and  the  central  north  coast  of  Jamaica 
(Goodfriend  and  Mitterer,  1988)  have  been  noted,  but 
these  species  have  survived  elsewhere  on  the  island.  Local 
endemics,  as  Hemitrochus  bowdenensis  may  have  been, 
will  be  more  sensitive  to  environmental  changes;  when 
more  widespread  species  undergo  local  extirpations,  local 
endemics  may  undergo  extinction.  The  forests  of  south- 
eastern St.  Thomas  have  been  almost  completely  cut 
down  and  replaced  by  agriculture — degraded  forest  re- 
mains in  only  a  very  few  areas.  One  vNonders  whether 
other  extinct  species  may  turn  up  in  the  recent  fossil 
record  in  this  area  and  other  areas  that  have  been  sim- 
ilarly degraded. 

ACKNOWLEDGMENTS 

I  am  indebted  to  Dr.  G.  Rosenberg  for  bringing  this 
material  to  my  attention  and  providing  information  on 


G.  A.  Goodfriend,  1992 


Page  59 


the  Schumo  and  Smith  Bowden  collection,  and  to  Dr.  G, 
M.  Davis  for  use  of  the  collections  of  the  Academy  of 
Natural  Sciences  of  Philadelphia,  loan  of  the  samples, 
and  permission  to  carry  out  amino  acid  analysis  on  a 
shell  fragment.  Dr.  K.  Emberton  and  an  anonymous  re- 
viewer provided  useful  comments  that  were  incorporat- 
ed into  the  manuscript.  Dr.  P.  E.  Hare  provided  pho- 
tographic equipment. 


LITERATURE  CITED 

Baker,  H,  B,  1943.  Some  Antillean  helicids.  Nautilus  56(3): 
81-91. 

Bartsch,  P.  1942.  Thecyclopiiorid  niollusksof  the  West  Indies, 
exclusive  of  Cuba.  Bulletin  of  the  United  States  National 
Museum  181:43-141,  -l-pl.  8-18  and  41 

Belli,  H.  M.  and  P.  J.  Bermudez  196.5  Zonation  based  on 
planktonic  foraminifera  of  Middle  Miocene  to  Pliocene 
warm-water  sediments.  Boletino  de  Informacion  de  la  As- 
sociacion  Venezueiana  de  Geologia,  Mineria,  v  Petrologia 
8:121-149. 

Belli,  H.  M.  and  I  Premoii  Silva.  1973  Oiigocene  to  Recent 
planktonic  foraminifera  and  stratigraphy  of  the  Leg  15 
sites  in  the  Caribbean  Sea.  /;i.-  Edgar,  N.  T,  et  al.  (eds.). 
Initial  reports  of  the  Deep  Sea  Drilling  Project,  Vol.  15. 
U.S.  Government  Printing  Office,  Washington,  DC,  p.  475- 
497. 

Chubb,  L.  J.  1958.  Higher  Miocene  rocks  of  southeast  Ja- 
maica. Geonotes  l(l/2):25-31. 

Goodfriend,  G.  .\.  1987.  Late  Holocene  morphological  changes 
in  a  Jamaican  land  snail:  evidence  for  changes  in  rainfall, 
/n-  Berger,  W.  H.  and  L.  D.  Labeyrie  (eds.).  Abrupt  cli- 
matic change.  D.  Reidel  Publishing  Co.,  Dordrecht,  p. 
123-126. 

Goodfriend,  G.  A.  1989.  Quaternary  biogeographical  history 
of  land  snails  in  Jamaica.  In:  Woods,  C.  A.  (ed).  Bioge- 
ography  of  the  West  Indies:  past,  present,  and  future. 
Sandhill  Crane  Press,  Gainesville,  FL,  p  201-216, 


Goodfriend,  G  A.  1991  Patterns  of  racemization  and  epi- 
merization  of  amino  acids  in  land  snail  shells  over  the 
course  of  the  Holocene.  Geochimica  et  Cosmochimica  Acta 
55:293-302. 

Goodfriend,  G  .\.  and  R.  M.  Mitterer.  1988.  LateQuaternar> 
land  snails  from  the  north  coast  of  Jamaica:  local  extinc- 
tions and  climatic  change.  Palaeogeography,  Palaeocli- 
matology,  Palaeoecology  63:293-311. 

Jung,  P.  1989  Revision  of  the  Strombina-group  (Gastropoda: 
Columbeiiidae),  fossil  and  living.  Schweizerische  Palaon- 
tologische  Abhandlungen  111. 

Kimball,  D.  1947.  A  new  Pleurodonte  from  the  Miocene, 
Bowden,  Jamaica.  Nautilus  61(2):37-39. 

Morrison,  J.  P.  E.  1955.  Notes  on  American  cyclophorid  land 
snails,  with  two  new  names,  eight  new  species,  three  new 
genera,  and  the  family  Amphicyclotidae,  separated  on 
animal  characters.  Journal  of  the  Washington  Academy 
of  Sciences  45(5):  149-162. 

Nordsieck,  H.  1987.  Revision  des  Systems  der  Helicoidea. 
Archiv  fiir  Molluskenkunde  118(l'/3):9-50. 

Pilsbry,  H.  A.  1939.  Land  Mollusca  of  North  America  (north 
of  Mexico),  Vol.  I,  part  I  Monographs  of  the  Academy  of 
Natural  Sciences  of  Philadelphia,  3:I-XVII,  1-.573,  i-ix. 

Simpson,  C.  T.  1895.  Distribution  of  the  land  and  fresh-water 
mollusks  of  the  West  Indian  region,  and  their  evidence 
with  regard  to  past  changes  of  land  and  sea.  Proceedings 
of  the  I'nited  States  National  Museum  17:423-451. 

Turner,  R  D  1958.  The  genus  Hemitrochus  in  Puerto  Rico. 
Occasional  Papers  on  Mollusks,  Museum  of  Comparative 
Zoology,  Harvard  University  2(22):  153-178. 

Vaught,  K.  C  1989.  A  classification  of  the  living  Mollusca. 
Abbott,  R.  T.  and  K,  J.  Boss  (eds.)  American  Malacologists, 
Inc.,  Melbourne,  Florida 

Woodring,  W.  P.  1925.  Miocene  mollusks  from  Bowden,  Ja- 
maica. Pelecypods  and  scaphopods.  Carnegie  Institution 
of  Washington  Publications  366:I-IV,  1-222,  pi.  1-28. 

Woodring,  W.  P.  1928  Miocene  mollusks  from  Bowden,  Ja- 
maica Part  II  Gastropods  and  discussion  of  results.  Car- 
negie Institution  of  Washington  Publication  385:I-VII,  1- 
564. 


THE  NAUTILUS  106(2):60-67,  1992 


Page  60 


Dreissena  polymorpha  (Zebra  Mussel):  Colonization  of  Soft 
Substrata  and  Some  Effects  on  Unionid  Bivalves 


R.  Douglas  Hunter 
John  F.  Bailey 

Department  of  Biological  Sciences 

Oakland  University 

Rochester,  MI  48309-4401  USA 


ABSTRACT 

Zebra  mussel  {Dreissena  polymorpha)  and  native  bivalve 
(Unionidae)  densities  were  determined  at  three  sites  in  southern 
Lake  St.  Clair  (Ontario)  using  SCUBA  Lake  St.  Clair  bottom 
in  this  area  is  mostly  soft  (silty  clay)  and  unionids  were  virtually 
the  only  hard  surfaces  present.  Dreissena  abundance  and  bio- 
mass  increased  from  west  to  east:  152  individuals/m-  and  26.1 
g/m-  live  mass  (west  site),  to  11,655/m-  and  845.8  g/m-  (east 
site).  Density  of  live  unionids  showed  a  reverse  trend:  2.3/m- 
and  129.5  g/m-  (west  site),  to  0.1/m-  and  16.6  g/m-  (east  site). 
At  the  central  and  east  sites  all  that  was  visible  of  any  unionid 
(live  or  empty  shell)  was  a  mound  of  Dreissena.  In  a  few  of 
those  samples,  recruitment  was  so  intense  that  the  colony  had 
grown  laterally  from  the  original  settled  surface,  resulting  in  a 
loose  mat  of  Dreissena  over  the  bottom.  Among  the  three  sites 
there  was  a  strong  negative  correlation  between  Dreissena  bio- 
mass  and  abundance  of  live  unionids  (r=  — 1.0,  P<0.01,  N=3). 
Most  of  the  more  heavily  colonized  unionids  showed  damage 
to  the  posterior  valve  edges.  These  data  demonstrate  that  the 
zebra  mussel  can  not  only  colonize  lakes  consisting  almost  en- 
tirely of  soft  substrata  but  can  reach  densities  of  10-20  thou- 
sand/m-  partially  by  lateral  extension  of  colonies  from  an  orig- 
inal "seed"  surface.  At  such  densities  they  will  have  a  major 
impact  on  the  benthos,  especially  indigenous  unionid  bivalves, 
which  are  likely  to  face  virtual  elimination. 

Key  words:  Dreissena  polymorpha,  unionids,  soft  substrata. 
Lake  St.  Clair. 


1991),  rapid  dispersal  by  passive  means  either  involving 
drift  by  veliger  larvae  or  boat /ship  dispersal,  growth  to 
se.xual  maturity  in  one  season,  and  others  (Mackie,  1991). 
In  addition,  it  is  ecologically  unique,  i.e.  it  has  a  niche 
which  is  not  represented  in  North  American  fresh  waters. 
Specifically,  there  is  no  other  attached  macrofouling  in- 
vertebrate of  any  ecological  significance  in  the  Great 
Lakes.  In  some  areas  such  as  Lakes  St.  Clair  and  Erie, 
relatively  high  primary  productivity  and  otherwise  op- 
timal trophic  conditions  have  further  contributed  to  its 
success.  Information  on  tlie  distribution  and  abundance 
of  Dreissena  in  Lake  St.  Clair  has  been  reported  by 
Hebert  et  al.,  1991,  and  its  biology  reviewed  by  Mackie, 
1991.  Hebert  et  al.  (1989)  and  Hunter  and  Bailey  (1991) 
made  the  observation  that  Dreissena  settled  on  and  at- 
tached readily  to  a  variety  of  hard  surfaces  including  the 
shells  of  native  clams  (Bivalvia;  Unionidae),  of  which 
there  are  at  least  18  species  in  Lake  St.  Clair  (Nalepa 
and  Gauvin,  1988).  Since  settled  individuals  attach  by 
means  of  a  byssus,  they  require  a  hard  substratum,  hence 
it  is  often  assumed  that  a  soft-bottom  lake  or  river  will 
not  support  significant  zebra  mussel  colonization. 

The  purpose  of  this  study  was  to  document  the  success 
of  Dreissena  in  colonizing  one  region  of  a  soft-bottom 
lake.  Lake  St.  Clair,  and  to  provide  preliminary  evidence 
of  its  impact  on  the  communit\  of  unionids  in  that  area. 


INTRODUCTION 

Lake  St.  Clair  is  part  of  the  Great  Lakes  drainage  system 
between  Lake  Huron  and  Lake  Erie.  It  is  a  relatively 
shallow  lake,  with  a  mean  depth  of  3.0m,  a  length  of 
43km,  and  a  width  of  40km  (Leach,  1991).  Most  au- 
thorities believe  that  the  zebra  mussel  (Driessena  poly- 
morpha Pallas,  1771)  was  first  introduced  into  North 
America  in  Lake  St.  Clair  in  1985  or  1986  (Hebert  et 
al..  1989,  1991;  Mackie,  1991).  Since  then,  it  has  spread 
rapidly  and  is  now  found  in  all  five  of  the  (;reat  Lakes. 
Reasons  for  its  remarkable  success  include  relatively  high 
fecundity  (ca.  10''  eggs/female  spawning  event;  Sprung, 


MATERIALS  AND  METHODS 

Three  sites  in  southern  Lake  St.  Clair  were  sampled  for 
zebra  mussels  and  unionids  on  29  and  30  September, 
1990  (Fig.  1).  All  three  sites  were  located  on  42°22'3"  N 
latitude  which  is  roughly  parallel  to  and  about  5  miles 
north  of  the  southern  shoreline  of  Lake  St.  Clair  in  Ca- 
nadian waters  (Ontario).  The  site  farthest  west  (hence- 
forth the  "west  site")  was  at  82°47'30"  W;  the  central 
and  east  sites  were  at  82°40'00"  W  and  82°32'30"  W, 
respectively.  Total  distance  between  adjacent  sites  was 
about  10.3  km  (6.4  mi;  F'ig.  1).  These  sites  were  the  same 
as  locations  15,  18,  and  21  of  Pugsley  et  al.,  1985,  and 


R.  D.  Hunter  and  J.  F.  Bailey,  1992 


Page  61 


two  of  them,  the  w  est  and  east  sites,  were  also  sampled 
b\  Nalepa  and  Gauvin  (1988).  The  depth  at  each  site 
was  6.0  ±  0.61  m. 

The  bottom  of  Lake  St.  Clair  is  nearly  all  soft  sediment, 
the  main  component  of  which  is  muddy  sand,  especially 
in  the  central  part,  with  areas  of  gravel  sand  closer  to 
shore  (Leach,  1991).  In  the  areas  sampled  here,  the  sur- 
ficial  sediment  was  virtually  the  same  from  site  to  site; 
silty  clay  overlain  by  a  thin  layer  of  detritus.  The  lake 
bed  was  relatively  flat  with  few  low  undulations.  Mac- 
rophytes  were  almost  non-e.\istent  and  nearly  the  only 
visible  solid  substrata  consisted  of  mounds  of  zebra  mus- 
sels. 

Zebra  mussel  and  unionid  sampling  were  done  using 
SCUBA  in  two  dive  teams  working  independenth ,  with 
two  members  per  team.  An  aluminum  scjuare  frame  of 
0.25  m-  area  was  used  to  obtain  a  series  of  samples  by 
casting  (throwing  forward)  the  frame,  then  by  touch, 
collecting  all  hard  objects  to  a  depth  of  approximately  2 
cm  into  the  sediment.  All  objects  within  each  quarter  m- 
were  separately  bagged,  brought  to  the  surface,  labeled, 
and  returned  to  the  laboratory  for  further  analysis.  Sam- 
pling was  not  statistically  random,  however  due  to  the 
poor  visibility  (30-50  cm),  bottom  features,  including 
mounds  of  zebra  mussels,  were  not  visible  to  the  divers 
when  the  quarter  m-  frames  were  cast.  Therefore  this 
procedure  was  relatively  free  of  sampler  bias.  Each  dive 
team  moved  about  1-2  m  further  over  the  bottom  be- 
tween successive  samples.  Sampling  protocol  requireti 
that  a  total  of  ten  "hits"  were  needed  for  each  dive  team 
in  order  to  finish  a  site.  A  hit  was  a  sample  containing 
at  least  one  zebra  mussel.  Underwater  records  were  kept 
of  total  casts  so  that  misses  could  later  be  used  for  density 
calculations.  Hence  the  data  reported  here  are  based  on 
a  different  number  of  casts  (=samples)  at  each  site,  with 
each  site  having  20  hits. 

In  the  laboratory,  each  quarter  m-  sample  was  indi- 
vidually e.xamined.  All  hard  surfaces  (mostly  unionid 
shells)  were  scraped  free  of  zebra  mussels  b)  cutting  the 
byssal  threads.  Removed  mussels  were  rinsed  free  of  sed- 
iment and  the  rinse  water  collected  and  passed  through 
a  0.5  mm  sieve  in  order  to  recover  small  individuals.  The 
smallest  zebra  mussels,  mostly  0.2-0.5  mm  shell  width, 
were  not  retained  by  this  process,  because  the  large  vol- 
umes of  animals  and  sediment  made  working  with  small- 
er meshes  impractical.  Care  was  taken  to  separately  re- 
cord clusters  of  zebra  mussels  that  were  not  attached  to 
any  other  hard  surface.  After  draining,  the  zebra  mussels 
from  each  quarter  m-  sample  were  pooled  for  weighing. 
Live  unionids  were  also  cleaned,  weighed,  and  prepared 
for  identification.  All  weights  reported  herein  are  total 
(shell  -I-  soft  tissues)  live  weight.  A  random  subsample 
of  zebra  mussels  from  each  site  was  preserved  in  10% 
formalin  (neutralized  with  CaCO,)  and  used  for  shell 
length  measurements.  Length  measurements  were  to  the 
nearest  0.1  mm.  From  these  data,  density  vs.  size  class 
plots  could  be  done  by  site. 

Dreissena  shell  to  tissue  mass  was  determined  by  dry- 
ing to  a  constant  mass  at  70°C,  animals  that  had  been 


St  Clair/ 


MICHIGAN 


42°  30 


ONTARIO  82   30 

Figure  I.    Location  of  the  three  sample  sites  in  Lake  St.  Clair. 
The  shipping  channel  is  shown  as  a  dashed  line. 


sorted  by  shell  length  .  Whole  dry  animals  were  decal- 
cified using  5%  HCl  for  2  hours  or  until  CO2  evolution 
stopped.  This  method  gives  identical  results  to  "ashing" 
methods  (Hunter  and  Lull,  1976).  The  remaining  non- 
shell  material  (internal  tissues  +  periostracum)  was  dried 
and  weighed  giving  "tissue"  dry  mass.  Computed  from 
these  values  was  the  shelhtissue  mass  ratio  w  hich  is  simply 
shell  mass  divided  by  tissue  mass,  a  number  which  is  a 
useful  inde.x  of  the  general  condition  of  the  animal. 

RESULTS 

A  total  of  104  quarter  m-  samples  were  taken.  At  the 
west  site,  out  of  43  quarter  m-  samples  (=casts),  20  had 
Dreissena  (=hits)  and  23  did  not  (=misses),  i.e.,  53.5% 
of  the  43  casts  contained  no  zebra  mussels.  Numbers  for 
the  other  two  sites  were:  33  casts,  13  misses  for  the  central 
site  (39.4%);  and  28  casts,  8  misses  for  the  east  site  (28.6%). 
Table  1  gives  the  abundance  and  biomass  for  Dreissena 
and  for  unionids  at  the  three  sites.  Dreissena  density  was 
greatest  at  the  east  site  averaging  11,655  individuals/m- 
and  845.8  g/m-;  and  lowest  at  the  west  site,  averaging 
152  individuals/m-  and  26.1  g/m-.  Each  of  the  sites 
differed  significantly  in  biomass  from  the  other  two  (one- 
factor  ANOVA,  P=0.0001;  Scheffe's  F-test,  P<0.01).  In 
terms  of  abundance,  the  west  and  central  sites  were  not 
significantly  different  but  the  other  comparisons  (west 
and  east;  central  and  east)  were  significantlv  different 
(one-factor  ANOVA,  P<0.000L  Scheffe's  F-test,  P<0.01). 
Both  biomass  and  abundance  increased  from  west  to 
central  to  east  site.  The  maximum  biomass  of  Dreissena 
recorded  for  any  sample  was  at  the  east  site  at  1409  g/m^ 
which  corresponds  to  19,417  individuals/m^. 


Page  62 


THE  NAUTILUS,  Vol.  106,  No,  2 


15 


I   10 

u 
O 
Si 

E 

Z      5 


■  Wood  &  other 

D  Druses 

E  Dead  unionids 

S  Live  unionids 


S\S-.V;\.A. 


■'•^■■'•^■^■^ 


•yrrrjrjrji 


^.>.^.^.^.i». 


•./.^.^.^.^. 


S''.«\-%'\*^ 


••^•^'f'^•^• 


'■■.'■■iX'i-i-i-: 


'•'■•i-i-iX-i-iX-i-^ 


'•■.■  '.••■.■  ••.•  •'.••'.*'''.■  'if  ^ 


••  J-  .  ^i^Z^'f'  f3 


• '.'  '.•^:  ^.-  ^.•^.•'.-  '.•^.* 


■ .  ^ .  ^•^•^-  .»«^  ■ 


West 


Central 
Site 


East 


Figure  2.  Mean  density  of  hard  substrata  in  Lake  St.  Clair  by 
type  of  material  and  by  site.  Druses  are  discreet  clusters  of 
zebra  mussels. 


In  contrast  to  Dreissena  density  trends,  density  of  live 
unionids  was  highest  at  the  west  site  and  decreased  mov- 
ing to  the  centra!  and  east  sites  (Table  1).  Unionid  density 
was  greatest  at  the  west  site  averaging  2.3  individuals/ 
m-  and  129.5  g/m-.  Out  of  28  quarter  m-  samples  at  the 
east  site,  only  one  live  unionid  was  found.  At  the  west 
site  there  were  24  live  individuals  found  over  43  quarter 
m-  samples.  The  data  for  the  east  site  were  non-normal 
due  to  the  presence  of  zeros  for  all  but  one  sample.  For 
purposes  of  statistical  analysis,  a  small  number  (0.01)  was 
added  to  all  values  which  were  then  log  (In)  transformed. 
This  transformed  data  set  was  then  statistically  analyzed 
for  differences  between  sites.  For  both  biomass  and  abun- 
dance, there  were  significant  differences  among  sites  (one- 
factor  ANOVA,  P<0.002),  and  the  east  site  differed  sig- 
nificantly from  the  other  two  (Scheffe's  F-test,  P<0.01). 

It  was  readily  apparent  b\'  inspection  of  the  lake  bot- 
tom earlier  in  the  season,  when  visibility  was  2.5-3.5  m, 


that  hard  substrata  in  this  general  area  of  the  lake  con- 
sisted of  small,  discrete  objects  imbedded  in  the  soft 
bottom.  Later  observations  indicated  they  were  fairly 
common,  ranging  from  about  2.4  objects/m-  at  the  west 
site  to  14  objects/m-  at  the  east  site,  and  they  were 
sufficiently  small  so  as  not  to  cover  a  very  large  proportion 
of  a  square  meter  except  in  a  few  of  the  east  samples. 
With  the  exception  of  3  pieces  of  wood,  and  two  snail 
shells  (Elimia  livescens  Menke  1830),  all  of  these  hard 
substrata  were  either  unionids,  unionid  shells,  or  inde- 
pendent clusters  of  zebra  mussels  (Fig.  2).  The  densit\ 
of  live  unionids  at  the  three  sites  has  been  presented  in 
Table  1  and  is  shown  in  Fig.  2  relative  to  other  hard 
substrata.  The  number  of  dead  unionids  (shells  only  with 
both  valves  usually  connected),  shows  a  reverse  trend  to 
that  of  the  living  unionids.  There  was,  on  average,  0.1 
shell  at  the  west  site,  1.4  at  the  central,  and  7.4  at  the 
east  site.  Likewise  the  number  of  Dreissena  clusters 
(druses)  that  were  independent  from  other  hard  substrata 
also  increased  from  west  to  east  (0  west,  2.6  central,  and 
6.1  east;  Fig.  2). 

Figure  3  illustrates  the  negative  correlation  between 
Dreissena  biomass  and  unionid  abundance.  This  corre- 
lation is  significant  P<0.01,  r=-1.0,  N=3.  The  plot  of 
unionid  density  also  shows  the  ratio  of  live  to  dead  (L: 
D)  animals  by  site.  At  the  west  site  there  were  25  live 
individuals  for  ever\  dead  unionid.  This  ratio  was  0.9:1 
at  the  central  site,  whereas  at  the  east  site  there  was  only 
a  single  live  unionid,  giving  an  L:D  ratio  of  0.02:1. 

Based  on  subsamples  of  fixed  material,  size  distribu- 
tions of  Dreissena  were  expressed  on  a  per  m-  basis  (Fig. 
4).  It  is  evident  that  at  both  the  central  and  east  sites 
there  was  substantial  recent  recruitment  based  on  the 
large  group  of  juveniles  from  about  1  to  5  mm  shell 
length.  Fifty-two  percent  of  the  central  sample  and  62% 
of  the  east  sample  were  made  up  of  juveniles  5  mm  or 
less. 

In  contrast,  at  the  west  site  there  was  a  pronounced 
absence  of  Dreissena  in  these  smaller  juvenile  size  classes 
(Fig.  4).  The  smallest  animal  in  the  \\est  subsample  was 
2.6  mm  and  only  3%  of  the  subsample  was  less  than  5 
mm  shell  length.  Larger  size  classes  were  well-repre- 
sented especially  in  the  central  and  east  subsamples.  For 
example  at  the  central  site,  34%  of  the  subsample  was 
comprised  of  mussels  measuring  12  mm  or  greater,  com- 


Table  1.    Mean  abundance  and  biomass  (±SE)  for  Dreissciui  and  lor  unionids  at  three  sites  in  southern  Lake  St.  Clair.  Only  a 
single  live  unionid  was  found  in  the  east  samples. 


Site  in  Lake  St.  Clair 


West 


Central 


East 


Dreissena  density 
Number/m^ 
Biomass  (g/m^) 

Unionid  density 
Nuinber/in- 
Hiomass  (g/m-) 


152  ±  36,9 
26,1  ±  6,34 

2,3  ±  0.49 
129.5  ±  27,79 


2,847  ±  620,9 

402.1  ±  87.69 

L3  ±  0  25 

73  4  ±  14,69 


1L655 

±  2,000.4 

845.8 

±  145.17 

0.1 

16  6 

R.  D.  Hunter  and  J.  F.  Bailey,  1992 


Page  63 


8 
7 
6 

CM 

Is 

n  4H 

2  2- 

1 

0 


D  Live 
■  Dead 
L:D    Ratio 


Unionids   ,  0-02:i 


25:1 


West        Central         East 


1000 


800- 


^-  600 


O) 


400- 


200- 


Dreissena 

Mean  ±  SE 


k\\\\\\\\\N 


West 


Central 
Site 


East 


Figure  3.  N4ean  abundance  of  live  and  dead  (empty  siiells) 
unionids  by  site  (upper  plot)  and  mean  biomass  of  Dreissena 
(lower  plot)  by  site  in  Lake  St.  Clair. 


pared  to  12%  at  the  east  and  9%  at  the  west  sites.  Although 
ages  were  not  determined  from  this  material,  it  is  likely 
that  the  majority  of  Dreissena  measuring  12  or  more 
mm  were  at  least  one-year-old. 

In  addition  to  the  above  differences,  the  condition  of 
the  animals  differed  among  the  sites.  There  was  a  ten- 
dency for  the  central  and  especially  the  east  animals  to 
have  areas  of  eroded  shell,  ie.  patches  where  the  peri- 
ostracum  was  clearly  missing  revealing  the  underlying 
whitish  CaCOj.  Although  no  data  were  recorded,  there 
was  also  a  higher  proportion  of  empty  Dreissena  shells 
at  the  east  site  suggesting  that  mortality  rates  were  sub- 
stantially higher  there.  Not  only  were  west  site  densities 
far  lower  (there  were  major  areas  of  live  unionid  shell 


200 


150- 


100- 


0  5  10         15        20         25         30         35 


200 


0  5  10         15         20         25         30         35 


800 


600 


400 


200 


5  10         15         20         25         30 

Shell  Length  (mm) 


35 


Figure  4.    Density  of  Dreissena  by  shell  length  size  class  at 
three  sites  in  Lake  St.  Clair. 


surface  unoccupied  b\  zebra  mussels)  but  the  Dreissena 
there  had  shells  with  fully  intact  periostraca. 

Figure  5  shows  the  relationship  between  shell  length 
and  shelLtissue  mass  ratio  for  each  site.  The  general  ten- 
dency is  that  as  the  mussels  grow  (increase  in  length) 
their  shelhtissue  ratio  also  increases,  with  a  fairly  linear 
relationship  between  the  variables.  Smaller  size  animals 
(<10  mm  shell  length)  do  not  differ  much  between  sites 
and  their  shelhtissue  ratios  are  mostly  within  the  range 
of  7-10.  In  contrast,  larger  animals  (>20  mm)  show- 
widely  differing  shelhtissue  ratios  at  different  sites  (Fig. 
5).  For  example,  a  25  mm  animal  from  the  east  site  would 
have  a  dry  tissue  mass  averaging  34  mg  and  a  dry  shell 


Page  64 


THE  NAUTILUS,  Vol.  106,  No.  2 


22 
20  - 

18  - 

16 
14 
r.    12 


I    10 
8 


-  ^  -  West 

0 

---a-   Centra 

0     East 

0         ^, 

D 

- 

D        0 

a  -  - 

1 

0 

1                1 

10 


15   20   25   30 


Shell   length   (mm) 

Figure  5.   Shell  to  tissue  mass  ratio  vs  shell  length  for  Dreissena 
at  three  sites  in  Lake  St.  Clair 


mass  of  552  mg  giving  a  shell:tissue  ratio  of  16.2.  Animals 
at  the  central  and  west  sites  of  the  same  25  mm  length 
would  have  54  and  86  mg  dry  tissue  and  812  and  821 
mg  dry  shell  for  a  shelbtissue  ratio  of  15.0  and  9.5  re- 
spectively. When  groups  of  mussels  over  20  mm  shell 
length  were  compared  across  the  three  sites,  the  east  and 
central  sites  were  not  significantK'  different  in  shelhtis- 
sues  mass  ratio,  however  the  west  site  differed  from  the 
other  two  (one-factor  ANOVA,  P<0.0001;  Sheffes  F-test, 
P<0.01).  In  other  words,  larger  sized  Dreissena  at  the 
east  and  central  sites  are  "tissue-deficient"  compared  to 
west  site  animals. 

It  was  evident  from  close  inspection  of  the  unionid 
shells  whether  of  live  or  dead  material,  that  many  of 
them  showed  damage  to  their  posterior  salve  edges  which 
was  a  direct  result  of  dense  aggregations  of  Dreissena. 
An  Anodonta  grandis  that  is  heavily  infested  with  zebra 
mussels  is  shown  in  Fig.  6.  The  posterior  one  third  to  one 
half  of  this  and  other  species  is  typically  covered  with  a 
thick  and  solid  mass  of  Dreissena.  often  more  than  2  cm 
thick.  Dreissena  attachment  of  this  magnitude  renders 
normal  valve  movement  virtually  impossible  and  activ- 
ities normally  occurring  through  the  gape  (feeding,  re- 
production, respiration,  locomotion,  etc.)  are  severly  cur- 
tailed. The  resulting  damage,  illustrated  in  Figs.  7  and 
8  with  Lampsilis  siliquoidea,  involved  substantial  de- 
formation of  the  shell  so  that  the  normally  gradual  curve 
of  the  posterior  valve  border  was  jagged  or  indented. 
The  majority  of  specimens  collected  had  levels  of  damage 
that  fell  between  the  extremes  shown  in  Figs.  7  and  8. 
In  some  specimens,  this  damage  extended  internally  so 
that  the  nacre  was  rough,  irregular,  and/or  discolored. 
In  a  few  individuals  the  damage  was  so  extensive  that  a 
gap  of  2-5  mm  remained  at  the  posterior  edge  even 
through  the  valves  were  fully  closed  (Figs.  7  and  8).  For 


Figure  6.  Specimen  of  Anodonta  grandis  from  Lake  St.  Clair 
that  has  been  heavily  colonized  b\  Dreissena  polyrr^orpha.  Such 
colonized  indi\  iduals  ranged  from  about  5  5-9.5  cm  in  length. 
Anodonta  is  shown  posterior  end  up. 


Lampsilis  siliquoidea,  the  only  species  for  which  there 
was  sufficient  material,  22.5%  of  the  shells  had  no  dam- 
age, 55/c  had  moderate  damage,  and  22.5%  had  rela- 
tively heavy  damage. 

Nine  species  of  uilionids  were  found  among  the  three 
sites  (Table  2).  For  six  of  the  species,  at  least  one  live 
individual  was  obtained,  but  three  species,  Lampsilis 
ventricosa,  Truncilla  truncata.  and  Elliptiodilatata.  were 
represented  onK  by  empty  shells.  The  unionid  fauna 
from  these  sites  was  dominated  by  Lampsilis  siliquoidea. 
such  that  this  species  comprised  79.4%  of  all  the  live 
unionids  collected.  When  empty  shells  alone  are  consid- 
ered, then  L.  siliquoidea  comprised  67.8%  of  the  unionid 
communit\-,  suggesting  that  its  dominance  has  recently 
increased.  For  all  of  the  species  found,  the  number  of 
dead  individuals  was  greater  than  or  equal  to  that  of  the 
live  indi\iduals. 

DISCUSSION 

.\  number  of  European  investigators  have  reported  on 
unionids  as  a  substratum  for  Dreissena.  Sebestyen  (1938: 
178)  concluded  that  there  was  "no  doubt  that  Dreissena 
has  a  decided  ill  effect  on  unionidae. "  In  that  same  study 
it  was  observed  that  the  native  unionids  (6  species)  w  ere 


R.  D.  Hunter  and  ].  F.  Bailey,  1992 


Page  65 


7  • 

4 

Ik 

mi 

Rki 

I 

i    m 

I^^mI 

PVmI 

■'*^ 

\  ■  H 

K,     ^1 

r  l^^fo 

I 

\  ifl 

H^'iirl-.^afla 

^^■L  ^n  ^^^1 

■ 

kJ^ 

k..Imi 

^^B             V         A^l 

f_ 

Figures  7-8.  Dorsal  (7.)  and  posterior  (8. )  views  of  two  Lamp- 
silis  siliqtioidea  shells  from  Lake  St.  Clair  from  which  all  at- 
tached Dreissena  have  been  removed.  The  shells  on  the  left 
are  nearly  normal,  whereas  the  shells  on  the  right  show  con- 
siderable damage.  One  (8.)  appears  fuzzy  due  to  remaining 
byssal  threads.  Shells  are  between  6  and  7  cm  in  length. 


in  danger  of  disappearing  entirely  from  Lake  Balaton, 
Hungary.  Sebestyen  (1938)  suggested  that  the  interaction 
resulted  in  interference  with  locomotion  and  reduction 
in  food  suppK  for  the  unionids.  In  contrast,  Lewandowski 
(1976)  concluded  that  Dreissena  had  no  unfavorable  ef- 
fects. In  spite  of  this  finding,  he  cited  other  European 
studies  that  identified  the  interaction  as  one  of  a  com- 
petitive nature.  There  was  evidence  that  Dreissena  caused 
a  slight  negative  effect  on  growth  in  shells  of  \ounger 
individuals  and  that  in  older  unionids,  the  shells  of  Dreis- 
sena-infested  individuals  were  thicker  than  shells  of 
Drei.ss^na-free  indixiduals.  The  body  (soft-tissue)  weight 


Nun- 

iber/m- 

Live 

Dead 

1.04 

1.73 

0.12 

0.23 

0.04 

0.15 

0.04 

0.04 

0.04 

0.04 

0.04 

0.04 

0 

0.23 

0 

0  04 

(.1 

0  04 

Table  2.  Mean  abundance  of  unionids  by  species  for  all  three 
Lake  St.  Clair  sites  combined.  Dead  =  empty  shells.  Total  area 
sampled  =  26  m-,  so  that  a  density  of  0.04/m^  corresponds  to 
one  individual 


Lanipsilis  siliquoidea  (Barnes,  1823) 
Leptodea  jragihs  (Rafinesque,  1820) 
Proptera  alata  (Say,  1817) 
Anodonta  grandis  (Say,  1829) 
Ligttmia  nasitta  (Say,  1817) 
Fusconaia  flava  (Rafinesque,  1820) 
Lampsilis  vcntricosa  (Barnes.  1823) 
Truncilla  truncata  Rafinesque,  1820 
Elliptio  dilalaln  Rafinesque,  1820 


was  not  significantly  altered  by  infestation.  Lewandowski 
(1976)  also  observed  posterior  shell  deformations  in  some 
of  the  Anodonta  piscinalis  and  speculated  that  the  si- 
phonal  region  was  particularly  attractive  to  post-veliger 
Dreissena. 

It  is  likely  that  the  intensitv  of  the  infestations  with 
which  Lewandowski  was  working,  (20  Dreissena  on  each 
unionid,  specificalK  Anodonta  piscinalis)  was  below  the 
level  required  to  cause  an  observable  impact.  The  above 
densities  are  comparable  to  those  reported  here  for  the 
west  site  but  are  much  lower  than  occur  in  the  central 
or  east  sites  in  southern  Lake  St.  Clair. 

Hebert  et  al.  (1989)  observed  that  Dreissena  is  often 
located  on  unionid  shells  in  Lake  St.  Clair.  They  consid- 
ered Dreissena  to  be  an  "interference  competitor"  of 
native  mussels  (Hebert  et  al..  1989;  1991).  Field  exper- 
iments in  Lake  Erie  have  shown  that  Dreissena  has  both 
species  and  sex-specific  effects  on  unionids  (Haag  et  al., 
1991). 

Two  bits  of  data  presented  in  this  studv'  are  strongly 
suggestive  that  Dreissena  is  having  a  negative  impact  on 
the  community  of  unionids  in  those  areas  of  Lake  St. 
Clair  where  Dreissena  densities  are  relatively  high.  This 
evidence  includes  the  negative  correlation  between 
Dreissena  biomass  and  unionid  densitv ,  as  well  as  dam- 
age to  unionid  shells  beneath  Dreissena  accumulations 
(Hunter  and  Bailey,  1991 ).  Negative  impact  of  Dreissena 
on  unionids  was  also  shown  by  Hebert  et  al.,  (1991)  who 
reported  that  heaviK-  infested  unionids  had  half  the  lipid 
reserves  as  unaffected  individuals.  Mackie  (1991)  con- 
cluded that  heavy  infestations  of  Dreissena  cause  several 
kinds  of  negative  effects  on  unionids,  some  of  which 
interfere  with  normal  locomotion.  Lhiequivocally  dem- 
onstrating that  zebra  mussels  cause  death  of  unionids  is 
difficult  since  direct  observation  of  death  is  impractical 
and  the  cause  is  likely  to  be  multifactorial.  Unionids  also 
die  of  a  varietv  of  other  causes  resulting  in  emptv  shells 
on  which  Dreissena  can  settle.  Hence,  observation  of 
unionid  shells  colonized  by  Dreissena  is  not  necessarily 
evidence  that  they  were  the  cause  of  death.  Observation 
of  empty  unionid  shells  bearing  deformation  to  the  pos- 
terior valve  edges  of  the  sort  illustrated  in  Figs.  7  and  8, 


Page  66 


THE  NAUTILUS,  Vol.  106,  No.  2 


is  strongly  suggestive  that  Dreissena  has  played  a  role  in 
the  death  since  there  is  little  else  that  will  cause  this  kind 
of  damage  (R.  Hoeh,  pers.  comm.). 

This  study  also  suggests  that  the  general  health  or 
condition  of  zebra  mussels  is  inversely  related  to  their 
densit\'.  Specifically,  where  the  population  density  is  high, 
as  is  the  case  at  the  central  and  east  sites,  the  larger 
animals  are  emaciated.  The  result  is  a  shell  to  tissue  mass 
ratio  that  is  high  compared  to  that  of  animals  from  low 
density  situations,  i.e.,  under  such  conditions  adult-sized 
shells  contain  an  undersized  mass  of  tissue.  As  larger 
(older)  individuals  are  settled  on  and  overgrown  by  more 
recent  recruits,  it  is  likely  they  experience  increasingly 
suboptimal  conditions,  resulting  in  an  emaciated  animal. 
Tissue  degrowth  under  poor  trophic  conditions  is  known 
to  occur  in  gastropods  (see  Russell-Hunter  and  Eversole, 
1976)  and  it  is  likely  to  be  occurring  in  Drcisscna-in- 
fested  unionids  as  well. 

Stanczykovvska  (1975)  concluded  that  as  the  Dreissena 
population  density  increased,  their  condition  decreased. 
Her  findings  were  based  largely  on  size  and  weight  data, 
i.e.,  animals  of  specific  age  groups  in  dense  populations 
(e.g.,  700/m-)  tended  to  be  smaller  in  both  length  and 
weight  than  those  from  low  densitv  conditions  (e.g.,  30/ 

Size-distribution  data  suggest  that  there  are  certainly 
two  and  probably  three  year-classes  present  at  each  site. 
This  contention  agrees  with  other  studies  w  hich  indicate 
that  Dreissena  in  both  Lakes  Erie  and  St.  Clair  can  grow 
to  well  over  10  mm  shell  length  in  one  year  (Nichols  et 
al.,  1991;  Mackie,  1991).  Thus  the  largest  individuals 
sampled  (i.e.,  20  mm  shell  length)  are  likely  to  be  in 
their  second  year  w  ith  a  few  larger  individuals  possibly 
three  years  old.  The  timing  of  recruitment  of  the  most 
recent  cohort  is  distinctly  different  at  the  west  site  com- 
pared to  the  other  two  (Fig.  4).  At  the  west  site  there  is 
an  absence  of  small  juveniles,  whereas  they  are  well 
represented  at  both  the  central  and  east  sites.  The  modal 
size  for  juveniles  (defined  here  as  individuals  <10  mm 
shell  length)  at  both  the  central  and  east  sites  was  2  mm. 
Forty-three  percent  of  the  central  sample  and  35%  of 
the  east  sample  consisted  of  juveniles  less  than  3  mm 
shell  length.  It  would  appear  that  a  relatively  intense  but 
brief  recruitment  of  post-veligers  occurred  at  the  west 
site  perhaps  4-8  weeks  before  these  samples  were  taken, 
resulting  in  a  relatively  well-defined  cohort  of  about  5- 
10  mm  in  late  September  (as  seen  in  Fig.  4).  In  contrast, 
a  more  recent  recruitment  is  in  evidence  at  the  other 
two  sites  beginning  perhaps  four  weeks  previously  and 
possibly  still  continuing  or  just  ending  as  of  late  Septem- 
ber. It  appears  that  there  was  little  recruitment  activity 
at  the  central  site  during  the  time  when  recruitment  was 
intense  at  the  west  site  (Fig.  4). 

These  contrasting  recruitment  patterns  are  likeK  to  be 
due  to  differences  in  veliger  density  in  different  major 
water  masses  and  to  wind-induced  irregularities  influ- 
encing the  extent  of  these  water  masses.  Water  on  the 
western  and  northwestern  side  of  the  lake  ("Huron  wa- 
ter) is  largely  from  the  Lake  Huron/St.  Clair  River  flow 


system  (Leach,  1980,  1991).  Huron  water  is  relatively 
veliger-poor  due  to  very  low  Dreissena  densit\  in  that 
area  in  1990,  which  would  result  in  low  recruitment 
around  the  west  site.  Water  in  eastern  and  southeastern 
Lake  St.  Clair  ("St.  Clair"  water)  is  more  stable  and 
productive  due  to  nutrient  enrichment  from  Ontario  riv- 
ers (Leach,  1980,  1991).  In  this  water  mass  Dreissena 
reached  much  higher  densities  (eg.  central  and  east  sites) 
than  elsewhere  in  the  lake.  Changing  wind  speed  and 
direction  move  the  boundries  of  these  water  masses  to 
some  degree.  If  the  St.  Clair  water  mass  extended  over 
the  west  site  for  a  few  da>s  in  summer,  then  it  could 
produce  the  recruitment  pulse  that  is  seen  in  September 
as  a  distinct  cohort  of  settled  juveniles. 

Two  of  the  sites  sampled  in  this  study  (west  and  east) 
were  approximately  the  same  as  sampled  by  Nalepa  and 
Gauvin  (1988)  in  their  study  of  unionid  distribution  in 
Lake  St.  Clair.  Since  their  samples  were  taken  in  Sep- 
tember, 1986,  they  contained  no  zebra  mussels.  The  av- 
erage unionid  densitv  thev  reported  for  the  southern  half 
of  Lake  St.  Clair  was  2^6  ±  1.91/m-  (.x  ±  s,  N=15) 
compared  to  the  average  for  the  present  study  of  1.2  ± 
1.10/m2  (N=3).  Nalepa  and  Gauvin  (1988)' found  18 
species  of  unionids  of  which  nine  were  the  same  as  those 
in  Table  2.  Of  the  nine  species  listed  in  Table  2,  four  were 
represented  by  empty  shells  only.  Although  they  col- 
lected material  from  a  much  larger  area  it  is  interesting 
to  note  that  seven  out  of  nine  species  common  to  the  two 
studies,  declined  as  a  percentage  of  the  total  numbers 
collected.  Only  Lampsilis  siliquoidea  (=L.  radiata  sili- 
qiioidea  of  Nalepa  and  Gauvin,  1988)  increased  substan- 
tially as  a  proportion  of  the  total  unionid  community. 
That  species  comprised  45%  of  Nalepa  and  Gauvin's 
unionid  abundance  whereas  it  was  79%  of  our  samples. 
Although  the  goals  of  these  two  studies  differ,  it  is  possible 
that  L.  siliquoidea  may  be  less  affected  by  Dreissena 
colonization  than  other  species  hence  has  become  an  even 
more  dominant  member  of  the  unionid  community  than 
it  was  before.  In  their  study,  Nalepa  and  Gauvin  con- 
cluded that  the  diversity  and  composition  of  unionids  in 
Lake  St  Clair  appeared  to  be  relati\ely  unchanged  since 
the  turn  of  the  century.  With  the  introduction  of  the 
zebra  mussel  at  very  nearly  the  same  time  and  location 
as  their  stud)-,  the  unionid  communit\  will  now  un- 
doubtedK  change  very  substantialK .  Because  no  indig- 
enous species  appears  to  be  able  to  tolerate  or  escape 
Dreissena  colonization,  it  is  likeK  that  the  unionid  com- 
munit)  in  Lake  St.  Clair  will  be  virtually  eliminated  in 
a  few  \ear's  time. 

ACKNOWLEDGEMENTS 

This  study  was  supported  in  part  by  the  Michigan  Sea 
Grant  College  Program  (Project  No.  M /PM-3H).  The 
cheerful  enthusiasm  and  careful  sampling  of  SCUBA 
divers  Luke  Clyburn,  Phyllis  Higman,  and  Mary  Black- 
burn is  gratefully  acknowledged.  Joe  Leach  (Ontario 
Ministr\  of  Natural  Resourses)  and  Bob  Haas  (Michigan 
DNR)  provided  thoughtful  discussion  on  Lake  St.  Clair 


R,  D.  Hunter  and  J.  F.  Bailey,  1992 


Page  67 


water  masses.  Special  thanks  are  due  to  LCDR  Luke 
CK'burn  (NSCC)  for  providing  the  boat  transportation 
and  dive  support.  Randy  Hoeh  (Univ.  of  Michigan,  Mu- 
seum of  Zoology)  kindly  identified  or  confirmed  our  iden- 
tification of  the  unionids. 

LITERATURE  CITED 

Garton,  D.,  and  VV.  Haag.  1990.  Reproduction  and  recruit- 
ment of  Drcissena  during  the  first  invasion  year  in  western 
Lake  Erie.  Abstract.  In.  Zebra  mussels:  the  Great  Lakes 
E.xperience,  Feb.,  19,  Univ.  of  Guelph,  Guelph,  Ontario, 
p4. 

Haag,  W  R  .  D  J  Berg,  and  D  W  Garton  1991  Drcissena 
polynunplia  colonies  encrusting  native  unionid  bivalves 
produce  species-specific  and  .sex-specific  effects.  Journal  of 
Shellfish  Research  10:2.59. 

Hebert,  P.D.N. ,  B.W.  Muncaster,  and  G.L.  Mackie.  1989. 
Ecological  and  genetic  studies  on  Drcissena  polijrnorpha 
(Pallas):  A  new  mollusc  in  the  Great  Lakes,  Canadian 
Journal  of  Fisheries  and  Aquatic  Science  46:1587-1591. 

Hebert,  P.D  N,,  C.C.  Wilson,  M.H.  Murdoch,  and  R,  Lazar. 
1991,  Demography  and  ecological  impacts  of  the  invad- 
ing mollusc  Drcissena  poli/nwrpha.  Canadian  Journal  of 
Zoology  69:405-409, 

Hunter,  R,D.  and  VV.W.  Lull.  1976.  A  comparison  of  two 
methods  for  estimating  the  weight  of  inorganic  materials 
in  molluscs.  Malacological  Review  9:118-120. 

Hunter,  R.D,  and  J.F.  Bailey.  1991,  Colonization  of  soft  sub- 
strata by  zebra  mussels:  role  of  and  cost  to  native  bivalves. 
Journal  of  Shellfish  Research  10249-250 

Leach,  J  H.  1980.  Limnological  sampling  intensity  in  Lake 
St.  Clair  in  relation  to  distribution  of  water  masses.  Journal 
of  Great  Lakes  Research  6:141-145. 

Leach.  J.H.  1991.  Biota  of  Lake  St.  Clair:  habitat  evaluation 
and  environmental  assessment.  Hydrobiologia  219:187- 
202. 


Lewandowski,  K  1976.  Unionidae  as  a  substratum  for  Dreis- 
sena  polymorpha  Pall.  Polish  Archives  of  Hydrobiology 
23:409-420. 

Mackie,  G.L.  1991.  Biology  of  the  exotic  zebra  mussel,  Dreis- 
sena  pohjmorpha,  relative  to  native  bivalves  and  its  po- 
tential impact  in  Lake  St.  Clair  H\drobiologia  219:251- 
268. 

Nalepa,  T.F.  and  J.M.  Gauvin.  1988.  Distribution,  abundance, 
and  biomass  of  freshwater  mussels  (Bivalvia:  Unionidae) 
in  Lake  St.  Clair.  Journal  of  Great  Lakes  Research  14:41 1- 
419. 

Nichols,  S.J.,  A.M.  Bitterman,  and  F,  Ely,  1991,  Growth  rates 
of  Drcissena  polymorpha  in  the  St.  Clair  River  and  Lakes 
St.  Clair,  Erie,  and  Ontario  from  June  to  November,  1990. 
Journal  of  Shellfish  Research  10:  250. 

Pugsley,  C.W.,  P.D.N,  Hebert,  G,W.  Wood,  G.W.  Brotea,  and 
T.W.  Obal.  1985.  Distribution  of  contaminants  in  clams 
and  sediments  from  the  Huron-Erie  corridor.  I  -  PCBs  and 
octachlorostv  rene  Journal  of  Great  Lakes  Research  11: 
275-289. 

Russell-Hunter.  WD,  and  AG.  Eversole.  1976.  Evidence  for 
tissue  degrowth  in  starved  freshwater  pulmonale  snails 
(Helisoma  trivolvis)  from  tissue,  carbon,  and  nitrogen 
analyses.  Comparative  Biochemistry  and  Phvsiologv  54A: 
447-453. 

Sebestyen,  O.  1938.  Colonization  of  two  new  fauna-elements 
of  Pontus-origin  (Drcissena  polymorpha  Pall,  and  Coro- 
phium  curvispinum  GO,  Sars  forma  devium  Wundsch) 
in  Lake  Balaton,  Verhandhmgen  Internationale  Vereini- 
gung  Limnologie  8:169-181, 

Sprung,  M,  1991,  Costs  of  reproduction:  a  study  on  metabolic 
requirements  of  the  gonads  and  fecundity  of  the  bivalve 
Drcissena  polymorpha  Malacologia  33:  63-70. 

Stanczykowska,  A.  1975.  Ecosystem  of  the  Mikolajskie  Lake. 
Regularities  of  the  Drcissena  polymorpha  Pall  (Bivalvia) 
occurrence  and  its  function  in  the  lake,  Polish  Archives  of 
Hydrobiology  22:73-78. 


THE  NAUTILUS  106(2):68-71,  1992 


Page  68 


New  Ecphoras  (Gastropoda:  Thaididae:  Ecphorinae)  from  the 
Calvert  Formation  of  Maryland  (Langhian  Miocene) 


Edward  J.  Petuch 

Department  of  Geology 
Florida  Atlantic  University 
Boca  Raton,  FL  33431 


ABSTRACT 

Three  new  ecphoras  are  described  from  the  relatively  unstudied 
Shattuck  Zones  12  and  14  of  the  Plum  Point  Member  of  the 
Calvert  Formation  (Langhian  Miocene)  of  Maryland.  The  new 
species,  Ecphora  {Ecphora)  chesapeakensis  n.sp.,  Ecphora  (Ec- 
phora)  turneri  n.sp.,  and  Ecphora  (Trisecphora)  scientistensis 
n.sp.,  fill  in  gaps  in  the  evolutionary  lineages  of  the  Ecphora 
ganlnerae  species  complex,  the  Ecphora  choptankensis  species 
complex,  and  the  Ecphora  (Trisecphora)  tricostata  species 
complex,  respectively. 

Key  words:  Ecphora,  Gastropoda;  Calvert  Formation;  Mio- 
cene; Marvland. 


INTRODUCTION 

Since  the  publication  of  the  "Field  Guide  to  the  Ec- 
phoras" (Petuch,  1989),  three  new  species  of  Ecphora 
Conrad,  1843  have  been  brought  to  my  attention.  These 
were  collected  in  the  relatively  unstudied  Beds  12  and 
14  (Zones  12  and  14  of  Shattuck,  1904:xxvii)  of  the  Cal- 
vert Formation  (Plum  Point  Member),  along  the  Calvert 
Cliffs  of  western  Chesapeake  Bay,  in  Calvert  County, 
Maryland.  As  the  preservation  of  the  moUuscan  fossils  of 
Beds  12  and  14  is  not  good,  the  result  of  extensive  leach- 
ing by  groundwater,  most  specimens  of  upper  Calvert 
ecphoras  are  collected  in  a  fragmentary  state.  In  spite  of 
this  difficult  collecting  of  extremely  fragile  and  shattered 
specimens,  Mr.  Joseph  Turner  of  Baltimore,  Maryland, 
through  great  diligence  and  patience,  has  managed  to 
extract  a  number  of  unusual  forms  from  these  virtually 
unworkable  beds.  Through  his  generosity  in  the  donation 
of  numerous  study  specimens,  I  was  able  to  determine 
that  three  ecphoras  were  new  to  science,  and  these  are 
described  here.  This  paper  is  considered  an  addendum 
to  "Field  Guide  to  the  Ecphoras." 

The  three  new  species  fill  gaps  in  the  evolutionary 
lineages  of  three  main  groups  of  ecphoras.  One  new 
species,  here  named  Ecphora  (Ecphora)  turneri  n.sp.,  is 
morphologically  intermediate  between  E.  [Ecphora) 
wardi  I'etuch,  1989  (figure  5)  from  Bed  10  of  the  Calvert 
Formation  (Langhian  Stage,  Miocene)  and  £.  [Ecphora) 
choptankensis  vokesi  Petuch,  1989  (figure  6)  from  Bed 


16  of  the  Choptank  Formation  (Serravallian  Stage,  Mio- 
cene). Likewise,  a  new  three-ribbed  ecphora,  here  named 
Ecphora  (Trisecphora)  scientistensis  n.sp.,  is  interme- 
diate between  E.  [Trisecphora)  eccentrica  Petuch,  1989 
(figure  9)  of  Bed  10  of  the  Calvert  Formation  and  E. 
(Trisecphora)  smithae  Petuch,  1988  (figure  11)  from  Bed 
16  of  the  Choptank  Formation.  The  third  new  species, 
here  named  Ecphora  (Ecphora)  chesapeakensis  n.sp.,  is 
the  oldest  known  member  of  the  E.  (Ecphora)  gardnerae 
Wilson,  1987  species  complex.  This  new  ecphora  is  an- 
cestral to  £.  (Ecphora)  williamsi  Ward  and  Gilinsky, 
1988  (figure  2)  from  Bed  19  of  the  Choptank  Formation, 
which  previously  had  been  thought  (Petuch,  1989)  to 
have  been  the  original  progenitor  of  the  wide-ribbed  E. 
gardnerae  species  group. 

In  this  paper,  I  use  the  morphological  criteria  for  ec- 
phora species-level  determinations  that  are  outlined  in 
my  earlier  works  (Petuch,  1988,  1989).  The  holotypes  of 
the  new  taxa  are  deposited  in  the  invertebrate  paleon- 
tology collection  of  the  Florida  Museum  of  Natural  His- 
tory, University  of  Florida,  Gainesville,  Florida,  and  bear 
UF  numbers. 

SYSTEMATICS 

Class  Gastropoda 

Subclass  Prosobranchia 

Order  Caenogastropoda 

Superfamily  Muricacea 

Family  Thaididae 

Subfamily  Ecphorinae  Petuch,  1988 

Genus  Ecphora  Conrad,  1843 


chesapeakensis  new  species 


Ecphora  [Ecphora 
(figures  3,  4) 

Materials  examined:  Holotijpe:  Length  (incomplete 
and  reconstructed)  28  mm,  in  Shattuck  Zone  14,  Plum 
Point  Member,  Calvert  Formation,  at  Scientists  C'liffs, 
(Calvert  County,  Maryland,  Chesapeake  Bay,  UF  23798; 
Paratype:  length  (incomplete  and  reconstructed)  35  mm, 
.same  locality  as  holotype,  Petuch  collection. 

Description:  Shell  cylindrical  in  shape,  slightly  inflated; 
shoulder  rountled;  body  whorl  ornamented  with  4  large, 


E.  J.  Petuch,  1992 


Page  69 


Figures  1-11.  Ecphoras  from  the  Miocene  of  Maryland.  1.  Ecphora  (Ecphora)  calvertensis  Petuch,  1988,  dorsal  view  of  43  mm 
specimen,  Shattuck  Zone  12,  Calvert  Formation.  2.  Ecphora  (Ecphora)  willianui  Ward  and  Gilinsky,  1988,  dorsal  view  of  57  mm 
specimen,  Shattuck  Zone  19,  Choptank  Formation.  3.  Ecphora  (Ecphora)  chesapeahensu  n.sp.,  ventral  view  of  holotype,  length 
(incomplete)  28  mm,  UF  23798,  Shattuck  Zone  14,  Calvert  Formation.  4.  Ecphora  (Ecphora)  chesapeakensis  n.sp.,  ventral  view 
of  paratype,  length  (incomplete)  35  mm,  Shattuck  Zone  14,  Calvert  Formation  5.  Ecphora  (Ecphora)  wardi  Petuch,  1989,  ventral 


Page  70 


THE  NAUTILUS.  Vol    106,  No.  2 


wide,  rounded,  adherent  cordlike  ribs  that  are  sHghtK 
"T"-shaped  in  cross  section;  ribs  sculptured  with  1-4 
thin,  impressed  spiral  threads;  shoulder  rib  largest  and 
widest,  curving  upward  ( posteriorward )  to  produce  can- 
aliculate spire  whorls;  wide  shoulder  rib  slightK  in- 
curved, producing  distinctly  rounded  appearance;  areas 
between  ribs  relatively  smooth,  with  only  few  very  fine 
spiral  threads;  siphonal  canal  well  developed,  orna- 
mented with  numerous  large  spiral  threads;  umbilicus 
narrow. 

Etymology :  Named  for  the  Chesapeake  Bay,  which  bor- 
ders the  type  locality. 

Discussion:  Ecphora  chesapeakensis  is  closest  to  E.  wil- 
liarwii  Ward  and  Gilinsky,  1988  (figure  2)  from  the  youn- 
ger Choptank  Formation,  and  appears  to  be  its  direct 
ancestor.  The  new  species  differs  from  its  Choptank  de- 
scendant in  being  a  smaller,  less  inflated  shell  with  thin- 
ner, less  rounded  ribs.  The  ribs  of  E.  williamsi  are  lower 
and  more  adherent,  while  those  of  E.  chesapeakensis 
project  farther  from  the  body  whorl.  Both  species  have 
similar  wide,  rounded,  incurved  shoulder  ribs.  Ecphora 
cheaapeakenais  is  ai.so  similar  to  £.  calvertenaiH  Petuch, 
1988  (figure  Ij  from  Shattuck  Zone  12,  but  differs  in 
having  distinctly  rounded,  cordlike  ribs  instead  of  the 
sharply-flanged,  "T"-shaped  ribs  of  the  older  Calvert 
species.  This  new  ecphora  constitutes  a  morphfjlogical 
link  between  the  generalized  E.  calvertemis  and  the  E. 
gardnerae  species  complex  of  the  later  Miocene.  Ecphora 
cheaapeakennia  is  confined  to  Shattuck  Zone  14  (Bed  14, 
Calvertj. 

Ecphf/ra  (Ecphora)  turneri  new  species 
(figures  7,  H) 

Material  examirH'<l:  llololype:  Length  (incomplete)  38 
rnrn,  in  Shattuck  Zone  14,  Plum  Point  Member,  Calvert 
FoTinayioii,  approximately  I  km  rjortli  of  (iovernor  Bun, 
Calvert  fJiifs,  Calvert  Oiurrty,  Maryland,  ( ;(iesa[jcakc 
Bay,  UF  2)465;  ParaUjpe:  length  44  rrun,  sann-  locality 
as  holotype,  Peluch  collection. 

DeHcriplion:  Shf;ll  cylindrical  in  shape,  with  sharply 
angled  shoulder;  body  whorl  ornamented  with  4  thin, 
narrow,  hladelike  ribs;  edge  of  ribs  rounded;  subsutural 
areas  flattened,  (jroducing  stepjjed,  s<alariiorin  spire;  ar- 
eas between  rib.s  smooth,  without  spiral  s(  iilptnre;  si- 
phonal  canal  (elongated;  utnbilic us  n;irrow  Uiit  well  rie 
veloped. 


Etymolog>':  The  ta.xon  honors  Mr.  Joseph  Turner  of 
Baltimore,  Maryland,  who  generously  donated  a  large 
suite  of  research  material  from  Shattuck  Zones  12  and 
14  of  the  Calvert  Formation. 

Discussion:  Ecphora  turneri  is  closest  to  £.  choptanken- 
sis  vokesi  Petuch,  1989  (figure  6)  from  Shattuck  Zone  16 
of  the  Choptank  Formation,  but  differs  in  being  a  much 
smaller,  much  more  cylindrical  and  slender  shell,  with 
lower,  less  projecting  ribs.  The  umbilicus  of  E.  turneri 
is  also  much  narrower  than  that  of  E.  choptankensis 
vokesi,  and  the  new  species  has  a  proportionally  longer 
siphonal  canal.  Ecphora  turneri  is  confined  to  Shattuck 
Zone  14  (Bed  14,  Calvert). 

Subgenus  Trisecphora  Petuch,  1988 

Ecphora  (Trisecphora)  scientistensis 
new  species 
(figure  10) 

Material  examined:  Holotype:  Length  (incomplete, 
fragmentary)  29  mm,  in  Shattuck  Zone  12,  Plum  Point 
Member,  Calvert  Formation,  at  Scientists  Cliffs,  Calvert 
County,  Maryland,  Chesapeake  Bay,  UF  23799. 

Description:  Shell  cylindrical,  slightly  inflated  in  shape; 
shoulder  sharply  angled;  subsutural  area  flattened,  pla- 
nar; spire  whorls  .scalariform,  ste[)ped;  body  whorl  or- 
namented with  3  large  cordlike  ribs;  ribs  rounded  on 
edges,  sculptured  with  1  or  2  faint,  shallow  impressed 
spiral  threads;  areas  between  ribs  smooth,  without  spiral 
sculpture. 

Etymology:  Named  for  the  Scientists  (>liffs,  ("alvert 
County,  Maryland,  the  type  locality. 

Discussion:  Ecphora  (Trisecphora)  scientistensis  is 
(•los<-st  to  /•;.  (Trisecphora)  eccentrica  Petuch,  1989  (fig- 
ure 9)  from  Shattuck  Zone  10  of  the  Calvert  I'ornKilion, 
but  differs  in  being  a  more  cylindrical  shell,  by  lacking 
fine  spiral  threaded  .sculpturing  between  the  ribs,  and  by 
having  a  lower,  ;idherent  spire  that  does  not  become 
detaclied  and  uncoiled.  'I"he  ribs  of  /•,'.  (Trisecphora) 
scientistensis  are  also  thicker  than  those  of  E.  (Trisec- 
phora) eccentrica.  The  new  si)ecies  is  also  somewhat 
similar  to  E.  (Trisecphora)  tricostata  Martin,  1904,  also 
from  Zone  10,  but  that  well  known  species  luis  w  ider  ribs 
that  are  "T"-sliai)ecl  in  cross  section  and  al.so  has  a  more 
inflated,  globo.se  shell.  Ecphora  (Trisecphora)  tricostata, 
like  /','.  (Trisecphora)  eccentrica,  has  detached,  uncoiled 
whorls,  wliile  the  whorls  of  E.  (Trisecphora)  scientislen- 


vicw  of  lioffjtypc,  len(;lli  70  iiiiii,  Slialliick  Zone  10,  Calvert  I'oriniitioii  (tiikcn  from  Ward  and  (illiiisky,  1988:  plate  I.  figure  4) 
h.  Erphora  (Ecphora)  clioptankciisis  ixikcsl  I'i^Iik  li  \'IH'),  vcnlral  view  of  paratype,  fenglli  ()2  nun,  Siialluck  Zone  !(>,  Ofioplank 
(''orriiiition  7,  K.  Ecphora  (Ecphora)  Inrncrt  w  .p  doisal  .imi  ventral  views  of  liololype,  Icngtfi  (iiudinpiete)  38  nnn,  I'l''  214(i5, 
SlialtiKJc  /one  M,  Calvert  T'or/iialioii  ').  Etjihorii  {'t'rb.c(pli()r(i)  ccccnlrlra  retiicli,  lOKO,  cforsaf  view  ol  paiatvpe.  fcnglli  5(i  mm, 
Sfiiitliiek  /.one  10.  (lufvcrt  I'drnialion  10.  /'>7)/i«r«  C/V/.srr/i/ifyff/)  vi /cii^/.v/c/i.vi.v  nsp.  dorsal  view  ol  ' 
(raxnienlary)  2\)  mm.  Ill''  23700,  Sliallmk  /one  12,  <  advert  lormalioir   I 


viitw  ol  paratype,  leiigtli  75  mm,  Stiallii<k  /one  10,  ( liioplinjk  I'onnatioii 


lololvpe,  leiiglfi  (incomplete, 
l'.(  liliiiKi  ( '/'rrvcc/i/iora)  siiilllidf  I'dniii,   10S8.  dorsal 


E.  J.  Petuch.  199: 


«s  are  tightly  coiled  and  attached.  The  new  species  is  Petuch,  E.  J     19S9    Field  guide  to  the  Ecphoras.  CoastsI  Ed- 

confined  to  Shattuck  Zone  12  of  the  Calvert  Formation  ueation  and  Research  Foundati.:-     "     -    -:es%ille.  \"A_ 

(Bed  12.  Calvert).  140 p. 

Shattuck.  G  B,    1904.   Geological  and  pai^nJuccaC'^iical  relatiotis. 

with  a  review  of  eariier  investigations.  In.  Clark.  W   R. 

G   B.  ShattucL  and  W   H.  DalT  eds.  .  The  Miocece  de- 

LITER.\TIRE  CITED  poats  of  Mar>  land  Mar>  land  Geological  Sun.e> .  Miocene. 

p.  xiii-cniL 
Petuch.  E.  J.     19SS.    New  species  of  Ecphora  and  ecphorine  \\'ard.  I_  \V.  and  X.  L  Gilinsk> .    19SS.    Ecphonz  Gastropoda; 

thaidids  from  the  Miocene  of  Chesapeake  Ba>.  Mainland.  Muricidae'  from  the  Chesapeake  group  of  Maryland  and 

U.S-.\.  Bulletin  of  Paleomalacolog>  lvl':l-16.  Virginia.  Notuke  Naturae  469(15  Marcfa':l-il. 


THE  NAUTILUS  106(2):72-75,  1992 


Page  72 


Parallel  Development-Depth  Trends  in  Deep-Sea 
Turrid  Snails  from  the  Eastern  and  Western 
North  Atlantic 


Elizabeth  Potter 

Massatliusetts  Water  Resources 

Authority 
100  First  Avenue 
Charlestown,  MA  02129 


Michael  A.  Rex' 

Department  of  Biology 
University  of  Massachusetts 
Boston,  MA  02125 


Bouchet  (1976a)  first  pointed  out  that  a  surprisingly  high 
proportion  of  deep-sea  prosobranchs  bear  larval  shells 
that  indicate  planktotrophic  development,  a  finding  since 
supported  by  numerous  surveys  of  deep-sea  snail  faunas 
(e.g.  Bouchet  and  Waren,  1979,  1980,  1985;  Rex  and 
Waren,  1982;  Colman  et  al.,  1986a).  Studies  using  oo- 
cyte-size  distributions  (Rex  et  al.,  1979;  Colman  et  al., 
1986b),  stable  isotope  composition  of  larval  and  adult 
shells  (Bouchet  and  Pontes,  1981;  Killingley  and  Rex, 
1985)  and  direct  recovery  of  larvae  from  surface  plank- 
ton and  benthic  egg  capsules  (Bouchet,  1976b;  Bouchet 
and  Waren,  1980;  Gustafson  et  al.,  1991)  have  corrob- 
orated the  use  of  larval  shell  morphology  to  infer  mode 
of  development,  and  revealed  that  planktotrophic  larvae 
of  some  species  undergo  vertical  migration  in  the  water 
column.  Both  the  frequency  of  planktotrophic  devel- 
opment in  deep-sea  snails  and  ontogenetic  vertical  mi- 
gration violate  theoretical  predictions  about  larval  de- 
velopment in  this  cold,  remote  and  nutrient-poor 
environment  (Thorson,  1950;  Vance,  1973).  Rex  and 
Waren  (1982)  showed  that  the  incidence  of  plankto- 
trophic development  in  prosobranchs  actually  increases 
with  depth  in  the  western  North  Atlantic.  Bathymetric 
trends  in  developmental  mode  have  not  been  analyzed 
in  other  regions  of  the  deep  sea  to  establish  the  generality 
of  this  pattern.  Here,  we  compare  development-depth 
trends  between  the  eastern  and  western  North  Atlantic 
for  the  Turridae,  the  largest  family  of  deep-sea  proso- 
branchs. 

The  morphological  criteria  for  determining  the  mode 
of  larval  development  are  particularly  clear  in  turrids 
(Bouchet,  1990)  and  have  been  confirmed  by  direct  ob- 
servation in  coastal  representatives  (e.g.  Labour,  1934; 
Thorson,  1946;  Thiriot-Quievreux,  1972).  Planktotro[ihic 


'  Correspond  witfi  M   A    Rex 


larval  shells  have  high  spires,  2-5  whorls,  brown  color 
and  fine  reticulate  or  ribbed  sculpture.  The  shell  consists 
of  two  distinct  parts:  a  minute  Protoconch  I  at  the  apex 
which  is  deposited  before  hatching,  and  a  larger  Proto- 
conch II  grown  during  the  planktotrophic  phase  of  de- 
velopment (Robertson,  1974).  Commonly,  these  larval 
shells  are  of  the  sinusigera  type  (Robertson,  1974)  which 
have  a  distinctive  beak  on  the  aperture  that  projects 
between  the  velar  lobes  during  the  free-swimming  stage. 
In  contrast,  shells  of  nonplanktotrophic  larvae  have  a 
single  bulbous  whorl,  are  of  the  same  white  or  gray  color 
as  the  adult  shell,  and  either  lack  visible  sculpture  or 
have  very  simple  robust  sculpture.  Nonplanktotrophic 
larvae  are  provisioned  with  sufficient  food  to  undergo 
development  and  hatch  from  benthic  egg  capsules. 
Though  nonplanktotrophic  larvae  generally  seem  to  have 
less  dispersal  potential  than  planktotrophic  forms,  the 
nonplanktotrophic  larva  of  at  least  one  turrid  has  a  brief 
swimiTiing  phase  after  hatching  and  then  lives  demersally 
before  metamorphosis  (Shiniek,  1986). 

The  two  data  .sets  compared  are  from  Rex  and  Waren 
(1982)  and  Bouchet  and  Waren  (1980)  for  the  western 
and  eastern  North  Atlantic  respectively.  We  used  only 
those  species  for  which  larval  shells  are  known  and  which 
belong  to  genera  included  in  the  Turridae  by  Bouchet 
and  Waren's  taxonomic  revision.  This  included  39  west- 
ern species  (80%  with  planktotrophic  development)  and 
90  eastern  species  (74%  with  planktotrophic  develop- 
ment). Rex  and  Waren  (1982:  Table  1)  provide  relative 
abundance  data  for  species  in  20  epibenthic  sled  samples 
collected  from  south  of  New  England,  whereas  Bouchet 
and  Waren  report  depth  records  for  material  from  many 
sources  includeil  in  their  taxonomic  revision.  To  make 
the  two  data  bases  more  compatible,  we  determined  the 
known  depth  range  for  each  species  in  the  separate 
regions,  and  calculated  the  percentage  of  species  with 
planktotrophic  development  in  each  500  m  depth  inter- 
val from  the  continental  shelf  to  the  aby.s.sal  plain.  We 
then  regressed  the  frequencies  of  planktotrophic  devel- 


E.  Potter  and  M.  A.  Rex,  1992 


Page  73 


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Depth  (m) 

Figure  I .  Relationships  between  the  proportion  of  species  with 
planktotrophic  development  in  deep-sea  turrid  snails  and  depth, 
for  the  western  (WNA)  and  eastern  (ENA)  North  Atlantic. 
Regression  equations  and  their  statistics  for  WNA  and  ENA 
respectively  are;  Y  =  0.-161  +  0.t)0012X;  N=ll,  R^  =  0.801. 
F  =  36.155.  P<0.001;  Y  =  0  486  +  O.OOOIO.X;  N  =  I1.  R-  = 
0.798.  F  =  35.476.  P<0.001. 


opment  against  depth,  and  performed  an  analysis  of  co- 
variance  (ANCOVA)  to  test  for  differences  in  slope  and 
elevation  of  the  development-depth  relationships  be- 
tween the  two  regions. 

Turrids  from  both  regions  show  highly  significant  in- 
creases in  the  incidence  of  planktotrophic  development 
with  depth,  and  the  trends  are  very  similar  (Figure  1; 
see  caption  for  regression  ecinations  and  their  statistics). 
The  ANCOVA  reveals  that  the  regressions  for  the  eastern 
and  western  North  Atlantic  faunas  are  not  statistically 
different  in  either  slope  (F  =  0.855;  d.f.  =  1,18;  n.s.)  or 
elevation  (F  =  0.952;  d.f.  =  1,19;  n.s.).  When  variables 
used  in  a  regression  are  percentage  values,  such  as  here 
with  the  frequency  of  planktotrophy,  arcsine  transfor- 
mations can  be  used  to  correct  for  departures  from  nor- 
mality (Sokal  and  Rohlf,  1981).  When  this  is  done  the 
regressions  remain  highly  significant  (R-  =  0.736,  F  = 
25.040.  N  =  11.  P  <  0.001;  a'nd  R-  =  0.797,  F  =  35.406, 
N  =  11,  P  <  0.001  for  western  and  eastern  faunas  re- 
spectively), and  are  still  indistinguishable  in  slope  (F  = 
2.246;  d.f.  =  1,18;  n.s.)  and  elevation  (F  =  3.688;  d.f.  = 
1,19;  n.s.)  by  the  ANCOVA.  There  are  some  idiosyncra- 
sies in  the  trends.  For  example,  there  is  a  marked  increase 
of  about  40%  in  planktotrophy  at  2500  m  in  the  western 
fauna,  and  a  large  increment  of  about  20%  at  1000  m 
in  the  eastern  group  (Figure  1).  But  overall,  the  two 
faunas  show  convincing  increases  in  planktotrophic  de- 
velopment with  depth,  and  are  remarkably  alike  in  both 
the  level  of  planktotrophy  and  its  rate  of  increase  with 
depth. 

The  analysis  above  suggests  that  a  depth-related  in- 
crease in  the  proportion  of  turrid  species  with  plankto- 


trophic development  is  widespread  in  the  North  Atlantic. 
It  would  be  interesting  to  expand  this  research  to  include 
the  proportion  of  endemic  and  shared  species  in  eastern 
and  western  basins.  This  would  enable  us  to  explore,  for 
example,  whether  horizontal  geographic  range  is  asso- 
ciated with  mode  of  development  (Jablonski  and  Lutz, 
1983).  We  could  also  determine  what  proportion  of  the 
similarity  in  response  show  n  in  Figure  1  is  attributable 
to  shared  and  endemic  components  of  the  two  faunas. 
At  present,  however,  it  is  not  possible  to  critically  eval- 
uate similarity  in  species  composition  on  such  large  spa- 
tial scales.  There  has  been  no  taxonomic  synthesis  of 
deep-sea  turrids  for  the  North  Atlantic  as  a  whole.  Except 
for  abundant  and  well-known  species  (see  e.g.  Etter  and 
Rex,  1990),  it  is  still  unclear  which  species  are  actually 
common  to  both  regions.  Also,  the  large  difference  in  the 
number  of  species  represented  in  the  two  data  bases 
places  severe  limitations  on  any  calculated  faunal  simi- 
larity. The  degree  to  which  the  two  available  species  lists 
reflect  a  real  shift  in  the  regional  diversity  of  turrids  or 
merely  the  very  substantial  inter-regional  difference  in 
sampling  intensity  remains  to  be  established. 

It  is  intriguing  that  deep-sea  turrids  should  have  such 
a  high  proportion  of  species  with  planktotrophic  devel- 
opment, and  even  show  an  increase  in  planktotrophy 
with  depth,  in  an  environment  where  nonplanktotrophic 
development  appears  to  predominate  in  most  taxa. 
Bouchet  (1988)  proposed  that  modes  of  larval  develop- 
ment in  deep-sea  taxa,  including  snails,  represent  phy- 
logenetic  constraints,  and  are  of  no  immediate  adaptive 
significance.  His  arguments  are  especially  compelling  for 
groups  like  the  archaeogastropods,  which  are  found 
throughout  the  deep  sea  and  have  only  nonplanktotroph- 
ic development.  In  these  groups,  larval  development  may 
be  incidental  to  other  adaptive  features  directly  related 
to  geographic  distribution.  In  turrids,  developmental 
mode  is  a  fixed  species-wide  characteristic  (Bouchet  1989, 
1990).  Planktotrophy  is  an  ancestral  condition  that  has 
been  lost  in  many  lineages,  presumably  during  the  course 
of  speciation  and  adaptive  radiation.  The  loss  of  feeding 
larvae  in  snails  is  potentially  reversible  (Strathmann, 
1978),  but  there  is  no  evidence  that  species  have  reac- 
quired planktotrophic  development  during  the  evolu- 
tionary histor\  of  turrids  (Bouchet,  1990). 

However,  while  historical  evolutionary  events  are  re- 
sponsible for  establishing  the  overall  developmental 
makeup  of  deep-sea  snail  assemblages,  it  is  difficult  to 
accoimt  for  the  clear  and  consistent  depth  trends  shown 
in  Figure  1  in  purely  nonadaptive  terms.  Environmental 
gradients  associated  with  depth  have  caused  marked 
bathymetric  changes  in  the  density  (Rex  et  ai,  1990), 
species  diversity  (Rex,  1983)  and  species  composition 
(Rex,  1977)  of  deep-sea  snail  faunas  Individual  species 
also  exhibit  clinal  effects  with  depth  (Etter  and  Rex, 
1990),  particularly  across  the  upper  bathyal  region  where 
nonplanktotrophic  development  is  more  common  (Fig- 
ure 1).  Depth-correlated  selective  gradients  might  be 
responsible  for  both  driving  the  processes  of  population 
differentiation  and  speciation  that  ultimately  generate 


Page  74 


THE  NAUTILUS,  Vol.  106,  No.  2 


new  species,  and  for  actively  maintaining  their  geo- 
graphic distributions.  Rex  and  Waren  (1982)  suggested 
two  ways  in  which  modes  of  larval  development  in  tur- 
rids  might  be  adaptively  related  to  this  selective  gradient. 
Nonplanktotrophic  development  might  confer  an  ad- 
vantage at  upper  bathyal  depths  because  species'  bathy- 
metric  and  horizontal  ranges  are  much  more  restricted 
there.  Large-scale  dispersal  in  ocean  currents  would  car- 
ry planktotrophic  larvae  away  from  adult  habitats  and 
limit  successful  recruitment.  Conversely,  selection  may 
favor  increased  dispersal  ability  at  greater  depths  to  en- 
able turrids  to  track  progressively  more  rare  and  patchy 
pre>  resources. 

Clearly,  adaptation  cannot  provide  a  complete  expla- 
nation for  the  trends  shown  in  Figure  1,  because  species 
with  planktotrophic  and  nonplanktotrophic  develop- 
ment are  found  at  all  depths.  Latitudinal  gradients  in 
the  proportion  of  development  types  in  prosobranchs 
(Thorson,  1950)  present  a  similar  problem  of  interpre- 
tation. Adaptation  and  phyletic  constraint  are  both  plau- 
sible hypotheses  to  account  for  part  of  the  bathymetric 
variation  in  developmental  modes  of  deep-sea  turrids. 
Presently,  it  is  not  feasible  to  test  these  hypotheses  ex- 
perimentally b\  showing  whether  modes  of  development 
are  adaptive  to  the  ecological  conditions  at  different 
depths  in  the  sense  that  they  actually  improve  fitness. 
However,  a  phylogenetic  study  of  deep-sea  turrids  could 
demonstrate  whether  or  not  the  same  developmental 
modes  have  arisen  independentK  in  similar  selective  re- 
gimes. 

We  thank  Philippe  Bouchet  and  Ron  Etter  for  reading 
the  manuscript.  Ron  Etter  also  provided  assistance  in 
performing  the  ANCOVA.  This  research  was  supported 
by  the  Department  of  Biology,  University  of  Massachu- 
setts at  Boston. 


LITERATURE  CITED 

Bouchet,  P.    1976a.    Mise  en  evidence  destades  larvairesplaiic- 

loniques  chez  des  Gasteropodes  Prosobranches  des  etages 

hathyal  et  abyssal.  Bulletin  du  N4useum  national  d'Histoire 

naturelle  277;  947-972. 
Bouchet,   P.     19761)      Mi.se  en  evidence  d  uiie  migration  de 

larves  veligeres  entre  I'etage  abyssal  et  la  surface,  t^omptes 

Kendus  des  Seances  de  rAcademie  des  Sciences,  Paris,  283: 

821-824. 
Bouchet,  P.     1988.     Adaptive  and  non-adaptive  strategies  in 

larval  biology  of  deep-sea  snails.   Abstracts  of  the  Fifth 

Deep-Sea  Biologv  Svin[)osiuni,  Centre  de  Brest,  France, 

IFREMEK,  p.  46 
Bouchet,  P.     1989.     .'\  review  ol   ijoci  ilogoin    in  gastropods. 

Journal  of  Molluscan  Studies  .55;  67-78. 
Bouchet,  P.     1990.    'I'urrid  genera  and  mode  ol  iic\flopmciit; 

the  use  and  ahu.sc  ot  protocdmli  moipliologN    Malaroloiiia 

32;  69-77. 
Bouchet,  P.  and  J.-C.  Pontes.    1981.    Migrations  verticales  des 

larves  de  Gasteropodes  ahyssaux;  arRumcnIs  nouveanx  dus 

a  I  analy.se  isolopi(|ue  de  la  cocpiillc  larvaire  et  posllarvaire. 

(>omptes  Rendus  des  Seances  de  I  Academic  des  Sciences, 

Paris  (3)  292;  1005- 1008 
Bouchet,  P.  and  .■\.  Waren.    1979    Theahvssal  mollusraii  lauii;i 


of  the  Norwegian  Sea  and  its  relation  toother  faunas.  Sarsia 
64;  211-243. 

Bouchet.  P.  and  A.  Waren  1980.  Revision  of  the  north-east 
Atlantic  batliyal  and  abyssal  Turridae  (Mollusca,  Gastrop- 
oda), Journal  of  Molluscan  Studies,  Supplement  8;  1-119. 

Bouchet,  P  and  A  Waren.  1985.  Revision  of  the  northeast 
Atlantic  bathyal  arid  abyssal  Neogastropoda,  excluding 
Turridae  (Mollusca,  Gastropoda).  Bollettino  Malacologico, 
Supplemento  1;  123-296. 

Cx)lman,  J.G.,  PA.  Tyler  and  J.D.  Gage.  1986a.  Larval  de- 
velopment of  deep-sea  gastropods  (Prosobrancliia;  Neo- 
gastropoda) from  the  Rockall  Trough  Journal  of  the  Ma- 
rine Biological  Association  of  the  Lnited  Kingdom  66;  951- 
965, 

Colnian,  J.G.,  PA.  T\ler  and  J  D  Gage  1986b  The  repro- 
ductive biology  of  Colus  jcjjreysianus  (Gastropoda;  Pros- 
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Etter,  R.J.  and  M.A.  Rex.  1990,  Population  differentiation 
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Gustahson.  R.G.,  D  T  J  Littlewood  and  R,A,  Lutz  1991,  Gas- 
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Jahlonski,  D  and  R,A,  Lutz  1983.  Larval  ecology  of  marine 
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Killingley,  J.S,  and  M.A,  Re,\,  1985,  Mode  of  larval  devel- 
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Rex,  M.A.,  C.A.  Van  Linnnersen  and  R  D  Turner  1979  Re- 
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E.  Potter  and  M.  A.  Rex,  1992  Page  75 


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THE  NAUTILUS  106(2):76,  1992 


Page  76 


Drillia  macleani,  New  Name  for  Drillia  sinuosa  McLean  and 
Poorman,  1971  (Gastropoda:  Turridae) 


John  K.  Tucker 

Department  of  Biological  Sciences 
Illinois  State  University 
Normal,  IL  61761  USA 


During  the  preparation  of  a  catalog  of  the  family  Tur- 
ridae, I  found  the  name  Drillia  sinuosa  McLean  and 
Poorman,  1971,  which  was  applied  to  a  Recent  eastern 
pacific  species,  to  be  preoccupied  by  Drillia  sinuosa  Bel- 
iardi,  1877,  which  was  applied  to  an  unrelated  species 
from  the  Middle  Miocene  of  Italy.  In  order  to  rectify 
this  homonymy  and  to  acknowledge  the  seminal  work 
on  turrids  done  by  the  senior  author  of  the  preoccupied 
taxon,  Drillia  macleani  is  here  proposed  as  a  replacement 
name  for  the  eastern  Pacific  species. 

Drillia  macleani  Tucker,  nomen  novum 

Drillia  sinuosa  McLean  and  Poorman,  1971:96,  fig.  16., 
McLean  in  Keen,  1971  :#  1622. 

Not  Drillia  sinuosa  Bellardi,  1877:126,  pl.4,  figs.  18a,b. 


Remarks:  The  holotype  is  in  the  collection  of  the  Los 
Angeles  County  Museum  of  Natural  History  (LACM 
1491).  The  type  locality  is  off  the  southernmost  coast  of 
Isla  Santa  Cruz,  Galapagos  Islands,  Ecuador  (0°47'S, 
90°21'W),  in  150  m,  collected  10  June  1968  by  Andre 
and  Jacqueline  DeRoy. 

I  thank  Dr.  James  H.   McLean  and  an  anonymous 
referee  for  reading  the  manuscript. 

LITERATURE  CITED 

Bellardi,  L.  1877.  I  molluschi  dei  terreni  Terziarii  del  Pied- 
monte  e  della  Liguria,  2.  Torino.  364  p.,  7  pis. 

Keen,  A.  M.  1971.  Sea  shells  of  tropical  west  America,  second 
edition.  Stanford  University  Press,  Stanford,  1064  p. 

McLean,  J  H  and  R  Poorman,  1971  New  species  of  tropical 
eastern  Pacific  Turridae.  The  Veliger  14(1):89-113. 


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THIS  PUBLICATION  IS  PRINTED  ON  ACID-FREE  PAPER. 


rHE  NAUTILUS 


Volume  106,  Number  3 
August  21,  1992 
ISSN  0028-1344 

A  quarterly  devoted 
to  malacology. 


.t- 


AUG  2  C 1992 


Woods  Hols,  Mass. 


EDITOR-IN-CHIEF 
Dr.  M.  G.  Harasewych 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


ASSOCIATE  EDITOR     . 
Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
P.O.  Box  2255 
Melbourne,  FL  32902 


CONSULTING  EDITORS 
Dr.  Riidiger  Bieler 
Department  of  Invertebrates 
Field  Museum  of 
Natural  History 
Chicago,  IL  60605 


Dr.  Robert  T.  Dillon,  Jr. 
Department  of  Biology 
College  of  (Charleston 
Charleston,  SC  29424 


Dr.  William  K.  Emerson 

Department  of  Living  Invertebrates 

The  American  Museum  of  Natural 

History 

New  York,  NY  10024 


Dr.  Robert  Hershler 
Division  of  Mollusks 
National  Mu.seum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


Dr.  Richard  S.  Houbrick 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


Mr.  Richard  I.  Johnson 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Department  of  Malacology 
Los  Angeles  County  Museum  of 
Natural  History 
900  E.xposition  Boulevard 
Los  Angeles,  CA  90007 

Dr.  Arthur  S.  Merrill 
%  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Ms.  Paula  M.  Mikkelsen 
Harbor  Branch  Oceanographic 
Institution,  Inc. 
Ft.  Pierce,  FL  33450 

Dr.  Donald  R.  Moore 

Division  of  Marine  Geology 

and  Geophysics 

Rosenstiel  School  of  Marine  and 

Atmospheric  Science 

University  of  Miami 

4600  Rickenbacker  Causeway 

Miami,  FL  33149 

Dr.  Gustav  Paulay 
Marine  Laboratory 
University  of  (iuam 
Mangilao,  Guam  96923 

Mr.  Richard  E.  Petit 

P.O.  Bo.x  30 

North  Myrtle  Beach,  SC  29582 

Dr.  Edward  J.  Petuch 
Department  of  Geology 
Florida  Atlantic  University 
Boca  Raton,  FL  33431 


Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Geerat  J.  Vermeij 
Department  of  Geology 
University  of  California  at  Davis 
Davis,  CA  95616 


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THEt^NAUTILUS 


CONTENTS 


Volume  106,  Number  3 

August  21,  1992 

ISSN  0028-1344 


'larine   Rioi 


IBRAR 


James  F.  Quinn,  Jr. 


New  species  of  Calliostoma  Swainson,  1840  (Gastropoda: 

Trochidae),  and  notes  on  some  poorK  known  species  from 

the  western  Atlantic  Ocean  ^  '  ;?;'5;    Hol 


r.i,n.;.. .  71 


James  H.  McLean 


A  new  species  of  Pseudorimula  {Fissurellacea: 
Clypeosectidae)  from  hydrothermal  vents  of  the  Mid- 
Atlantic  Ridge 


115 


Jov  C.  Goodsell 

Prodissoconch  I  and  II  length  in  Mercenaria  Taxa    

119 

Arnold  G.  Eversole 

Harald  A.  Rehder 

Harpa  cabriti  Fischer.  1860,  a  replacement  name  for 

Harpa  ventricosa  Lamarck,  1816  (Gastropoda:  Harpidae)     

123 

James  H.  McLean 

Obituary:  Rae  Baxter,  1929-91 

125 

THE  NAUTILUS  106(3):77-114,  1992 


Page  77 


New  Species  of  CaUiostoma  Swainson,  1840 
(Gastropoda:  Trochidae),  and  Notes  on  Some 
Poorly  Known  Species  from  the 
Western  Atlantic  Ocean 


James  F.  Quinn,  Jr. 

Florida  Marine  Research  Institute 
Department  of  Natural  Resources 
100  Eighth  Avenue,  S.  E. 
St   Petersburg,  FL  33701-5095,  USA 


ABSTRACT 

Twent)  -seven  new  species  of  CaUiostoma  Swainson  {sensii  lata) 
are  described  from  the  western  Atlantic  Ocean.  Trochus  (Zi- 
zyphiniis) stirophorus  Watson  is  confirmed  to  be  a  CaUiostoma. 
with  C.  arestum  Dall  as  a  synonym,  and  Trochus  (Margarita) 
dnopherus  Watson  is  transferred  to  CaUiostoma  New  records 
and  observations  are  presented  for  C.  apicitium  Dall,  C.  in- 
diana  Dall,  C.  orion  Dall,  C.  aurora  Dall,  C  jernandezi  Princz, 
C,  atlantis  Clench  &  Aguayo,  C,  axelolssoni  Quinn  (new  name 
for  C.  olssoni  Bayer),  and  C.  adspersun^  (Philippi).  External 
morphologies  of  the  head-foot  of  13  species  are  briefly  de- 
scribed, and  illustrations  of  the  radulae  of  13  species  are  pre- 
sented, along  with  pertinent  remarks. 

Key  tvurds:  Trochidae,  Calliostomatinae,  CaUiostoma.  systeni- 
atics,  new  species,  new  records,  radulae 


INTRODUCTION 

The  subfamily  Calliostomatinae  is  by  far  the  most  spe- 
ciose  trochid  subfamily  in  the  western  Atlantic  Ocean. 
Species  occur  from  the  subarctic  waters  of  Canada  to  the 
subantarctic  waters  of  southern  Argentina  and  the  Ant- 
arctic waters  of  South  Georgia  Island  (Powell,  1951; 
Clench  &  Turner,  1960),  Calliostomatines  inhabit  a  great 
variety  of  habitats,  both  biotic  and  abiotic,  in  depths 
ranging  from  the  shallow  subtidal  to  se\eral  hundred 
meters  (Clench  &  Turner,  1960;  Reed  &  Mikkelsen,  1987; 
Quinn,  in  press). 

The  greatest  number  of  species  of  Calliostomatinae  is 
assigned  to  the  rather  ill-defined  genus  CaUiostoma 
Swainson,  1840.  Clench  and  Turner  (1960)  monographed 
the  western  Atlantic  species  of  the  genus  and  concluded 
that  41  names  represented  valid  species  (including  5  new 
species),  24  names  were  synonyms,  and  8  names  were 
not  referrable  to  the  genus.  Subsequent  studies  (Merrill, 


1970;  Bayer,  1971;  Princz,  1978;  Quinn,  1979,  in  press; 
Rios,  1985,  1990)  have  revealed  one  new  synonym  and 
two  new  species,  reinstated  two  species  excluded  by 
Clench  and  Turner,  and  resurrected  three  species  from 
synonymy.  Another  15  nominal  species  and  subspecies 
not  treated  by  Clench  and  Turner  (1960)  have  been 
described  from  off  southern  Argentina  (Philippi,  1845 
[in  1842-1851];  Martens,  1881;  Smith.  1881,  1915;  Roch- 
ebrune  &  Mabille,  1889;  Strebel,  1905,  1908);  a  few  of 
these  species  were  discussed  or  mentioned  by  Powell 
(1951),  who  later  (1960)  published  a  complete  list,  but 
most  of  these  species  remain  essentially  unknown.  In- 
cluding the  27  new  species  described  in  this  paper,  the 
2  here  reassigned  to  the  genus,  and  at  least  7  that  are 
currently  unnamed,  almost  100  valid  Recent  species  of 
CaUiostoma  are  now  known  from  the  western  Atlantic 
(Table  1),  In  contrast,  only  36  species  of  CaUiostoma  are 
known  to  occur  in  the  eastern  Pacific  (Table  2)  (see  Dall, 
1909;  McLean,  1970,  1971,  1984;  Rehder,  1971;  McLean 
&  Andrade,  1982).  The  number  of  valid  Recent  species 
of  CaUiostoma  now  known  in  the  Americas  is  somewhat 
greater  than  the  number  of  species  described  from  the 
Neogene  formations  of  the  Americas. 

At  least  six  other  calliostomatine  species  occur  only  off 
Argentina  and  in  the  Falkland  and  South  Georgia  Islands 
(Powell,  1951,  1960).  Powell  assigned  two  species  each 
to  Falsimargarita  Powell,  1951,  and  Photinastoma  Pow- 
ell, 1951,  and  one  species  each  to  Venustatrochus  Powell, 
1951,  and  Photiniila  Adams  and  Adams,  1854.  Clench 
and  Turner  (1960)  assigned  CaUiostoma  blakei  Clench 
and  Aguayo,  1938,  to  Photitnila,  but  characters  of  the 
animal  and  radula  show  that  the  species  should  be  as- 
signed to  the  Lirulariinae  Hickman  and  McLean,  1990 
(Quinn,  in  preparation). 

During  the  preparation  of  monographs  of  the  Tro- 
chidae of  the  Gulf  of  Me.xico  (Quinn,  in  press)  and  the 


Page  78 


THE  NAUTILUS,  Vol.  106,  No.  3 


adjoining  Straits  of  Florida  (Quinn,  1979),  I  examined 
specimens  of  many  calliostomatine  species  collected  from 
throughout  the  Caribbean  Sea  and  southwestern  Atlantic 
Ocean,  most  obtained  subsequent  to  (blench  and  Turners 
1960  monograph.  Many  of  the  species  represented  in 
that  material  were  clearly  undescribed,  some  were  found 
to  be  species  complexes,  and  others  represented  signifi- 
cant range  extensions  of  pre\iously  described  but  poorly 
known  species.  The  purposes  of  this  paper  are  to  provide 
descriptions  of  the  new  species,  to  clarify  the  identities 
of  some  species,  and  to  report  new  distributional  data 
for  species  of  Calliostoma  in  the  western  Atlantic  Ocean. 
The  external  morphologies  of  the  head-foot  of  13  of  the 
species  are  briefly  described,  and  radulae  of  13  species 
are  illustrated.  However,  because  so  few  species  of  west- 
ern .\tlantic  Calliostoma  have  had  their  anatomy  and 
radula  illustrated,  and  because  character  states  are  so 
poorly  understood  for  species  worldwide,  speculations  on 
relationships  based  on  the  present  observations  are  pre- 
mature. Of  particular  interest  would  be  optical  micro- 
scope observations  to  clarif\  characteristics  of  these  rad- 
ulae as  espoused  by  Hickman  (1977). 

Numerous  genus-level  taxa  have  been  proposed  for 
perceived  species  groups  within  Calliostomatinae.  Al- 
though some  authors  incorporated  features  of  the  exter- 
nal anatomy  and  radula  (e.g.,  Powell,  1951;  Clench  & 
Turner,  1960),  most  of  these  names  were  based  on  shell 
characters  alone,  and  some  names  seem  to  reflect  a  re- 
gional bias  of  the  authors;  however,  the  morphological 
limits  of  these  nominal  taxa  are  nebulous,  and  a  world- 
wide survey  of  genus-level  taxa  is  needed.  Therefore,  the 
species  discussed  in  the  present  paper  are  not  formally 
assigned  to  nominal  subgenera,  but  the  following  species 
groups  are  recognized  as  relatively  distinct:  the  C.  ptil- 
chrum  group  [Calliostoma  s.s.,  or  Eucasta  Dall,  1889a); 
the  C.  jujulnnum  group  {Elmerlinia  Clench  &  Turner, 
1960);  the  C.  bairdii  group  (Kombologion  Clench  &  Tur- 
ner, 1960);  the  C.  sayantim  group;  the  C.  circumcinctum 
group  (?=Otuhaia  Ikebe,  1942);  the  C.  atlantis  group. 
Many  western  Atlantic  species  cannot  be  satisfactorily 
assigned  to  any  species  group,  but  a  forthcoming  revision 
of  the  southwestern  Pacific  calliostomatine  species  (B.  A. 
Marshall,  in  litt.)  may  help  clarify  some  of  these  uncer- 
tainties. 

For  brevity,  authors  and  dates  of  publication  for  spe- 
cies included  in  Tables  1  and  2  are  not  repeated  in  the 
Remarks  section  of  the  species  accounts.  Species  not  in- 
cluded in  those  tables  are  cited  with  their  authors  and 
dates  whenever  mentioned  in  the  text.  Bibliographic  ref- 
erences of  species  included  in  Tables  1  and  2  are  not 
included  in  the  Literature  Cited  section  unless  the  species 
are  also  included  in  the  text;  most  of  these  citations  are 
readily  available  in  Clench  and  Turner  (1960),  Keen 
(1971),  and  Quinn  (in  press). 

Acronyms  for  specimen  repositories  are  as  follows: 
AMNH  (American  Museum  of  Natural  History,  New 
York);  ANSP  (Academy  of  Natural  Sciences  of  Phila- 
delphia); BM(NH)  (The  Natural  History  Museum,  Lon- 
don); DMNH  (Delaware  Museum  of  Natural  History, 


Wilmington);  FSBC  I  (Invertebrate  Collection,  Florida 
Marine  Research  Institute,  St.  Petersburg);  HMNS  (Hous- 
ton Museum  of  Natural  History,  Texas);  LACM  (Natural 
History  Museum  of  Los  Angeles  County,  California); 
MCZ  (Museum  of  Comparative  Zoology,  Harvard  Uni- 
versity, Cambridge,  Massachusetts);  MORG  (Museu 
Oceanographico  da  Fundafao  L'niversidade  do  Rio 
Grande,  Rio  Grande,  Rio  Grande  do  Sul,  Brazil);  UF 
(Florida  Museum  of  Natural  History,  University  of  Flor- 
ida, Gainesville);  UMML  (Rosenstiel  School  of  Marine 
and  Atmospheric  Science,  University  of  Miami,  Florida); 
USNM  (National  Museum  of  Natural  History,  Smithson- 
ian Institution,  Washington,  DC). 


SYSTEMATICS 

Superfamily  Trochoidea  Rafinesque,  1815 

Family  Trochidae  Rafinesque,  1815 

Subfamily  Calliostomatinae  Thiele,  1924 

Genus  Calliostoma  Swainson,  1840 

Type  species   (subsequent   designation;    Herrmannsen, 

1846:154):  Trochus  conulns  Linnaeus,  1758. 

Calliostoma  hilare  new  species 
(figures  1,  2) 

Description:  Shell  medium-sized,  attaining  height  of 
14.5  mm,  width  of  11.5  mm.  conical,  nonumbilicate, 
finely  sculptured.  Protoconch  375  nm  maximum  diam- 
eter, 1  whorl.  Teleoconch  whorls  8.6,  flat-sided;  first  w  horl 
with  weak  axial  riblets  and  2  strong  spiral  cords;  axial 
riblets  disappearing  on  second  whorl;  spiral  sculpture 
increasing  by  intercalation  to  7  cords  and  4  threads; 
supraperipheral  spiral  cords  finely  beaded,  beads  crowd- 
ed, rounded;  periphery  of  first  4  whorls  sharply  carinate, 
composed  of  abapical  spiral  cord  set  w  ith  sharpK  conical 
beads,  periphery  of  subsequent  w  horls  narrowK  round- 
ed, composed  of  two  adpressed  spiral  cords  set  with 
rounded  beads.  Base  weakly  convex,  with  15  narrow, 
finely  beaded  spiral  cords,  cords  becoming  broader  and 
more  coarseK  beaded  adaxialK .  Aperture  subquadrate. 
lips  thin;  columella  almost  straight,  thickened,  slightly 
twisted,  terminating  in  weak,  rounded  denticle.  Shell 
ground  color  golden  yellow  with  greenish  iridescence, 
with  light  golden  brown  and  white  spots  alternating  along 
periphery;  w  hite  spots  spiralK  elongate,  brow  n  spots  ax- 
ialK  elongate. 

Holotype:  USNM  860249,  height  14.5  mm.  width  11.5 
tnm. 

Type  locality:  Off  northwestern  tip  of  Little  Bahama 
Bank,  John  Elliott  Pillsbury  Station  P-198,  27°30'N, 
79°10'W,  in  242  m. 

Distribution:  This  species  is  known  onl\  from  the  ho- 
lotype, an  empt\-  shell,  collected  in  242  m  off  the  north- 
western tip  of  the  Little  Bahama  Bank. 

Remarks:    The  holotype  of  C  hilare  is  most  similar  to 


J.  F.  Quinn,  Jr.,  1992 


Page  79 


Table  1.    Cieograpliif  anil  hatliynietric  distributions  of  calliostoinatine  species  in  the  western  Atlantic  Ocean. 

Species 

C'alliostoma  Swainson,  1840 
jujulnnuui  (CInielin,  1791) 
javanicnm  (Lamarck,  1822) 
occidentale  (Mighels  &  A.  Adams,  1842) 
gemmosum  (Reeve,  1842) 
nudum  nudum  (Philippi,  184.5) 
tampaense  (C^onrad,  1846) 
jucunduni  (Gould,  1849) 
pulchriim  (C   B   Adams.  18.50) 
adspcrsum  (Pliilippi,  1851) 
euglyptuni  (A,  Adams,  18.55) 
dnophcrum  (Watson,  1879) 
stirophorum  (Watson,  1879) 
coppingeri  Smith,  1880 
bairdii  Verriil  &  Smith,  1880 
apicinum  Dall,  1881 
circumcinctum  Dall,  1881 
echinatum  Dall,  1881 
orion  Dall,  1881 
roscoluni  Dall.  1881 
sapidum  Dall,  1.881 
ijucatccanum  Dall,  1881 
consimilis  (Smith,  1881) 
aurora  Dall,  1888 
benedicti  Dall,  1889 
briinneum  (Dall,  1889) 
cinctellum  Dall,  1889 
Indiana  Dall,  1889 
psyche  Dall,  1889 
sayanum  Dall,  1889 
dozei  Rochebrime  &  Mabille,  1889 
nudum  roseotinclum  Rochebrune  &  Mabille,  1889 
optimum  Rochebrune  &  Mabille,  1889 
senilis  Rochebrune  &  Mabille,  1889 
irisans  Strebel,  1905 
kophamcli  Strebel,  1905 
mocbiusi  Strebel,  1905 
nudum  flaviducarnea  Strebel,  1905 
marionae  Dall,  190(5 
militare  Ihering,  1907 
andersoni  Strebel,  1908 
falklandicum  Strebel,  1908 
modestulum  Strebel,  1908 
nordenskjoldi  Strebel,  1908 
venustulum  Strebel,  1908 
depiclum  Dall,  1927 
hahbrectum  Dall,  1927 
hendersoni  Dall,  1927 
sarcodum  Dall,  1927 
amazonicum  Finlay,  19.30 
bigelowi  Clench  &  Aguayo,  1938 
schroederi  Clench  &  Aguayo,  1938 
hassler  Clench  &  Aguayo,  19.39 
atlantis  Clench  &  Aguayo,  1940 
carcellesi  Clench  &  .Aguayo,  1940 
cubanum  Clench  &  Aguayo,  1940 
torrei  Clench  &  Aguayo,  1940 
fascinans  Schwengel  &  McGinty,  1942 
barbouri  Clench  &  Aguayo,  1946 
adelae  Schwengel,  1951 
bullisi  Clench  &  Turner,  1960 


)eptli  (m) 

Distribution 

0-10 

4 

0-40 

4 

.30-1,800 

1,2- 

10-100 

6 

? 

7 

0-10 

3- 

20-30 

7 

0-70 

3,  4,  5,  ?6 

0-40 

5,6 

10-30 

3 

640 

6* 

700 

3-,4- 

10-80 

6,7 

70-450 

2 

100-200 

4- 

200-300 

4- 

70-1.50 

4 

20-60 

4 

40-80 

3 

100-250 

?3-,  4 

1.5-60 

3 

? 

7 

180-1,0.50 

4 

365 

3-* 

20-70 

4 

?-300 

3-,  4* 

.50-100 

4 

100-400 

3- 

150-300 

3- 

? 

7 

? 

7 

? 

7 

? 

7 

? 

7 

? 

7 

90 

7 

? 

7 

40-120 

3 

30-65 

6,7 

? 

7 

1.5-2.50 

7 

65-350 

7 

? 

7 

40 

7 

1-7 

6 

535 

3* 

150-3.50 

3- 

1-10 

4 

■? 

7 

,375-4.30 

4* 

250-450 

4- 

65 

6* 

?-600 

4- 

50 

7* 

900 

4* 

700 

4* 

70-120 

3 

25-100 

3-,  4 

0-2 

3- 

70 

5* 

Page  80 


THE  NAUTILUS,  Vol.  106,  No.  3 


Table  1.    ('ontiiiiied. 


Species 


Depth  (m) 


Distributii 


jeanneae  Clench  &  Turner,  1960 
Oregon  Cleiieh  &  Turner,  1960 
roseicateri  ( blench  &  Turner,  1960 
springvri  Clench  &  Turner,  1960 
fcrnandezi  Princz,  1978 
alternttm  Quinn,  1992 
argcnttim  Quinn,  1992 
allantoides  Quinn,  1992 
(lulinim  Quinn,  1992 
axclolssoni  Quinn,  1992 
hertmulcnsc  Quinn,  1992 
hruniicophttan  Quinn,  1992 
ciiithjphiltini  Quinn,  1992 
coronatnm  Quinn,  1992 
cuhense  Quinn,  1992 
defnle  Quinn,  1992 
dentatum  Quinn,  1992 
fucosum  Quinn,  1992 
guesti  Quinn,  1992 
hihirc  Quinn,  1992 
hirttini  Quinn,  1992 
moscatcUii  Quinn,  1992 
ptirpureurn  Quinn,  1992 
rota  Quinn,  1992 
rude  Quinn,  1992 
rugosum  Quinn,  1992 
scalenum  Quinn,  1992 
scurru  Quiim,  1992 
senii.sudvr  Quinn,  1992 
serratnhim  Quinn,  1992 
tenebrosum  Quinn,  1992 
vinosum  Quinn,  1992 
oiscardii  Quinn,  1992 

Venitstatrochus  Powell,  1951 

georgianus  Powell,  1951 
Falsimargarita  Powell,  1951 

gemma  (Smith,  1915) 

iris  (Smith,  1915) 

Photinula  Adams  &  Adams,  185-1 
coeridescens  (Kiufi,  &  Broderip,  1831) 

Photinaatoma  Powell,  1951 
tacniata  taeniata  (Wood,  1828) 
taeniata  nivea  (Cooper  &  Preston,  1910) 
'^gamma  (Rorhehrune  &  Mahiile.  1889) 


200-350 
274-641 
200-450 

4.5-145 

5.5-146 
350-450 
400-600 

10-50 
200-300 

82 

50-80 

150-350 

770-800 

0-20 

? 

15-55 
5-9 
183-219 
250 
520 
50-80 
0-60 
27 

60-90 
? 

25-80 

20-90 

70-155 

120-190 

? 

18-22 
40-45 

120-200 

200-400 
225-450 

0-200 


0-80 
0-100 

? 


4* 

3- 

4- 

3- 

5 

5 

4 

4* 

5 

4-,6- 

3* 

6* 

3 

6* 

4- 

4* 

3- 

5* 

4* 

4* 

4* 

6 

5- 

6* 

5- 

4* 

3 

5 

5- 

5- 

6* 

6 

6* 


*  Known  only  from  the  type  lot  or  locality;  1  =  northern  boreal;  2  =  "Virginian"  (Cape  Cod  to  Cape  Hatteras);  3  =  Carolinian; 
4  =  Caribbean;  5  =  northern  ,South  America;  6  =  Brazil-northern  Argentina;  7  =  Patagonian;  8  =  Subantarctic;  '—  '  indicates 
restricted  range  within  region. 


shells  of  C.  rude  in  shape,  size,  and  sculpture,  but  it 
differs  by  having  a  slightly  larger  protoconch  (375  ^m 
Ds.  350  nm);  by  having  the  ()eriplierul  spiral  cord  strong 
and  sharply  beaded  on  whorls  2-4,  becoming  weaker 
and  with  rounded  beads  on  later  whorls,  whereas  the 
peripheral  cord  of  C.  rude  is  inconspicuous  on  the  first 
three  whorls,  becoming  very  strong  and  coarsciv  beaded 
on  subsequent  whorls;  and  by  having  more  numerous 


but  weaker  spiral  cords.  Calliostoma  hilarc  is  also  similar 
to  C.  serratulum,  but  the  shell  of  the  former  has  a  smaller 
protoconch  (375  nm  vs.  400-425  ^m),  is  proportioiiateK- 
narrower,  has  more  numerous  spiral  cords  that  are  more 
weakly  beaded,  and  has  a  periphery  that  is  composed  of 
two  subequal,  finely  beaded,  adpressed  cords  rather  than 
a  strong,  coarsely  beaded  upper  cord  and  a  weaker  lower 
cord. 


J.  F.  Quinn,  Jr.,  1992 


Page  81 


Table  2.    (ieograpliic  ami  balln  metric  distrihiitions  of  calliostomatine  species  in  the  eastern  Pacific  Ocean. 


Species 


Depth  (m) 


Distrilmtiiin 


Calliostoma  Swainson,  1840 
anniildtum  (Liglitfoot.  ITcSfi) 
caridlUulatum  (Lightfoot,  1786) 
anlonii  (Koch,  1843) 
exiniium  (Reeve,  1843) 
ligatum  (Gould.  1849) 
leatmm  (C.  B.  Adams,  1852) 
fonkii  (Philippi,  1860) 
gemrnulalum  Carpenter,  1864 
splcndcns  Carpenter,  1864 
supragranosum  Carpenter,  1864 
varirgatum  C^arpenter,  1864 
aequisculptnm  Carpenter,  1865 
triculor  Gahh,  1865 
gloriosum  Dall,  1871 
palmeri  Dall,  1871 
platinum  Dall,  1890 
tiirhiuum  Dall,  1895 
iridium  Dall,  1896 
nephcloidc  Dall,  1913 
honita  Strong,  Hanna.  &  Hertlein,  1933 
rema  Strong,  Hanna,  &  Flertlein.  19.33 
marshalli  Lowe,  1935 
mcleani  Shasky  &  Campbell,  1964 
gordanum  McLean,  1970 
jacquelinae  McLean,  1970 
keenae  McLean,  1970 
sanjaimense  McLean,  1970 
santacruzanum  McLean,  1970 
veleruae  McLean,  1970 
insignis  Olsson,  1971 
joanneae  Olsson,  1971 
pillsburyae  Olsson,  1971 
chilena  Rehder.  1971 
delli  McLean  &  Andrade,  1982 
bernardi  McLean,  1984 
titanium  McLean,  1984 


"offshore 

25-750 

0-3 

0-40 

0-3 

0-3 

?-457 

0-3 

"offshore  ' 

0-3 

25-750 

1-3 
15-60 
0-40 
0-45 
80-750 
60-120 
230-280 
50-125 
35-75 
20-45 
0 
.5-10 
128 
146 

.5.5-110 
137 
45 

60-100 
50-80 
53 

57-64 
200-7.50 
200-4.50 
12.5-175 
200-300 


offshore' 


1 

1 

3 

3 

1 

3 

3,4 

2 

2 

2 

1 

3 

2,3 

2 

3 

1 

2 

3 

3 

3 

3 

3 

3 

2* 

3* 

2,3 

2* 

3* 

3 

3 

3* 

3* 

3,4 

4- 


*  Known  link   from  the  t\pe  lot  or  locality; 
restricted  range  within  region. 


rthern  boreal;  2  =  Californian;  3  =  Panamic;  4  =  Peruvian; 


indicates 


Calliostoma  serratulum  new  species 
(figures  3-6) 

Calliostoma  sp.  4:  Bayer  et  al.  1970;A29,  A137,  A146. 

Description:  Shell  medium-sized,  attaining  height  of 
16.7  iTim,  width  of  16,4  mm,  conical,  nonumbilicate, 
coarsely  sculptured.  Protoconch  400-42.5  p.m  ma.ximum 
diameter,  1  whorl.  Teleoconch  whorls  about  7,  flat;  first 
2  whorls  with  low,  rounded  axial  riblets  and  2  strong, 
beaded  spiral  cords;  a.\ial  riblets  gradualK  disappearing 
on  whorls  3-5;  spiral  sculpture  increasing  by  interca- 
lation to  5-8  cords  and  0-5  threads,  of  which  abapical 
2  cords  forming  periphery;  periphery  sharply  carinate; 
adapical  peripheral  cord  strong,  projecting,  set  with 
strong,  conical  beads,  beads  crossed  by  2  fine  spiral  threads 
producing  squarish  apices;  abapical  peripheral  cord 
weaker,  weakly  beaded.  Base  weakly  convex,  with  13- 
17  narrow,  weakly  beaded  spiral  cords,  adaxial  2-3  cords 


stronger,  more  widely  spaced,  and  more  distinctly  bead- 
ed. Aperture  subquadrate,  lips  thin,  weakly  crenulate; 
columella  concave  above,  straight  below ,  thickened,  ter- 
minating in  weak,  rounded  denticle.  Shell  ground  color 
golden  yellow  with  spots  and  axial  streaks  of  brown; 
beads  white;  basal  spiral  cords  with  elongate  spots  of 
brown,  darkest  on  adaxial  2-3  cords. 

Holotype:  USNM  860250,  height  12.5-F  mm,  width 
12.3  mm. 

Type  locality:  Southw  est  of  Cartagena,  Colombia,  John 
Elliott  Pillsbiry  Station  P-375,  9°59.0'N,  76°02.0'W, 
in  135-130  m. 

Paratypes:  1,  UMML  30.3539;  off  Santa  Marta,  Colom- 
bia, John  Elliott  Pillsbiry  Station  P-399,  9°01.3'N, 
76°40.2'W,  119-179  m;  17  Julv  1966;  10-ft  otter  trawl— 
1,  UMML  30.6935;  1,  FSBC'l  44069;  off  Peninsula  de 


Page  82 


THE  NAUTILUS.  Vol.  106,  No.  3 


.A 


/ 


ji 


"\ 


S, 


V 


/ 


■•'1 


v. 


■k_ 


^' 


8 


N 


10 


Figures  1,  2.  Calliostotna  hilare  new  species,  liolotype,  I'SNM  860249;  height  14  3  mm,  width  1 1.5  mm  Figures  3-6.  Calliostoma 
serratuhim  new  species,  3,  i.  Holotype,  USNM  860250;  height  12.5+  mm,  width  12, .i  mm,  5.  6.  Parat\  pe,  L'MML  30,35.39;  height 
10.1  mm,  width  100  mm.  Figures  7,  8.  Calliostoma  rude  new  species,  ho]()t>pe,  L'SNM  860251;  height  13.3  mm,  width  10  6  mm. 
Figures  9.  10.  Calliostoma  sp,  cf.  rude,  specimen  Irom  off  Barbados,  Sunderland  collection;  height  9,7  mm,  width  7.3  mm. 


Paraguana,  Venezuela,  John  Elliott  Pillsbi  ry  Station 
P-757,  11°39.6'N,  69°22.1'W,  161-187  m;  27  July  1968; 
10-ft  otter  trawl. 

Other  material:  4  broken,  UMML  30.8366;  off  Santa 
Marta,  Colombia,  John  Elliott  Pillsbury  Station 
P-785,  11°16.9'N,  74°17.0'\V,  176-165  m;  31  July  1968; 
10-ft  otter  trawl. 


Distribution:  Calliostoma  serratuhim  is  known  only 
from  empty  shells  traw  let!  from  off  the  nortiieastern  coast 
of  Colombia  and  northwestern  coast  of  Venezuela  in  1 19- 
187  m. 

Remarks:  Shells  of  C.  serratuhim  are  most  similar  to 
those  of  C.  rude  but  differ  by  having  a  much  larger 
protoconch  (400-425  nm  vs.  350  fim),  by  being  propor- 


J.  F.  Quinn,  Jr.,  1992 


Page  83 


tionately  wider,  and  b\  having  the  adapical  peripheral 
cord  set  with  uniformly  sized  beads  that  are  squarish  in 
profile. 

Calliostoma  rude  new  species 
(figures  7,  8) 

Description:  Shell  medium-sized,  attaining  height  of 
13.3  mm,  width  of  10.6  mm,  conical,  nonumbilicate, 
coarsely  sculptured.  Protoconch  350  ixm  maximum  di- 
ameter, 1  whorl.  Teleoconch  whorls  8.3,  flat;  first  w  horl 
with  low,  rounded  axial  riblets  and  2  strong,  beaded  spiral 
cords;  axial  sculpture  absent  after  first  w  horl;  spiral  sculp- 
ture increasing  by  intercalation  to  5-6  cords  and  1-2 
threads,  of  which  abapical  2  cords  forming  periphery; 
periphery  sharpK  carinate;  adapical  peripheral  cord 
strong,  strongly  beaded,  beads  triangular,  every  third  or 
fourth  bead  markedK-  larger;  abapical  peripheral  cord 
weaker,  strongK  undulate.  Base  weakly  convex,  with  11- 
13  strong,  narrow,  fineK  beaded  spiral  cords  .-Aperture 
subcjuadrate,  lips  thin,  fineK  crenulate;  columella  almost 
straight,  thickened,  terminating  in  weak,  rounded  den- 
ticle. Shell  ground  color  ivor\  w  ith  greenish  and  reddish 
iridescence;  streaks  and  flammules  of  golden  brown  pres- 
ent above  periphery;  periphery  with  spots  of  darker 
brow  n;  base  with  light  goklen-brown  spots  arranged  on 
cords  to  form  obscure,  crescentic  radial  streaks. 

Holoiype:  USNM  860251,  height  13.3  mm,  width  10.6 
mm 

Type  locality:  Off  Cayenne,  French  Guiana,  John  El- 
liott PiLLSBURY  Station  P-650,  6°07.0'N,  52°19.0'W,  in 
84-91  m. 

Paratypes:  2,  UMML  30.5844;  1,  FSBC  I  44076;  from 
same  lot  as  holotype. — 2,  MCZ  302593;  off  Georgetow  n, 
Guyana,  Chain  Cruise  35,  Station  35  +  36,  8°10.5'- 
8°lb.0'N,  57°48'W,  53-60  m;  28  .\pril  1963. 

Distribution:  Calliostoma  rude  is  known  only  from  two 
collections,  one  off  Georgetown,  Guyana,  in  53-60  m, 
and  the  other  from  the  type  localit}  off  Cayenne,  French 
Guiana,  in  84-91  m;  the  latter  collection  contained  a 
single  live-collected  specimen.  The  field  notes  on  the  type 
localit)  cite  shell  rubble,  and  the  associated  invertebrates 
suggest  that  hard-bottom  outcrops  are  nearby. 

Remarks:  Comparisons  of  shells  of  Calliostoma  rude 
with  those  of  the  similar  C.  serratuhim  and  C.  hilare 
are  discussed  in  the  remarks  for  the  latter  two  species. 
The  adapical  peripheral  cord  of  shells  of  C  rude  is  sub- 
equal  to  the  other  spiral  cords  on  the  first  three  whorls, 
but  it  becomes  markedly  stronger  on  the  fourth  and 
subsequent  whorls.  This  characteristic  separates  C.  rude 
from  all  other  peripherally  carinate  species  in  which 
peripheral  carination  is  present  on  the  first  w  horls  and 
either  remains  strong  on  all  whorls  (e.g.,  C.  cincteltum) 
or  disappears  on  the  last  one  or  two  whorls  (e.g.,  C. 
echinatum).  The  alternation  of  two  or  three  small  beads 
with  a  much  larger  triangular  bead  is  also  unique  to  C. 
rude.  Four  specimens  collected  from  off  the  western  coast 


of  Barbados  (Figures  9,  10;  Sunderland  collection,  ex  F 
Sander)  are  very  similar  to  the  types  of  C,  rude  but  are 
relativeK  smaller  and  narrower  and  have  some  subtle 
sculptural  differences  that  suggest  that  the  Barbados  shells 
may  be  a  separate  species. 

Calliostoma  cnidophilum  new  species 
(figures  11-14,  107) 

Description:  Shell  medium-sized,  attaining  height  of 
14.1  nmi,  width  of  12.1  mm,  conical,  nonumbilicate, 
finely  sculptured.  Protoconch  375  /um  maximum  diam- 
eter, 1  whorl.  Teleoconch  whorls  about  8,  flat;  first  whorl 
with  low,  rounded  axial  riblets  and  2  strong,  beaded  spiral 
cords;  axial  riblets  disappearing  on  second  whorl;  spiral 
sculpture  increasing  b\  intercalation  to  10-12  cords  and 
0-2  threads,  cords  fineK  beaded;  peripher\  of  first  2-2.5 
w  horls  sharpK  carinate,  set  with  sharply  conical  beads; 
periphery  becoming  narrowly  rounded  on  subsequent 
whorls,  beads  becoming  finer,  rounded;  peripheral  cord 
splitting  into  2  subequal,  adpressed  cords  on  fifth  w  horl; 
periphery  obscureK  multiangulate  when  viewed  from 
base.  Base  weakly  convex,  with  14-20  narrow,  sharp, 
very  finely  beaded  spiral  cords.  Aperture  subquadrate, 
lips  thin,  fineK'  crenulate;  columella  short,  weakly  con- 
cave, thickened.  Shell  ground  color  golden  yellow  with 
greenish  iridescence;  periphery  with  regularly  spaced, 
spirally  elongate  pinkish  spots.  Animal  (in  alcohol)  white; 
cephalic  tentacles  long,  slender,  gradualK  tapering,  right 
longer  than  left,  ocular  peduncles  long,  slender,  with 
large,  black  eye  at  tips;  epipodium  with  4  pairs  of  ten- 
tacles decreasing  in  size  posteriorly,  neck  lobes  well-de- 
veloped, thin,  semicircular,  smooth. 

Holotype:  USNM  860265,  height  13.7  mm,  width  11.4 
mm. 

Type  locality:  Off  Ilets-a-Goyaves,  Guadeloupe,  Lesser 
.\ntilles,  Johnson-Sea-Link  II  Station  JSL-lI-1845, 
16°10'00"N,  61°49'08"W,  in  306  m. 

Paratypes:  2,  UMML  30.8370;  1,  FSBC  1  44070;  off 
Dominica.,  John  Elliott  Pillsbury  Station  P-931, 
15°31.2'N,  6ri2.3'W,  174-357  m;  15  July  1969;  5-ft 
Blake  trawl. 

Other  material:  1,  Sunderland  collection;  off  St.  James, 
Barbados,  152  m;  ex  F.  Sander. 

Distribution:  Calliostoma  cnidophilum  is  known  from 
the  Lesser  Antilles  in  depths  of  152-357  m;  living  spec- 
imens were  collected  in  306  m  and  174-357  m. 

Remarks:  Shells  of  Calliostoma  cnidophdum  are  most 
similar  to  the  holotype  of  C.  hdare  but  differ  by  being 
proportionately  broader;  by  having  an  obscurely  mul- 
tiangulate peripheral  profile  when  viewed  from  the  base; 
by  having  finer,  more  numerous  spiral  cords  above  the 
periphery;  by  having  the  spiral  cords  more  finely  beaded; 
and  by  having  a  shorter,  thicker  columella.  The  radula 
(formula  00.5.1. 5. oo;  figure  107)  differs  from  that  illus- 
trated by  Clench  and  Turner  for  C.  pulchrum  (1960:pl. 


Page  84 


THE  NAUTILUS,  Vol.  106,  No.  3 


16 


19 


J.  F.  Quinn,  Jr..  1992 


Page  85 


3,  fig.  3)  by  having  the  rachidian  teeth  with  much  broader 
bases  and  cusps  and  b\'  having  narrower  cusps  on  the 
lateral  teeth. 

Observations  made  from  the  submersible 
Johnson-Se.\-Link  II  indicate  that  C.  cnidophiliim  may 
feed  on  octocorals  (M.  G.  Harasewych.  personal  com- 
munication; see  also  Harasewych  ct  al..  1992.  fig.  12). 

Calliostoma  semisuave  new  species 
(figures  15.  16) 

Description:  Shell  medium-sized,  attaining  height  of 
13.4  mm.  width  of  11.4  mm.  conical,  nonumbilicate. 
finely  sculptured.  Protoconch  400  ^"1  maximum  diam- 
eter. 1  whorl.  Teleoconch  whorls  8.5.  flat;  first  4  whorls 
with  low,  rounded  axial  riblets  and  2-4  strong  beaded 
spiral  cords;  axial  sculpture  disappearing  on  fifth  whorl; 
spiral  sculpture  increasing  b\  intercalation  to  11  cords 
and  2-3  threads,  cords  strongly  beaded  on  first  5  whorls, 
beads  gradually  weakening  on  subsequent  whorls;  pe- 
riphery sharply  carinate.  with  single  strong,  smooth,  pe- 
ripheral cord.  Base  weakly  con\ex,  with  16-18  narrow, 
smooth  to  ver\  weaklv  beaded  spiral  cords.  Aperture 
subquadrate,  lips  thin;  columella  almost  straight,  thick- 
ened, terminating  in  sharp  denticle.  Shell  ground  color 
cream  to  light  golden  \  ellow ,  with  diffuse  spots  and  flam- 
mules  of  light  golden  brown;  peripher\  with  distinct, 
squarish  spots  of  orange-brown. 

Holotype:  USNM  860252,  height  13.4  mm,  width  11.4 
mm. 

Type  locality:  West  of  Isla  La  Tortuga,  N'enezuela,  John 
Elliott  Pillsbury  Station  P-7o6,  10°57  0'\.  65°52.0'W. 
in  69-155  m. 

Paratype:  1.  UMML  30.5915;  off  Pointe  Isere.  French 
Guiana.  John  Elliott  Pillsbiry  Station  P-65S.  7°10.0'N. 
53°36.0'W.  in  135-126  m;  9  July  1968;  10-ft  otter  trawl. 

Distribution:  Calliostoma  semisuave  is  known  from  dead 
shells  collected  off  eastern  X'enezuela  and  northwestern 
French  Guiana  in  depths  of  69-155  m. 

Remarks:  The  shells  of  Calliostoma  semisuave  are  most 
similar  to  those  of  C.  cnidophiliim  but  are  proportion- 
ately narrower,  have  a  larger  protoconch.  have  the  bead- 
ing of  the  spiral  cords  becoming  very  weak  on  the  last 
several  whorls,  and  have  a  smooth  peripheral  cord  on 
the  last  several  whorls. 

Other  organisms  listed  on  the  field  data  sheets  for  the 
two  collections  of  C.  semisuave  include  sponges,  alcyo- 
narians.  Millepora.  Haliotis,  and  chitons,  suggesting  that 
this  species  li\es  on  or  near  hard-bottom  outcrops  in 
depths  of  about  70-150  m. 


Calliostoma  hrunneopictum  new  species 

(figures  17-20) 

Description:  Shell  small,  attaining  height  of  8.5  mm, 
w  idth  of  6.9  mm.  conical,  narrowly  umbilicate  or  non- 
umbilicate. finely  sculptured,  Protoconch  365-375  ^i.va 
maximum  diameter.  1  whorl.  Teleoconch  whorls  6.7.  flat 
to  weakly  convex;  first  whorl  with  low,  rounded  axial 
riblets  and  2  strong,  beaded  spiral  cords;  axial  sculpture 
absent  after  first  whorl;  spiral  sculpture  increasing  b%' 
intercalation  to  8-12  cords  and  1-3  threads,  cords  finely 
beaded,  beads  conical;  peripher\  rather  sharp  on  first  5 
whorls,  becoming  narrowly  rounded  on  last  2  whorls. 
Base  weakly  convex.  v\ith  12-16  narrow,  fineh'  beaded 
spiral  cords,  adaxial  2-3  cords  slightly  stronger  and  more 
w idel\  spaced.  Umbilicus  ver\  narrow,  chink-like,  par- 
tialK  or  completeK  filled  by  columella.  Aperture 
subquadrate  to  subovate.  lips  thin,  finely  crenulate;  col- 
umella concave,  thickened,  terminating  in  obscure, 
rounded  denticle.  Shell  ground  color  ivory  with  greenish 
and  reddish  iridescence,  irregular  spots  and  flammules 
of  brown  above  periphery,  squarish  spots  of  brown  reg- 
ularly spaced  on  peripher\ .  and  obscure,  crescentic  streaks 
of  brown  on  base. 

Holotype:  MORG  29.291.  height  8.5  mm,  width  6.9 
mm. 

Type  locality:  Off  Santana  Island.  Estado  de  Rio  de 
Janeiro.  Brazil,  in  50-80  m. 

Paratypes:  1,  MCZ  302592;  1.  FSBC  I  44068;  from  same 
lot  as  holot\pe. 

Other  material:  2.  Coltro  collection;  from  same  lot  as 
holot\pe.  —  1.  Sunderland  collection;  off  Rio  de  Janeiro. 
Brazil,  50-70  m;  trawled  by  shrimpers. 

Distribution:  At  present.  Calliostoma  hrunneopictum 
is  know  n  onl\  from  off  Rio  de  Janeiro.  Brazil,  in  depths 
of  50-80  m.  Living  specimens  are  also  known  from  50- 
80  m. 

Remarks:  Shells  of  Calliostoma  hrunneopictum  are  most 
similar  to  those  of  C.  roseolum  but  differ  by  having  a 
larger  protoconch  (365-375  (um  r.s.  325  nm);  b\  having 
more  numerous,  more  finely  beaded  spiral  cords;  b\  hav- 
ing an  umbilical  chink;  and  by  having  crescentic  streaks 
of  brown  on  the  base.  Differences  between  C.  hrunneo- 
pictum and  the  similar  C.  viscardii  are  discussed  in  the 
Remarks  section  of  the  latter  species. 


Calliostoma  viscardii  new  species 
(figures  21.  22.  108) 


Figures  11-14.  Calliostoma  cnidophilum  new  species.  11,  12.  Holotype,  USNM  860265;  height  13,7  mm,  width  11.4  mm  13, 
14.  Specimen  from  off  Barbados,  Sunderland  collection;  height  11,2  mm,  width  9,2  mm.  Figures  15,  16.  Calliostoma  semisuave 
new  species.  holot\pe.  USNM  860252;  height  13,4  mm,  width  11,4  mm.  Figures  16-20.  Calliostoma  hrunneopictum  new  species. 
17,  18.  Holotype,  MORG  29  291;  height  8.5  mm.  width  6.9  mm.  19.  20.  Paratype,  FSBC  I  44068;  height  7.5  mm,  width  6  0  mm 
Figures  21.  22.  Calliostoma  viscardii  new  species,  holotype.  MORG  29,292;  height  10,9  mm.  width  8,2  mm. 


Page  86 


THE  NAUTILUS,  Vol.  106,  No.  3 


Description:  Shell  small  to  medium-sized,  attaining 
height  of  13.0  mm,  width  of  10.8  mm,  conical,  non- 
umbilicate,  finely  sculptured.  Protoconch  300-335  ^m 
maximum  diameter,  1  whorl,  tip  violet.  Teleoconch 
whorls  8.25,  flat  to  weakly  convex;  first  whorl  with  2 
strong,  smooth  to  weakly  beaded  spiral  cords,  initially 
lacking  axial  sculpture  but  developing  low,  rounded  rib- 
lets  near  end  of  whorl,  riblets  persisting  on  subsequent 
whorls  as  low  folds  between  spiral  cords;  spiral  sculpture 
increasing  b\-  intercalation  to  7-8  cords  and  2-6  threads, 
of  which  abapical  2  spiral  cords  forming  periphery,  cords 
rather  coarsely  beaded,  beads  rounded,  slightly  spirally 
elongate  on  last  2  whorls;  periphery  rather  sharp  on  first 
5  whorls,  becoming  narrowly  rounded  on  last  2  whorls, 
peripheral  cords  adpressed.  Base  weakly  convex,  with 
13-16  narrow,  weakly  beaded  spiral  cords.  Aperture 
subquadrate  to  subovate,  lips  thin,  finely  crenulate;  col- 
umella concave,  thickened,  terminating  in  obscure, 
rounded  denticle.  Shell  ground  color  tan  with  greenish 
and  reddish  iridescence,  patches  and  flammules  of  brown 
and  occasional  white  spots  on  and  above  periphery,  and 
elongate  spots  of  brown  on  basal  cords.  Animal  (in  al- 
cohol): foot  dark  brown  with  white  papillae;  mantle  edge 
with  irregular  band  of  brown,  mantle  mottled  black  and 
white  posteriorly;  head  mottled  black  and  white;  cephalic 
tentacles  short,  stout,  tapering  rapidK  to  narrow  tip;  eye- 
stalks  short  (but  about  40%  tentacle  length),  stout,  with 
large,  black  eye  at  tips;  snout  mottled  with  brown  and 
white,  longer  than  broad,  slightly  longer  than  cephalic 
tentacles,  slightly  tapered  at  tip,  tip  with  fringe  of  fine, 
rather  long  papillae;  epipodium  with  4  pairs  of  short, 
stout  tentacles;  left  neck  lobe  seems  to  be  smooth,  right 
lobe  finely  digitate. 

Holotype:  MORG  29.292,  height  10.9  mm,  width  8.2 
mm. 

Type  locality:  Between  Ihla  de  Sao  Sebastiao  and  Ihla 
de  Buzias,  off  Sao  Paulo,  Estado  de  Sao  Paulo,  Brazil,  in 
40-45  m. 

Paratypes:  1,  USNM  860253;  1,  FSBC  1  44066;  from 
same  lot  as  holotype. 

Other  material:  3,  Coltro  collection;  off  Guarapari,  Es- 
tado de  Espirito  Santo,  Brazil,  20-30  m;  trawl;  July 
1991. — 6,  Coltro  collection;  off  Ihla  de  Sao  Sebastiao, 
Estado  de  Sao  Paulo,  Brazil,  30-35  m;  dredge;  J.  &  M. 
Coltro  &  L.  F.  Viscardi,  collectors. 

Distribution:  Calliostoma  viscardii  is  known  from 
southeastern  Brazil  (Guarapari  to  Sao  Paulo)  in  depths 
of  20-45  m;  living  specimens  have  been  collected  in 
depths  of  30-45  m. 

Remarks:  Shells  of  this  species  resemble  those  of  Cal- 
liostoma brunneopictiim  but  differ  by  having  a  smaller 
protoconch  (300-335  nm  vs.  365-375  ^m)  with  a  violet 
tip;  by  having  stronger,  more  widely  spaced  spiral  cords 
whose  beads  are  large  and  rounded  rather  than  small 
and  conical;  by  having  the  whorl  periphery  composed 
of  a  pair  of  close-set  spiral  cords  abapical  to  the  cord 


that  is  the  continuation  of  the  abapical  cord  on  the  first 
whorl,  whereas  the  abapical  cord  on  the  first  whorl  of 
C.  brunneopictiim  forms  the  periphery  on  all  whorls; 
and  b\  having  less  well-defined  beading  on  the  basal 
spiral  cords.  Calliostoma  viscardii  is  also  similar  to  C. 
roseolum,  but  the  shells  have  a  larger  protoconch  (335- 
350  (um  vs.  300-335  fim);  lack  the  prominent,  broadly 
rounded  whorl  periphery;  have  stronger,  more  widely 
spaced  spiral  cords  above  the  periphery;  and  have  slightly 
larger  but  less  discrete  beading.  The  radula  of  C.  viscardii 
(formula  oo.5.1.5.oo;  figure  108)  does  not  differ  signifi- 
cantl)'  from  that  of  C.  piilchrum  illustrated  b\  Clench 
and  turner  (1960:pl.  3,  fig.  3). 

This  species  is  named  viscardii  to  recognize  the  con- 
tributions of  Sr.  Luiz  Francisco  Viscardi  to  Brazilian  mal- 
acology. 

Calliostoma  bermiidense  new  species 

(figures  23,  24) 

Description:  Shell  small  to  medium-sized,  attaining 
height  of  12.6  mm,  width  of  8.9  mm,  conical,  nonum- 
bilicate,  finely  sculptured.  Protoconch  315-325  ^m  max- 
imum diameter,  1  whorl.  Teleoconch  whorls  about  8,  flat 
to  weakly  concave;  first  whorl  with  low,  rounded  axial 
riblets  and  2  strong,  beaded  spiral  cords;  axial  sculpture 
weakening  rapidly  at  end  of  second  whorl,  absent  there- 
after; spiral  sculpture  increasing  by  intercalation  to  8- 
10  cords  and  1-4  threads,  of  which  abapical  2-3  cords 
forming  periphery,  cords  strongly  beaded;  periphery  of 
first  3-4  whorls  sharply  angulate,  that  of  subsequent 
whorls  broadly  rounded,  not  distinctly  set  off  from  base. 
Base  flat  to  weakly  convex,  with  10-14  narrow,  weakly 
beaded  spiral  cords  of  which  1-3  markedK  weaker.  .Ap- 
erture subquadrate  to  ovate,  thickened  within,  lips  thin, 
crenulate;  columella  short,  almost  straight,  thickened. 
Shell  ground  color  yellowish  brown,  spiral  cords  brown, 
often  broken  into  dashed  lines  on  base,  periphery  often 
with  diffuse  patches  of  brown  to  red-brown,  apical  3-4 
whorls  pinkish  brown. 

Holotype:    DMNH  187591,  height  11.9  mm,  width  8.9 


Type  locality:    Off  Castle  Roads,  Bermuda,  in  82  m. 

Paratypes:  9,  DMNH  187592;  2,  USNM  860269;  1,  FSBC 
I  42541;  all  from  same  lot  as  holot\pe. 

Distribution:  Calliostoma  bermiidense  is  known  only 
from  the  type  localit> ;  no  living  specimens  are  known. 

Remarks:  Shells  of  Calliostoma  bermiidense  are  most 
similar  to  those  of  C.  roseolum  but  differ  by  being  rel- 
atively broader;  by  having  narrower,  more  widely  spaced 
and  more  finely  beaded  spiral  cords;  b\-  having  a  flatter 
whorl  profile  after  the  fourth  whorl;  b\  having  a  pro- 
portionateK-  shorter  and  thicker  columella;  b\  having 
brown  or  pinkish-brown  apical  whorls;  and  by  having 
brown  spiral  cords. 


J.  F.  Quinn,  Jr.,  1992 


Page  87 


Calliostoma  deniatum  new  species 
(figures  25,  26) 

Description:  Shell  small,  attaining  height  of  8.4  mm, 
width  of  7.4  mm,  conical,  nonumbilieate,  finely  sculp- 
tured. Protoconch  335-350  /jm  ma.ximum  diameter.  1 
whorl.  Teleoconch  whorls  8,  flat  to  weakly  concave;  first 
2  whorls  with  low,  rounded  axial  riblets  and  2  strong, 
beaded  spiral  cords;  axial  sculpture  on  remaining  whorls 
of  low,  rounded,  discontinuous  axial  threads,  most  dis- 
tinct at  peripher\;  spiral  sculpture  increasing  by  inter- 
calation to  4-8  strongly  beaded  cords,  of  which  abapical 
2-3  spiral  cords  forming  periphery;  periphery  angulate, 
adapical  peripheral  cord  strongest,  abapical  2  (when  3 
present)  weaker,  closeK'  appressed,  set  with  axially  elon- 
gate beads.  Base  flat,  with  10-14  narrow,  finely  beaded 
spiral  cords.  Aperture  subquadrate,  lips  thin,  crenulate; 
columella  straight,  thickened.  Shell  ground  color  ivory 
with  diffuse  axial  flammules  of  golden  brown  above  pe- 
riphery, and  2-3  spiral  row  s  of  discrete,  spirally  elongate 
spots  of  golden  or  reddish  brown  on  base. 

Holotype:  USNM  859358,  height  5.7  mm,  width  5.2 
mm. 

Type  locality:  Off  Freeport,  Texas,  approximately 
28°05'N,  94°35'W,  in  51  m. 

Paratypes:  1,  MCZ  297052;  1,  LACM  2293;  1,  HMNS 
3946;  N\V  Gulf  Survey  station,  off  Freeport,  Texas,  51 
m;  trawl;  A.  Kight,  collector.— 1,  HMNS  15866;  2,  FSBC 
I  32301;  XW  Gulf  Survey  station,  2  mi  south  of  #1  buoy, 
Heald  Bank,  22  m;  22  August  1966;  H.  Geis  and  W. 
Pierce,  collectors.  Other  paratypes  and  material  (44  lots) 
are  listed  by  Quinn  (in  press). 

Distribution:  Calliostoma  dentatum  is  confined  to  the 
northwestern  Gulf  of  Mexico,  from  the  Mississippi  River 
to  off  Port  Isabel,  Texas,  and  is  usually  collected  from 
depths  of  15-55  m;  living  specimens  are  known  from 
depths  of  12-51  m. 


Turner  (1960:pl.  3,  fig.  3)  by  having  four  rather  than  five 
lateral  teeth. 

Calliostoma  tenebrosum  new  species 
(figures  27,  28) 

Description:  Shell  medium-sized,  attaining  height  of 
13.6  mm.  width  of  11.5  mm,  conical,  nonumbilieate, 
finely  sculptured.  Protoconch  slightly  chipped,  about  325 
;um  maximum  diameter,  1  whorl.  Teleoconch  whorls  7.9, 
flat;  first  3  whorls  with  low,  rounded  axial  riblets  and  2- 
4  strong,  beaded  spiral  cords;  axial  riblets  weakening  on 
fourth  whorl,  becoming  weak,  irregular  folds  on  subse- 
quent whorls;  spiral  sculpture  increasing  by  intercalation 
to  7  cords  and  1  thread,  of  which  abapical  3  cords  form- 
ing periphery,  beads  on  cords  strong,  rounded;  periphery 
rather  broadly  rounded,  adapical  2  peripheral  cords 
strongest,  subequal,  separated  by  rather  broad  groove. 
Base  very  weakly  convex,  with  13  narrow,  finely  beaded 
spiral  cords.  Aperture  subquadrate,  moderately  thick- 
ened within,  lips  thin,  crenulate;  columella  very  weakly 
concave,  thickened,  terminating  in  rounded  tubercle. 
Shell  ground  color  tan  with  spots  of  white  and  spots  and 
axial  streaks  of  dark  brown. 

Holotype:  ANSP  300356,  height  13.6  mm,  width  11.5 
mm. 

Type  locality:  Off  Potengi  River,  Natal,  Estado  Rio 
Grande  do  Norte,  Brazil,  depth  unrecorded. 

Distribution:  This  species  is  known  only  from  the  ho- 
lotype, a  hermit-crabbed  shell,  collected  from  a  bottom 
of  black  mud  in  an  unrecorded  depth. 

Remarks:  Calliostoma  tenebrosum  is  readily  distin- 
guished from  other  species  of  the  C.  pulchrum  group  by 
the  relatively  large,  darkly  colored  shell  that  has  the 
whorl  periphery  composed  of  three  widely  spaced  spiral 
cords;  the  beads  on  the  spiral  cords  are  unusually  large 
for  this  species  group. 


Remarks:  Calliostoma  dentatum  is  closely  related  to 
C.  pulchrum,  C.  roseolum.  and  the  Plio-Pleistocene  C. 
bowdenense  Woodring,  1928.  Shells  of  C.  dentatum  dif- 
fer from  those  of  C.  pulchrum  by  having  a  larger  pro- 
toconch .X  =  346  |um  vs.  300  ^m;  Quinn,  in  press);  by 
having  fewer  spiral  cords  both  above  and  below  the  pe- 
riphery; and  by  having  the  beads  on  the  cords  stronger, 
more  sharply  defined,  and  lateralK'  compressed.  Shells 
of  C.  dentatum  differ  from  those  of  C.  roseolum  by 
having  a  larger  protoconch  x  =  346  ^m  vs.  324  jum),  by 
being  relatively  narrower,  b\'  having  a  narrower  and 
more  angulate  periphery,  and  by  having  laterally  com- 
pressed beads  on  the  spiral  cords.  Shells  of  C.  dentatum 
differ  from  those  of  C.  bowdenense  by  having  more 
numerous  spiral  cords  above  the  periphery;  by  having  a 
narrower,  more  sharply  angulate  periphery;  and  by  hav- 
ing weaker,  more  weakly  beaded  spiral  cords  on  the  base. 
The  radula  of  C.  dcntatun}  (formula  go. 4.1. 4. oo)  differs 
from  that  of  C.  pulchrum  illustrated  by  Clench  and 


Calliostoma  moscatellii  new  species 
(figures  29,  30,  109) 

Calliostoma  carcellesi.  Rios,  1970:25,  pi.  4  (partim);  1975:23, 
pi.  6,  fig.  61  (partim).  (Non  Calliostoma  carcellesi  Clench 
&  .'^guayo,  1940). 

Calliostoma  {Neocalliostoma)  carcellesi:  Rios,  1985:22,  pi,  10, 
fig,  88  (partim). 

Description:  Shell  medium-sized,  attaining  height  of 
27.7  mm,  width  of  17.9  mm,  conical,  nonumbilieate, 
rather  coarsely  sculptured.  Protoconch  375  ^m  maxi- 
mum diameter,  1  whorl.  Teleoconch  whorls  about  10, 
flat  to  weakly  convex;  first  whorl  with  weak,  rounded 
axial  riblets  and  2  strong,  beaded  spiral  cords;  axial  riblets 
replaced  on  subsequent  whorls  by  low,  rounded,  crowded 
plicae  in  interspaces  between  spiral  cords,  plicae  strong- 
est near  suture  and  below  peripheral  cord;  spiral  sculp- 
ture increasing  by  intercalation  to  5-8  cords  and  4-9 
threads,  of  which  abaxial  2-3  strong  cords  and  2-3  weak 


Page  88 


THE  NAUTILUS,  Vol.  106,  No.  3 


27 


28 


32 


J.  F.  Quinn,  Jr.,  1992 


Page  89 


cords  or  threads  forming  peripher\ ,  cords  rather  coarsely 
beaded,  beads  rounded,  spirally  elongate;  periphery  nar- 
rowly rounded.  Base  weakK  convex,  w  ith  24-27  narrow, 
smooth  to  weakly  beaded  spiral  cords  and  threads.  Ap- 
erture subquadrate,  lips  thin,  creiiulate;  columella  short, 
straight,  thickened.  Shell  ground  color  light  yellowish  or 
pinkish  tan  with  greenish  iridescence,  diffuse  patches  of 
tan  to  reddish  brown,  and  occasional  discrete  spots  of 
light  brown;  2-5  basal  spiral  cords  with  elongate  spots 
of  brown;  protoconch  and  first  2  whorls  lilac.  Animal  (in 
alcohol)  white  except  brown  sides  of  snout;  cephalic  ten- 
tacles long,  broad,  tapering  abruptK-  near  tip,  ocular 
peduncles  short  (about  20%  tentacle  length),  with  large, 
black  eye  at  tips;  snout  broad  at  base,  tapering  to  rounded 
tip,  tip  with  fringe  of  ver\  small,  short  papillae;  epipo- 
dium  with  1  tentacle  on  left,  2  on  right,  left  larger  than 
right,  neck  lobes  well  de\eloped,  thin,  semicircular, 
smooth. 

Holotype:  MORG  29.293,  height  25.9  mm,  width  17.7 
mm. 

Type  locality:  Off  Ilha  de  Santana,  Estado  de  Rio  de 
Janeiro,  Brazil,  in  approximately  50-80  m. 

Paratypes:  1,  USNM  860254;  1,  MCZ  302591;  1,  ANSP 
389337;  1,  FSBC  I  44075;  1,  UP  189458;  all  from  same 
lot  as  holotype. 

Other  material:  5,  Coltro  collection;  from  same  lot  as 
holotype. 

Distribution:  Numerous  specimens  of  C.  moscatellii, 
including  live-collected  specimens,  have  been  obtained 
from  depths  of  50-200  m  off  Rio  de  Janeiro,  Brazil,  by 
shrimpers. 

Remarks:  Shells  of  Calliostoma  moscatellii  are  ver\' 
similar  to  those  of  C.  carcellesi  (figures  31,  32)  but  differ 
by  being  markedly  narrower;  by  having  a  smaller  pro- 
toconch (375  ^m  vs.  400-425  fim);  by  having  fewer  basal 
spiral  cords,  most  of  which  are  weakly  but  discretely 
beaded;  by  having  more  strongly  spirally  elongate  beads 
on  the  supraperipheral  spiral  cords;  by  having  lilac  apical 
whorls;  and  by  having  a  distinct  color  pattern.  Callios- 
toma moscatellii  is  also  similar  to  C.  jucundum  but  has 
narrower  shells  with  more  numerous,  narrower  spiral 
cords  that  bear  much  smaller,  spirally  elongate  beads. 
The  radula  of  C.  moscatellii  (formula  00.6.I.6.00;  figure 
109)  resembles  that  of  C.  pulchrum  illustrated  by  Clench 
and  Turner  (1960:pl.  3,  fig.  3)  but  has  a  broader  rachidian 
tooth,  and  the  cusps  of  the  rachidian  and  lateral  teeth 
are  more  finelv  denticulate. 


This  species  is  named  for  Sr.  Renato  Moscatelli,  in 
recognition  of  his  support  of  malacological  publications 
in  Brazil. 

Calliostoma  apicii^um  Dall,  1881 
(figures  33,  34) 

Catliosionm  apicinum  Dall,  1881:46;  1889b:162,  pi.  24,  figs.  3, 
3a.—  Piisbrv,  1890:379,  pi.  60,  figs.  1,  2.— Johnson,  1934: 
69. 

Calliostoma  {Calliostoma)  apicintim.  Dall,  1889a:366,  pi.  24, 
figs.  3,  3a. 

Calliostoma  roseolum.  Clench  &  Turner,  1960:19,  pi.  15,  figs. 
1,2  (partim).— Abbott,  1974:43  (partim).— Quinn,  1979: 
26  (partim),—  Sander  &  Lalli,  1982:table  4.  (Non  Cal- 
liostoma roseolum  Dall,  1881). 

Description:    See  Dall  (1881:46). 

Lectotype:  (by  implication  of  holotype;  Clench  &  Tur- 
ner, 1960):  USXM  95013,  height  7.7  mm,  width  6.7  mm. 

Type  locality:  Off  Barbados,  Blake  station  (number  and 
coordmates  unrecorded),  in  183  m. 

Paralectotypes:  1,  USNM  95012;  off  Barbados,  Blake 
Station  290,  13ni'54"N,  59°39'45"W,  134  m;  9  March 
1879. — 2,  MCZ  7564;  Blake  station  (number  and  coor- 
dinates unrecorded),  183  m. 

Other  material:  1,  ANSP  353528;  1,  FSBC  I  44077;  off 
Holetown,  Barbados,  175-225  m;  ex  F.  Sander. — 5,  Sun- 
derland collection;  off  St.  James,  Barbados,  100  m;  dredge; 
ex  F.  Sander. — 1,  Sunderland  collection;  off  St.  James, 
Barbados,  152  m;  1988;  ex  F.  Sander. — 6,  Sunderland 
collection;  off  St.  James,  Barbados,  175-225  m;  ex  F. 
Sander. — Several  other  specimens,  all  from  Barbados,  in 
private  collections. 

Distribution:  This  species  is  known  only  from  off  Bar- 
bados in  depths  of  100-225  m;  1  have  only  seen  one  live- 
collected  specimen  from  a  depth  of  100  m. 

Remarks:  This  species  was  synonymized  with  Callios- 
toma roseolum  b\  Clench  and  Turner  (1960),  who 
thought  that  the  shells  described  by  Dall  as  C.  apicinum 
were  merely  juveniles  of  C.  roseolum.  Shells  of  C.  api- 
cinum. however,  are  distinguished  from  those  of  C.  ro- 
seolum  by  retaining  flat-sided,  peripherally  subangulate 
whorls  as  adults  (7-1-  whorls)  rather  than  by  ha\ing  con- 
cave whorls  with  broadly  rounded  peripheries  after  the 
third  or  fourth  whorl;  by  having  a  larger  protoconch 
(360-375  ^lm  vs.  300-330  ^m)  that  has  a  purplish-brown 
tip;  by  having  the  apical  two  teleoconch  whorls  brown 
or  purplish  brown;  by  having  more  widely  spaced,  smooth 


Figures  23,  24.  Calliostoma  bermudctise  new  species,  holotype,  DMNH  187591;  height  11.9  mm,  width  8.9  mm.  Figures  25, 
26.  Calliostoma  dentatum  new  species,  hoiot\pe,  I'SXM  859358;  height  5.7  mm,  width  5,2  mm.  Figures  27,  28.  Calliostoma 
tcncbrosum  new  species,  holot\'pe,  ANSP  300356;  height  13,6  mm,  width  11,5  mm.  Figures  29,  30.  Calliostoma  moscatellii  new 
species,  holotype.  MORG  29  293;  height  25,9  mm,  width  17  7  mm  Figures  31,  32.  Calliostoma  carcellesi  Clench  &  ,\guayo,  1940, 
holotype,  MCZ  104719;  height  20,5  mm,  width  17.5  mm. 


Page  90 


THE  NAUTILUS,  Vol.  106,  No.  3 


Figures  33,  34.  Calliostoma  apicintim  Dall,  1881,  specimen  from  off  Barbados,  FSBC  I  44077;  height  10.0  mm,  width  8.2  mm. 
Figures  35, 36.  Calliostoma  delnle  new  species,  holotype,  USNM  860255;  height  8. 1  mm,  width  7.0  mm.  Figures  37, 38.  Calliostoma 
Indiana  Dall,  1889,  specimen  from  off  Honduras,  FSBC  I  44071;  height  10.4  mm,  width  9.0  mm.  Figures  39,  40.  Calliostoma 
orion  Dall,  1881,  specimen  from  John  Elliott  Pillsbury  ,Station  P-425,  UMML  30.;3635;  height  16  2  mm,  width  14  4  nmi. 


or  very  weakly  beaded  spiral  cords  on  the  outer  two- 
thirds  of  the  base;  by  often  having  a  chink-like  umbilicus; 
and  by  having  a  shorter  columella  that  often  has  a  sub- 
conical,  medial  swelling.  Calliostoma  apicinum  seems 
to  be  most  closely  related  to  the  sympatric  C.  debile  (see 
Remarks  under  the  latter  species  for  comparisons).  Spec- 
imens that  I  can  assign  unequivocally  to  C.  apicintim 
seem  to  come  only  from  Barbados.  Some  other  specimens 
w  ith  similar  morphologies  from  the  Bahama  Islands  (MCZ 
uncatalogued)  and  northwestern  Cuba  (M("Z  7566)  are 
not  here  considered  conspecific  with  the  Barbados  spec- 


imens. A  paralectotype  of  C.  apicinum  (MCZ  7565)  from 
off  Havana,  Cuba,  is  definitely  not  this  species;  the  spec- 
imen has  a  uniquely  undulate  peripheral  spiral  cord  on 
several  of  the  adapical  teleoconch  whorls  and  probably 
represents  an  undescribed  species. 

Calliostoma  dehilc  new  species 
(figures  35,  36) 

Description:  Shell  small  to  medium-sized,  attaining 
height  of  10.3  mm,  width  of  8.7  mm,  conical,  nonum- 


J.  F.  Quinn,  Jr.,  1992 


Page  91 


bilicate,  finely  sculptured.  Protoconch  350-365  A^ni  max- 
imum diameter,  1  whorl.  Teleoconch  whorls  7.7,  flat; 
first  2  whorls  with  low,  rounded  a.xial  riblets  and  2  strong, 
beaded  spiral  cords,  abapical  cord  stronger  on  first  4 
whorls,  set  with  strong,  conical  beads;  a.\ial  sculpture 
absent  on  subsequent  whorls;  spiral  sculpture  increasing 
by  intercalation  to  7-10  cords  and  0-3  threads,  of  which 
aba.xial  3,  sometimes  2,  cords  forming  periphery,  supra- 
peripheral  cords  fine,  sharp,  finely  beaded;  periphery 
subcarinate,  slighth  projecting,  peripheral  cords  stronger 
than  supraperipheral  cords,  set  with  conical  beads.  Base 
flat,  with  10-14  narrow,  flattened,  smooth  to  weakly  ru- 
gose spiral  cords  and  2-3  threads  or  weak  cords  near 
periphery.  Aperture  subquadrate,  lips  thin,  weakly  cren- 
ulate;  columella  straight,  thickened.  Shell  ground  color 
ivory,  periphery  light  pinkish  brown  or  with  pale  to 
bright  spots  of  pinkish  or  \  ellowish  brou  n  or  clear  rose. 

Holotype:  USNM  860255,  height  8.1  mm,  width  7.0 
mm. 

Type  locality:    Off  St.  James,  Barbados,  in  175-225  m. 

Paratype:  1,  FSBC  I  45776;  off  St.  James,  Barbados,  152 
m;  dredge;  ex  F.  Sander. 

Other  material:  4,  Sunderland  collection;  off  St.  James, 
Barbados,  152  m;  1988;  dredge;  ex  F.  Sander. — 6,  Sun- 
derland collection;  off  St.  James,  Barbados,  175-225  m; 
dredge;  ex  F.  Sander. 

Distribution:  Calliostoma  debile  is  known  only  from 
empt\  shells  collected  off  Barbados  in  152-225  m. 

Remarks:  Shells  of  Calliostoma  debile  are  most  similar 
to  those  of  the  sympatric  C.  apicinitm  but  differ  by 
having  a  slightK  smaller  protoconch;  b\  lacking  the  brow  n 
or  purplish-brown  apical  whorls;  by  having  a  more  pro- 
jecting periphery,  usually  composed  of  three  (sometimes 
two)  close-set  spiral  cords;  by  usually  having  two  to  three 
spiral  threads  or  weak  cords  between  the  circumbasal 
cord  and  the  outermost  strong  basal  spiral  cord;  by  having 
distinct  radial  threads  that  finely  bead  the  basal  spiral 
cords;  by  having  a  less-thickened  columella  that  lacks 
any  indication  of  a  medial  swelling;  and  by  lacking  strong 
ridges  within  the  aperture. 


Calliostoma  indiana  Dall,  1889 
(figures  37,  38) 

Calliostoma  (Eucasta)  indiana  Dall,  lS89a;368,  pi.  32,  figs,  3, 

5.—  .Abbott,  1974:46,  fig.  333, 
Calliostoma  indiana:  CHench  &  Turner.  1960:52,  pi  34,  fig.  1. 

Description:    See  Dall  (1889a:368)  and  Clench  and  Tur- 
ner (1960:52). 

Holotype:     USNM  214273,  height  7.9  mm,  width  6.9 


Type  locality:    Off  Grenada,  Lesser  Antilles,  Blake  Sta- 
tion 247,  12°05'25"N,  61°47'15"W,  in  311  m. 

Other  material:     1,  FSBC  1  44071;  off  Honduras,  Cape 


Hatteras  station,  15°32.36'N,  8r39.80'W,  50-85  m;  8 
April  1987;  rock  dredge. 

Distribution:  This  species  is  now  known  from  two  spec- 
imens, the  holotype  from  Grenada  and  the  Cape 
Hatteras  specimen  from  off  Honduras.  Based  on  know  1- 
edge  of  the  distributions  of  other  species  of  Calliostoma 
(Clench  &  Turner,  1960;  Quinn,  herein,  in  press),  I  be- 
lieve C.  indiana  probabK  inhaliits  deep  fore-reef  areas 
(50-150  m)  of  Central  America  and  the  Greater  and 
Lesser  Antilles. 

Remarks:  The  new  shell  reported  here  is  larger  than 
the  holotype  (height  10.4  mm,  width  9.0  mm),  has  five 
rather  than  four  strong  supraperipheral  cords,  and  has 
ten  rather  than  seven  basal  spiral  cords;  otherwise  the 
two  shells  are  very  similar.  Based  on  shell  characters,  this 
species  clearly  belongs  to  the  C.  pidchrum  species  com- 
plex and  is  most  similar  to  C.  echinatiim.  C.  orion,  C. 
sopidiim,  and  C.  cinctetliim. 


Calliostoma  orion  Dall,  1889 
(figures  39,  40) 

Calliostoma  orion  Dall,  lS89a:367,  pi.  28,  fig.  2;  1889b:162.— 
Pilsbrv,  1890:383,  pi.  48.  fig.  18.— Johnson,  1934:70.— 
Clench  &  Turner,  1960:54,  pi.  35,  figs.  1,  2.— Quinn,  1981: 
151,  figs,  1-13. 

Description:  See  Dall  (1889a;367)  and  Quinn  (1981: 
151). 

Holotype:  USNM  214272,  height  4.6  mm,  width  4.3 
mm. 

Type  locality:  Off  Havana,  Cuba,  Blake  Station  62,  in 
146  m. 

Other  material:  2,  FSBC  I  31518;  off  western  end  of 
Isla  de  Roatan,  Honduras,  19.8  m;  15  September  1982; 
SCUBA  collection  by  W.  C.  Jaap. —  1,  Crnkovic  collec- 
tion; off  western  end  of  Isla  de  Roatan,  Honduras,  ap- 
proximatelv  30  m;  1990;  SCUBA  collection  bv  L.  A. 
Crnkovic  — 1,  UMML  30.5688;  off  Isla  de  Roatan,  Hon- 
duras, John  Elliott  Pillsbury  Station  P-629, 15°58.2'N, 
86°09.0'W,  40  m;  21  March  1968;  40-ft  otter  trawl—  1, 
UF  36324;  Isla  de  Providencia,  Colombia  (off  Nicara- 
gua), "shallow  water";  August  1971;  C.  R.  Gilbert,  col- 
lector.—1,  UMML  30.3635;  off  Punta  Manzanillo,  Pan- 
ama, John  Elliott  Pillsbury  Station  P-425,  9°38.9'N, 
79°15.3'W,  70-64  m;  19  July  1968;  10-ft  otter  trawl.— 
3,  Sunderland  collection;  off  Barbados;  ex  F.  Sander. 

Distribution:  Calliostoma  orion  is  now  known  from 
almost  the  entire  Caribbean  coast  of  Central  America, 
Cuba,  the  Bahama  Islands,  and  Barbados. 

Remarks:  1  discussed  the  ecology,  shell  and  radular 
morphologies,  and  other  distributional  records  in  a  pre- 
vious paper  (Quinn,  1981).  The  present  specimens  in- 
crease the  maximum  known  size  to  20.2  mm  high,  17.1 
mm  wide;  extend  the  maximum  depth  from  w  hich  living 
specimens  have  been  obtained  from  43  m  to  64-70  m; 


Page  92 


THE  NAUTILUS,  Vol.  106,  No.  3 


and  add  new  collection  localities  off  Honduras,  Nicara- 
gua, and  Panama. 

Calliostoma  jernandezi  Princz,  1978 
(figures  41,  42) 

Calliostoma  jernandezi  Princz,  1978:1.52-154,  figs.  1,  2. 

Description:  Shell  large,  attaining  height  of  28. .5  mm, 
width  of  25.5  mm,  conical,  umbilicate,  finely  sculptured. 
Protoconch  350-375  ^m  maximum  diameter,  1  whorl. 
Teleoconch  whorls  more  than  10,  flat  to  concave;  first  2 
whorls  flat-sided,  with  low,  rounded  a.xial  riblets  and  2- 
4  finely  beaded  spiral  cords;  axial  sculpture  absent  on 
subsequent  v\horls;  spiral  cords  increasing  by  intercala- 
tion to  17-22,  of  which  abapical  3-4  cords  forming  pe- 
riphery, all  cords  subequal  and  with  well-defined,  close- 
set,  rounded  beads;  periphery  narrowly  rounded.  Base 
flat  to  weakly  convex,  with  17-28  finely  beaded  spiral 
cords,  those  near  periphery  narrow,  sharp,  cords  becom- 
ing broader  and  flatter  adaxially,  often  splitting  into  2 
subequal  cords;  circumumbilical  1-2  cords  strongest,  tu- 
berculate.  Umbilicus  funnel-shaped,  17%-24%  maxi- 
mum shell  diameter,  wall  almost  vertical,  white.  Aper- 
ture subquadrate,  thickened  within  and  with  strong,  sharp 
ridges,  lips  thin,  crenulate;  columella  weakly  sigmoid, 
thickened,  terminating  in  blunt,  rounded  tubercle.  Shell 
ground  color  cream  to  straw  with  irregular,  diffuse  patch- 
es of  light  brown  above  periphery;  periphery  with  reg- 
ular series  of  squarish,  rose-brown  to  brown  spots;  inter- 
spaces of  supraperipheral  spiral  cords  golden  brown;  base 
finely  mottled  with  cream  and  yellow-brown. 

Holotype:  Collection  of  "Familia  Fernandez  en  Ma- 
racay,  Estado  Aragua,  Venezuela"  (Princz,  1978:152), 
height  14.7  mm,  width  15.8  mm. 

Type  locality:  "Grottos"  at  Las  Cuevas  Beach,  Trinidad, 
depth  not  reported. 

Other  material:  1,  UMML  30.7162;  off  Peninsula  de  la 
Guajira,  Colombia,  John  Elliott  Pillsbury  Station 
P-769,  12°31.0'N,  71°41.0"W,  14.3-146  m;  28  July  1968; 
10-ft  otter  trawl.— 1,  UMML  30.6424;  off  Peninsula  de 
Paria,  Venezuela,  John  Elliott  Pillsbury  Station  P-709, 
11°08.8'N,  62°46.1'W,  46  m;  19  July  1968;  10-ft  otter 
trawl.— 1,  UMML  30.6454;  off  Peninsula  de  Paria,  Ven- 
ezuela, John  Elliott  Pillsbury  Station  P-708, 11°24.7'N, 
62''40.5'W,  69-73  m;  19  July  1968;  10-ft  otter  trawl.— 
1,  UMML  30.6373;  off  Peninsula  de  Paria,  Venezuela, 
John  Elliott  Pillsbury  Station  P-707,  ir21'N,  62°21'W, 
78  m;  19  July  1968;  10-ft  otter  trawl— 1,  UMML  30.6373; 
off  Peninsula  de  Paria,  Venezuela,  John  Elliott 
Pillsbury  Station  P-705,  10°45'N,  62°00'W,  77-86  m;  18 
July  1968;  10-ft  otter  trawl.— 2,  MCZ  273512;  George- 
town, Guyana,  Ch.mn  Cruise  35,  Station  354-36,  8°10.5'- 
8°10.0'N,  57'='48'W,  97-110  m;  28  April  1963—2,  USNM 
866503;  1,  FSBC  I  44072;  1,  UF  189459;  5,  UMML 
30.5840;  off  Pointe  Isere,  French  Guiana,  John  Elliott 
Pillsbury  Station  F'-650,  6°07'N,  52°19'W,  84-91  m;  8 
July  1968;  10-ft  otter  trawl. 


Distribution:  Calliostoma  jernandezi  occurs  from  Su- 
riname  northward  and  westward  to  off  Cabo  de  la  \'ela, 
Colombia,  in  depths  of  about  45-  245  m;  living  specimens 
have  been  collected  from  depths  of  84-146  m. 

Remarks:  Although  the  description  presented  by  Princz 
(1978)  is  incomplete,  the  specimens  recorded  here  seem 
to  be  referrable  to  Calliostoma  jernandezi.  The  shells 
of  this  species  are  very  similar  to  those  of  C.  scalenum 
but  differ  by  having  a  larger  protoconch  (375  ^m  vs. 
320-325  fim);  by  having  fewer,  stronger,  more  evenly 
sized  spiral  cords;  by  having  stronger,  more  closely  spaced, 
and  more  s\  mmetrical  beading;  by  having  an  umbilical 
wall  that  is  almost  vertical  rather  than  sloping  steeply 
inward;  and  by  having  a  different,  lighter  color  pattern. 
The  spiral  cords  at  the  periphery  and  on  the  base  of  shells 
of  C.  jernandezi  tend  to  increase  in  number  by  fission 
of  existing  cords  rather  than  by  intercalation  of  new 
cords;  this  tendency  is  rare  among  species  of  Calliostoma 
that  I  have  examined. 

Data  obtained  from  the  original  field  data  sheets  in- 
dicate that  C.  jernandezi  occurs  on  shell-hash  and  coral- 
rubble  bottoms,  often  in  association  with  sponges  and 
octocorals. 


Calliostoma  scalenum  new  species 
(figures  43,  44,  110,  111) 

Description:  Shell  large  to  ver\-  large,  attaining  height 
of  40.6  mm,  width  of  34.2  mm,  conical,  umbilicate,  finely 
sculptured.  Protoconch  320-325  yum  maximum  diameter, 
1  whorl.  Teleoconch  whorls  about  10,  first  4-5  whorls 
flat-sided,  subsequent  whorls  weakly  to  strongly  concave; 
first  3  whorls  with  low,  rounded  axial  riblets  and  2-6 
finely  beaded  spiral  cords;  axial  sculpture  absent  on  sub- 
sequent whorls;  spiral  cords  increasing  by  intercalation 
to  25-30,  of  which  abapical  8-12  cords  forming  periph- 
ery, alternating  in  size,  finely  beaded,  beads  close-set, 
rounded,  conical;  periphery  narrowly  rounded.  Base  flat 
to  weakly  convex,  with  as  many  as  42  narrow,  weakly 
beaded  spiral  cords,  those  on  abaxial  half  alternating  in 
size,  those  on  adaxial  half  subequal  except  2-3  stronger, 
tuberculate  circumumbilical  cords.  Umbilicus  funnel- 
shaped,  14%-23%  maximum  shell  diameter,  wall  smooth, 
white,  often  with  \  ellow-brown  flush,  .\perture  subquad- 
rate, thickened  within  and  with  strong  ridges,  lips  thin, 
crenulate;  columella  sigmoid,  thickened,  terminating  in 
blunt,  rounded  tubercle.  Shell  ground  color  light  chestnut 
to  red-brown  with  irregular,  diffuse  axial  flammules  of 
darker  ground  color  and  white;  light  and  dark  flammules 
more  distinct  and  closely  spaced  on  periphery,  .\nimal 
(in  alcohol)  white;  foot  with  numerous,  scattered,  small, 
opaque,  white  spots;  cephalic  tentacles  long,  tapered, 
bases  with  short  ocular  peduncles  bearing  small,  black 
eye  at  tips;  epipodium  with  4  pairs  of  tentacles,  neck 
lobes  well-developed,  thin,  semicircular,  smooth-edged. 

Holotype:    USNM  859356,  height  31.5  mm,  width  25.9 


J.  F.  Quinn,  Jr.,  1992 


Page  93 


Type  locality:  Stetson  Bank,  southeast  of  Galveston, 
Texas,  28°09'54"N,  94°18'00"W,  in  21-27  m. 

Paralypes:  2,  HMNS  15022;  2,  USNM  859357;  1,  MCZ 
297051;  1,  ANSP  367152;  1,  AMNH  225980;  1,  LACM 
2292;  1,  UF  110222;  2,  FSBC  I  32315;  34,  HMNS  15850; 
all  from  same  lot  as  holot>pe.— 3,  ANSP  338470;  off 
Freeport,  Texas,  NW  Gulf  Survey  station,  28°10'N, 
94°55W,  51  m;  dredge;  A.  Kight,  collector 

Other  material:  More  than  200  lots  listed  b>  Quinn  (in 
press). 

Distribution:  Calliostoma  scalenum  inhabits  offshore 
waters  on  calcareous  substrates  from  North  Carolina 
southward  through  the  Florida  Keys  and  throughout  the 
Gulf  of  Mexico  in  depths  of  25-80  m. 

Remarks:  Shells  of  Calliostoma  scalenum  have  been 
identified  pre\iously  as  C,  jujuhinum  \\  ithoiit  exception 
(see  Quinn,  in  press,  for  complete  synonymy).  Callios- 
toma scalenum  differs  from  C.  jujuhinum  by  having 
shells  that  are  larger  and  more  narrowly  conical;  that 
have  a  slightK'  larger  protoconch  (320-325  ^m  vs.  315 
jj.m)\  that  lack  fine,  coUabral  threads  after  the  fourth  or 
fifth  whorls;  that  have  two  rather  than  three  distinct  sizes 
of  spiral  cords  with  discrete,  symmetrical  rather  than 
spiralK  elongate  beads;  that  have  a  greater  number  of 
spiral  cords,  particular!}  on  the  base  (35-42  vs.  25-30 
total);  and  that  lack  very  dark-colored  apical  whorls. 
Shells  of  C.  scalenum  are  also  similar  to  those  of  C. 
tampaense  but  differ  b\  having  straight-sided  apical 
whorls  that  lack  a  strong,  sharply  beaded  peripheral  ca- 
rina; by  having  finer,  more  numerous,  and  more  fineK' 
beaded  spiral  cords;  and  b\  being  proportionately  much 
narrower.  Both  C.  jujuhinum  and  C.  tampaense  are  also 
shallow-water  species,  neither  species  occurring  alive  in 
depths  exceeding  11m  (Quinn,  in  press).  The  radula  of 
C.  scalenum  (formula  00.6.I.6.00;  figures  110,  111)  is 
very  similar  to  that  of  C.  tampaense  illustrated  by  Clench 
and  Turner  (1960:pl.  5,  fig.  2,  as  C.  jujuhinum)  and  C. 
vinosum  (figure  114  herein)  in  having  six  pairs  of  lateral 
teeth,  of  which  the  outer  two  pairs  lack  cusps.  Radulae 
of  C.  jujuhinum  (figure  112)  and  C.  cuhense  (figure  1 13) 
are  also  very  similar  in  morphology  but  differ  in  having 
only  five  pairs  of  lateral  teeth.  One  character  common 
to  all  of  these  radulae  is  that  the  cusps  of  the  inner  lateral 
teeth  arise  from  the  outer  corner  of  the  tooth  bases;  this 
characteristic  has  not  been  explicitly  noted  before  and 
is  not  clear  in  anv  of  the  previously  published  line  draw- 
ings (Clench  &  Turner,  1960;pl.  5,'figs.  1,  2;  Calvo,  1987: 
fig.  26). 

Calliostoma  cuhense  new  species 
(figures  45,  46,  113) 

Description:  Shell  medium-sized,  attaining  height  of 
16.6  mm,  width  of  14.3  mm,  conical,  umbilicate,  rather 
coarsely  sculptured.  Protoconch  about  350  ^m  maximum 
diameter,  1  whorl.  Teleoconch  whorls  7-8.  flat  to  weakK 
concave;  first  2-3  whorls  with  low ,  sharp  axial  riblets  and 


2-4  strong,  beaded  spiral  cords;  axial  riblets  replaced  by 
low,  crowded,  rounded  axial  threads  in  interspaces  be- 
tween primar\  spiral  cords;  spiral  cords  increasing  by 
intercalation  to  6-12,  of  which  abapical  1-2  beaded  cords 
and  broad,  smooth  cord  forming  periphery,  usually  al- 
ternating in  size,  primary  cords  strongly  beaded,  sec- 
ondary cords  smooth  or  finely  beaded  by  axial  threads; 
periphery  narrowly  rounded,  smooth  peripheral  cord  of- 
ten dissected  by  1  (rarely  3-5)  fine,  spiral  stria.  Base  flat 
to  weakly  convex,  with  10-13  narrow,  smooth  or  weakl) 
beaded  spiral  cords.  Umbilicus  funnel-shaped,  18%-21% 
maximum  shell  diameter,  w  all  w  hite.  Aperture  subquad- 
rate.  thickened  w  ithin  and  w  ith  strong  ridges,  lips  thin, 
crenulate;  columella  sigmoid,  thickened,  terminating  in 
moderately  strong,  rounded  denticle.  Shell  ground  color 
tan  to  pinkish  brown  with  numerous  white  dots,  prin- 
cipally on  indi\  idual  beads  of  primary  spiral  cords  but 
broader  on  peripher\ ;  interspaces  between  spiral  sculp- 
ture golden  brown,  usually  at  periphery  and  on  base, 
occasionally  above  periphery.  Animal  (reconstituted  in 
trisodium  phosphate)  uniformly  tan;  cephalic  tentacles 
long,  slender,  gradually  tapering;  eyestalks  short  (about 
10%-15%  tentacle  length),  stout,  with  large  black  eye  at 
tips;  snout  longer  than  broad,  with  fringe  of  fine,  short 
papillae;  epipodium  w  ith  3  pairs  of  long  tentacles;  neck 
lobes  well-developed,  left  lobe  finely  digitate,  right  lobe 
smooth-edged. 

Holotype:  ANSP  196933,  height  14.7  mm,  width  13.7 
mm. 

Type  locality:  Off  Pini  Pini,  Cardenas  Bay,  Matanzas, 
Cuba,  in  1 1  m. 

Paratypes:  4,  USNM  438265;  ToM.is  Barrera  Expe- 
dition Station  224,  Cabo  San  Antonio,  Pinar  del  Rio, 
Cuba,— 12,  ANSP  196926;  off  Cardenas  Bay,  Matanzas, 
Cuba,  11  m;  11  February  1954;  V.  Conde.  collector. — 
9,  ANSP  357161;  2.  FSBC  1  32405;  from  same  lot  as 
holotype. — 2.  MCZ  204705;  Peninsula  de  Hicacos,  Ma- 
tanzas, Cuba;  1940;  V.  Conde,  collector.— 4,  MCZ  216120; 
Cayo  Fragoso,  Villa  Clara,  Cuba;  R.  Humes,  collector. — 
1,  MCZ  235536;  off  Cayo  Fragoso,  Villa  Clara,  Cuba;  R. 
T.  Abbott,  collector.— 8,  MCZ  129623;  Cayo  Frances, 
Villa  Clara,  Cuba;  ex  P.  J.  Bermudez  collection. — 3,  ANSP 
216122;  Caibarien,  Villa  Clara,  Cuba;  R.  Humes,  collec- 
tor. 

Distribution:  This  species  seems  to  be  restricted  to  the 
northern  coast  of  Cuba;  live-collected  specimens  were 
obtained  from  off  Pini  Pini  in  11  m. 

Remarks:  Shells  of  Calliostoma  cuhense  most  resemble 
those  of  C.  adelae  in  shape  and  size  and  in  having  rather 
coarse  sculpture  relative  to  other  species  of  the  C.  ju- 
juhinum species  complex.  Shells  of  C.  cuhense  differ 
from  those  of  C  adelae  by  having  more  numerous,  un- 
equally sized  spiral  cords  both  above  and  below  the  pe- 
riphery; b\  having  the  basal  spiral  cords  closeK  spaced, 
separated  b\  narrow.  \'-shaped  grooves,  rather  than 
widely  spaced  and  separated  by  broad,  flat-bottomed 


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/\ 


51 


J,  F,  Quinn,  Jr.,  1992 


Page  95 


grooves;  by  having  distinct,  fine  collabral  threads  be- 
tween the  supraperipheral  spiral  cords;  and  by  having  a 
rather  broad  and  smooth  rather  tlian  narrow  and  strongly 
beaded  peripheral  cord.  The  shells  of  C  cuhense  from 
the  ToM.4s  Barrera  Expedition  (USNM  438265)  are  ap- 
parently those  that  Henderson  (1916:185)  recognized  as 
"an  apparently  new  Calliostoma"  from  dredgings  off 
Cabo  Cajon  (near  Cabo  San  Antonio)  on  a  bottom  of 
coral  sand  with  patches  of  seagrass  and  sponge,  the  area 
swept  by  strong  currents. 

The  intestine  of  one  specimen  [ex  FSBC  1  32405)  was 
packed  with  amorphous  organic  material,  hydroid  stalks, 
and  several  different  forms  of  foraminiferans. 

Calliostoma  ptirpurciim  new  species 
(figures  47-30) 

Description:  Shell  medium-sized,  attaining  height  of 
23.9  mm,  width  of  20.2  mm,  conical,  umbilicate,  rather 
coarsely  sculptured.  Protoconch  unknown.  Teleoconch 
whorls  about  9,  flat  to  weakK  convex;  first  1-2  whorls 
worn  on  ail  specimens  but  apparently  with  axial  riblets 
and  2-4  spiral  cords;  axial  sculpture  absent  on  subsequent 
whorls;  spiral  sculpture  increasing  by  intercalation  to  8- 

14  cords  and  1-8  fine  threads,  of  which  abapical  3-5 
cords  forming  periphery;  peripher\  broadly  rounded. 
Base  flat  to  weakly  convex,  with  13-18  coarsely  beaded 
spiral  cords.  Umbilicus  funnel-shaped,  18%-26/o  maxi- 
mum shell  diameter,  wall  white  to  intense  violet.  Ap- 
erture subquadrate,  thickened  within  and  with  strong, 
sharp  ridges,  lips  thin,  crenulate;  columella  weakly  sig- 
moid, thickened,  often  with  1-3  small,  sharp  denticles 
or  ridges  on  lower  third,  terminating  in  strong,  rounded 
tubercle.  Shell  ground  color  tan  to  reddish  brown,  mot- 
tled with  cream  to  yellowish  brown. 

Holotype:  USNM  860247,  height  21.1  mm,  width  19.6 
mm. 

Type  locality:  Off  Cabo  de  la  Vela,  Colombia,  in  61- 
91  m. 

Paratypes:  3,  USNM  860248;  1,  FSBC  I  44074;  from 
same  lot  as  holotype. 

Other  material:  5,  Deynzer  collection;  from  same  lot 
as  holotype.— 1,  UMML  30.7204;  off  Cabo  de  la  Vela, 
Colombia,  John  Elliott  Pillsbury  Station  P-773, 
12°17,0'N,  72°15.0'W,  60-64  m;  29  July  1968;  10-ft  otter 
trawl— 3  broken,  FSBC  I  32742;  about  5  km  north  of 
La  Guardia,  Isia  de  Margarita,  Venezuela,  beach  drift; 
19  November  1987;  W.  G.  &  C.  B.  Lyons,  collectors.— 

15  broken,  FSBC  I  32747;  6  km  north  of  La  Guardia, 


Isla  de  Margarita,  Venezuela,  beach  drift;  18  November 
1987;  W.  G.  &  C.  B.  Lyons,  collectors.  — 1,  UMML 
30.7269;  off  Galera  Point,  Trinidad,  John  Elliott 
Pillsbury  Station  P-840,  10°40.5'N,  60°37.5'W,  33-37 
m;  1  July  1969;  10-ft  otter  trawl. 

Distribution:  Calliostoma  purpiireum  is  known  from 
off  northeastern  Colombia,  Isla  de  Margarita,  Venezuela, 
and  Trinidad;  most  specimens  have  been  collected  from 
33-91  m,  but  broken  shells  are  also  known  from  beach 
drift. 

Remarks:  Shells  of  Calliostoma  purpiireum  are  very 
similar  to  those  of  C.  jujubinum  but  differ  by  lacking 
collabral  threads  between  the  spiral  cords,  by  having  the 
spiral  cords  narrower  and  more  finely  beaded,  and  by 
having  the  umbilical  wall  almost  vertical  rather  than 
strongK  sloped.  Most  specimens  of  C  purpureum  have 
one  or  more  small,  sharp  denticles  on  the  columella, 
whereas  specimens  of  C.  jujubinum  very  rarely  have 
only  a  single  denticle;  I  know  of  no  other  species  that 
has  a  denticulate  columella.  Additionally,  whereas  spec- 
imens of  C.  jujubinum  may  have  the  umbilical  wall 
flushed  with  pinkish  brown,  I  have  never  seen  any  spec- 
imen of  that  or  any  other  species  of  western  Atlantic 
Calliostoma  with  the  violet  color  or  the  intensity  of  any 
other  color  displayed  on  the  umbilical  wall  of  specimens 
of  C.  purpureum  except  the  red-brown  in  C.  brunneum 
and  C.  barbouri.  Shells  of  C.  purpureum  may  have  been 
reported  from  Curasao  by  de  Jong  and  Coomans  (1988) 
as  C,  tampaense. 

Almost  all  of  the  specimens  collected  at  Isla  de  Mar- 
garita consisted  of  only  the  last  one  to  three  whorls; 
because  the  damage  to  these  shells  is  almost  identical  to 
that  illustrated  by  Vermeij  (1978:  fig.  2.10),  this  damage 
may  have  been  the  result  of  predation  by  xanthid  crabs. 
This  species  has  been  collected  principally  in  areas  with 
large  concentrations  of  sponges  and  was  probably  the 
species  from  Islas  Los  Roques,  Venezuela,  that  Work 
(1969)  observed  eating  an  encrusting  sponge. 

Calliostoma  fiicosum  new  species 

(figures  51,  52) 

Description:  Shell  medium-sized,  attaining  height  of 
12.8  mm,  width  of  12.6  mm,  conicoturbinate,  umbilicate, 
rather  coarseK  sculptured.  Protoconch  eroded.  Teleo- 
conch whorls  about  7,  weakly  concave;  first  whorl  eroded, 
next  whorl  with  3  strong,  strongly  beaded  spiral  cords; 
spiral  cords  increasing  by  intercalation  to  7,  of  which 
abapical  2  cords  forming  periphery;  periphery  broadly 
rounded,  adapical  peripheral  cord  beaded,  abapical  pe- 


Figures  41,  42.  Calliostoma  jernandezi  Princz,  1978,  specimen  from  John  Elliott  Pillsbury  Station  P-650,  UMML  30.5840; 
height  21.5  mm,  width  19,;3  mm.  Figures  43,  44.  Calliostoma  scalenum  new  species,  holotype  from  Stetson  Bank,  SE  of  Galveston, 
Texas,  USNM  859356;  height  31.5  mm,  width  25.9  mm.  Figures  45,  46.  Calliostoma  cubense  new  species,  holotype  from  Pini 
Pini,  Cuba,  ANSP  196933;  height  14.7  mm,  width  13.7  mm.  Figures  47-50.  Calliostoma  purpureum  new  species.  47, 48.  Holotype, 
USNM  860247;  height  21.1  mm,  width  19.6  mm.  49,  50.  Paratype,  USNM  860248;  height  15.8  mm,  width  15.4  mm.  Figures  51^ 
52.  Calliostoma  fucosum  new  species,  holotype,  USNM  860259;  height  12.8  mm,  width  12.6  mm. 


Page  96 


THE  NAUTILUS,  Vol.  106,  No.  3 


ripheral  cord  smooth  with  shallow  medial  stria.  Base 
weakK  convex,  with  9  broad,  coarseK  beaded  and  2 
narrow,  smooth  spiral  cords;  circumumbilical  cord 
strongest  and  most  strongly  beaded.  Umbilicus  20%  max- 
imum shell  diameter,  wall  white  with  faint  flush  of  pink. 
Aperture  subquadrate,  lips  thin,  weakK  crenulate;  col- 
umella weakly  sigmoid,  somewhat  thickened,  terminat- 
ing in  sharp  denticle.  Shell  ground  color  predominantly 
orange-brown  with  spots  and  flammules  of  white  and 
darker  orange-brown. 

Holotype:  USNM  860259,  height  12.8  mm,  width  12.6 
mm. 

Type  locality:  Off  Cabo  de  la  Vela,  Colombia,  John 
Elliott  Pillsbury  Station  P-774,  11°56.5'N,  72°17.9'W, 
in  5-9  m. 

Distribution:  This  species  is  onl\  known  from  the  ho- 
lot\pe  shell. 

Remarks:  The  holotype  of  Calliostoma  fucosiim  resem- 
bles a  small  C.  euglypfum,  but  the  shell  is  umbilicate, 
proportionately  broader,  has  stronger  and  coarser  sculp- 
ture, and  has  a  much  brighter  color  pattern. 

The  specimen  of  C.  fucosum  was  trawled  in  shallow- 
water  from  a  bottom  composed  principally  of  calcareous 
red  algae  (UMML,  unpublished). 

Calliostoma  vinosiim  new  species 
(figures  53-56,  114) 

Calliostoma  (Elmerlinia)  Inillisi:  Rios,   1970:24,  pi.  5.   (Non 

Calliostoma  bidlisi  Clench  &  Turner,  1960). 
Calliostoma  barbouri.  Rios,  197.5:23,  pi. 5,  fig.  60;  1985:22,  pi. 

10.  fig.  87.  (Non  Calliostoma  barbouri  Clench  &  Aguavo, 

1946). 
Calliostoma  javanicum:  Leal,  1991:45, 353  (partim).  (Non  Tro- 

cliiis  javanicum  Lamarck,  1822), 

Description:  Shell  medium-sized,  attaining  height  of 
19.8  mm,  width  of  21.3  mm,  conical,  umbilicate,  finely 
sculptured.  Protoconch  325-350  /ttm  maximum  diameter, 
1  whorl.  Teleoconch  whorls  8,  flat  to  weakly  convex;  first 
whorl  initially  with  2  strong  spiral  cords  and  low,  round- 
ed axial  riblets;  axial  sculpture  absent  after  first  one-third 
whorl;  spiral  sculpture  increasing  by  intercalation  to  9- 
15  beaded  cords  and  0-6  smooth  to  finely  beaded  threads, 
of  which  abapical  2  cords  forming  periphery,  beads 
rounded;  surface  of  first  4-5  whorls  microscopically  frost- 
ed; periphery  narrow,  subcarinate,  abapical  peripheral 
cord  often  with  shallow,  median  groove.  Base  flat  to 
weakly  concave,  with  12-14  narrow  ,  finely  beaded  spiral 
cords  and  0-3  spiral  threads.  Umbilicus  funnel-shaped, 
18%-22%  maximum  shell  diameter,  wall  smooth,  white 
or  flushed  with  yellow.  Aperture  subquadrate,  lips  thin, 
crenulate;  columella  weakly  sigmoid,  weakly  thickened, 
terminating  in  sharp  denticle.  Shell  ground  color  deep 
red-brown  with  crescentic  streaks  of  yellow-brown  above 
periphery;  base  golden  brown  with  6-9  spiral  lines  of 
red-brown.  Animal  (in  alcohol):  foot  red-brf)w  ti  w  ith  large, 
white  papillae;  mantle  with  narrow  band  of  white  and 


red-brown  spots  at  edge  behind  which  is  series  of  dark 
brown  spots  extending  posteriorly  as  gradually  fading 
streaks;  cephalic  tentacles  red-brown  with  darker  median 
stripe,  white  near  base,  left  tentacle  stout,  about  length 
of  snout,  right  tentacle  longer  and  more  slender;  snout 
long,  broad,  mottled  red-brown  and  w  hite,  with  anterior 
fringe  of  long  papillae;  epipodium  with  4  pairs  of  red- 
brown  tentacles,  neck  lobes  well-developed,  semicircu- 
lar, left  lobe  smooth,  right  lobe  finely  fringed. 

Holotype:  MORG  29.294,  height  13.5  mm,  width  15.3 
mm. 

Type  locality:  Off  Guarapari,  Estado  de  Espirito  Santo, 
Brazil,  in  18-22  m. 

Paratypes:  1,  USNM  860256;  1,  FSBC  I  44067;  both 
from  same  lot  as  holotype.  — 1,  MORG  15.043;  off  Cabo 
Orange,  Estado  de  Amapa,  Brazil,  .Almir.^nte  Saldanh.\ 
Station  2029,  103  m;  30  November  1968. 

Other  material:  1,  MNHN  uncatalogued; 
Marion-Dufresne  Cruise  MD-55,  Station  DC-40, 
20°40'S,  34°41'W,  60  m;  17  May  1987;  dredge— 2, 
MNHN  uncatalogued;  Marion-Dufresne  Cruise  MD- 
55,  Station  DC-47,  20°42'S,  32°13'W,  94-105  m;  19  May 
1987;  dredge. — 2,  MNHN  uncatalogued;  Marion- 
Dufresne  Cruise  MD-55,  Station  DC-42,  20°55'S, 
34°01"W,  60  m;  17  May  1987;  dredge —7,  MNHN  un- 
catalogued; Marion-Dufresne  Cruise  MD-55,  Station 
DC-15,  2r37'S,  40°18'W,  37  m;  11  May  1987;  dredge. 

Distribution:  Calliostoma  vinosttm  is  known  in  north- 
eastern Brazil  from  the  Amazon  River  to  just  north  of 
Rio  de  Janeiro,  in  18-105  m. 

Remarks:  Shells  of  Calliostoma  vinosum  are  most  sim- 
ilar to  those  of  C.  barbouri  in  shape,  size,  and  color  but 
differ  b\  having  a  somewhat  smaller  protoconch  (325- 
350  ^m  vs.  350-400  nm);  b\  having  flat-sided  rather  than 
convex  whorls;  by  having  fewer,  stronger,  more  coarsely 
beaded  spiral  cords,  particularly  on  the  base;  by  having 
axial  riblets  that  disappear  early  on  the  first  whorl  rather 
than  persisting  onto  the  second  whorl;  b\  lacking  col- 
labral  threads  between  the  primary  spiral  cords;  and  by 
having  the  first  4-5  whorls  more  distincth  frosted. 

Most  of  the  specimens  examined  for  this  study  were 
trawled  from  bottoms  on  which  bryozoans  and  coralline 
algae  were  abundant  (J.  &  M.  Coltro,  personal  com- 
munication; Leal,  1991). 


Calliostoma  alternum  new  species 
(figures  57,  58) 

Description:  Shell  medium-sized,  attaining  height  of 
15.6  mm,  width  of  13.9  mm,  conical,  umbilicate,  fineK 
sculptured.  Protoconch  about  350  /um  maximum  diam- 
eter, 1  whorl.  Teleoconch  whorls  about  8.5,  flat;  first  4 
whorls  with  low,  rounded  axial  riblets  and  2-7  beaded 
spiral  cords;  axial  sculpture  reduced  to  fine  collabral 
threads  or  absent  on  subsequent  whorls;  spiral  sculpture 


J.  F.  Quinn,  Jr.,  1992 


Page  97 


increasing  by  intercalation  to  7-11  beaded  cords  and  5- 
6  threads,  of  which  abapical  3  cords  forming  periphery; 
periphery  narrowly  rounded.  Base  weakly  convex,  with 
17  narrow,  finely  beaded  spiral  cords.  Umbilicus  22%- 
24%  maximum  shell  diameter,  wall  almost  vertical,  white. 
Aperture  subquadrate,  lips  thin,  weakly  crenulate;  col- 
umella weakly  sigmoid,  rather  thin,  terminating  in  small, 
rounded  denticle.  Shell  ground  color  uniformK  ivory  or 
v\ ith  \erv  faint  peripheral  light  brow n  maculations. 

Holotype:  USNM  860257,  height  15.6  mm,  width  13.9 
mm. 

Type  locality:  Northeast  of  Caracas,  Venezuela,  John 
Elliott  Pillsbury  Station  P-737,  10°44.0'N,  66°07.0'W, 
in  60-73  m. 

Paratypes:  1,  UMML  30.8374;  off  Peninsula  de  la  Gua- 
jira,  Colombia,  John  Elliott  Pillsbi  ry  Station  P-769, 
12°31.0'N,  71°41.0'\V,  143-146  ni;  28  Julv  1968;  10-ft 
otter  trawl. — 1,  UN4ML  30.6155;  off  Suriname,  John  El- 
liott Pillsbury  Station  P-684,  7°19.0'N,  56°51.0'W,  55- 
59  m;  14  July  1968;  10-ft  otter  trawl. 

Distribution:  Shells  of  this  species  are  known  from 
northeastern  Colombia,  Venezuela,  and  Suriname,  in  55- 
146  m. 

Remarks:  Shells  of  Calliostoma  alternitm  are  most  sim- 
ilar to  those  of  C.  aulicum  but  differ  b\  being  evenly 
conical  rather  than  coeloconoid;  by  having  a  more  round- 
ed, less  strongly  projecting  periphery;  by  having  fewer, 
stronger,  more  coarsely  beaded  spiral  cords,  most  of  which 
are  separated  by  a  fine  spiral  thread;  by  having  axial 
riblets  that  persist  on  the  first  four  whorls  rather  than 
only  on  the  first  two;  and  by  having  the  abapical  of  the 
two  primary  spiral  cords  remaining  conspicuously  strong 
on  first  five  whorls. 

The  specimens  of  C.  altcrnum  were  collected  on  bot- 
toms composed  of  carbonate  rock  covered  by  calcareous 
algae  and  sponges  (UMML,  unpublished). 


Calliostoma  aulicum  nev\  species 
(figures  59-62) 

Description:  Shell  medium-sized,  attaining  height  of 
15.9  mm,  width  of  16.1  mm,  conical,  umbilicate,  finely 
sculptured.  Protoconch  about  350  m  maximum  diameter, 
1  whorl.  Teleoconch  whorls  7.3,  weakly  concave;  first 
two  whorls  with  low,  rounded  axial  riblets  and  2-3  bead- 
ed spiral  cords;  axial  sculpture  absent  after  second  whorl; 
spiral  cords  strongly  beaded,  increasing  by  intercalation 
to  10-12,  of  which  abapical  3  strongest  cords  forming 
periphery  on  last  whorl,  beads  rounded;  periphery  nar- 
rowly rounded.  Base  flat  to  weakly  convex,  with  17-19 
narrow,  finely  beaded  spiral  cords.  I'mbilicus  funnel- 
shaped,  17%-20%  maximum  shell  diameter,  wall  white. 
Aperture  subquadrate,  thickened  and  stronglv  ridged 
within,  lips  thin,  crenulate;  columella  weakly  sigmoid, 
thickened,  terminating  in  narrow  denticle.  Shell  ground 
color  ivory  with  few,  faint  patches  of  golden  brown 


Holotype:  USNM  860258,  height  15.9  mm,  width  16.1 
mm. 

Type  locality:  Off  Colon,  Panama,  John  Elliott 
Pillsbury  Station  P-451,  9°22.0'N,  79°56.0'W,  in  ap- 
proximately 12  m. 

Paratype:  1,  UMML  30.8375;  off  Suriname,  John  El- 
liott Pillsbury  Station  P-669,  6°39.0'N,  55°15.5'W,  33 
m;  10  July  1968;  10-ft  otter  trawl. 

Other  material:  1  fragment,  UMML  30.7238;  off  Ori- 
noco River,  Venezuela,  John  Elliott  Pillsbury  Station 
P-835,  9°36.0'N,  60°10.0"W,  48  m;  30  June  1968;  10-ft 
otter  trawl.— 1  fragment,  UMML  30.7306;  off  Galera 
Point,  Trinidad,  John  Elliott  Pillsbury  Station  P-840, 
10°40.5'N,  60°37.5'W,  33-37  m;  1  July  1968;  10-ft  otter 
trawl. 

Distribution:  Calliostoma  aulicum  is  known  from  off 
Panama  and  eastern  Venezuela  in  12-48  m. 

Remarks:  Shells  of  this  species  are  very  similar  to  those 
of  Calliostoma  tampaense  but  differ  by  having  a  slightly 
larger  protoconch  (350  nm  vs.  325  nm);  bv'  having  a 
more  broadlv'  rounded,  more  strongly  projecting  periph- 
ery; by  being  much  thicker;  by  having  more  strongly 
beaded  basal  spiral  cords;  and  by  lacking  a  distinct  color 
pattern.  If  the  holotype  of  C.  aulicum  represents  the  size 
of  a  fully  grown  adult,  as  the  thickened  and  ridged  ap- 
erture seems  to  indicate,  then  C.  aulicum  is  much  smaller 
than  C.  tampaense,  shells  of  which  attain  a  height  of 
about  30  mm  (Quinn,  in  press). 


Calliostoma  hirtum  new  species 
(figures  63,  64) 

Description:  Shell  medium-sized,  attaining  height  of 
17.9  mm,  width  of  19.8  mm,  conical,  umbilicate,  finely 
sculptured.  Protoconch  missing.  Teleoconch  whorls  about 
7,  flat  to  weakly  concave;  first  remaining  v\horl  worn 
smooth;  next  whorl  with  4  beaded  spiral  cords,  cords 
increasing  to  13,  of  which  abapical  2  cords  forming  pe- 
riphery on  last  whorl,  beads  conical;  periphery  carinate, 
adapical  peripheral  cord  stronger  than  abapical  periph- 
eral cord.  Base  weakly  concave  to  weakly  convex,  with 
17-18  narrow,  weakly  beaded  spiral  cords.  Umbilicus 
funnel-shaped,  20%  maximum  shell  diameter,  wall  al- 
most vertical,  white  or  yellow-brov\n.  Aperture  subquad- 
rate, lips  thin,  crenulate;  columella  weakly  sigmoid,  thin. 
Shell  ground  color  light  yellow-brown  with  diffuse  flam- 
mules  of  cream  and  light  brown;  interspaces  between 
several  spiral  cords  golden  brown. 

Holotype:  USNM  860260,  height  17.9  mm,  width  19.8 
mm. 

Type  locality:  Off  Navidad  Bank,  northeast  of  the  Do- 
minican Republic,  John  Elliott  Pillsbury  Station  P- 

1160,  20°01'N,  68°51'W,  in  521  m. 

Paratype:    1,  UMML  30.8371;  from  same  lot  as  holotype. 


Page  98 


THE  NAUTILUS,  Vol.  106,  No.  3 


53 


J.  F.  Quinn,  Jr.,  1992 


Page  99 


V^    ) 


-;7 


Figures  63,  66.    Calliostonia  atlantis  C\euch  &  Aguayo,  1940,   holotype,  MCZ  135164;  height  32.7  mm,  width  33.6  mm. 


Distribution:  This  species  is  known  only  from  the  type 
locality. 

Remarks:  The  shells  of  Calliostoma  hirtnni  are  most 
similar  to  those  of  C.  javanicum  but  differ  by  having  a 
stronger,  more  sharply  beaded  peripheral  cord;  by  lack- 
ing axial  sculpture  in  the  interspaces  between  the  spiral 
cords;  and  by  having  more  numerous,  sharper  spiral  cords, 
particularly  on  the  outer  part  of  the  base. 

Calliostoma  atlantis  Clench  &  Aguayo,  1940 
(figures  65,  66) 

Calliostoma  (Calliostoma)  atlantis  Clench  &  Aguavo,  1940: 

81,  pi.  13,  fig.  4. 
Calliostoma  atlantis:  Clench  &  Turner,  1960:62,  pi.  44;  Har- 

asewych,  1989:27,  pi.  17. 
Calliostoma  (Kombologion)  atlantis:  Abbott,  1974:4.5. 

Description:  See  Clench  and  Aguayo  (1940:81)  and 
Clench  and  Turner  (1960:62). 

Holotype:  MCZ  135164,  height  32.7  mm,  width  33.6 
mm. 

Type  locality:  Northwest  of  Mariel,  Cuba,  Atlantis 
Station  3306,  23°04'30"N,  82°37'00"W,  in  604  m. 

Other  material:  1,  USNM  869000;  off  Great  Inagua 
Island,  Bahama  Islands,  Johnson-Sea-Link  I  Station  JSL- 
1-2323,  21°01'45"N,  74°43'48"W,  628  m;  15  October  1988. 

Distribution:  The  specimen  of  this  species  reported  here 
is  only  the  second  collected  and  extends  the  range  from 


northwestern  Cuba  to  off  Great  Inagua  Island  northeast 
of  the  eastern  tip  of  Cuba.  This  specimen  was  collected 
by  M.  G.  Harasewych,  using  the  submersible 
Johnson-Sea-Link  I  from  a  large  boulder  in  628  m,  a 
depth  very  similar  to  that  from  which  the  holotype  was 
collected  (604  ni).  The  animal  was  reported  to  be  bright 
red  (Harasewych,  1989:27). 

Remarks:  Based  on  shell  characters,  Calliostoma  at- 
lantis, C.  torrei,  C.  dnopherum,  and  the  new  species  C. 
rugosum,  C.  atlantoides,  C  rota,  and  C.  coronatum 
form  a  distinctive  species  group.  All  seven  species  have 
shells  w  ith  strongly  bicarinate  whorls,  at  least  on  the  first 
several  whorls;  have,  at  the  beginning  of  the  first  teleo- 
conch  whorl,  an  adapical  spiral  cord  that  becomes  weak 
or  disappears  on  subsequent  whorls;  have  a  weak  an- 
gulation or  strong  spiral  cord  below  the  peripheral  cord; 
have  the  inner  lip  of  the  columella  weakly  or  strongly 
reflected  over  the  umbilicus,  or  thickened  and  filling  the 
umbilicus  with  callus;  and  have  the  columella  rounding 
into  the  outer  lip.  Other  species  that  may  belong  to  this 
group  include  C.  cuhanum  Clench  &  Aguayo,  1940,  and 
the  eastern  Atlantic  C.  grimaldii  Dautzenberg  &  Fischer, 
1896;  C.  leptophyma  Dautzenberg,  1927;  C.  normani 
Dautzenberg,  1927;  and  C.  caroli  Dautzenberg,  1927. 

Calliostoma  dnopherum  (Watson,  1879), 
new  combination 
(figures  67-69) 

Trochus  (Margarita)  dnopherns  Watson,   1879:711;  1886:90, 
pi.  5,  fig.  3. 


Figures  53-56.  Calliostoma  vinosum  new  species.  53,  54.  Holotype,  MORG  29.294;  height  13.5  mm,  width  15.3  mm.  55,  56. 
Paratype,  \40RG  15  043;  height  10.2  mm,  width  11.9  mm.  Figures  57,  58.  Calliostoma  alternum  new  species,  hoiotvpe,  USNM 
860257;  height  15.6  mm,  width  13,9  mm.  Figures  59-62.  Calliostoma  auliciun  new  species,  55,  56.  Hoiotvpe,  USNM  860258; 
height  15,9  mm,  width  16,1  mm  57,  58.  Parat\pe  from  John  Elliott  Pillsbiry  Station  P-669,  L'MML  30,8375;  height  10.8  mm, 
width  11.2  mm.  Figures  63,  64.  Calliostoma  hirtiim  new  species,  holotype,  USNM  860260;  height  17,9  mm,  width  19.8  mm. 


Page  100 


THE  NAUTILUS,  Vol.  106,  No.  3 


67 

Figures  67-69.    Calliostoma  dnopherum  (Watson,  1879).  Lectotype  of  Trochus  {Margarita)  dnophcrus.  BM(NH)  1887.2.9.333; 
height  7.2  mm.  width  8.8  mm. 


Margarites  dnopherus:  Lange  de  Morretes,  1949:58. 
"Margarites"  dnopherus:  Rios,  198.5:19,  pi.  8,  fig.  72. 

Description:    See  Watson  (1879:711;  1886:90). 

Lectotype  (here  selected):  BM(NH)  1887.2.9.333,  height 
7.2  mm,  width  8.8  mm. 

Type  locality:  Southeast  of  Recife,  Brazil,  Challenger 
Station  122,  9''05'S,  34°49'W,  in  640  m. 

Paralectotypes:  4  (broken  or  fragments),  BM(NH) 
1887.2.9.334-335;  from  same  lot  as  holotype. 

Other  material:  1,  Coltro  collection;  off  Ihla  de  Sao 
Sebastiao,  Estado  de  Sao  Paulo,  Brazil,  600  m;  dredge; 
1991. 

Distribution:  This  species  is  now  known  from  off  Recife 
to  off  Sao  Paulo,  Brazil  (a  range  extension  of  appro.xi- 
mately  2000  km),  in  600-640  m. 

Remarks:  The  shape  and  sculpture  of  the  shells  of  this 
species  are  very  similar  to  those  of  C  rota,  C.  atlan- 
toides,  and  C.  coronatum  {q.  v.);  therefore,  the  species 
is  here  transferred  to  Calliostoma  sensu  lata.  Shells  of 
C.  dnopherum  differ  from  those  of  C.  rota  by  being 
somewhat  higher  and  more  globose,  by  having  a  larger 
protoconch  (825-850  ^m  vs.  400-425  /im),  by  having  a 
strong  spiral  cord  midway  between  the  shoulder  spiral 
cord  and  the  circumbasal  cord,  by  liavirig  a  strong  spiral 
cord  just  beneath  the  circumbasal  cord,  and  by  having 
an  open  umbilicus.  Differences  between  C.  dnopherum 
and  C.  atlantoides  and  C.  coronatum.  are  discussed  in 
the  Remarks  in  the  species  accounts  of  the  latter  two 
species. 

The  syntype  lot  originally  contained  five  specimens, 
one  large,  live-collected  specimen  and  four  very  small 
shells.  The  large  shell  is  in  excellent  condition,  although 
the  animal  was  removed  at  some  point  and  the  oper- 
culum glued  to  cotton  in  the  aperture.  Of  the  four  small 
shells,  one  has  completely  disintegrated,  and  the  other 
three  are  in  various  stages  of  disintegration. 

The  recently  collected  specimen  (Cloltro  collection)  is 
very  similar  to  the  lectotype  but  is  slightly  .smaller  (7.0 
mm  height,  8.5  mm  width),  lacks  fine  threads  between 
the  supraijeripheral  spiral  cords,  has  one  fewer  basal 


spiral  cord,  and  has  the  inner  lip  of  the  columella  rather 
strongly  expanded  to  almost  cover  the  umbilicus. 

Calliostoma  rota  new  species 
(hgures  70-73,  117,  118) 

Description:  Shell  small,  attaining  height  of  8.1  mm, 
width  of  9.9  mm,  depressed  turbinate,  nonumbilicate. 
Protoconch  400-425  ^m  maximum  diameter,  1  whorl. 
Teleoconch  whorls  5.5,  rapidly  expanding,  carinate;  first 
whorl  initially  with  2  smooth  spiral  cords;  adapical  cord 
strong  on  first  2  whorls,  gradually  weakening  to  fine 
thread  on  next  3  whorls,  finely  undulate  to  beaded  on 
last  3  whorls;  abapical  cord  strong,  smooth  to  weakly 
undulate  on  all  whorls,  forming  carinate  whorl  shoulder; 
subsutural  cord  appearing  on  first  quarter-whorl,  becom- 
ing strong,  finely  beaded  by  third  whorl;  last  whorl  u  ith 
strong,  smooth,  circumbasal  spiral  cord  forming  periph- 
eral carina;  interspaces  between  spiral  cords  concave  with 
fine,  smooth  or  finely  beaded  spiral  threads  (last  whorl 
with  11-12  between  subsutural  and  shoulder  cords.  8- 
14  between  shoulder  and  peripheral  cords);  axial  sculp- 
ture absent  except  fine  growth  lines.  Base  almost  flat, 
with  17-21  weak  to  strong,  flat,  smooth  spiral  cords  and 
0-4  fine  threads  in  interspace  between  peripheral  cord 
and  outermost  ba.sal  cord;  outermost  cord  sometimes 
torniing  very  weak  subperipheral  angulation.  Umbilicus 
tilled  with  callus.  Aperture  subquadrate;  outer  lip  thin, 
angulate;  inner  lip  thickened,  forming  umbilical  plug; 
columella  rather  short,  concave  in  upper  third,  straight 
l)elov\ .  Shell  ground  color  ivory  to  |)inkisli  tan  with  green 
and  pink  iridescence,  with  regularly  spaced  spots  of  light 
to  dark  yellow-brown  on  subsutural,  shoulder,  periph- 
eral, and  1-2  inner  basal  cords,  basal  cords  sometimes 
lacking  spots. 

iloloiypc:  MOiU;  29.295,  height  8.1  nun,  width  9.3 
mm. 

Type  loraliiy:  Off  illuibcki,  illui  de  Sao  Sebastiao,  Es- 
tado (Ic  S;h)  I';ui1(i,  Hr;i/ii,  in  20  30  ni, 

Paratype:  I,  MCZ  258057;  oil  Hio  de  J.ineiro,  Estado 
de  Rio  de  Janeiro,  Brazil,  from  stom;uli  ol  st;irfish,  in 
ai)oul   16  m;  Bernard  'I'ursch  colleetor. 


J.  F.  Quiim.  Jr  .  1992 


^^ 


Figures  70-73.  Calliostoma  rota  new  species  70.  71.  Holotr>^.  MORG  29.295;  height  S.l  mm.  width  9.3  mm.  72,  73.  Parat>^, 
MCZ  258057;  height  7.6  mm.  width  9  9  mm.  Figures  74.  75.  Calliostoma  atlantoides  new  species,  holot\pe,  USNM  S60261;  height 
9.1  mm,  width  10  0  mm  Figures  76.  77.  Calliostoma  cvronatum  new  species,  holotype,  MC2  2T-1.56S;  height  4.2  mm.  width 
4.2+  mm 


Other  material:  1,  Coitro  collection;  from  same  lot  as 
holot\-pe. 

Distribution:  This  species  is  known  only  from  off  Rio  de 
Janeiro  and  Sao  Paulo.  Brazil,  in  20-46  m;  the  single 
li\lng  specimen  was  from  46  m. 

Remark*:  Shells  of  Calliostoma  rota  are  most  similar 
to  those  of  C.  dnopherum  but  differ  by  ha\ing  more 
whorls,  b>  ha\  ing  a  much  smaller  protoconch,  by  lacking 
a  strong  spiral  cord  between  the  shoulder  and  circum- 


basal  cords,  by  lacking  a  strong  subperipheral  spiral  an- 
gulation, and  by  having  the  umbiliciis  filled  with  callus. 
The  radula  of  C.  rota  i^formula  12-13.7.1.7.12-13;  figures 
117,  lis)  has  bvo  principal  characters  that  depart  from 
the  typical  calliostoma  tine  morphology:  1"  the  reduced 
number  of  elements  in  the  marginal  tooth  field;  and  2^ 
the  single,  imcusf)ed  tooth  base  representing  the  outer 
lateral  tooth.  The  number  of  pairs  of  calliostomatine 
marginal  teeth  is  rarely  reported,  but  in  those  instances 
where  the  number  of  pairs  is  known  ^e.g.,  Calvo,  1987). 


Page  102 


THE  NAUTILUS,  Vol.  106,  No.  3 


the  number  is  25  or  more  except  in  Astele  Swainson, 
1855,  the  type  species  of  which  has  about  20  pairs  (Clench 
&  Turner,  1960:76);  however,  in  C.  rota  there  are  only 
12  or  13  pairs.  The  presence  of  a  single,  uncusped  outer 
lateral  tooth  has  not  been  reported  pre\iously  in  the 
Calliostomatinae,  although  this  condition  bridges  the  gap 
between  the  majority  of  species  that  have  all  lateral  teeth 
cusped  and  those  in  the  C.  jujubimim  species  group  that 
have  the  outer  two  pairs  of  lateral  teeth  uncusped.  How- 
ever, although  the  anterior  two-thirds  of  the  radula  ex- 
amined here  had  the  uncusped  lateral  tooth,  a  few  teeth 
from  the  posterior  third  retained  a  membranaceous,  ru- 
dimentary cusp.  The  holotype  and  Coltro  specimen  were 
dredged  from  muddy  sand  bottom. 

Calliostoma  atlantoides  new  species 
(6gures74,  75,  115,  116) 

Description:  Shell  small,  attaining  height  of  9.1  mm, 
width  of  10.0  mm,  turbinate,  narrowK  umbilicate.  Pro- 
toconch  1.10  mm  maximum  diameter,  1  whorl.  Teleo- 
conch  whorls  3.4,  rapidly  expanding,  carinate;  first  whorl 
initially  with  2  smooth,  spiral  cords,  adapical  cord  dis- 
appearing b\  end  of  first  whorl,  abapical  cord  strength- 
ening and  forming  strong  midwhorl  carination  on  sub- 
sequent whorls;  third  spiral  cord  appearing  between 
adapical  cord  and  suture  near  beginning  of  first  whorl, 
strengthening  and  forming  rather  strong  subsutural  an- 
gulation; both  subsutural  and  midwhorl  angulations  set 
with  narrow,  rounded,  spirally  elongate  beads;  last  whorl 
with  strong,  smooth  spiral  cord  forming  periphery;  in- 
terspaces between  spiral  angulations  weakly  concave,  with 
fine,  smooth  spiral  threads  (last  whorl  with  2  between 
suture  and  subsutural  angulation,  16  between  subsutural 
and  midwhorl  angulations,  and  12  between  midwhorl 
and  peripheral  angulations);  axial  sculpture  of  low, 
rounded  riblets  restricted  to  first  1.25  whorls.  Base  di- 
vided into  narrow,  weakly  concave  peripheral  zone  and 
broad,  weakly  convex  central  zone  by  strong  spiral  cord; 
peripheral  zone  with  6  spiral  threads,  central  zone  with 
about  18  weak,  smooth  spiral  cords.  Umbilicus  funnel- 
shaped,  broad,  about  25%  maximum  shell  diameter,  wall 
strongly  constricted  to  narrow  pore.  Aperture  obscurely 
subquadrate,  almost  elliptical;  outer  lip  thin,  slightly  an- 
gulate;  inner  lip  thin  and  expanded  above  to  partially 
cover  umbilicus,  becoming  narrower  and  thicker  below; 
columella  concave  in  upper  and  lower  quarters,  almost 
straight  in  middle  half,  narrow  above  but  broad  and 
weakly  concave  at  base,  with  prominent  adapical  nacre- 
ous tongue.  Shell  ground  color  silvery  white  w  ith  brilliant 
pink  and  green  iridescence. 

Holotype:  USNM  860261,  height  9.1  mm,  width  10  0 
mm. 

Type  locality:  West  of  St.  Lucia,  Lesser  Antilles,  John 
Elliott  Pillsbury  Station  P-904,  13°45.5'N,  61°05.7'W, 
in  417-589  m. 

Distribution:  This  species  is  known  only  from  the  t\  pe 
locality. 


Remarks:  In  shell  shape  and  sculpture,  Calliostoma 
atlantoides  is  most  similar  to  C.  coronatum  new  species 
but  differs  by  having  a  larger  protoconch,  by  having  the 
beads  on  the  subsutural  spiral  cord  weaker  and  more 
widely  spaced,  b\  having  numerous  spiral  threads  in  the 
interspaces,  by  having  a  strongK  beaded  rather  than 
finely  undulate  shoulder  spiral  cord,  by  having  weaker 
and  more  numerous  basal  spiral  cords,  and  by  having 
the  inner  columellar  lip  expanded  to  partially  cover  the 
umbilicus.  The  shell  of  C.  atlantoides  is  also  ver\  similar 
to  those  of  C.  rota  and  C.  dnophenim.  It  differs  from 
that  of  C.  rota  by  being  relatively  higher  and  more 
globose,  by  having  a  much  larger  protoconch,  by  having 
a  more  coarsely  beaded  subsutural  spiral  cord,  b\'  having 
a  strongly  beaded  rather  than  smooth  shoulder  spiral 
cord,  by  having  an  open  umbilicus,  and  by  being  silvery 
white  rather  than  slightly  brassy.  The  shell  of  C.  atlan- 
toides differs  from  those  of  C.  dnophenim  by  being 
relatively  higher;  by  having  a  larger  protoconch;  b\  hav- 
ing a  single,  beaded  spiral  cord  rather  than  two  smooth 
spiral  cords  below  the  subsutural  cord;  by  having  finer, 
more  numerous  spiral  threads  in  the  interspaces;  by  hav- 
ing more  numerous  basal  spiral  cords;  and  b\  having  the 
inner  lip  of  the  columella  thinner  and  more  strongly 
expanded  to  partially  cover  the  umbilicus.  Calliostoma 
atlantoides  is  similar  to  C.  atlantis  but  has  a  much  small- 
er, more  globose  shell;  has  a  much  larger  protoconch; 
has  the  shoulder  spiral  cord  located  much  higher  on  the 
whorl;  has  a  much  stronger  subperipheral  angulation; 
and  has  the  inner  lip  of  the  columella  thin  and  partially 
covering  the  umbilicus  rather  than  being  thickened  and 
completely  filling  the  umbilicus.  The  radula  of  C.  at- 
lantoides'({ormula  <20.5?.1.5^.<20;  Figures  115,  116) 
is  very  small  (length  3.1  mm)  and  delicate,  and  there 
seem  to  be  five,  thin-cusped  laterals  and  fewer  than  20 
pairs  of  marginal  teeth,  the  innermost  of  which  has  a 
heavily  buttressed  and  rather  finely  denticulate  cusp.  The 
seemingly  degenerate  radula  of  C.  atlantoides  is  similar 
to  that  illustrated  for  an  undescribed  species  of  Callios- 
toma illustrated  by  Hickman  and  McLean  (1990:fig.  71C). 

Calliostoma  coronatum  new  species 
(figures  76,  77) 

Description:  Shell  small,  attaining  height  of  4.2  mm, 
width  of  more  than  4.2  mm,  depressed  turbinate,  um- 
bilicate. Protoconch  850  nm  maximum  diameter,  1  whorl. 
Teleoconch  whorls  about  2.2,  rapidK  expanding,  cari- 
nate; first  whorl  initially  with  3  smooth  spiral  cords,  adap- 
ical cord  disappearing  at  whorl  1.3,  midwhorl  cord 
strengthening,  becoming  weakly  undulate  at  whorl  1.3 
and  forming  strong  midwhorl  angulation,  abapical  cord 
remaining  weak  on  all  whorls;  fourth  spiral  cord  ap- 
pearing between  adapical  cord  and  suture  at  whorl  0.3, 
rapidly  strengthening,  undulate  on  last  half  of  first  whorl, 
undulations  becoming  strong,  closel\-  .set,  upturned,  tri- 
angular beads;  last  whorl  with  strong,  smooth  spiral  cord 
forming  periphery;  interspaces  smooth  except  for  weak 
spiral  cord  between  midwhorl  and  peripheral  cords;  axial 


J.  F.  Quinn,  Jr..  1992 


Page  103 


Figures  78,  79.    Calliostotna  rugosum  new  species,  holot\pe.  USNM  860262;  height  23.5  mm,  width  24. S  mm   Figures  80,  81. 

Calliostoma  torrei  Clench  &  Aguayo,  1940,  holotype,  MCZ  135165;  height  40.8  mm,  width  36.8  mm. 


sculpture  absent  except  for  \o\\.  rather  broad,  rounded 
folds  on  last  whorl.  Base  divided  into  narrow,  smooth, 
concave  peripheral  zone  and  broad,  almost  flat  central 
zone  by  strong,  smooth  spiral  cord;  central  zone  with  12 
narrow,  smooth  spiral  cords,  adaxial  3  somewhat  stronger 
and  more  widely  spaced.  Umbilicus  rather  narrow,  less 
than  20%  maximum  shell  diameter,  constricted  v\ithin 
to  small  pore.  Aperture  probably  obscurely  subquadrate 
or  ovate,  outer  lip  broken;  inner  lip  thin,  narrow,  slightly 
reflected  over  umbilicus;  columella  convex,  thin. 

Holotype:  MCZ  274568,  height  4.2  mm,  width  4.2  + 
mm. 

Type  locality:    Off  Joao  Pessoa,  Brazil.  Ch.\in  Cruise  35, 

Station  12,  7°09.0'S,  34°25.5'W,  in  768-805  m. 


Distribution: 

locality. 


This  species  is  known  only  from  the  type 


Remarks:  Calliostoma  coronatum  is  similar  to  C.  dno- 
pherum  but  differs  by  having  a  relatively  smaller,  lower- 
spired  shell  that  has  a  much  weaker  spiral  cord  between 
the  shoulder  and  peripheral  cords,  that  lacks  spiral  threads 
between  the  primary  spiral  cords,  and  that  has  a  pro- 
portionately narrower  umbilicus. 

Calliostoma  rugosum  new  species 
(figures  78,  79) 

Description:  Shell  medium-sized  to  large,  attaining  es- 
timated height  of  25  mm.  estimated  width  of  31  mm. 
trochoid,  nonumbilicate.  Protoconch  worn,  at  least  550 
Hm  maximum  diameter.  1  whorl.  Teleoconch  whorls 
about  7.5  (last  0.5  whorl  broken),  flat  to  weakly  concave; 
first  2  whorls  worn  and  partially  decorticated;  whorls  3- 
4  with  5  strong,  subequal,  beaded  spiral  cords,  adapical 
cord  forming  weak  subsutural  angulation,  abapical  cord 
forming  upper  limit  of  flat  periphery  bearing  1  spiral 
thread;  subsutural  and  upper  peripheral  cords  each  split- 
ting into  2  subequal  spiral  cords  on  whorls  5-6;  spiral 
cords  between  subsutural  and  upper  peripheral  cords 
increasing  to  7  by  end  of  seventh  whorl;  peripheral  zone 
with  4  spiral  cords  by  end  of  seventh  whorl,  periphery 


becoming  rounded  on  last  whorl;  all  cords  set  w  ith  rather 
strong,  rounded,  generally  spirally  elongate  beads.  Base 
weakly  convex,  with  19  strong,  smooth,  subequal  spiral 
cords.  Umbilical  area  filled  with  irregularly  ridged  callus. 
.Aperture  subquadrate;  outer  lip  broken;  iiuier  lip  thick- 
ened; columella  concave,  broken  basallv. 


Holotype:     USNM  860262, 
24.8+  mm. 


height  23.5+   mm,  width 


Type  locality:    Straits  of  Florida,  J.^mes  M.  Gillis  Cruise 
7307,  Station  13,  depth  and  exact  locality  unknown. 


Distribution: 

localitv . 


This  species  is  known  only  from  the  type 


Remarks:  The  holoty  pe  of  Calliostoma  rugosum  is  very 
similar  to  that  of  C.  torrei  Clench  and  Aguayo,  1940 
(figures  80,  81),  from  off  Cardenas,  Cuba,  but  differs  by 
being  relativ  ely  much  broader;  by  having  a  weaker,  more 
abapical  upper  peripheral  cord;  by  having  coarser,  more 
rounded  spiral  cords;  and  by  having  larger,  rounded, 
spirally  elongate  beads  on  the  cords.  The  protoconch  of 
C.  rugosum  (estimated  to  be  more  than  550  ^m)  may 
be  larger  than  that  of  C.  torrei  (about  525  ^m),  and  the 
subsutural  and  upper  peripheral  cords  of  C.  rugosum 
split  into  two  rather  coarse  cords  rather  than  three  fine, 
sharp  cords  as  in  C.  torrei.  Both  C.  rugosum  and  C. 
torrei,  based  on  similarities  of  shape  and  sculpture,  seem 
to  be  closely  related  to  C.  caribbcanum  \\'eisbord,  1962, 
a  fossil  species  from  the  Pleistocene  Mare  Formation  of 
\enezuela. 

Calliostoma  argentum  new  species 

(figures  82-85) 

CaliiiKloma  tnarionac.  Sander  &  Laili,  1982:table  4.  (Non  Cal- 
liostoma marionae  Dail.  1906). 

Description:  Shell  medium-sized,  attaining  height  of 
25.6  mm,  width  of  28.2  mm,  conical,  umbilicate,  finely 
sculptured  to  almost  smooth.  Protoconch  about  375  jum 
maximum  diameter,  1  whorl  Teleoconch  whorls  about 
9.5,  flat  to  verv  weakly  convex;  first  2  whorls  with  low, 
rounded  axial  riblets  and  2-3  finely  beaded  spiral  cords; 


Page  104 


THE  NAUTILUS,  Vol.  106,  No.  3 


> 


J 


Figures  82-85.  CUitliostoma  argentuni  new  species.  82.  83.  Holotype,  USNM  860263;  height  and  width  18.1  mm.  84,  85. 
Specimen  from  off  Barbados,  Sunderland  collection;  height  22.5  mm,  width  24.2  mm  Figures  86,  87.  Calliostoma  jeanneae  Clench 
&  Turner,  1960,  holotype,  MCZ  228370;  height  11.2  mm,  width  13.2  mm.  Figures  88,  89.  Calliostoma  axelolssoni.  new  name, 
specimen  from  off  Mostardas,  Brazil,  MORG  18.738;  height  30  4  mm,  width  34  2  mm 


axial  sculpture  absent  on  subsecjuent  w  liorls;  spiral  cords 
increasing  by  intercalation  to  9-1 1,  beading  disappearing 
on  fourth  or  fifth  whorl  but  sometimes  reappearing  on 
eighth  or  ninth  whorl;  periphery  sharply  carinate,  smooth, 
sometimes  reflected  adapically.  Base  flat  to  weakly  con- 
vex, with  13-15  weak,  broad,  smooth  spiral  cords,  adaxial 
2-3  cords  slightly  stronger,  weakly  beaded;  circumum- 
bilical  cord  strong,  coarsely  beaded.  Umbilicus  funnel- 


shaped,  14%-17%  maximum  shell  diameter,  wall  almost 
vertical,  white.  Aperture  subquadrate,  slightly  thickened 
within  and  with  several  low,  rounded  ridges,  lips  thin; 
columella  rather  long,  weakK  concave,  slightly  thick- 
ened, terminating  in  small,  rounded  tubercle.  Shell  ground 
color  cream  to  light  tan  w  ith  pale  orange-brown  patches 
below  suture  and  spots  on  periphery;  base  uniformly 


J.  F.  Quinii,  Jr.,  1992 


Page  105 


Holotype:    USNM  860263,  height  and  width  18  1  mm. 

Type  locality:  Arrovvsmith  Bank,  (^uintana  Roo,  Mex- 
ico, John  Elliott  Pillsbury  Station  P-594,  21°00.5'N, 
86°23.0'W,  in  307-329  m. 

Paratypes:  1,  UMML  30.5583;  off  Arrowsmith  Bank, 
Mexico,  John  Elliott  Pillsbury  Station  P-584, 
21°02.0'N,  86°24.0'W,  353-347  m;  23  May  1967;  10-ft 
otter  trawl.— 1,  ANSP  353529;  off  Holetovvn,  Barbados; 
F.  Sander,  collector. 

Other  material:  1  fragment,  UMML  30.8372;  off  St. 
Vincent,  John  Elliott  Pillsbury  Station  P-877, 
13°16.7'N,  6r05.6"W,  348-466  m;  6  July  1969;  5-f t  Blake 
trawl. — 3,  Sunderland  collection;  off  St.  James,  Barbados, 
175  m;  dredge;  F.  Sander,  collector. 

Distribution:  Shells  of  Calliostoma  argentum  are  known 
from  off  the  Yucatan  Peninsula.  Mexico,  and  St.  Vincent 
and  Barbados,  Lesser  Antilles,  in  depths  of  175-466  m. 

Remarks:  Shells  of  Calliostoma  argentum  are  most  sim- 
ilar to  the  holotype  of  C.  jeanneae  (figures  86,  87),  from 
off  Havana,  Cuba,  but  differ  by  being  relatively  narrow- 
er; by  having  spiral  cords  that  are  strongly  beaded  until 
the  middle  of  the  fourth  or  fifth  whorls  rather  than  be- 
coming smooth  near  the  beginning  of  the  third  whorl; 
b>  lacking  a  strong  spiral  cord  just  above  the  peripher\ 
on  the  third  through  sixth  whorls;  by  having  an  umbilicus 
that  is  open  rather  than  filled  with  callus;  by  having  a 
strongly  beaded  rather  than  smooth  circumumbilical  cord; 
and  by  having  a  longer,  less  thickened  and  oblique  col- 
umella. Perhaps  of  less  importance  are  differences  in  the 
color  patterns:  that  of  C.  argentum  consists  of  cream  to 
light  tan  spiral  cords  separated  by  darker-colored  striae 
and  pale  orange-brown  patches  below  the  sutures  and 
somewhat  darker  spots  on  the  periphery;  that  of  C.  jean- 
neae consists  of  rows  of  golden-brown  spots  on  the  spiral 
cords  above  and  below  the  periphery  and  brighter  spots 
of  the  same  color  on  the  periphery.  Both  C.  argentum 
and  C.  jeanneae  are  similar  to  the  eastern  Pacific  C. 
platinum,  C.  chilena,  and  C.  titanium,  and  all  five  spe- 
cies may  be  closely  related  to  C.  metalium  Woodring, 
1957,  from  the  late  Miocene  to  early  Pliocene  Chagres 
Sandstone  of  Panama. 


Calliostoma  axelolssoni  new  name 
(figures  88,  89,  119) 

Calliostoma  olssoni  Ba\er,  1971:118,  fig.  4  (left). — Kaicher, 

1980:card  no.  2239.  (Non  Calliostoma  olssoni  Maury,  1925). 
Calliostoma  (Kombologion)  rosewateri:  Rios,  1975:23,  pi  5, 

fig.  59—  Calvo,  1987:63,  65,  fig.  28.  (Non  Calliostoma 

rosewateri  Clench  &  Turner,  1960). 
Calliostoma  [Kombologion)  bairdi  rosewateri:  Rios,  1985:22, 

pl.9,  fig.  85. 

Description:  See  Ba\er  (1971:118)  for  description  of 
shell.  Animal  (in  alcohol)  white;  mantle  edge  smooth; 
cephalic  tentacles  moderately  long,  slender,  gradually 
tapering,  right  longer  than  left,  ocular  peduncles  rather 


long  (about  35%  tentacle  length),  broad,  with  large,  black 
eye  at  tips;  snout  long,  broad,  slightly  expanded  at  tip, 
tip  fringed  with  small,  short  papillae;  epipodium  with  4 
pairs  of  tentacles  decreasing  in  size  posteriorly,  neck  lobes 
well-developed,  semicircular,  smooth. 

Holotype:    USNM  700002,  height  16.8  mm,  width  21.0 


Type  locality:  Southwest  of  St.  N'incent,  Lesser  Antilles, 
John  Elliott  Pillsbury  Station  P-876,  13°13.9'N, 
6r04.7'W,  in  231-238  m. 

Other  material:  1,  UMML  30.8373;  off  Dominica,  John 
Elliott  Pillsbury  Station  P-931,  15°31.2'N,  61°12.3'W, 
174-357  m;  15  July  1969;  5-ft  Blake  trawl.— 1,  MORG 
18.738;  W.  Besnard  station,  off  Mostardas,  Rio  Grande 
do  Sul,  Brazil,  230  m;  1972. 

Distribution:  This  species  is  now  known  from  off  Dom- 
inica and  St.  Vincent  in  the  Lesser  Antilles,  and  off  south- 
ern Brazil,  in  174-357  m. 

Rema.»-ks:  Calliostoma  olssoni  Bayer,  1971,  is  preoc- 
cupied by  C.  olssoni  Maury,  1925,  a  species  (probably 
assignable  to  Calliomphahis  Cossmann,  1888)  from  the 
Mio-Pliocene  formations  of  Trinidad  (Maury,  1925;  Jung, 
1969).  With  F.  M.  Bayer's  permission,  1  offer  a  replace- 
ment name  that  preserves  his  original  intent  to  honor 
the  late  Dr.  Axel  A,  Olsson. 

The  Brazilian  specimen  (misidentified  as  C.  rosewateri 
by  Rios,  1975,  1985)  (figures  88,  89)  is  much  larger  (height 
30.4  mm,  w  idth  34.2  mm)  than  the  holot>  pe,  has  a  nar- 
rower umbilicus  that  has  been  partially  filled  with  callus, 
has  three  weak  circumumbilical  and  six  weak  subperi- 
pheral  spiral  cords,  and  has  a  more  diffuse  and  subdued 
color  pattern  except  for  the  pink  spots  on  the  periphery. 
The  Dominican  specimen  is  very  similar  to  the  holotype 
but  is  shghtly  larger  (height  19.9  mm,  width  24.1  mm), 
has  a  slightly  narrower  umbilicus,  and  has  a  much  less 
vivid  color  pattern.  The  radula  from  this  specimen  (figure 
119)  conforms  well  with  Calvo's  illustration  (1987:fig. 
28)  but  the  lateral  tooth  cusps  are  longer  than  he  de- 
picted. 

Based  on  shell  characters,  C.  axelolssoni  seems  to  be 
closely  related  to  C.  bigeloivi,  C  brunneum,  and  C. 
hendersoni  —  the  four  species  forming  a  rather  distinct 
subgroup  of  Calliostoma.  This  subgroup  may  be  most 
closely  related  to  the  C.  bairdii  species  group. 

Calliostoma  aurora  Dall,  1888 
(figures  90,  91,  120) 

Calliostoma  aurora  Dall,  1888:68,  fig.  285;  1889a:366,  pi.  37, 
fig.2— Clench  &  Turner,  1960:64,  pi.  45,  figs.  1,  2. 

Description:  Shell:  see  Dall  (1889a:366)  and  Clench  and 
Turner  (1960:64).  Animal  (reconstituted  in  trisodium 
phosphate):  foot  mottled  with  red-brown,  mantle  edge 
with  light,  transverse  band  and  long,  median  longitudinal 
streak  of  red-brown;  cephalic  tentacles  very  long,  slen- 
der; eyestalks  very  short  (less  than  10%  tentacle  length) 


Page  106 


THE  XAUTILIS.  \ol.  106,  No  3 


Figure?  90.  91.    Calliostoma  aurora  Dall,  1888.  specimen  from  off  Guadeloupe,  USXM  860504;  height  25.4  mm,  width  27  8  mm. 
Figures  92.  93.  Calliostoma  guesti  new  species,  holotype,  DMNH  96994;  height  and  width  27.5  mm. 


with  large,  black  eye  at  tips;  snout  slightly  longer  than 
broad,  tip  fringed  by  rather  long,  stout  papillae;  epipo- 
dium  with  3  or  4  pairs  of  very  long,  slender  tentacles; 
neck  lobes  very  well-developed,  seemingK  smooth-edged. 

Holotype:   MCZ  73808.  height  21.0  mm,  width  26,5  mm. 

Type  locality:  Off  Grenada,  Lesser  .Antilles,  Bl.\ke  Sta- 
tion 26.5,  12°03'55"N,  6r49'40"W,  in  1054  m. 

Other  material:  1,  USNM  860504;  off  Great  Inagua 
Island,  Bahama  Islands.  Johnson-Se.^-Link  I  Station  JSL- 
1-2323,  2r01'45"N',  74''43'48"\V,  628  m;  15  October 
1988, — 1,  Dan  collection;  off  Barbados,  Lesser  .\ntilles, 
183  m. 

Distribution:  Calliostoma  aurora  is  now  known  from 
the  southeastern  Bahama  Islands  and  from  off  Barbados, 
in  depths  of  ia3-1054  m. 

Remarks:  The  present  specimens  agree  well  with  the 
holotype  in  most  shell  characters,  but  are  larger  (Great 
Inagua;  height  25.4  mm,  width  27.8  mm;  Barbados:  height 
23.5  mm,  width  27.2  mm;;  have  whorls  with  flatter  pro- 
files; and  have  numerous  very  fine  spiral  striae  on  the 
outer  two-thirds  of  the  base,  the  striae  becoming  deeper 
on  the  adaxial  third  and  forming  several  indistinct  and 
three  distinct  spiral  cords  near  the  center.  The  ground 
color  of  the  new  shells  is  a  dark  golden  brown  with 
alternating  spots  of  white  and  brown  on  the  periphery; 
the  base  of  the  Barbados  specimen  has  faint,  crescent- 
shaped  streaks  of  pale  brown;  and  the  umbilical  callus 
of  both  specimens  is  white. 

.Although  Clench  and  Turner  (1960)  compared  the 
shell  morphology  of  C,  aurora  to  that  of  C.  hairdii  and 
C.  psyche,  the  relationship  of  this  species  to  other  Cal- 
liostoma  species  is  unclear.  The  radula  of  C.  aurora 
(formula  oo,10,l,10.oo;  figure  120)  indicates  that  the  spe- 
cies is  not  very  closely  related  to  the  C.  hairdii  species 
group  or  to  any  other  western  Atlantic  species  group. 
The  most  notable  difference  is  that  C,  aurora  has  ten 
pairs  of  lateral  teeth,  whereas  most  other  species  have 
only  four  to  seven  pairs;  only  Calliostoma  militare  (22 
pairs;  Castellanos  &  Fernandez,  1976;  Calvo,  1987)  and 
Venustatrochus  georgianus  (16  pairs;  Powell,  1951)  have 
more  lateral  tooth  pairs.  The  inner  marginal  teeth  of  C. 
aurora  sltc  also  rather  unusual:  the  innermost  marginal 


is  rather  slender  with  a  sharply  curved  cusp  and  ten 
denticles,  of  which  the  terminal  one  is  unusualK'  long, 
and  the  ne.xt  several  teeth  have  a  very  long  terminal 
denticle  with  four  to  six  secondary  denticles  about  a  third 
of  the  wa>  down  the  shaft. 

The  Barbados  specimen  was  collected  alive  from  a 
depth  of  183  m,  which  is  shallower  than,  but  comparable 
to,  the  provenance  of  the  paratype  lot  (from  Barbados 
in  256  m),  suggesting  that  the  occurrence  of  the  holot>pe 
in  1054  m  was  adventitious.  The  specimen  from  Great 
Inagua  was  collected  from  atop  a  small,  blue  sponge  (M. 
G,  Harasewych,  personal  communication),  and  its  intes- 
tine was  packed  with  a  dark  brown,  rather  solid  mass  of 
organic  material  in  which  were  embedded  numerous 
sponge  spicules  and  poK  chaete  setae,  suggesting  that  this 
species  feeds  on  sponges, 

Calliostoma  guesti  new  species 

(figures  92,  93) 

Description:  Shell  medium-sized,  attaining  height  and 
width  of  27.5  mm,  conical,  nonumbilicate,  finely  sculp- 
tured. Protoconch  about  425  nm  maximum  diameter,  1 
whorl.  Teleoconch  whorls  more  than  9.1,  flat  to  weakly 
convex;  first  2  whorls  with  low,  rounded  axial  riblets  and 
2-3  beaded  spiral  cords;  axial  sculpture  absent  on  sub- 
sequent whorls;  spiral  cords  increasing  by  intercalation 
to  12,  finely  beaded  throughout;  periphery  sharp,  nar- 
rowly bicarinate,  peripheral  cords  adpressed,  adapical 
peripheral  cord  distinctly  stronger  than  abapical  periph- 
eral cord.  Base  weakly  convex,  with  numerous  fine  striae 
on  outer  three-fourths  and  3-4  weak,  weakly  beaded 
spiral  cords  near  center.  Umbilicus  filled  with  white  cal- 
lus. Aperture  subquadrate,  lips  thin,  weakly  crenulate; 
columella  rather  short,  weakly  concave,  thickened.  Shell 
ground  color  ivory  with  irregular,  diffuse  patches  of  gold- 
en brown  above  periphery,  somewhat  more  discrete  spots 
of  same  color  on  periphery;  base  with  very  faint,  cres- 
cent-shaped, radial  streaks  of  light  brown  and  sometimes 
a  few  spots  on  1-2  spiral  cords, 

Holotype:    DMNH  96994,  height  and  width  27.5  mm. 

Type  locality:  2.5  mi  off  south  shore  of  Bermuda,  in 
183-219  m. 


J.  F.  Quinn,  Jr..  1992 


Paee  107 


96 


97 


Figures  94-97.  Calliostoma  stirophorum  (Watson.  1879).  94.  9.5.  Holot\  pe  of  Trochus  (Zizyphinus)  stirophonis  Watson.  1879. 
BM(\H)  1887.2.9,210;  height  7.5  mm,  width  6  6  mm  96.  97.  Holotype  of  Calliostoma  arestum  Dall,  1927.  ISN'M  108412;  height 
5.4  mm.  width  5.0  mm. 


Paratypes:  3.  DMXH  1S75SS;  1.  USXM  S60270;  from 
same  lot  as  holot\  pe. 

Distribution:  This  species  is  known  onK  from  the  t\  pe 
locality. 

Remarks:  Shells  of  C.alliostoma  guesti  are  most  similar 
to  those  of  C.  psyche  but  differ  b\  being  relati\  eK  nar- 
rower; b\  having  more  fineK  beaded  spiral  cords;  b\ 
having  the  supraperipheral  spiral  cord  on  all  whorls 
sharply  beaded  and  stronger  than  the  subsutural  cord 
rather  than  being  smooth  on  the  first  two  to  three  whorls 
and  weaker  than  the  subsutural  cord;  b\  ha\ing  the  pe- 
riphery composed  of  tw  o  adpressed  spiral  cords,  of  which 
the  adapical  cord  is  distinctly  stronger,  rather  than  hav- 
ing the  cords  subequal  and  separated  b\  a  concave  in- 
terspace that  is  as  wide  or  w  ider  than  the  cords;  h\  ha\ing 
much  weaker  spiral  and  radial  sculpture  on  the  base; 
and  by  having  a  color  pattern  of  light  brow  n  rather  than 
rose,  particularly  on  the  peripher\ . 

Calliostoma  guesti  is  named  in  honor  of  Mr  .\rthur 
Guest  for  his  contributions  to  the  knowledge  of  the  mol- 
luscan  fauna  of  Bermuda. 

Calliostoma  stirophorum  (Watson,  1879) 
new  combination 
(figures  94-97) 

Trochus  {Zizitphinus)  stirophorus  Watson,  1879:695;  1886:59, 

pi.  6,  fig.  2. 
Calliostoma  arestum  Dall,   1927b  127. — Johnson,  1934:70, — 

Clench  &  Turner,  196079  — .\bbott,  1974:46, 

Description:    See  Watson  (1879:695;  1886:59), 

Holotypes:  BM{NH)  1887.2.9.210.  height  7.5  mm,  width 
6,6  mm  (Trochus  (Zizyphinus)  stirophorus);  USNM 
108412,  height  5.4  mm.  w  idth  5.0  mm  (Calliostoma  ares- 
tum). 

Type  localities:  Off  Culebra  Island.  \'irgin  Islands, 
c'h.allenger  Station  24.  18''38'30"N,  64°05'3b"W.  in  713 
m  (Trochus  (Zizyphinus)  stirophorus);  off  southern 
Georgia,  .-^lb.^tross  Station  2415.  30°44'00"N, 
79°26'00"W.  in  805  m  (Calliostoma  arestum). 


Distribution:  This  species  is  know  n  onlv  from  the  two 
t\  pe  specimens,  one  from  off  Fernandina.  Florida,  in  805 
m.  and  off  Culebra  Island,  Virgin  Islands,  in  713  m. 

Remarks:  Both  Watson  (1879,  1886)  and  Dall  (1927b) 
based  their  respective  species  descriptions  on  unique 
specimens  that  w  ere  empt\  and  somew  hat  damaged  w  hen 
collected.  The  specimen  of  Calliostoma  arestum  ifigures 
96.  971  is  slightK  smaller,  has  fewer  whorls,  and  is  more 
w  eathered  than  that  of  Trochus  (Zizyphinus)  stirophorus 
vfigures  94,  95),  but  both  shells  seem  to  represent  the 
same  species.  These  two  shells  are  most  similar  to  those 
of  C.  circumcinctum  but  differ  principalK  b>  ha\ing 
three  strong,  rounded,  beaded  spiral  carinae  above  the 
suture  rather  than  the  two  ver\  strong,  blade-like  carinae 
of  C.  circumcinctum;  the  shells  of  C.  stirophorum  are 
also  proportionateK  broader  than  those  of  C.  circum- 
cinctum. The  shells  of  both  C.  stirophorum  and  C.  cir- 
cumcinctum are  ver\  similar  to  those  of  Otukaia  hlacki 
(Dell.  1956)  from  New  Zealand:  Calliostoma  (Otukaia) 
delli  McLean  and  .\ndrade.  19S2;  from  off  Chile,  and 
Otukaia  eltanini  Dell.  1990.  from  .Antarctica,  and  could 
be  included  in  the  genus  or  subgenus  Otukaia  Ikebe. 
1942.  if  that  ta.xon  gains  general  acceptance.  Some  recent 
authors  (Powell,  1979;  McLean  &  .\ndrade,  19S2;  Dell, 
1990)  are  now  using  Otukaia  at  either  the  genus  or  sub- 
genus level. 


Calliostoma  adspersum  (Philippi,  1851) 
(figures  98-100,  121) 

Trochus  eximius  Philippi,  1844  Un  1842-1851).  vol.  1.  pt.  6: 
17.  pi   4.  fig  7   (Non  Trochus  eximius  Reeve,  1843). 

Trochus  adspersus  Philippi.  1851  ^in  1846-1S55):217.  pi.  32 
fig.  8. 

Calliostoma  aspersum.  Lange  de  Morretes.  1949:58. — Calvo. 
1987:63.  fig.  27.  (Unjustified  emendation). 

Calliostoma  adspersum:  Calvo.  1987:63.  fig.  27. 

Calliostoma  [Koml>ologion)  adspersum .  Clench  &  Turner,  1960: 
46.  pi.  30.  figs.  1.  2  (partim).— Rios.  1970:25.  pi.  4  (lower 
right)  (partim);  1975:23.  pi.  5.  fig.  58;  1985:21.  pi.  9,  fig. 
84;  1990:9,  10  (photographs). 


Page  108 


THE  NAUTILUS,  Vol.  106,  No.  3 


Figures  98-100.  Calliostoma  adspersum  (Philippi,  1851),  specimens  from  off  Cabo  de  la  Vela,  Colombia.  98,  99.  FSBC  1  34044; 
height  17.9  mm,  width  18.2  mm,  100.  Deynzer  collection;  height  16.3  mm,  width  15.6  mm  Figures  101,  102.  Calliosloma 
depictum  Dall,  1927,  specimen  from  off  Salvador,  Brazil,  FSBC  I  44110;  height  10.3  mm,  width  9.9  mm. 


Description:  See  Clench  and  Turner  (1960:46)  and  Rios 
(1990:9)  for  descriptions  of  shells.  Animal  (in  alcohol): 
foot  red-brown;  mantle  with  narrow  band  of  white  at 
edge,  behind  which  is  broad,  irregularly  edged  band  of 
brown;  cephalic  tentacles  long,  slender,  evenly  tapered, 
ocular  peduncles  stout,  about  25%  tentacle  length,  with 
large,  black  eye  at  tips;  snout  long,  broad,  expanded  at 
tip,  red-brown,  with  anterior  fringe  of  long  papillae; 
epipodium  w  ith  3  pairs  of  red-brown  tentacles,  those  on 
left  larger,  neck  lobes  well-developed,  semicircular,  left 
lobe  finely  fringed,  right  lobe  smooth. 

Holotype:  Philippi's  type  material  could  not  be  located 
in  the  Museum  fiir  Naturkunde  der  Humboldt-Univer- 
sitat  zu  Berlin  (R.  Kilias,  in  litt.),  and  although  it  may 
be  present  in  Chile,  I  consider  it  to  be  lost. 

Type  locality:  Restricted  by  Clench  and  Turner  (1960; 
47)  to  Praia  do  Leste,  Ihia  Guaiba,  Estado  de  Rio  de 
Janeiro,  Brazil. 

Other  material:  1,  UMML  30.8368;  off  Cabo  de  la  Vela, 
Colombia,  John  Elliott  Pillsbury  Station  P-780, 
11°39.0'N,  73°08.5'W,  18-27  m;  30  July  1968;  10-ft  otter 
trawl.— 1,   FSBC  I  34044;  21,   Devnzer  collection;  off 


Cabo  de  la  Vela,  Colombia;  1990;  shrimp  trawlers. — 1, 
UMML  30.7156;  off  Cabo  de  la  \'ela,  Colombia,  John 
Elliott  Pillsbury  Station  P-767,  12°I6.1'N,  71°03.3'W, 
24-26  m;  28  July  1968;  10-ft  otter  trawl.— L  UMML 
30.7040;  off  Peninsula  de  Paraguana,  Venezuela,  John 
Elliott  Pillsbury  Station  P-758,  11°42.2'N,  69°40.0'W, 
15-18  m;  27  July  1968;  10-ft  otter  trawl— 1,  UMML 
30.6888;  off  Tucacas,  Venezuela,  John  Elliott  Pillsbury 
Station  P-750,  10°36.1'N,  68n2.2'W,  22-26  m;  25  July 
1968;  10-ft  otter  trawl —2,  UMML  30.6617;  off  Isla  de 
Margarita,  N'enezuela,  John  Elliott  Pillsbi:ry  Station 
P-721,  11°06.5'N,  64°22.5'\V,  26-27  m;  21  July  1968;  10- 
ft  otter  trawl— 1,  UMML  30.6183;  off  Georgetown,  Guy- 
ana, John  Elliott  Pillsbury  Station  P-686,  7°00.0'N, 
57°08.0'W,  27-26  m;  15  July  1968;  10-ft  otter  trawl  — 
2,  UMML  30.5931;  off  Cayenne,  French  Guiana,  John 
Elliott  Pillsbury  Station  P-655,  6°07.0'N,  53°39.0'W, 
26  m;  9  July  1968;  10-ft  otter  trawl.— 4,  FSBC  I  44109; 
off  Guarapari,  Estado  de  Espirito  Santo,  Brazil,  18-20 
m;  March  1991;  ex  J.  &  M.  Coltro. 

Distribution:  Calliostoma  adspersum  w  as  thought  to  be 
endemic  to  Brazil  (Clench  &  Turner,  1960;  Rios,  1970, 
1975)  until  Rios  (1985)  reported  its  occurrence  in  Suri- 


J.  F.  Quinn,  Jr.,  1992 


Page  109 


105 


106 


Figures  103-106.    Calliostoma  scurra  new  species.  103,  104.  Holotype,  USNM  860264;  height  14.1  mm.  width  13  3  mm.  105. 
106.  Paratype  from  John  Elliott  Pillsblry  Station  P-834,  FSBC  I  44073;  height  13.9  mm,  width  12.7  mm. 


Figures  107-1  II.  Radulae  of  Calliostoma  species.  107.  Callidstiiuta  <  nulnphilum  new  species,  ex  paratype,  FSBC  I  44070;  95  x. 
108.  Calliostoma  viscardii  new  species,  ex  holotype,  MORG  29,292,  310  x .  109.  Calliostoma  moscatellii  new  species,  ex  specimen 
from  off  Rio  de  Janeiro,  Brazil;  65  x ,  1 10,  111.  Calliostoma  scalenum  new  species,  ex  specimen  from  Campeche  Bank,  Yucatan, 
FSBC  I  32311;  235  x  and  435  x,  respectively  112.  Calliostoma  jujtibinum  (Gmelin,  1791),  ex  specimen  from  Cat  Island,  Bahama 
Islands,  FSBC  I  15698;  125  x  113.  Calliostoma  cubense  new  species,  ex  paratype  from  off  Peninsula  de  Hicacos,  Cuba,  FSBC  1 
32405;  245 X.  114.  Calliostoma  vinosum  new  species,  ex  holotype,  MORG  29.294;  280x. 


Page  110 


THE  NAUTILUS,  Vol.  106,  No.  3 


Figures  115-122. 


Radulae  of  Calliostoma  species 


115, 


116.  Calliostoma  atlantoides  new  species,  ex  holotype,  USNM  860261; 
75  X  and  160  x,  respectively  117,  118.  Calliostoma  rota  new  species,  ex  paratype,  MCZ  2580.57;  .50  x  and  115x,  respectively. 
119.  Calliostoma  axelolssoni  new  name,  ex  specimen  from  off  Dominica,  UMML  30.8.373;  65  x.  120.  Calliostoma  aurora  Dall. 
1888,  ex  specimen  from  Guadeloupe,  USNM  860504;  70x.  121.  Calliostoma  adspersum  (Philippi,  1851),  ex  specimen  from  off 
Guarapari,  Brazil,  FSBC  1  44108;  95  x.  122.  Calliostoma  depietum  Dall,  1927,  ex  specimen  from  off  llhabela,  Sao  Paulo,  Brazil, 
FSBC  1  44109;  190  X. 


name.  The  present  material  establishes  the  presence  of 
the  species  along  the  entire  northern  coast  of  South  Amer- 
ica, principally  in  depths  of  about  15-30  m. 

Remarks:  As  noted  by  Rios  (1975,  1985,  1990),  the 
names  Calliostoma  adspersum  and  C.  depietum  (figures 
101,  102)  do  not  represent  the  same  species,  as  was  con- 
tended by  Clench  and  Turner  (1960).  Shells  of  C.  ad- 
spersum attain  a  larger  size;  have  all  whorls  with  a  dis- 
tinct shoulder  angulation  and  fiat-sided  periphery;  have 
more  numerous,  more  unequally  sized  spiral  cords;  and 
have  more  diffuse  color  patterns  than  those  of  C.  depie- 
tum. One  juvenile  specimen  (UMML  30.7156;  height  6.5 
nun,  width  7.3  mm)  differs  from  larger  shells  by  having 
an  open  umbilicus  whose  wall  narrows  rapidlv  from  1.6 
mm  to  a  pore-like  opening  onK  0.3  mm  wide;  larger 
specimens  have  the  umbilicus  filled  with  callus.  The  rad- 
ula  of  C.  adspersum  (figure  121)  differs  from  that  of  C. 
depietum  (figure  122)  by  having  a  more  broadly  cusped 
rachidian,  by  having  fewer  pairs  of  lateral  teeth  (4  vs. 


5,  respectively),  and  by  having  m.ore  numerous  (about 
40  vs.  about  25)  marginal  tooth  pairs  whose  cusps  are 
narrower  and  have  more  slender  denticles. 

This  species  seems  to  occur  most  frequentK  in  areas 
of  "hard  bottom"  in  the  western  part  of  its  range  (John 
Elliott  Pillsbury  field  data  log)  and  on  sandy  mud 
bottom  in  the  eastern  and  southern  part  (Rios,  1985;  J. 
&  M.  Coltro,  personal  communication).  Calliostoma  de- 
pietum lives  in  depths  of  1-7  m  in  sandy  areas  aroimd 
rocks  and  corals  (Rios,  1990),  where  it  is  often  found 
living  under  an  unidentified  sea  urchin  (J.  &.  M.  Coltro, 
personal  communication);  the  species  is  also  found  on 
Sargassum  (Rios,  1985,  1990). 


Calliostoma  seurra  new  species 
(figures  103-106) 

Description:     Shell  medium-sized,  attaining  height  of 
14.1   mm,  width  of   13,3  mm,  conical,  nonumbilicate. 


J.  F.  Qiiinii,  Jr.,  1992 


Page  111 


rather  coarsely  sculptured.  Protoconch  40U— 125  ^lm  ma.\- 
imum  diameter,  1  whorl.  Teleoconch  whorls  about  7, 
weakly  convex;  first  4  whorls  with  niunerous  low ,  round- 
ed axial  riblets  and  2-5  fineK  beaded  spiral  cords;  axial 
sculpture  absent  on  subsequent  whorls;  spiral  cords  in- 
creasing by  intercalation  to  10-15,  usualK  alternating  in 
size,  set  with  crowded,  rounded  beads;  periphery  weakly 
angulate,  composed  of  pair  of  adpressed  spiral  cords, 
adapical  cord  stronger,  beaded,  abapical  cord  smooth. 
Base  convex,  with  12-19  strong,  narrow,  smooth  spiral 
cords,  interspace  between  abaxial  2  cords  usually  with 
1-5  fine  spiral  threads  and  next  adaxial  interspace  usually 
with  1  thread.  Umbilicus  filled  with  callus,  sliallnw  de- 
pression sometimes  present.  Aperture  subquadrate, 
thickened  within,  often  with  channel  corresponding  to 
periphery,  lips  thin,  weakly  crenulate;  columella  very 
weakly  concave,  greatly  thickened,  with  tongue  of  nacre 
extending  outward  to  umbilical  callus.  Shell  ground  color 
cream  with  spots  and  irregular  patches  ol  reddish  brow  n 
above  periphery  and  irregularly  crescentic  streaks  of  same 
color  on  base. 

Holotype:  USNM  860264,  height  14.1  mm,  width  13.3 
mm. 

Type  locality:  Off  mouth  of  Orinoco  River,  Venezuela, 
John  Elliott  Pillsburv  Station  P-834,  9°04.1'N, 
60°10.7'W,  in  33-35  m. 

Paratypes:  1,  USNM  860265;  1,  FSBC  I  44073;  1,  UP 
189457;  1,  MCZ  302590;  1,  ANSP  389338;  1,  LACM 
2253;  4,  UMML  30.7225;  all  from  same  lot  as  holotype. 

Other  material:  1,  UMML  30.6529;  off  Isla  de  Mar- 
garita, Venezuela,  John  Elliott  Pillsbury  Station  P-718, 
ll''22.5'N,  64°08.6'W,  60  m;  20  July  1968;  10-ft 
otter  trawl.— 1,  UMML  30.6374;  off  Peninsula  de  Paria, 
Venezuela,  John  Elliott  Pillsblry  Station  P-705, 
10°45.0'N,  62°00.0'W,  77-86  m;  18  Julv  1968;  10-ft  otter 
trawl.— 6,  UMML  30.6359;  Gulf  of  Paria,  Venezuela, 
John  Elliott  Pillsbury  Station  P-704,  10°34.3'N, 
61°57.0'W.  18  m;  18  July  1968;  10-ft  otter  trawl.— 1, 
UMML  30.6339;  off  Orinoco  River,  Venezuela,  John  El- 
liott Pillsbury  Station  P-696,  8°38.0'N,  58°56.0'W,  55- 
59  m;  16  July  1968;  10-ft  otter  trawl.— 1  +  1  fragment, 
UMML  30.5979;  off  Parimaribo,  Suriname,  John  El- 
liott Pillsbury  Station  P-663,  6°29.0'N,  54°41,0"W,  24 
m;  10  July  1968;  10-ft  otter  trawl. 

Distribution:  Calliostoma  scurra  has  been  collected  from 
off  Suriname  northwestward  to  off  Isla  de  Margarita, 
Venezuela,  in  depths  of  18-86  m;  live-collected  speci- 
mens are  known  from  depths  of  18-35  m 

Remarks:  Calliostoma  scurra  does  not  seem  to  be  close- 
K  related  to  any  other  western  Atlantic  species  except, 
perhaps,  C.  adspersum.  Shells  of  the  latter  species  differ 
from  those  of  the  former  by  having  a  broad,  flat  periph- 
ery; by  having  distinct  axial  rugae  in  the  interspaces 
between  the  primary  spiral  cords;  by  having  finer,  more 
numerous,  more  distinctly  beaded  supraperipheral  spiral 


cords;  by  having  a  more  oblique  columella;  and  by  hav- 
ing a  different  color  pattern. 

The  field  data  indicate  that  this  species  lives  on  bottoms 
composed  of  mud,  shell  hash,  and  coral  rubble. 

ACKNOWLEDGMENTS 

I  thank  the  following  for  access  to  collections  under  their 
care:  George  M.  Davis  and  Robert  Robertson  (ANSP); 
Richard  S.  Houbrick,  M.  G.  Harasewych,  and  the  late 
Joseph  Rosewater  (USNM);  Kenneth  J.  Boss,  Silvard  P. 
Kool.  and  Ruth  D.  Turner  (MCZ);  Riidiger  Bieler  and 
Russell  Jensen  (DMNH);  Nancy  A.  and  the  late  Gilbert 
L.  Voss  (UMML);  John  D.  Taylor  and  Kathie  M.  Way 
(BM[NH]);  James  H.  McLean  (LACM);  Fred  G.  Thomp- 
son and  Kurt  Auffenberg  (UF);  Constance  E.  Boone  and 
the  late  Thomas  E.  Pulle>  (HMNS);  and  Eliezer  de  C. 
Rios  (MORG).  I  am  especialK  grateful  to  Kevan  and 
Linda  Sunderland  for  their  hospitality,  for  allow  ing  me 
to  examine  their  collection  that  contains  much  important 
material,  and  for  generously  donating  type  or  \oucher 
specimens.  Jose  and  Marcus  Coltro  were  instrumental  in 
obtaining  Brazilian  material  and  generousK'  donated  type 
material.  Jose  Leal  (UMML)  also  provided  material  col- 
lected from  seamounts  off  southeastern  Brazil.  Arthur 
Guest  and  Jack  Lightbourn  kindly  provided  the  type 
material  of  the  Bermudan  species.  Albert  E.  and  Beverly 
A.  Deynzer  generously  donated  material  of  Calliostoma 
purpureum.  Harry  G.  Lee,  Donald  Dan,  and  Leslie  A. 
Crnkovic  kindly  allowed  me  to  examine  specimens  in 
their  collections.  Rudolf  Kilias  (Museum  fiir  Naturkunde 
der  Humboldt-Universitat  zu  Berlin)  provided  infor- 
mation on  Philippi's  type  material.  Sally  D.  Kaicher  kind- 
ly provided  photographs  of  the  types  of  Watson's  species. 
Thomas  H.  Perkins.  Da\  id  K.  Camp.  William  G.  Lyons 
(all  Florida  Marine  Research  Institute),  and  two  anon- 
ymous review  ers  provided  helpful  comments  on  previous 
versions  of  this  paper. 

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Woodring,  VV.  P.  1928.  Miocene  moUusks  from  Bowden,  Ja- 
maica. Part  II.  Gastropods  and  discussion  of  results.  Car- 
negie Institution  of  Washington,  Publication  385:vii  -I- 
564. 


Woodring,  W.  P.  1957.  Geology  and  paleontology  of  Canal 
Zone  and  adjoining  parts  of  Panama.  Geology  and  de- 
scription of  Tertiary  mollusks  (Gastropods:  Trochidae  to 
Turritellidae).  United  States  Geological  Survey,  Profes- 
sional Paper  306-A:iv-H45  p. 

Work,  R.  C.  1969.  Systematics,  ecology,  and  distribution  of 
the  mollusks  of  Los  Roques,  Venezuela.  Bulletin  of  Marine 
Science  19(3):614-711. 


THE  NAUTILIIS  106(3):115-118,  1992 


Page  115 


A  New  Species  of  Pseiidori77iula 
(Fissurellacea:  Clypeosectidae)  from 
Hydrothermal  Vents  of  the 
Mid-Atlantic  Ridge 


James  H.  McLean 

Los  Angeles  C^iuinh   Miiseiiin  of 

Natural  Histor\ 
900  Exposition  Houle\ard 
Los  Angeles,  CA  90007,  USA 


ABSTRACT 

Pseudorimula  midatlantica  new  species  is  described  from  the 
Snake  Pit  hydrothermal  lield  on  the  Mid-Atlantic  Ridge  It  is 
the  second  member  of  its  genus,  the  type  species  being  known 
Irom  the  Mariana  Trough  h)drothermal  vents  in  the  mid-Pa- 
cific It  differs  from  the  type  species  in  its  hypertrophied  de- 
velopment of  the  gonad,  which  displaces  part  of  the  space 
normally  occupied  by  the  foot  on  the  left  side;  correspondingly, 
the  posterior  shell  muscle  of  the  type  species  is  merged  with 
the  right  shell  muscle  in  P.  midatlantica.  Other  differences  are 
that  it  has  three  rather  than  six  pairs  of  epipodial  tentacles. 
The  new  species  also  provides  evidence  of  faunal  interchange 
between  widely  separated  ridge  systems. 

Key  words:  .'\rchaeogastropoda,  Fissurellacea,  Clypeosectidae; 
hydrothermal- vent  limpets;  Mid-Atlantic  Ridge. 


INTRODUCTION 

The  slit  limpet  genus  Pseudorimula  McLean,  1989,  was 
based  on  a  single  species  from  hydrothermal  vents  at  the 
Marina  Trough  in  the  mid-Pacific.  Here  I  add  to  the 
genus  a  second  species  from  the  Mid-Atlantic  Ridge,  a 
ridge  system  for  which  other  components  of  the  fauna 
are  largely  undescribed.  First  indications  of  biota  on  the 
Mid-Atlantic  Ridge  came  from  camera  tows  and  dredg- 
ings  by  the  NOAA  vessel  Researcher  at  a  hydrothermal 
field  at  26°N  (Rona  et  a/.,  1986);  moilusks  were  not  re- 
ported. Mollusks  from  the  Mid-Atlantic  Ridge  were  first 
collected  in  1988  by  observers  on  the  deep-submersible 
Nautile  at  the  Snake  Pit  hydrothermal  field  at  23°N. 
Spreading  centers  at  these  two  sites  on  the  Mid-Atlantic 
Ridge  are  diverging  at  a  slower  rate  than  those  of  the 
East  Pacific  (Rona  et  a/.,  1986;  Tunnicliffe,  1991). 

Unusual  features  of  the  biota  of  the  Snake  Pit  vents 
were  noted  by  Mevel  et  al.  (1989):  "The  characteristic 
feature  of  these  hydothermal  sites  is  the  amazing  density 
of  shrimps  agglutinated  on  the  chimneys;  around  the 
vents,  the  fauna  consists  of  sea  anemones,  polychaetes. 


gastropods,  galatheids,  mussels  and  zoarcid  fish.  Tiie  Snake 
Pit  differs  from  the  Pacific  sites  mostly  by  the  absence 
of  vestimentiferan  worms,  alvinellid  and  serpulid  poly- 
chaetes and  cephalopods.  ' 

This  new  species  of  Pseudorimula  came  to  my  atten- 
tion after  the  original  paper  (McLean,  1989)  was  in  press. 
It  adds  new  limits  to  the  morphology  known  in  the  family 
and  provides  an  example  of  interchange  between  widely 
separated  ridge  systems.  It  is  also  the  first  mollusk  to  be 
documented  from  the  Mid-Atlantic  Ridge. 


MATERIALS  AND  METHODS 

Specimens  were  collected  by  the  French  expedition  HY- 
DROSNAKE  to  the  Mid-Atlantic  Ridge,  June-July  1988, 
and  forwarded  to  me  by  Michel  Segonzac  of  the  Centre 
National  de  Tri  d'Oceanographie  Biologique  (CENTOB, 
IFREMER,  Brest). 

The  illustrated  radula  was  extracted  from  a  preserved 
specimen  after  dissolution  of  tissues  with  room  temper- 
ature 10%  NaOH  for  48  hours,  washed  in  distilled  water, 
air  dried  and  coated  with  gold  palladium  for  SEM  ex- 
amination. 

Abbreviations  for  museums  mentioned  in  the  text  are 
MNHN,  Museum  National  d'Histoire  Naturelle,  Paris 
LACM,  Los  Angeles  County  Museum  of  Natural  History 
LISNM,  National  Museum  of  Natural  History,  Washing- 
ton. 


Suborder  VETIGASTROPODA  Salvini-Plawen,  1980 
Superfamily  FISSURELLACEA  Fleming,  1922 
Family  CLYPEOSECTIDAE  McLean,  1989 

Clypeosectids  differ  from  fissurellids  in  having  a  distinct 
radular  plan,  a  reduced  epipodium,  a  different  pattern 
of  shell  musculature,  and  differences  in  the  internal  anat- 
omy, as  discussed  in  more  detail  by  McLean  (1989)  and 
Haszprunar  (1989).  Haszprunar  (1989)  provided  the  an- 


Page  116 


THE  NAUTILUS,  Vol.  106,  No.  3 


Figures  1-7.  PseudorUmda  midatlantica  McLean,  sp.  nov.  NaiUile  dive  HSIO.  Snake  Pit  hydrothermal  field,  Mid-Atlantic  Ridge, 
3,478  ni.  Hololype,  MNHN.  Anterior  at  top  in  vertical  views.  1-3.  External,  internal,  and  left  lateral  views  of  shell.  Length  8.1 
mm.  4.  Dorsal  view  of  detached  body  showing  left  and  right  shell  muscles.  5.  Ventral  view  of  body  attached  to  shell,  showing 
protruding  gonad  on  left  side  of  foot.  Epipodial  tentacles  are  concealed  by  the  foot  in  this  view  6,  7.  SEM  views  of  radula  of 
paratype.  Scale  bar  for  6  =  20  ^m;  scale  bar  for  7  =  10  fim. 


atomicai  evidence  to  justify  the  erection  of  a  second 
family  in  the  superfamily  Fissurellacea. 

On  radular  characters,  the  Plssurellidae  differ  from 
the  Clypeosectidae  in  having  a  massive  piuricuspid  tooth 
tfiat  separates  the  lateral  field  of  teeth  from  the  marginal 
teeth.  The  piuricuspid  tooth  is  lacking  in  the  C'lypeosec- 
tidae,  in  which  the  lateral  and  marginal  teeth  are  strik- 
ingly similar  in  morphology.  Clypeosectus  McLean,  1989, 
was  established  for  two  species  from  eastern  Pacific  vents 
that  have  shells  with  an  open  slit  deflected  to  the  right. 
Pseudorimula.  which  is  convergent  in  shell  morphology 
with  the  fi.ssurellid  genus  Rimtda  de  France,  1827,  has 
the  slit  closed  at  the  margin. 

Genus  Pseudorimula  McLean,  1989 

Pseudorimula  McLean,  1989:22.  Type  species;  Pseudo- 
rimula marianae  McLean,  1989. 


Pseudorimula  midatlantica  McLean,  spec.  nov. 
(figures  1-7) 

Pseudorimula  sp.  McLean;  Martin  &  Hessler,  1990:10; 
Tunnidiffe,  1991:349. 

Description:  Shell  (figures  1-3)  relatively  large  for  fam- 
ily, maximum  length  8. 1  mm.  Surface  coated  with  rusty 
mineral  deposits,  under  which  periostracum  yellowish 
brown,  tightly  adhering,  projecting  slightly  past  shell 
margin.  Outline  of  aperture  oval,  margin  of  aperture 
nearly  in  same  plane;  highest  elevation  of  shell  at  about 
one-half  its  length.  Profile  moderately  high,  height  of 
h()lot\pe  0.34  times  length.  Apical  whorl  at  two-thirds 
shell  length  from  anterior  end,  slightly  deflected  to  right; 
protoconch  diameter  150  nm.  First  teieoconch  whorl 
smooth,  rounded;  slit  arising  two  protoconch  diameters 
away.  Juvenile  shells  with  open  slit;  in  mature  shells  slit 
open  about  one-third  length  of  anterior  slope,  strongly 


J.  H.  McLean,  1992 


Page  117 


deflected  to  right.  Borders  of  foramen  raised,  except  an- 
teriorly, where  slit  is  sealed  in  mature  shells  and  its  trace 
slightly  depressed.  Selenizone  weakly  depressed  below 
slit  border,  additions  to  selenizone  extending  straight 
across.  Sculpture  of  about  30  well-defined  primary  ribs 
with  one  to  three  secondary  ribs  of  lesser  prominance 
arising  in  interspaces.  Concentric  sculpture  of  fine  growth 
lines,  raised  into  sharp  lamellar  scales  on  crossing  primary 
ribs.  Shell  interior  opaque.  Muscle  scar  well  marked  on 
shell  interior;  left  and  right  arms  swollen  at  anterior  tips, 
but  of  uneven  thickness  posteriorly;  right  arm  longer  and 
retaining  its  thickness  to  its  termination  near  midpoint; 
left  arm  shorter  anteriorly  and  posteriorly,  connecting  to 
right  arm  through  narrow  band.  Anterior  pallial  attach- 
ment scars  well  marked,  extending  close  to  suture  bring- 
ing two  anterior  portions  into  contact  anterior  to  fora- 
men. Suture  with  zig-zag  outline.  Shell  strengthened  by 
thickened  callus  adjacent  to  suture  and  surrounding  fo- 
ramen. 

Dimensions  of  holotype:  Length  8.1,  width  6.4,  height 
2.5  mm. 

External  anatomy  (figures  4,  5):  Anterior  end  of  foot 
with  double  anterior  edge  marking  opening  of  anterior 
pedal  gland;  foot  posterior  rounded;  left  side  of  foot 
displaced  posterio-laterally  by  projecting  gonad.  Ce- 
phalic tentacles  contracted  from  preservation.  Two  pos- 
terior pairs  of  epipodial  tentacles,  on  body  wall  midway 
between  foot  edge  and  thick  border  of  mantle  margin 
(posterior  tentacles  of  right  side  in  cavity  abutting  pro- 
jecting gonad);  single  anterior  pair,  all  with  thick,  joined 
bases,  each  with  narrow  projecting  tips.  Mantle  skirt 
deeply  emarginate,  corresponding  to  foramen  and  seam 
in  shell,  edge  of  emargination  with  projecting  papillae. 
Mantle  skirt  above  head  thin,  nearly  transparent.  Shell 
muscles  without  inturned  hooks;  left  muscle  short  in  re- 
lation to  right  and  truncate,  right  muscle  extending  more 
anteriorly  than  left,  having  thick  posterior  arm  extending 
not  quite  to  midpoint;  left  and  right  muscles  joined  by 
thin  connective  muscle  near  margin.  Right  ctenidium 
smaller  than  left;  both  ctenidia  reduced,  having  filaments 
only  on  inner  sides  of  axes. 

Radula  (figures  6,  7):  Radular  ribbon  nearly  symmet- 
rical. Rachidian  tooth  with  long  overhanging  tip,  edges 
deeply  serrate;  shaft  of  rachidian  short  but  broad  at  base. 
Four  pairs  of  lateral  teeth,  similar  in  morphology  to 
rachidian  except  much  narrower;  outer  edges  with  fine 
serration,  inner  edges  smooth;  size  of  overhanging  tips 
decreasing  gradually  away  from  rachidian.  Marginal  teeth 
numerous,  with  broad  tips,  edges  finely  and  sharply  ser- 
rate, serrations  similar  to  those  of  laterals.  Marginals  and 
outer  laterals  with  one  long  denticle  on  outer  edge  of 
shaft  near  overhanging  tip. 

Type  locality:  Snake  Pit  hydrothermal  field,  Mid-At- 
lantic Ridge  (23°22'N,  47°57'W),  3,478  m. 

Type  material:  Holotype  and  20  paratypes  from  Natt- 
tile  dive  HSIO,  28  June  1988.  Holotype  MNHN;  16  para- 
types MNHN;  2  paratypes  LACM  2424,  2  paratypes 


USNM  859485.  The  holotype  is  the  only  specimen  in 
which  both  body  and  shell  are  in  good  condition,  al- 
though the  shell  is  heavily  coated  with  mineral  deposits. 
All  other  paratypes  are  smaller  and  completely  decal- 
cified so  that  they  can  not  be  measured.  Bodies  are  mostly 
separated  from  the  shell  remnants  and  are  somewhat 
mangled,  although  it  is  evident  that  the  gonad  protrudes 
on  the  left  side  in  each.  The  radula  was  prepared  from 
a  detached  body. 


DISCUSSION 

Pseiidorimula  midatlantica  differs  from  P.  marianae  in 
having  the  right  shell  muscle  longer  than  the  left  rather 
than  having  three  separate  muscles,  a  hypertrophied  go- 
nad that  displaces  the  foot  on  the  left  side,  a  single  pair 
rather  than  two  pairs  of  anterior  epipodial  tentacles  and 
two  rather  than  four  pairs  of  posterior  epipodial  tentacles. 
The  two  species  can  hardly  be  distinguished  in  shell 
profile  or  sculpture,  although  the  holotype  shell  of  P. 
midatlantica  has  an  opaque  interior,  in  contrast  to  the 
transparent  condition  in  P.  marianae.  This,  however, 
may  be  due  to  differences  in  preservation,  including  the 
treatment  conditions  that  led  to  the  decalcification  of  all 
paratype  specimens  of  P.  midatlantica. 

In  the  original  description  of  the  genus  Pseudorimula, 
I  noted  that  the  outer  lateral  teeth  are  morphologically 
similar  to  the  inner  marginal  teeth,  so  much  so  that  it  is 
difficult  to  determine  which  are  lateral  teeth  and  which 
are  marginal  teeth.  For  P.  midatlantica  (figures  6,  7)  I 
identify  four  pairs  of  lateral  teeth  by  their  more  acute 
tips,  compared  to  the  broader  terminations  of  the  mar- 
ginal teeth.  Earlier  (McLean,  1989:  figs.  13C,D)  I  stated 
that  P.  marianae  has  five  pairs  of  lateral  teeth,  but  now 
revise  that  to  four  pairs  based  on  re-examination  of  the 
original  illustration. 

Both  of  the  two  known  species  of  Pseudorimula  have 
differing  features  that  set  each  apart  from  all  other  lim- 
pets: In  P.  mariana  the  shell  muscle  is  inexplicably  di- 
vided into  three  units,  quite  unlike  the  usual  horseshoe- 
shaped  muscle  configuration  in  limpets  of  many  families. 
In  P.  midatlantica  the  gonad  is  so  hypertrophied  that  it 
displaces  the  foot  on  the  left  side;  the  shell  muscle  on 
the  right  side,  where  the  foot  remains  large,  is  corre- 
spondingly longer.  Which,  then,  of  these  two  conditions 
is  the  more  derived?  We  can  assume  that  an  ancestral 
species  would  have  the  usual  horseshoe-shaped  shell  mus- 
cle and  a  gonad  of  normal  size  contained  within  the  body 
cavity.  Indeed,  such  a  species  may  yet  be  discovered 
living  on  unexplored  ridge  systems.  It  is  easy  to  under- 
stand the  origin  of  the  asymmetrical  muscle  pattern  of 
P.  midatlantica  as  an  adjustment  to  gonad  hypertro- 
phication  (in  response  to  need  for  greater  reproductive 
output)  that  eliminates  space  for  muscle  attachment  on 
the  left  side,  but  there  is  no  easy  explanation  for  the 
presence  of  three  separate  shell  muscles  in  P.  marianae. 
If,  however,  the  hypertrophied  gonad  of  P.  midatlantica 
were  to  revert  to  a  normal  size,  the  large  right  muscle 
would  already  be  in  place  and  the  posterior  muscle  could 


Page  118 


THE  NAUTILUS,  Vol.  106,  No.  3 


then  be  pinched  off  from  the  posterior  tip  of  the  right 
muscle,  once  there  was  no  longer  the  need  for  a  large 
right  muscle. 

The  occurrence  of  the  two  species  of  Pseudorimula  at 
such  widely  separate  habitats  as  the  Mariana  Trough  and 
the  Mid-Atlantic  Ridge  is  noteworthy  but  not  unique. 
Martin  and  Hessler  (1990)  discussed  "a  growing  body  of 
evidence  that  there  is  a  faunal  connection  between  the 
Mariana  vent  area  and  the  northern  Mid- Atlantic  Ridge." 
They  mentioned  similarities  in  the  bythograeid  crabs  and 
gave  three  examples  of  genera  represented  in  the  two 
habitats,  including  Pseudorimula  (as  a  personal  com- 
munication from  me),  the  shrimp  genus  Chorocaris 
(therein  proposed),  and  a  new  genus  of  mussel  (reported 
by  Grassle,  1989).  Martin  and  Hessler  proposed  that  "the 
hydrothermal  areas  of  the  western  Pacific  and  northern 
Mid-Atlantic  Ridge  were  at  one  time  connected  via  a 
series  of  active  vent  areas,  not  necessarily  active  simul- 
taneously, that  extended  from  the  Mid-Atlantic  Ridge 
south  to  the  Atlantic-Indian  Ocean  Ridge,  north  along 
the  Southwest  Indian  Ocean  Ridge,  Mid-Indian  Ocean 
Ridge,  and  Southeast  Indian  Ocean  Ridge,  and  finally 
north  through  the  various  spreading  centers  of  the  Indo- 
West  Pacific." 

Faunal  connection  between  the  Mariana  Vents  and 
those  of  the  eastern  Pacific  were  discussed  by  Hessler 
and  Lonsdale  ( 1991a, b).  For  vent  limpets  this  applies 
only  at  the  family  level  in  the  Clypeosectidae,  as  well  as 
the  Neomphalidae  (McLean,  1990),  but  is  exemplified 
at  the  species  level  by  Lepetodrilns  elevatus  McLean, 
1988,  reported  by  McLean  (1990)  to  occur  at  the  Mariana 
vents  as  well  as  the  east  Pacific  vents. 

ACKNOWLEDGMENTS 

I  thank  Michel  Segonzac  (CENTOB,  Brest),  and  Phi- 
lippe Bouchet  of  the  Museum  National  d  Histoire  Na- 
turelie,  Paris,  for  allowing  me  to  describe  the  new  species. 
(.1.  (Clifton  Coney  operated  the  SEM  to  produce  the  rad- 
ular  illustrations.  I  thank  Anders  Waren,  Swedish  Mu- 
semn  of  Natural  History,  and  two  anonymous  reviewers 
for  reading  the  manuscript. 


LITERATURE  CITED 

Grassle,  J.  F.  1989.  A  plethora  of  une.xpected  life.  Oceanus 
31:41-46. 

Haszprunar,  G.  1989.  New  slit-limpets  (Scissurellacea  and 
FLssurellacea)  from  hydrothermal  vents.  Part  2.  Anatomy 
and  relationships.  Natural  History  Museum  of  Los  Angeles 
County,  no.  408,  17  p. 

Hessler,  R.  R.  and  P.  F.  Lonsdale.  1991a.  Biogeography  of 
Mariana  Trough  hydrothermal  vent  communities.  Deep- 
Sea  Research  .38:185-199. 

Hessler,  R.  R.  and  P.  F.  Lonsdale.  1991b.  The  biogeography 
of  the  Mariana  Trough  hydrothermal  vents.  In:  J.  Mauch- 
ling  and  T  Nemoto  (eds. ),  Marine  biology,  its  accomplish- 
ment and  future  prospect.  Kokusen-sha  (Japan),  p  165- 
182. 

Martin,  J.  W  and  R  R  Hessler.  1990.  Chorocaris  vandoverae. 
a  new  genus  and  species  of  hydrothermal  vent  shrimp 
(Crustacea,  Decapoda,  Bresiliidae)  from  the  Western  Pa- 
cific. Natural  History  Museum  of  Los  Angeles  County, 
Contributions  in  Science,  no.  417,  lip. 

McLean,  J.  H.  1989.  New  slit-limpets  (Scissurellacea  and  Fis- 
surellacea)  from  hydrothermal  vents  Part  1.  Systematic 
descriptions  and  comparisons  based  on  shell  and  radular 
characters.  Natural  History  Museum  of  Los  Angeles  Coun- 
ty, Contributions  in  Science,  no.  407,  29  p. 

McLean,  J.  H.  1990.  A  new  genus  and  species  of  neomphalid 
limpet  from  the  Mariana  vents  with  a  review  of  current 
understanding  of  relationships  among  Neomphalacea  and 
Peltospiracea.  The  Nautilus  104(3):77-86. 

Mevel,  C,  J.  Auzende,  M.  Cannat,  J.  Donval,  J.  Dubois,  Y. 
Fouquet,  P.  Gente,  D.  Grimaud,  J.  A.  Karson,  M.  Segonzac, 
and  M  Stievenard.  1989.  La  ride  du  Snake  Pit  (dorsale 
medio-Atlantique,  23°22'N)  resultats  preliminaires  de  la 
compagne  HYDROSNAKE.  Comptes  Rendus  des  Seances 
de  1  Academic  des  Sciences,  Paris,  series  2,  308:545-552. 

Rona,  P.  A.,  G.  Klinkhammer,  T.  A.  Nelsen,  J.  H.  Trefry,  and 
H.  Elderfield.  1986.  Black  smokers,  massive  sulphides 
and  vent  biota  at  the  Mid-Atlantic  Ridge.  Nature  321:33- 
.37, 

Tunnicliffe,  V  1991.  The  biology  of  hydrothermal  vents: 
ecology  and  evolution.  Annual  Review  of  Oceanography 
and  Marine  Biology,  Margaret  Barnes,  ed.,  29:319-407. 


THE  NAUTILUS  106(3):119-122,  1992 


Page  119 


Prodissoconch  I  and  II  Length  in  Mercenaria  Taxa^ 


Joy  G.  Goodsell 
Arnold  G.  Eversole 

Department  of  Aqiiaculture, 

Fisheries  and  Wildlife 
Clemson  University 
Clemson,  SC  29634-0362,  I'SA 


ABSTRACT 

Mercenaria  mercenaria  larvae  raised  from  71)  aiul  SO  ^m  di- 
ameter eggs  were  reared  for  4.S  hours,  until  the  prodissoconch 
I  boundary  was  clearly  visible.  Prodissoconch  I  lengtlis  of  larvae 
from  80  ^m  eggs  were  significantly  greater  than  tho.se  from  70 
^m  eggs  Larvae  from  crosses  using  similarly  sized  eggs  of  A/. 
campechiensis  and  M.  campechiensis  texana  females  and  sperm 
from  M.  campechiensts.  M.  cairipechiensis  texana  and  M.  mer- 
cenaria males  were  reared  through  settlement  There  were  no 
significant  differences  in  prodissoconch  I  lengths;  however, 
prodissoconch  II  lengths  of  larvae  sired  by  A/,  mercenaria  were 
significantly  longer  than  those  sired  by  either  subspecies  of  M. 
campechiensis.  Differences  in  shell  growth  (AP  =  prodisso- 
conch II  —  prodissoconch  I  length)  also  followed  taxonomic 
lines. 

Key  words:  Prodissoconch,  egg,  Mercenaria  mercenaria.  Mer- 
cenaria campechiensis.  Mercenaria  campechiensis  texana. 


INTRODUCTION 

Relationships  between  egg  size  and  shell  length  at  the 
first  shelled  stage  (i.e.,  prodissoconch  I  or  protoconch  I) 
have  been  reported  for  both  bivalves  and  gastropods 
(Thorson,  1950;  Robertson,  1971;  Jablonski  &  Lutz,  1983; 
Lima  &  Lutz,  1990).  Ockelmann  (1962)  suggested  the 
usefulness  of  the  length  of  prodissoconch  I  for  estimating 
the  egg  size  of  bivalve  species  whose  eggs  were  unavail- 
able because  of  extinction  or  the  inaccessability  of  their 
habitat.  Estimates  of  egg  size  could  also  be  used  to  cat- 
egorize the  developmental  strategy  of  a  bivalve  as  lethici- 
trophic  ( 150-200 /zm  diameter  eggs),  mi.xed  (90-140  ^m) 
or  planktotrophic  (40-85  ^m)  (Ockelmann,  1962).  Small- 
er differences  in  the  diameter  of  eggs  (<  50  ^m)  within 
Mercenaria  taxa  and  other  planktotrophic  species  have 
been  observed  (Bricelj  &  Malouf,  1980;  Barber  &  Blake, 
1983;  Heffernan  et  al..  1989;  Goodsell,  1991).  Intraspe- 
cific  differences  in  egg  size  have  been  attributed  to  the 
gametogenic  state  of  the  spawning  female  (Gallagher  & 


'  Technical  contribution  No.  3188  of  South  Carolina  Exper- 
iment Station,  Clemson  University. 


Mann,  1986).  Small  differences  in  egg  size  (<10  fim) 
may  be  reflected  in  the  prodissoconch  I  length  of  Mer- 
cenaria, consequently  allowing  inferences  from  larval 
assemblages  about  the  condition  of  the  female  parent 
and  the  survival  potential  of  its  eggs. 

Developmental  markers,  such  as  prodissoconch  I,  re- 
main visible  with  continued  growth  (figure  1)  and  their 
size  can  be  compared  to  later  shell  features  (e.g.,  prodis- 
soconch II).  In  Mercenaria  these  shell  markers  delineate 
shell  margin  at  the  first  shelled  stage  and  at  metamor- 
phosis to  a  benthic  existence.  Although  molluscan  larvae 
have  been  shown  to  delay  metamorphosis  until  suitable 
environmental  conditions  are  found  (Bayne,  1965;  Crisp, 
1974)  no  corresponding  increase  in  shell  length  associated 
with  the  delay  has  been  noted  (Loosanoff,  1959). 

Pechenik  and  Lima  (1980)  demonstrated  an  inverse 
correlation  between  growth  rate  and  length  of  larval 
period  of  the  common  Atlantic  slippersnail,  Crepidula 
jornicata  (Linnaeus,  1758).  They  hypothesized  that  length 
of  larval  life  in  the  slippersnail  is  limited  by  the  rate  of 
development  toward  a  "predetermined  endpoint'  and 
that  such  an  endpoint  might  also  exist  in  bivalves.  Early 
studies  by  Loosanoff  (1959)  demonstrated  that  when  the 


Figure  I.  Scanning  electron  photomicrograph  of  a  M.  cam- 
pechiensis juvenile  753  Mm  in  shell  length  illustrating  prodis- 
soconch I  (P  I)  and  prodissoconch  II  (P  II)  boundaries. 


Page  120 


THE  NAUTILUS,  Vol.  106,  No.  3 


length  of  the  larval  period  was  increased  by  decreasing 
the  culture  temperature  of  M.  mercenaria  (Linnaeus, 
1758)  larvae  there  was  no  corresponding  increase  in  shell 
length,  supporting  Pechenik  and  Lima  s  "endpoint"  ar- 
gument. The  endpoint  of  larval  bivalve  life  may  be  as- 
sociated with  the  loss  of  feeding  ability  that  occurs  with 
the  degeneration  of  the  velum  and  ciliary  feeding  tracts 
(Bayne,  1965).  Differences  in  prodissoconch  II  length 
may  prove  useful  in  detecting  the  difference  in  larval 
endpoints  among  Mercenaria  taxa. 

During  the  course  of  investigations  into  the  genetics 
of  Mercenaria  we  noted  that  eggs  produced  by  any  sin- 
gle, laboratory  conditioned  female  during  a  given  spawn- 
ing event  were  remarkably  consistent  in  diameter.  The 
availability  of  these  consistently  sized  eggs  from  known 
parents  allowed  us  (I)  to  examine  the  relationship  be- 
tween egg  diameter  and  prodissoconch  I  length  in  M. 
mercenaria,  (2)  to  compare  the  prodissoconch  I  and  II 
lengths  of  offspring  from  crosses  involving  M.  campe- 
chiensis  (Gmelin,  1791)  and  M.  campechiensis  texana 
(Dall,  1902)  females  with  M.  campechiensis,  M.  cam- 
pechiensis texatia  and  M.  mercenaria  males;  and  (3)  to 
determine  whether  or  not  prodissoconch  II  lengths  reflect 
taxonomic  differences  in  parental  stocks. 

MATERIALS  AND  METHODS 

Collection  of  adult  Mercenaria  and  controlled  spawning 
were  achieved  using  methods  previously  described  by 
Goodsell  et  al.  (1992).  Briefly,  adults  were  collected  from 
geographically  isolated  areas.  M.  mercenaria  were  col- 
lected from  Milford,  Connecticut;  M.  campechiensis  from 
a  site  near  Appaiachicola,  Florida;  and  M.  campechiensis 
texana  from  a  location  near  Galveston,  Texas.  Adults 
were  conditioned  at  19  °C  for  six  weeks  prior  to  spawning 
and  were  induced  to  spawn  individually  using  thermal 
shock  and  the  addition  of  pasteurized  sperm.  Eggs  were 
counted  (40  x  )  and  100  eggs  from  each  spawning  female 
were  measured  with  an  ocular  micrometer  at  100  x 
using  a  compound  microscope.  Eggs  categorized  as  70 
fim  had  a  minimum  diameter  of  70  tim,  maximum  di- 
ameter of  75  ^m,  mean  <73  and  standard  deviation 
<2.0.  Those  categorized  as  80  ixm  had  a  minimum  di- 
ameter of  80  nm,  maximum  diameter  of  85  /xm,  mean 
<83  and  standard  deviation  <2.0.  Sperm  concentrations 
were  estimated  by  spectrophotometry  (Bricelj,  1979).  To 
insure  optimal  gamete  condition,  all  eggs  were  fertilized 
within  one  hour  of  spawning  at  the  rate  of  1,000  sperm 
ml'  (Goodsell,  1991). 

Egg  Size  and  Prodissoconch  I  Length  in 

M.  MERCENARIA 

Fertilized  70  and  80  ixm  eggs  of  M.  mercenaria  were 
reared  for  48  hours  in  1  ^m  filtered,  UV  treated  seawater 
at  a  density  <20  eggs  ml  '.  A  mixed  algal  diet  (25,000 
cells  ml"')  was  added  to  the  culture  at  24  hours.  After 
48  hours  larvae  were  viewed  with  light  microscopy  ( 100  x  ) 
to  ensure  that  the  prodissoconch  I  boundary  was  clearly 


visible.  Larvae  were  collected  on  a  44  /um  mesh  sieve 
and  treated  with  sodium  hypochlorite  and  distilled  water 
to  disarticulate  the  valves  and  remove  soft  tissues.  Dis- 
articulated valves  from  each  species  were  mounted,  shell 
margin  downward,  on  double  stick  tape  attached  to  a 
glass  slide.  Prodissoconch  I  lengths  (n  =  30)  from  each 
egg  size  category  were  measured  with  an  ocular  mi- 
crometer on  a  compound  microscope  (lOOx).  Analysis 
of  variance  by  general  linear  model  was  used  to  examine 
the  effect  of  egg  diameter  on  prodissoconch  I  length  (SAS 
Institute,  1985). 


Prodissoconch  I  and  II  Length  of 
Mercenaria  Taxa 

Larvae  were  reared  for  15  days  after  prodissoconch  I 
was  achieved  at  a  density  of  one  larva  ml  ',  to  minimize 
the  effects  of  crowding.  Simultaneous  spawning  attempts 
resulted  in  crosses  between  female  M.  campechiensis  (n 
=  3)  and  M.  campechiensis  texana  (n  =  2)  and  male  M. 
campechiensis  (n  =  2),  M.  campechiensis  texana  (n  = 
1 )  and  M.  mercenaria  (n  =  3).  No  M.  mercenaria  females 
spawned  during  the  trials.  Egg  diameters  and  prodisso- 
conch I  and  II  lengths  were  measured.  Egg  diameters  (n 
=  100)  from  each  spawning  female  were  compared  with 
an  analysis  of  variance  by  general  linear  model  to  insure 
that  egg  sizes  did  not  vary  significantly  between  species. 
A  measure  of  growth  after  the  deposition  of  the  prodisso- 
conch I  boundary  was  calculated  for  each  larva  (AP  = 
prodissoconch  II  length  —  prodissoconch  I  length).  Pro- 
dissoconch I  and  II  lengths  and  AP  for  each  parental 
cross  were  compared  with  analysis  of  variance  by  general 
linear  model  and  differences  among  crosses  were  sepa- 
rated using  least  square  difference  (SAS  Institute,  1985). 


RESULTS 

Egg  Diameter  and  Prodissoconch  I  Length  in 

M.  MERCENARIA 

Embryos  derived  from  80  um  eggs  of  M.  mercenaria 
(mean  =  80.9  ^"11,  SE  ±  0.32  um)  produce  larvae  with 
significantly  greater  (P  <  0.0001 )  prodissoconch  I  lengths 
(109.7  ^m  ±  0.7;  range  102-119)  than  prodissoconch  I 
lengths  (101.9  jum  ±  0.7;  88-109)  of  those  larvae  derived 
from  70  ^m  eggs  (70.8  ^m  ±  0.15). 


Prodissoconch  I  and  II  Length  of 
Mercenaria  taxa 

Prodissoconch  I  lengths  from  crosses  using  eggs  of  M. 
campechiensis  and  M.  campechiensis  texana  (egg  di- 
ameter =  75.4  ixm  ±  0.10)  were  not  significantly  differ- 
ent (table  1).  However,  significant  differences  (P  <  0.05) 
in  prodissoconch  II  length  were  found  among  the  crosses. 
Metamorphosing  larvae  from  crosses  involving  M.  mer- 
cenaria males  were  significantK  longer  than  from  males 
of  M.  campechiensis  texana  and  M.  campechiensis.  While 


J.  G.  Goodsell  and  A.  G.  Eversole,  1992 


Page  121 


not  statistically  significant,  it  is  interesting  to  note  that 
tlie  offspring  of  female  M.  campechiensis  were  longer 
than  offspring  of  female  A/,  campechiensis  texana.  No 
difference  in  prodissoconch  II  lengths  of  larvae  from  the 
male  parents  of  the  two  M.  campechiensis  taxa  was 
detected  (Table  1).  As  expected,  the  growth  (AP)  differ- 
ences among  crosses  were  similar  to  those  differences 
observed  with  the  prodissoconch  II  lengths  Crosses  in- 
volving male  M.  mercenaria  demonstrated  the  greatest 
amount  of  shell  growth. 

DISCUSSION 

Differences  in  prodissoconch  I  length  was  detected  with 
differences  in  egg  size  as  small  as  10  nm  in  M.  merce- 
naria, however,  when  similarly  sized  eggs  were  used  no 
significant  differences  in  prodissoconch  I  length  were 
found  among  the  closely  related  Mercenaria  taxa.  Prodis- 
soconch I  lengths  appear  to  vary  consistently  with  egg 
size  probably  from  some  minimum  egg  size.  Gallagher 
and  Mann  (1986)  suggested  that  a  minimum  threshold 
lipid  level  (=  egg  size)  was  necessary  to  achieve  normal 
embryogenesis.  Knowledge  of  the  relationship  between 
egg  size  and  prodissoconch  I  length  might  provide  a 
technique  for  predicting  the  threshold  size  of  eggs  pro- 
duced by  a  female  and  the  survival  potential  of  its  larvae. 

Loosanoff  (1959)  observed  that  the  shell  lengths  of 
metamorphosing  M.  mercenaria  and  M.  campechiensis 
larvae  from  mass  spawnings  were  similar,  suggesting  that 
these  taxa  shared  a  similar  larval  endpoint  (Pechenik  & 
Lima,  1980).  In  contrast  with  Loosanoff's  (1959)  earlier 
study,  we  found  differences  in  the  mean  prodissoconch 
II  length  (=  shell  length  at  metamorphosis)  among  off- 
spring sired  by  M.  mercenaria  and  M.  campechiensis. 
These  differences  in  prodissoconch  II  length  between  the 
two  species  lend  support  to  the  placement  of  the  texana 
subspecies  within  M.  campechiensis.  Examination  of  the 
distance  between  the  prodissoconch  I  and  II  boundaries 
(AP)  demonstrated  that  differences  in  prodissoconch  II 
lengths  were  a  function  of  growth  differences  among  the 
taxa  after  the  prodissoconch  I  boundary  had  been  de- 
posited. The  sampling  technique  in  our  study  was  dif- 
ferent from  that  of  Loosanoff  (1959).  Loosanoff  s  (1959) 
results  were  based  on  mass  spawnings  with  no  attempt 
to  limit  the  range  of  egg  diameter.  Furthermore,  in  Loo- 
sanoff's (1959)  study,  shell  length  at  metamorphosis  was 
determined  by  measuring  the  length  of  intact  larvae  as 
soon  as  they  were  observed  to  have  lost  their  vela.  In  our 
study,  shell  length  at  metamorphosis  was  determined  by 
measuring  length  at  the  prodissoconch  Il/dissoconch 
boundary  on  disarticulated  values. 

Prodissoconch  I  length  does  appear  to  be  a  good  in- 
dicator of  egg  diameter.  Also,  prodissoconch  II  lengths 
(predetermined  endpoint)  follow  taxonomic  lines  indi- 
cating genetic  input  as  suggested  by  Pechenik  and  Lima 
(1980).  Finally,  prodissoconch  I  and  II  lengths  should 
prove  valuable  in  determining  the  condition  of  known 
parents  and  the  relative  genetic  contributions  of  the  par- 
ents in  reciprocal  crosses. 


Table  1 .  Mean,  standard  error,  range  and  results  of  least  square 
difference  test-"  for  prodissoconch  I  (P  I)  and  II  (P  II)  lengths 
(^m)  and  growth  (AP),  measured  as  prodissoconch  II  —  pro- 
dissoconch I,  by  cross.  Means  not  sharing  the  same  letter  are 
significantK  different 


Prodissoconch  I  length 

LSD 

(;:ross'' 

n 

P  1 

+ 

SE 

Range 

Gi 

rouping 

TC 

100 

104.1 

± 

0.34 

91-107 

A 

TT 

50 

103.9 

+ 

0.54 

99-112 

A 

CT 

7,5 

103.7 

+ 

0.47 

91-116 

A 

TM 

150 

102.4 

+ 

0..35 

91-116 

A 

c:m 

225 

101.8 

+ 

0.29 

91-116 

A 

cc 

1.50 

101.4 

-1- 

0.37 

91-116 

A 

Prodissoconch  II  1 

ength 

LSD 

Cross 

11 

P  II 

+ 

SE 

Range 

Gi 

rouping 

CM 

225 

260  0 

± 

0.91 

224-290 

A 

TM 

150 

256.2 

± 

0.72 

22.3-273 

A 

B 

CT 

75 

246.0 

+ 

1.27 

21.5-282 

B     C 

TT 

50 

243.1 

+ 

1.27 

224-265 

C 

CC 

150 

243.0 

+ 

0.87 

215-273 

C 

TC 

100 

240  1 

± 

0  99 

199-261 

C 

Growth 

(AP) 

LSD 

Cross 

n 

AP 

+ 

SE 

Range 

Grouping 

CM 

225 

158.3 

+ 

0.89 

124-191 

A 

TM 

1.50 

1.53.8 

+ 

0.80 

116-169 

A 

B 

CT 

75 

142.4 

± 

1  34 

112-182 

B     C 

CC 

1.50 

141.6 

+ 

0.85 

107-174 

C 

TT 

50 

139.2 

+ 

1.24 

124-165 

C 

TC 

100 

1.36.0 

+ 

1.03 

91-162 

C 

^  Alpha  =  0.05,  df  =  8. 

^'  Abbreviations;  M.  campechiensis  pure  strain  (CC),  AT  cam- 
pechiensis texana  pure  strain  (TT),  hybrid  from  M.  campe- 
chiensis female  x  M.  mercenaria  male  (CM),  M.  campechien- 
sis texana  female  x  M.  mercenaria  male  (TM),  M. 
campechiensis  female  x  M.  campechiensis  texana  male  (CT), 
reciprocal  hybrid  (TC)  in  descending  order  of  size. 


ACKNOWLEDGMENTS 

The  authors  wish  to  thank  William  Arnold,  Robert  Bis- 
ker,  Thomas  Bright,  Paul  Chanley,  James  Moore  and 
Randal  Walker  for  their  aid  in  the  collection  of  the  var- 
ious brood  stocks.  Staff  support  and  culture  facilities  were 
provided  by  the  Waddell  Mariculture  Center,  Bluffton, 
South  Carolina.  Dr.  Lawrence  Grimes  assisted  in  the 
statistical  analyses.  The  research  was  supported  b\  grants 
from  South  Carolina  Sea  Grant  Consortium  and  South 
Carolina  Agricultural  Experiment  Station. 

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THE  NAUTILUS  106(3):  123-124,  1992 


Page  123 


Harpa  cabriti  Fischer,  1860,  a  replacement  name  for 
Harpa  ventricosa  Lamarck,  1816  (Gastropoda:  Harpidae) 


Flarald  A.  Rehder 

Department  (it  Iinertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Wasiiingtoii,  DC;  20560,  USA 


ABSTRACT 

Harpa  ventricosa  Lamarck,  1816,  is  preoccupied  In  Harpa 
ventricosa  Lamarck,  1801,  a  name  that  is  here  (letermined  to 
he  an  objective  junior  synonym  of  Harpa  major  Hodiiig,  1798 
The  name  Harpa  ventricosa  Lamarck,  1816,  used  for  a  western 
Indian  Ocean  species,  should  be  replaced  by  Harpa  cabriti 
Fischer,  1860,  a  name  based  on  an  immature  specimen  formerly 
considered  a  junior  synonym  of  Harpa  ventricosa  Lamarck, 
1816,  non  Lamarck,  1801. 

Key  words:  Harpidae,  Harpa,  nomenclature 


In  1986  Richard  E.  Petit  and  I  submitted  an  application 
to  the  International  (,'oininission  of  Zoological  Nomen- 
clature (ICZN)  proposing  that  the  specific  name  Harpa 
articuiaris  Lamarck,  1822,  be  conserved  and  placed  on 
the  Official  List  of  Specific  Names  in  Zoology  (Rehder 
&  Petit,  1987). 

When  Miss  Ruth  A.  Cooper  of  the  ICZN  was  drafting 
the  Opinion  on  this  case  she  encountered  a  problem  with 
our  use  of  the  name  Harpa  ventricosa  Lamarck,  1816, 
noting  that  there  was  an  earlier,  validly  published,  Harpa 
ventricosa  Lamarck,  1801  (Lamarck,  1801:79).  That  por- 
tion of  the  case  involving  Harpa  ventricosa  Lamarck, 


Figure  1.    Holot>pe  of  Harpa  cahnti  Fischer,  1860.  BM(NH)  1899.8.22.126.  No  locality  data.  Scale  bar  =  3  cm. 


Page  124 


THE  NAUTILUS,  Vol.  106,  No.  3 


1816,  and  Harpa  urnifortnis  Perry.  1811,  was  therefore 
withdrawn  from  the  petition,  being  considered  a  purely 
la.\onomic  question,  and  Opinion  1518,  conserving  the 
specific  name  Harpa  articularis  Lamarck,  1822,  was 
published  in  1989. 

Through  a  now  inexplicable  oversight  I  had  over- 
looked, in  my  monograph  of  the  family  Harpidae  (Reh- 
der,  1973),  the  problem  of  homonymy  involved  in  Harpa 
ventricosa  Lamarck,  1801,  versus  Harpa  ventricosa  La- 
marck, 1816.  The  present  paper  was  written  to  clarify 
this  matter. 

In  1801  Lamarck  published  his  "Systeme  des  Animaux 
sans  Vertebres",  wliich  in  essence  is  a  treatise  on  the 
cla.ssification  of  the  invertebrates,  listing  the  genera  of 
the  various  groups,  each  genus  being  briefly  diagnosed, 
with  one  or  more  species  cited  as  examples.  The  species 
are  without  diagnoses  but  have  bibliographic  references 
to  previous  descriptions  or  figures. 

His  genus  Harpa  (Lamarck,  1801:79)  is  exemplified 
by  one  taxon,  Harpa  ventricosa  Lamarck,  a  new  name 
for  the  species  illustrated  by  Lister  (1770:pl.  992,  fig.  55), 
and  by  Martini  (1777:pl.  119,  fig.  1090).  Lister's  figure 
represents  Harpa  harpa  (Linne,  1758),  as  noted  by  La- 
marck (1822:256)  and  Dodge  (1956:196-198).  Martini's 
figure  is  representative  of  Harpa  nmjor  Roding,  1798, 
and  was  the  figure  on  which  Roding  based  his  species.  I 
therefore  propose  that  the  specimen  illustrated  in  Mar- 
tini's figure  serve  as  the  type  specimen  of  Harpa  ven- 
tricosa Lamarck,  1801.  I  did  this  in  the  case  of  Roding's 
name  (Rehder,  1973:247).  This  makes  Lamarck's  name 
an  objective  junior  synonym  of  Harpa  major  Roding, 
1798. 

Harpa  ventricosa  Lamarck,  1816,  isa  junior  homonym 
but  not  a  junior  synonym  of  Harpa  ventricosa  Lamarck, 
1801.  Therefore,  the  species  that  had  been  called  Harpa 
ventricosa  Lamarck,  1816,  by  me  (Rehder,  1973:251) 
and  numerous  other  authors  beginning  with  Lamarck 
(1822:255),  is  in  need  of  a  replacement  name. 

I  therefore  propose  that  this  species,  which  is  restricted 
to  the  western  part  of  the  Indian  Ocean  and  the  Red 
Sea,  bear  the  name  of  Harpa  cabriti  Fischer,  1860.  The 
type  specimen  of  H.  cabriti,  which  I  listed  in  the  syn- 
onymy of  Harpa  ventricosa  Lamarck,  1816  (Rehder, 
1973:252),  is  a  juvenile  or  immature  specimen,  and  is  in 


the  collections  of  The  Natural  History  Museum,  London. 
Through  the  kindness  of  Ms.  Kath>  Way  I  am  able  to 
illustrate  this  holotype  [BM(NH)  1899.8.22.126]  (fig.l). 

Harpa  urniformis  Perry,  1811,  although  listed  in  the 
synonymy  of  H.  ventricosa  Lamarck  by  Deshayes  (1843), 
is  not  identifiable  and  is  not  considered  to  be  a  senior 
synonym  of  H.  cabriti  Fischer. 

LITERATURE  CITED 

Deshayes,  G.  P.  1843.  Histoire  naturelle  des  Animaux  sans 
Vertebres.  Ed.  2.  Volume  9.  Paris.  725  pp. 

Dodge,  H.  1956.  A  historical  review  of  the  mojlusk.s  of  Lin- 
naeus. Part  4.  The  genera  Bticcinum  and  Stromlms  of  the 
class  Gastropoda  Bulletin  of  the  American  Museum  of 
Natural  History  1 1 1(3):  15:3-312. 

Fischer,  P.  1860.  Descriptions  d'especes  nouvelles.  Journal  de 
Conchyliologie  8(2):208-211,  pi.  4,  figs.  1-2,  6-8. 

International  Commission  on  Zoological  Nomenclature  [ICZN] 
1989  Opinion  1518  Harpa  articularis  Lamarck.  1822 
(Mollusca:  Clastropoda):  specific  name  conservation.  Bul- 
letin of  Zoological  Nomenclature  46(1):.53. 

Lamarck,  j.  B  P.  A.  de  M.  de  1801.  Systeme  des  Animaux 
sans  Vertebres.  Deterville,  Paris,  viii  -I-  4.32  p. 

Lamarck,  J.  B.  P  A.  de  M.  de  1816.  [in  1791-1823].  Tableau 
Encyclopedique  et  Methodique  des  trois  Regnes  de  la  Na- 
ture. Vingt-Troisieme  Partie.  Mollusqueset  Polypes  Aivers. 
Liste  des  objets  representes  dans  les  planches  de  cette  liv- 
rai.son.  Panchouche,  Paris.  16  pp.  pls.391-488. 

Lamarck,  J  B.  P.  A  de  M.  de  1822  [in  181.5-1822].  Histoire 
Naturelle  des  Animaux  sans  Vertebres.  PrivateK  pub- 
lished, Paris,  vol.  7  [of  7],  711  p. 

Lister,  M  1770.  Historiae  sive  Synopsis  Methodicae  C^onchy- 
liorum.  .  .  Editio  Altera.  Recensuit  et  indicibus  auxit  Gu- 
liehnus  Huddesford.  Clarendon  Press,  Oxford,  iv  +  [96] 
pp.  439  pis.  [1082  figs.]. 

Martini,  F,  H.  W.  1777.  [in  1769-1829].  NeuesSystematisches 
Conchvlien-Cabinet.  vol  3  [If  12],  vi  +  434  p.  pis.  66- 
121. 

Perry.  G.    181 1.    Conchology,  or  the  Natural  History  of  Shells; 
Containing  a  new  arrangement  of  Genera  and  Species. 
W.  Miller,  London,  4  pp.  61  pis. 

Rehder,  H.  A.  1973.  The  family  Harpidae  of  the  world.  Indo- 
Pacific  Mollusca  .3(16):207-274. 

Rehder,  H.  A.  and  R.  E.  Petit,  1987.  Case  2548.  Harpa  ar- 
ticularis Lamarck,  1822  (Mollusca,  Gastropoda):  proposed 
conservation  of  the  specific  name.  Bulletin  of  Zoological 
Nomenclature  44(l):19-20. 


THE  NAUTILUS  106(3):125-126,  1992 


Page  125 


Obituary: 

Rae  Baxter,  1929-91 


James  H.  McLean 

Los  Angelt'S  Oiunty  Museum  of 

Natural  Histor\ 
900  Exposition  Boulf\ard 
Los  Angeles,  CA  90007,  USA 


Alaskan  malacology  suffered  a  great  loss  with  the  death 
of  Rae  Baxter,  the  most  knowledgeable  expert  on  Alaskan 
mollusks,  on  22  Marcli  1991  at  the  age  of  61.  Rae  was  a 
Field  Associate  in  Malacolog>  of  the  Los  Angeles  County 
Museum  of  Natural  History.  He  is  survived  by  his  wife 
Sera,  daughter  Lynx,  and  son  Brant. 

Rae  was  born  on  30  March  1929  in  Compton,  Cali- 
fornia. He  earned  a  Bachelor  of  Science  degree  in  fish- 
eries from  Humboldt  State  University,  California,  in  1955. 
During  his  days  at  Humboldt,  he  became  a  student  and 
collector  of  Pacific  coast  mollusks.  While  there  he  met 
Sera  Miller,  a  native  of  Washington  State,  whom  he  mar- 
ried in  1955.  Finding  that  northern  climates  were  to  their 
liking,  Rae  and  Sera  moved  to  Alaska  after  graduation. 

From  1956  to  1962,  Rae  worked  for  the  US  Fish  and 
Wildlife  Service  out  of  Anchorage.  Winters  during  this 
period  were  spent  camping  and  trapping,  during  which 
Rae  and  Sera  learned  much  about  cold  weather  survival 
techniques.  In  1960,  the  Baxters  spent  five  months  trav- 
eling in  their  sport  boat  from  Kachemak  Ba\  through 
southeastern  Alaska  and  British  C^olumbia  to  the  Stilla- 
quamish  River  in  Washington,  camping  on  shore  each 
night.  Mollusks  were  collected  at  every  opportunity  and 
mailed  back  to  Alaska.  Sediment  samples  often  were 
dried  over  the  same  stove  used  for  cooking. 

In  1962  Rae  joined  the  Alaska  Department  of  Fish  and 
Game,  being  stationed  first  at  Cordova.  During  this  pe- 
riod the  Baxters  built  an  A-frame  dwelling  for  use  as  a 
base  for  commercial  salmon  fishing  at  Kasitsna  Bay,  on 
the  southern  shore  of  Kachemak  Bay  in  Lower  Cook 
Inlet.  Sera  was  in  charge  of  this  venture,  which  continued 
each  summer,  with  the  help  of  their  children.  In  1966 
Rae  was  transferred  to  the  Fish  and  Game  laboratory  at 
Bethel,  on  the  Kuskwokwim  River  in  western  Alaska. 
The  family  lived  at  Bethel  during  the  off  season,  return- 
ing to  Kasitsna  Bay  each  summer,  until  1983,  when  Rae 
retired  and  returned  to  Kasitsna  Bay  year  round.  In  1989 
Rae  finished  building  a  laboratory  for  his  new  work  as 
a  marine  biological  consultant,  and  especially  for  re- 
search on  his  mollusk  collection,  for  which  he  had  as- 
sembled a  good  working  library. 


Although  Rae's  work  for  the  state  of  Alaska  concerned 
the  commercially  important  fishes  and  invertebrates,  his 
compelling  interest  was  the  systematics  and  distribution 
of  the  molluscan  fauna  of  Alaska.  Over  many  years  of 
field  work  throughout  Alaska,  Rae  became  familiar  with 
the  mollusks  of  the  entire  state,  including  the  land  and 
freshwater  species  as  well  as  marine  species.  He  es- 
pecialK  knew  the  molluscan  fauna  of  Kachemak  Bay 
and  would  often  set  out  alone  in  an  open  skiff  for  inter- 
tidal  collecting  and  dredging.  His  spare  time  that  was 
not  devoted  to  collecting  and  studying  mollusks  was  spent 
examining  collections  and  gathering  distributional  rec- 
ords on  visits  to  the  Los  Angeles  County  Museum  of 
Natural  History  and  the  US  National  Museum  of  Natural 
History.  After  his  retirement,  he  spent  many  summers 
aboard  research  vessels  on  long  cruises  in  the  Bering  Sea, 
C^hukchi  Sea,  and  Gulf  of  Alaska,  working  as  a  consultant 
to  the  National  Marine  Fisheries  Service  on  Alaskan  fish- 
ery assessment  surveys.  One  of  these  cruises  was  aboard 
the  Soviet  ship  Novokotovsk  in  the  northwestern  Bering 
Sea  during  the  summer  of  1990.  Mollusks  that  would 
otherwise  have  been  lost  were  retrieved  by  Rae  on  all 
the  cruises. 

At  the  time  of  his  death  he  was  completing  a  detailed 
key  to  the  fishes  of  Alaska  for  publication  by  the  Amer- 
ican Fisheries  Society.  Upon  completion  of  that  project 
he  was  looking  forward  to  being  free  to  pursue  his  study 
of  Alaskan  mollusks,  including  the  description  of  a  num- 
ber of  species  that  he  had  recognized  as  new.  He  had 
started  to  construct  specimen  cases  so  that  his  collection, 
until  then  labeled  with  station  numbers  and  species  code 
numbers,  could  finally  be  assembled  as  a  working  ref- 
erence collection.  Rae's  work  on  mollusks  was  not  to  be 
finished,  but  he  has  left  a  legacy  both  in  his  publications 
and  his  collection. 

In  1983  Rae  published  the  first  edition  of  his  distri- 
butional checklist.  The  Mollusks  of  Alaska,  a  major  proj- 
ect that  was  greatly  facilitated  b>  the  advent  of  personal 
computers,  which  enabled  him  to  produce  the  finished 
text.  Two  subsequent  editions  (1985,  1987)  were  more 
detailed,  treating  a  final  total  of  1,016  species.  Two  spe- 


Page  126 


THE  NAUTILUS,  Vol.  106,  No.  3 


m 

1 

MM3k 

1 

H 

H 

Rae  Baxter  at  Kasitsna  Bay,  Alaska,  in  August  197.3,  screening  mud  dredged  from  his  skiff  (left)  and  picking  salmon  from  a  set  net 
(right). 


cies  were  described  or  coauthored  by  Baxter:  Macoma 
dexioptera  in  1977  and  Spiromoelleria  kachemakensis 
in  1984.  Mollusks  named  in  his  honor  were  Monadenia 
fidelis  baxteriana  Talmadge,  1954,  Anatoma  baxteri  Mc- 
Lean, 1984,  and  Cocculina  baxteri  McLean,  1986. 

Rae  was  curious  about  all  living  things.  He  was  a  keen 
observer  and  was  called  upon  for  information  by  many 
biologists  in  Alaska.  He  often  supplied  live  specimens  for 
the  aquaria  at  the  Pratt  Museum  in  Homer.  He  was 
always  ready  to  assist  the  research  of  many  workers  around 
the  world  with  specimens  and  information  on  Alaskan 
mollusks.  Molluscan  material  from  his  field  work  in  re- 
cent years  was  sent  to  the  Los  Angeles  County  Museum 
of  Natural  History,  where  it  has  greatly  strengthened  the 
Alaskan  holdings.  In  accordance  with  his  wishes,  his  en- 
tire mollusk  collection  has  been  transferred  to  the  Los 
Angeles  County  Museum  of  Natural  History,  where  it  is 
to  be  integrated  into  the  research  collection  and  made 
available  for  study. 

I  was  privileged  to  stay  with  the  Baxter  family  at 
Kasitsna  Bay  for  a  week  in  August  of  1973,  a  wonderful 


experience  both  for  the  exposure  to  the  rich  molluscan 
fauna  and  the  zeal  with  which  Rae  sought  to  understand 
it.  Rae  was  a  kind  and  generous  man  who  is  sorely  missed. 

PUBLICATIONS  BY  RAE  BAXTER 
ON  MOLLUSKS 

Baxter,  R.  1977.  A  new  Alaska  Macoma  (Mollusca:  Bivalvia). 
The  Veliger  19(3);272-276. 

Baxter,  R.  1983.  Mollusks  of  .\laska,  a  listing  of  all  mollusks, 
freshwater,  land  and  marine,  reported  from  the  State  of 
Alaska,  with  known  locations  of  type  specimens,  maximum 
sizes,  and  marine  depths  inhabited.  Privately  published, 
xvii  -I-  69  p 

Baxter,  R.  1985.  Mollusks  of  Alaska,  2nd  ed.  Privately  pub- 
lished, 114  +  xxxiii  p. 

Baxter,  R.  1987.  Mollusks  of  Alaska,  3rd  ed.  Shells  &  Sea  Life, 
Bayside,  California.  163  p. 

Baxter,  R.  and  J.  H  McLean.  1984.  The  genera  Moelleria 
Jeffreys,  1865,  and  Spiromoelleria,  gen.  nov.,  in  the  North 
Pacific,  with  description  of  a  new  species  of  Spiromoelleria 
(Gastropoda:  Turbinidae).  The  Veliger  27(2):219-226. 


INSTRUCTIONS  TO  AUTHORS 


THE  NAUTILUS  publishes  papers  on  all  aspects  of  the 
biology  and  systematics  of  mollusks.  Manuscripts  de- 
scribing original,  unpublished  research  as  well  as  review 
articles  will  be  considered.  Brief  articles,  not  exceeding 
1000  words,  will  be  published  as  notes  and  do  not  re- 
quire an  abstract.  Notices  of  meetings  and  other  items 
of  interest  to  malacologists  will  appear  in  a  news  and 
notices  section. 

Manuscripts:  Each  original  manuscript  and  accompa- 
nying illustrations  should  be  submitted  in  triplicate.  Text 
must  be  typed  on  one  side  of  8'/2  x  11  inch  white  paper, 
double  spaced  throughout  (including  literature  cited,  ta- 
bles and  figure  captions),  with  at  least  1  inch  of  margin 
on  all  sides.  All  pages  must  be  numbered  consecutively. 
If  printed  on  a  word  processor,  the  right  margin  should 
be  ragged  rather  than  justified.  Authors  should  follow 
the  recommendations  of  the  Council  of  Biology  Editors 
Style  Manual,  which  is  available  from  the  Council  of 
Biology  Editors,  Inc.,  9650  Rockville  Pike,  Bethesda,  MD 
20814,  U.S.A.  The  first  mention  of  a  scientific  name  in 
the  text  should  be  accompanied  by  the  taxonomic  au- 
thority, including  year.  Latin  names  and  v\ords  to  be 
printed  in  italics  must  be  underlined;  leave  other  indi- 
cations to  the  editor.  Metric  and  Celsius  units  are  to  be 
used. 

The  sequence  of  sections  should  be:  title  page,  abstract 
page,  introduction,  materials  and  methods,  results,  dis- 
cussion, acknowledgements,  literature  cited,  tables,  fig- 
ure captions,  figures.  The  title  page  should  include  the 
title,  author's  name(s)  and  address(es).  The  abstract  page 
should  contain  the  title  and  abstract,  which  should  sum- 
marize in  250  words  or  less  the  scope,  main  results  and 
conclusions  of  the  paper.  The  abstract  may  be  followed 
by  a  maximum  of  8  key  words.  All  references  cited  in 
the  text  must  appear  in  the  literature  cited  section  and 
vice  versa.  In  the  literature  cited  section,  all  authors 
must  be  fully  identified  and  listed  alphabeticalh'.  Follow 
a  recent  issue  of  THE  NAUTILUS  for  bibliographic  style, 
noting  that  journal  titles  must  be  unabbreviated.  Infor- 
mation on  plates  and  figures  should  be  cited  only  if  not 
included  in  the  pagination.  Tables  must  be  numbered 
and  each  placed  on  a  separate  sheet.  A  brief  legend  must 
accompany  each  table.  Captions  for  each  group  of  illus- 
trations should  be  typed  on  a  separate  sheet  and  include 
a  key  to  all  lettered  labeling  appearing  in  that  group  of 
illustrations. 


All  line  drawings  must  be  in  black,  high  quality  ink, 
clearly  detailed  and  completely  labeled.  Photographs 
must  be  on  glossy,  high  contrast  paper.  All  figures  are 
to  be  consecutively  numbered  (figs.  1,  2,  3,  ...  ,  NOT 
figs.  la.  lb,  Ic,  .  .  .  NOR  plate  1,  fig.  1  .  .  .).  Illustrations 
must  be  arranged  in  proportions  that  will  conform  with 
the  width  of  a  page  (6H  inches  or  171  mm)  or  a  column 
(SVi  inches  or  82  mm).  The  maximum  size  of  a  printed 
figure  is  6%  by  9  incries  or  171  by  228  mm.  All  illus- 
trations must  be  fully  cropped,  mounted  on  a  firm,  white 
backing,  numbered,  labeled  and  camera  ready.  The  au- 
thor's name,  paper  title  and  figure  number(s)  should  ap- 
pear on  the  back.  Original  illustrations  must  be  between 
one  and  two  times  the  desired  final  size.  It  is  the  author's 
responsibility  that  the  line  weight  and  lettering  are  ap- 
propriate for  the  desired  reduction.  Original  illustrations 
will  be  returned  to  the  author  if  requested.  Color  illus- 
trations can  be  included  at  extra  cost  to  the  author. 

Voucher  Material:  Deposition  of  type  material  in  a 
recognized  public  museum  is  a  requirement  for  publi- 
cation of  papers  in  which  new  species  are  described. 
Deposition  of  representative  voucher  specimens  in  such 
institutions  is  strongly  encouraged  for  all  other  types  of 
research  papers. 

Processing  of  Manuscripts:  Upon  receipt,  every  manu- 
script is  acknowledged  and  sent  for  critical  review  by  at 
least  two  referees.  These  reviews  serve  as  the  basis  for 
acceptance  or  rejection.  Accepted  manuscripts  are  re- 
turned to  the  author  for  consideration  of  the  reviewers' 
comments.  A  finalized  version  of  the  manuscript  is  re- 
turned to  the  editor  and  sent  to  press.  Two  sets  of  proofs 
are  sent  to  the  author  for  correction.  Changes  other  than 
typesetting  errors  will  be  charged  to  the  author  at  cost. 
One  set  of  corrected  proofs  should  be  sent  to  the  editor 
as  soon  as  possible.  Authors  with  institutional,  grant  or 
other  research  support  will  be  billed  for  page  charges  at 
the  rate  of  $60.00  per  printed  page. 

An  order  form  for  reprints  will  accompany  the  proofs. 
Reprints  may  be  ordered  through  the  editor. 

Manuscripts,  corrected  proofs  and  correspondence  re- 
garding editorial  matters  should  be  sent  to:  Dr.  M.G. 
Harasewych,  Editor,  Division  of  Mollusks,  NHB  stop  118, 
National  Museum  of  Natural  History,  Smithsonian  In- 
stitution, Washington,  DC  20560,  USA. 


THIS  PUBLICATION  IS  PRINTED  ON  ACID-FREE  PAPER. 


rHE  NAUTILUS 


Volume  106,  Number  4 
Felnuary  5,  1993 
ISSN  0028-1344 

A  quarterly  devoted 
to  malacology. 


FEB  1 6  133: 


EDITOR-IN-CHIEF 
Dr.  M.  G.  Harasewych 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


ASSOCIATE  EDITOR 
Dr.  R.  Tucker  Abbott 
American  Malacologists,  Inc. 
P.O.  Box  2255 
Melbourne,  FL  32902 


CONSULTING  EDITORS 
Dr.  Riidiger  Bieler 
Department  of  Invertebrates 
Field  Museum  of 
Natural  History 
Chicago,  IL  60605 


Dr.  Robert  T.  Dillon,  Jr. 
Department  of  Biology 
College  of  Charleston 
Charleston,  SC  29424 


Dr.  William  K.  Emerson 

Department  of  Living  Invertebrates 

The  American  Museum  of  Natural 

History 

New  York,  NY  10024 


Dr.  Robert  Hershler 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


Dr.  Richard  S.  Houbrick 
Division  of  Mollusks 
National  Museum  of 
Natural  History 
Smithsonian  Institution 
Washington,  DC  20560 


Mr.  Richard  I.  Johnson 
Department  of  Mollusks 
Museum  of  (comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Aurele  La  Rocque 
Department  of  Geology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  James  H.  McLean 
Department  of  Malacology 
Los  Angeles  County  Museum  of 
Natural  History 
900  E.xposition  Boulevard 
Los  Angeles,  CA  90007 

Dr.  Arthur  S.  Merrill 
%  Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Ms.  Paula  M.  Mikkelsen 
Harbor  Branch  Oceanographic 
Institution,  Inc. 
Ft.  Pierce,  FL  33450 

Dr.  Donald  R.  Moore 

Division  of  Marine  Geology 

and  Geophysics 

Rosenstiel  School  of  Marine  and 

Atmospheric  Science 

University  of  Miami 

4600  Rickenbacker  Causeway 

Miami,  FL  33149 

Dr.  Gustav  Paulay 
Marine  Laboratory 
University  of  Guam 
Mangilao,  Guam  96923 

Mr.  Richard  E.  Petit 

P.O.  Box  30 

North  Myrtle  Beach,  SC  29582 

Dr.  Edward  J.  Petuch 
Department  of  Geology 
Florida  Atlantic  University 
Boca  Raton,  FL  33431 


Dr.  David  H.  Stansbery 
Museum  of  Zoology 
The  Ohio  State  University 
Columbus,  OH  43210 

Dr.  Ruth  D.  Turner 
Department  of  Mollusks 
Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  MA  02138 

Dr.  Geerat  J.  Vermeij 
Department  of  Geology 
University  of  California  at  Davis 
Davis,  CA  95616 


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TH  Et7NAUTI  LUS 


CONTENTS 


Volume  106,  Number  4 

February  5,  1993 

ISSN  0028-1344 


Harald  A.  Rehder 


A  new  species  of  Harpa  from  the  Leeward  Islands  of                      ^  ^      m 
Hawaii      i ._  .  .    127 

Pf'ro^roc'/u/,')  niai/r^-ri,  a  new  species  of  Pleurotomariid         \'■^-^•^,l„    M'M'     >  ■■  ^ - 
from  the  western  Atlantic  (Gastropoda:  Pleurotomariidae)  150 

The  genus  Fulgoraria  (Gastropoda:  Volutidae)  of  the 

northeastern  Kamchatka  Peninsula  and  Sakhalin  Island, 

with  notes  on  the  paleoecology  and  distribution  of  the 

subfamily  Fulgorariinae  in  the  Oligocene  of  the  northern 

Pacific    '  137 

A  new  species  of  Columbellid  Gastropod  from  the  Old 

World  tropics  147 

The  association  between  the  gastropod  Buccinanops 

cochlidium  (Dillwyn,  1817)  and  the  sea  anemone 

Phlyctenanthus  austratis  Carlgren,  1949  in  Patagonian 

shallow  waters  152 

Patterns  of  diversity  and  extinction  in  Transmarian 

Muricacean,  Buccinacean,  and  Conacean  Gastropods     155 

Publication  Dates  of  The  Nautilus      174 


M.  C.  Harasewych 
Timothy  M.  Askew 


Anton  E.  Oleinik 


William  K.  Emerson 


Cuido  Pastorino 


Edward  J.  Petuch 


Eugene  V.  Coan 
M.  G.  Harasewych 


Notice 


SMITHSONIAN    FUNDS    FOR     MALACOLOGY 
STUDENTS 

The  Division  of  Mollusks,  Department  of  Invertebrate 
Zoology,  National  Museum  of  Natural  History,  Smith- 
sonian Institution  announces  the  availability  of  the  Rose- 
water  Fellowship  (up  to  $750)  to  be  awarded  to  graduate 
students  of  systematic  malacology.  This  award  provides 
support  for  students  conducting  systematic  studies  of 
Mollusca  (leading  to  publication)  who  require  access  to 
collections  and  libraries  of  the  Division  of  Mollusks,  Na- 
tional Museum  of  Natural  History.  Funds  can  be  used 
for  travel,  subsistence,  and  research  costs.  Interested  stu- 
dents should  submit  a  succinct  proposal  (1-2  pages),  in- 
cluding budget  with  indication  of  any  matching  funds, 
and  a  supporting  letter  from  faculty  advisor(s).  Appli- 
cation deadline  is  March  15,  1993.  Award(s)  will  be  an- 
nounced on  April  15,  1993.  Applications  should  be  sent 
to: 

Dr.  M.  G.  Harasewych 
Division  of  Mollusks/NHB  stop  118 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC  20560  USA 


THE  NAUTILUS  106(4):127-129,  1993 


Page  127 


A  New  Species  of  Harpa  from  the  Leeward  Islands  of  Hawaii 


Harald  A.  Rehder 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  Histor\ 
Smithsonian  Institution 
Washington,  DC  20560  U.S.A. 


ABSTRACT 

Harpa  goodwini,  new  species,  is  described  from  two  localities 
in  the  Leeward  Islands  of  Hawaii,  French  Frigate  Shoals  and 
Maro  Reef,  where  it  occurs  in  82-228  m.  This  new  species  is 
most  closely  related  to  Harpa  cabritii  Fischer,  1860,  once  known 
as  H.  ventricosa  Lamarck,  1816  (not  Lamarck,  1801). 

Key  words:  Gastropoda;  Harpidae;  Harpa,  new  species;  Ha- 
waiian Islands. 


INTRODUCTION 

On  several  occasions  Mr.  Donald  Dan  has  turned  over 
to  me  for  study  specimens  of  Harpa  received  by  him 
from  correspondents.  One  of  these  lots  represents  a  strik- 
ing new  species  from  the  Leeward  Islands  of  the  Hawaii 
Group.  I  am  grateful  to  Mr.  Dan  for  the  privilege  of 
examining  this  material,  and  for  permitting  me  to  de- 
scribe this  new  species. 

ABBREVIATIONS  FOR  INSTITUTIONS 

AMNH — American  Museum  of  Natural  History.  New 

York 
AMS — Australian  Museum,  Sydney 
BM — Bernice  P.  Bishop  Museum,  Honolulu 
BM(NH) — British  Museum  (Natural  History),  London 
LACM — Los  Angeles  County  Museum  of  Natural  His- 
tory, Los  Angeles 
MCZ — Museum  of  Comparative  Zoology,  Cambridge 
MNHN — Museum  National  d'Histoire  Naturelle,  Paris 
MHNG — Museum  d'Histoire  Naturelle,  Geneva 
NSMT — National  Science  Museum,  Tokyo 
USNM — National  Museum  of  Natural  History,  Smith- 
sonian Institution,  Washington,  DC 

SYSTEMATICS 

Family  Harpidae  Bronn,  1849 
Genus  Harpa  Roding,  1798 
Harpa  goodwini,  new  species 
Figures  1,2;  Table  1. 


Description:  Adult  shell  (figs.  1,2)  60-74  mm  in  length, 
broadly  oval,  outline  of  last  whorl  arcuate.  Protoconch 
conical,  pale  pink,  consisting  of  2V8  to  S'/s  conve.x,  glossy 
whorls.  Postnuclear  whorls  3%-4  in  number,  first  l-l'/2 
whorls  convex,  pale  pink  in  color,  showing  fine  spiral 
cords  at  shoulder  and  below  that  cross  fine  axial  riblets; 
this  sculpture  increasingly  covered  in  subsequent  whorls 
by  the  upper  part  of  the  glaze  that  covers  the  ventral 
wall  at  succeeding  apertures  and  that  extends  up  to  the 
sharp  spines  of  the  axial  ribs  where  they  cross  the  shoulder 
cord.  These  spines  become  increasingly  prominent  and 
accentuate  the  flattened  channel  between  suture  and 
shoulder.  On  the  upper  whorls,  the  axial  riblets  in  the 
channel  are  fine  cords,  but  on  the  body  whorl  they  be- 
come broader  and  flatter.  Body  whorl  with  13-15  strong 
ribs  that  are  erect  with  the  moderately  acute  crest  slanted 
away  from  the  aperture  along  the  apical  half,  but  more 
rounded  and  flattened  on  the  lower  half  of  the  rib.  Ribs 
prominent  and  of  orange  color  in  the  subsutural  channel. 
At  the  shoulder,  ribs  form  acute,  triangular  spines  that 
are  orange  for  most  of  the  whorl,  but  become  white  near 
the  aperture.  The  intercostal  spaces  are,  as  in  most  species 
of  Harpa,  axially  finely  striate. 

Basic  color  of  fresh  specimens  is  pinkish  to  pinkish 
orange,  with  the  spire  whorls  appearing  yellowish  due 
to  the  glaze  covering.  Ribs  on  the  body  whorl  typically 
crossed  by  a  series  of  eight  pairs  of  dark  chestnut  lines 
that  are  generally  more  pronounced  on  the  last  four  or 
five  ribs  and  situated  as  follows:  two  pairs  between  the 
spine  at  the  upper  end  of  each  rib  and  Vt,  of  the  distance 
down  the  length  of  each  rib;  another  two  pairs  along  the 
center  of  the  w  horl;  below  that  a  single  line  (occasionally 
absent),  then  another  pair  of  lines,  and  at  the  base  three 
pairs  of  lines  (the  last  may  be  obscure  or  appear  as  a 
single  line),  making  a  total  of  17  lines.  Between  these 
four  groups  of  lines  are  moderateh  broad  bands  com- 
posed of  a  central  pale  pink  or  orange  pink  band  between 
two  white  bands.  The  intercostal  spaces  are  marked  with 
wavy,  chestnut  brown,  axial  lines,  and,  occasionally,  by 
deep  orange-rose  splotches  in  two  spiral  bands,  one  in 
the  center  of  the  whorl,  the  other  just  below  the  spinose 
shoulder.  Aperture  oval  with  top  of  outer  lip  meeting 
the  parietal  wall  at  a  right  angle,  the  outer  lip  evenly 


Page  128 


THE  NAUTILUS,  Vol.  106,  No.  4 


HA.  Rehder,  1993 


Page  129 


Table  1.    Ilarpa  goaduini.  new  species.  Measurements  of  shell  characters.  Linear  measurements  in  mm 


Length 


Width 


No.  ribs  on 
1)0(I\  whorl 


No.  whorls 


No.  nuclear 
whorls 


Holoty  pe 
Parat\pe  1 
Parat)  pe  2 
Parat)  pe  3 
Parat)  pe  4 
ParatN  pe  5 
Paratype  6 
Paratype  7 
Paratype  8 
Paratype  9 
Paratype  10 
Paratype  1 1 
Paratype  12 

Mean 

Range 

SD 


()3.S 
61.0 
73.4 
74.8 
70.2 
70.3 
69.2 
70.4 
70.1 
66.2 
61.4 
60  4 
60.0 

67.0 
60.0-74.8 

.50 


45.1 

42.7 

57.3 

55.2 

51.1 

52.5 

50.5 

48.1 

50.6 

44.7 

43.0 

43.7 

38.4 

47.9 

38.4-57.3 

\0 

13 
14 
14 
15 
15 
13 
15 
14 
15 
14 
15 
14 

14.3 
13-15 

0.7 


7.00 
7.00 
6.00 
6.16 
6.33 
6.25 
6.25 
6.33 
6.25 
6.12 
5.88 
6.00 
6.25 

6.29 

5.88-7.00 

0  .33 


3  33 

3.25 

2  + 

2.33 

2.50 

2.50 

2.50 

2.50 

2,25 

2.12 

2.12 

2.25 

2..50 

2.51 

2.12-3.33 

0.38 

arcuate.  Parietal  wall  of  aperture  gently  conve.x,  colu- 
mella nearly  straight,  or  slightly  concave,  anal  fasciole 
strongly  ridged  by  flattened  ends  of  the  ribs,  the  sinus 
fairly  deep.  Columella  and  parietal  walls  covered  by  a 
yellowish  glaze,  the  outer  lip  yellow  under  the  axial  ribs, 
with  deep  chestnut  hues  on  the  ribs  showing  tiirough  as 
chestnut  spots.  The  ventral  wail  has  three  chestnut  spots: 
a  strong,  elongate  central  spot  that  slants  apically  into 
the  aperture  above  the  top  of  the  coiinnella;  a  weak,  oval 
or  roughly  triangular  spot  between  tlie  ribs  at  the  top  of 
the  parietal  wall;  and  a  small  spot  at  the  base  of  the 
columella. 

Type  locality:  French  Frigate  Shoals,  Hawaiian  Islands, 
about  166°10'E,  23°45'N,  137  m.  Taken  with  hermit  crabs. 

Material  examined:  Holotype,  USNM  860312,  Paratype 
1,  USNM  860314,  from  the  type  locality;  Paratype  2, 
RM;  Paratype  3,  MCZ;  Paratype  4,  Goodwin  Collection; 
Paratype  5,  AMNH  226438;  Paratype  6,  RM(NH);  Para- 
tvpe  7,  USNM  860315;  Paratype  8,  MHNG;  Paratype  9, 
LACM;  Paratype  10,  AMS;  Paratype  11,  MNHN;'Para- 
type  12,  NSMT;  all  collected  between  French  Frigate 
Shoals  and  Maro  Reef,  Hawaiian  Islands,  in  82-228  m. 

Range:  Leeward  Islands,  Hawaii,  from  French  Frigate 
Shoals,  to  Maro  (Dowsett)  Reef. 

Habitat:  I  am  aware  of  only  13  specimens  of  this  species 
ha\ing  been  collected,  all  from  lobster  traps  set  in  depths 
ranging  from  82  m  to  228  m.  \\\  t\  pe  specimens  were 
brought  into  traps  by  hermit  crabs. 


Etymology:  This  new  species  is  named  after  Daniel  R. 
Goodwin  of  Honolulu,  Hawaii,  who  collected  all  of  the 
type  specimens. 

Comparative  remarks:  Harpa  gooduini  is  a  moderately 
large  species  characterized  by  its  broad  oval  shape,  its 
outline  resembling  that  of  Harpa  costata  (Linne,  1758) 
but  with  the  outline  of  the  last  whorl  more  evenly  ar- 
cuate, and  differing  by  possessing  an  o\erall  pinkish  or 
orange-pink  coloration  when  fresh,  with  dark  horizontal 
stripes  on  the  fewer,  moderately  broad  and  distant  ribs. 
Harpa  major  Roding,  1798  differs  from  this  new  species 
in  having  a  more  oval  shape,  the  whorls  without  the 
broad,  flattened  canal,  and  with  the  ribs  fewer  in  number 
and  lacking  the  dark  horizontal  lines. 

This  new  species  most  closely  resembles  H.  cahritii 
Fischer,  1860  (see  Rehder,  1973:251-252;  1992:123),  and 
has  the  same  general  arrangement  of  dark  chestnut 
blotches  on  its  ventral  side.  Harpa  gooduini  has,  how- 
ever, a  relatively  broader  shell,  w  ith  the  body  whorl  not 
medially  flattened  but  gently  rounded.  The  axial  ribs  of 
H.  cahritii  lack  the  horizontal  dark  bands  and  numerous 
dark  lines  of  H.  goodwini. 

LITERATURE  CITED 

Rehder,  H.  A.  1973.  The  family  Harpidae  of  the  world.  Indo- 
Pacific  Mollusca  3(16):207-274. 

Rehder,  H.  .\  1992.  A  new  name  for  Harpa  lentricosa  La- 
marck, 1816  (Gastropoda:  Harpidae).  The  Nautilus  106(3); 
12.3-124. 


Figures  1.  2.  Harpa  guoduini,  new  species.  1.  Holotype,  USNM  860312.  2.  Paratype  1,  USNM  860313,  both  from  French  Frigate 
Shoals,  Hawaiian  Islands,  about  166''10'E,  23°45'N,  137  m.  Both  specimens  were  brought  into  traps  by  hermit  crabs.  Scale  bar  = 
2.0  cm. 


THE  NAUTILUS  106(4):130-136,  1993 


Page  130 


Perotrochus  maureri,  a  New  Species  of  Pleurotomariid  from  the 
western  Atlantic  (Gastropoda:  Pleurotomariidae) 


M.  G.  Harasewych 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC:  20560  USA 


Timothy  M.  Askew 

Harbor  Branch  Oceanographic 

Institution,  Inc. 

5600  U.S.  I  North 

Fort  Pierce,  FL  34946  USA 


ABSTRACT 

Perotrochus  maureri.  a  new  species  of  pleurotomariid,  is  de- 
scribed from  the  upper  continental  slope  off  northeastern  Flor- 
ida and  the  Carolinas.  It  can  be  distinguished  from  its  sister 
species,  Perotrochus  amabilis  (Bayer,  1963),  and  from  the  more 
remotely  related  P.  atlanticus  (Rios  &  Mathews,  1968)  by  its 
smaller,  lower,  more  deeply  pigmented  shell  with  fewer  spiral 
cords  on  the  selenizone  and  shell  base,  as  well  as  on  the  basis 
of  its  radular  asymmetry  and  formula.  The  emergence  of  pen- 
insular Florida  is  hypothesized  to  have  separated  the  ancestral 
population  into  the  Carolinian  P.  maureri  and  P.  amabilis, 
which  appears  to  be  restricted  to  the  Gulf  of  Mexico. 

Key  words:  Pleurotomariidae;  Perotrochus.  new  species;  bathy- 
al;  vicariance;  radula. 


INTRODUCTION 

During  the  course  of  continuing  studies  on  the  ecology 
and  systematics  of  western  Atlantic  pleurotomariid  gas- 
tropods, we  had  occasion  to  re-examine  a  large  series  of 
specimens  that  we  collected  off  Charleston,  South  Car- 
olina and  previously  referred  to  Perotrochus  amabilis 
(Bayer,  1963)  (Askew,  1988:91;  Harasewych  et  al.  1988). 
Comparison  of  this  material,  as  well  as  additional  spec- 
imens from  off  Jacksonville,  Florida,  with  the  holotype 
and  additional  specimens  of  P.  amabilis  from  off  north- 
ern Cuba  and  throughout  the  Culf  of  Me.xico  revealed 
subtle  but  consistent  differences  in  size,  pigmentation  and 
sculpture.  We  ascribe  these  differences  to  allopatric  spe- 
ciation,  and  here  describe  as  new  the  member  of  the 
species  pair  ranging  from  northern  Florida  to  the  Car- 
olinas. 


MATERIALS  AND  METHODS 

The  type  material  was  collected  during  four  dives  aboard 
the  Research  Submersible  Nekton  Delta,  in  the  area  of 
the  "Charleston  Lumps,"  a  region  of  rugged  topography 
some  90  miles  east  of  Clharleslon,  .South  (Carolina,  on  May 
2-4,  1987.  Some  of  the  specimens  were  fixed  in  10%  sea 


water  formalin  and  stored  in  70%  ethanol  for  dissection; 
the  remainder  were  frozen  and  maintained  at  —  80°C. 

Additional  specimens  of  the  new  species,  as  well  as 
comparative  material  of  P.  amabilis  and  P.  atlanticus  in 
museum  collections  were  examined,  and  are  identified 
in  the  text  by  the  following  institutional  acronyms; 

AMNH — American  Museum  of  Natural  Historv,  New 

York 
AMS — Australian  Museum,  Sydney 
ANSP — Academy  of  Natural  Sciences,  Philadelphia 
BM(NH) — British  Museum  (Natural  History),  London 
FM — Fernbank  Museum  of  Natural  History,  Atlanta 
FMNH — Field  Museum  of  Natural  History,  Chicago 
FSBC  I — Florida  Marine  Research  Institute,  St.  Peters- 
burg 
HBOM — Harbor  Branch  Oceanographic  Museum,  Ft. 

Pierce,  Florida 
LACM — Los  Angeles  County  Museum  of  Natural  His- 
tory 
MCZ — Museum  of  Comparative  Zoology,  Cambridge 
MNHN — Museum  National  d'Histoire  Naturelle,  Paris 
USNM — National  Museum  of  National  History,  Smith- 
sonian Institution,  Washington,  DC 

SYSTEMATICS 

Perotrochus  maureri  new  species. 

Synonymy: 

Perotrochus  amabilis.  .\skew,  1988:91,  Harasewych  et  al.  1988, 
(Non  Perotrochus  amabilis  (Bayer,  1963)). 

Description:  Shell  (fig.  1)  medium-sized  (to  52  mm), 
thin,  trochoid,  non-umbilicate,  of  up  to  9%  whorls.  Spire 
coeloconoid,  straight  for  first  5  whorls,  becoming  in- 
creasingly concave  thereafter.  Protoconch  (figs.  2-3),  500 
^m  wide,  of  1  smooth,  glossy,  translucent  whorl.  Tran- 
sition to  teleoconch  abrupt,  delimited  by  slightly  flared 
protoconch  lip,  onset  of  spiral  and  axial  sculpture,  selen- 
izone. Selenizone  initialK  nearly  abutting  suture,  de- 
scending to  mid-whorl  b\  whorl  2,  below  mid-whorl  b\' 
whorl  6.  Axial  sculpture  of  pronounced  prosoc>  rt  riblets 


M.  G.  Harasewych  and  T.  M.  Askew,  1993 


Page  131 


Table  1.    Measurements  of  shell  characters  in  Pcrotrochiis  niaurcri.  P  amahilis  and  P  allanliciis.  Linear  measurements  in  mm 


Pvwt 

rochiia  maurcn 
(n  =  10) 

P  amahihs 

(n  =  8) 

J 

f,  allanliciis 

(n  =  3) 

Character 

Mean 

Range 

SD 

Mean 

Range 

SD 

Mean 

Range 

SD 

Maximum  dianu-tcr 
(MD) 

46.1 

34,4-59.5 

6,6 

72.2 

44.9-87.2 

12.1 

57.0 

56,3-58.2 

0.9 

Shell  height 

(SH) 

37.3 

34.7-46.9 

44 

60.58 

41.7-73.3 

6.1 

53.7 

52.7-54.8 

0.9 

SH/MD 

0.822 

.791-832 

001 

0.858 

.840-929 

0.03 

0  940 

.936-.  951 

0  00 

No.  whorls, 
teleoconch 

8.78 

8.25-9.75 

0,37 

9.76 

9.0-10.75 

0.54 

9,33 

9.0-9.75 

0.31 

No.  spiral  cords 
on  selenizone 

1.80 

1-3 

0.75 

2.88 

2-4 

0.60 

3.33 

3-4 

0.47 

No.  spiral  cords 
on  shell  hast- 

UMI 

1 6-22 

1  6 

2S2 

24-31 

2  3 

27  0 

23-31 

3.3 

(18-20  on  whorl  1 ),  above  and  below  selenizone,  aligned 
in  early  whorls.  Riblets  decrease  in  prominence,  being 
reduced  to  beads  on  spiral  cords  by  whorl  6.  Spiral  sculp- 
ture initially  of  fine  threads  that  cross  riblets  at  angle 
(20-30°)  to  converge  on  selenizone  from  above  and  be- 
low. Single,  continuous  spiral  cord  first  appears  above 
the  selenizone  on  whorl  2,  below  the  selenizone  on  whorl 
4.  Selenizone  with  opisthocyrt  ribs  that  are  more  nu- 
merous than,  and  unaligned  with,  prosocyrt  ribs.  Spiral 
sculpture  on  selenizone  of  fine  radial  threads  between 
adjacent  ribs  on  early  whorls,  single  medial  spiral  cord 
by  whorl  4.  Number  of  spiral  cords  above/on/  below  the 
selenizone  increasing  to  3-4/1/2-3  on  whorl  6,  6-8/1- 
3/3-4  on  whorl  8.  Suture  initially  grooved,  becoming 
flat  b\  whorl  5,  impressed  b\  whorl  7.  Aperture  hori- 
zontally ovately-rhomboidal.  Outer  lip  smooth,  portion 
below  slit  offset  from  portion  above  slit  by  30-33°  Slit 
narrow,  extending  posteriorly  84-92°  from  outer  lip.  Col- 
umella spiralK  coiled,  with  strong  sigmoid  flexure  near 
adapical  margin.  Umbilical  region  excavate  but  not  per- 
forate. Base  convex,  with  17-22  even  spiral  cords  be- 
tween periphery  and  umbilical  region,  which  is  nacreous 
due  to  resorption  of  outer  layers  of  shell.  Base  color  ivory, 
w  ith  broad  axial  bands  of  pale  brick  red,  and  narrower 
bands  of  dark  red.  Aperture  nacreous,  iridescent.  Oper- 
culum (fig. 4)  small,  (spanning  0.6  of  minor  axis  of  ap- 
erture) multispiral,  corneous. 

Anatomy:  As  the  anatomy  of  P.  maureri  agrees  in  most 
regards  with  that  of  P.  amahilis  as  described  b\'  Fretter 
(1964);  only  supplemental  observations  on  living  animals 
are  recorded.  The  foot,  head  and  tentacles  are  densely 
mottled  with  dark  brick  red.  When  the  animal  is  crawl- 
ing, the  posterior  portion  of  the  shell  is  supported  on  the 
operculum.  Tentacles  are  long,  cylindrical,  ventrally  di- 
rected. Left  tentacle  bilobed  in  one  of  the  five  specimens 
dissected.  The  jaws,  inner  lips  and  outer  lips  are  all  in- 
terconnected, being  formed  of  a  single  piece  of  sclero- 
protein.  The  heavily  papillated  mantle  edges  on  either 
side  of  the  slit  abut,  sealing  the  slit  except  for  a  small 
opening  along  the  posterior  %  of  its  length.  The  ctenidia 
do  not  project  beyond  the  mantle  edge  in  living  speci- 


mens. When  the  animal  is  disturbed,  the  hypobranchial 
gland  rapidly  secretes  large  volumes  of  a  whitish  fluid 
that  is  immiscible  with  and  denser  than  seawater.  This 
secretion  settles  around  and  adheres  to  the  shell. 

Radula:  Radulae  of  5  specimens  (39-52  mm  maximum 
diameter)  were  examined.  Radula  (figs.  5-10,  table  2) 
long  (75-82%  maximum  shell  diameter),  asymmetrical, 
left-skewed,  bifid  posteriorly,  composed  of  92-104  in- 
verted V-shaped  rows  of  teeth.  Hickman's  (1984)  ter- 
minology for  the  six  tooth  types  is  used  herein  and  cor- 
related to  other  terminologies  in  table  2.  Rachidian  Tooth 
(figs.  6,  8)  with  dorsal  surface  laterally  expanded,  forming 
flanges  that  apparently  serve  to  maintain  alignment  be- 
tween lateral  teeth.  Two  long.  Inner  Lateral  Teeth,  with 
laterally  expanded  dorsal  surfaces  and  broad,  strongly 
curved  distal  ends  (fig.  8,  arrows)  flank  the  rachidian 
tooth  on  each  side.  Adjacent  are  24-26  (number  increas- 
ing with  shell  size)  Outer  Lateral  Teeth  (figs.  6-8)  that 
are  shortest  opposite  Inner  Lateral  Teeth  of  adjacent  row 
(fig.  7),  and  become  progressively  broader  and  stouter, 
with  the  long  axis  of  the  basal  plates  of  the  outermost 
Outer  Lateral  Teeth  nearly  perpendicular  (*70°)  to  that 
of  the  innermost  Outer  Lateral  Teeth  and  to  the  radular 
axis  (fig.  8).  These  in  turn  are  flanked  by  21  large,  curved 
Sickle  Teeth  (figs.  5-7).  As  in  P.  amahilis,  the  innermost 
teeth  are  tricuspid  (fig.  7,  arrow),  but  the  cusp  on  the 
concave  surface  is  lost  in  subsequent  teeth  (Fretter,  1964: 
181).  The  transition  from  Sickle  Teeth  to  Filament-Tipped 
Teeth  is  gradual, the  first  Filament-Tipped  Tooth  dis- 
cerned b\'  the  presence  of  two  minute  bristles  on  either 
side  of  the  proximal  cusp  (Woodward,  1901:250).  The 
bristles  increase  in  number  and  become  larger,  while  the 
cusps  diminish  in  size  (fig.  9)  and  ultimateK-  are  lost  in 
the  outermost  Filament-Tipped  teeth.  The  outermost  8- 
9  teeth  (fig.  10),  referred  to  as  Paddle-Shaped  Teeth,  are 
broad,  flat,  and  blunt  ended,  the  preceding  1-2  teeth  are 
transitional  from  Filament-Tipped  teeth  to  Paddle- 
Shaped  Teeth  and  retain  vestiges  of  a  filaments  along 
the  inner  distal  ends.  The  Paddle-Shaped  Teeth  of  one 
row  overlap  the  outer  Filament-Tipped  Teeth  of  the 
adjacent,  more  proximal  row,  forming  a  telescoping  mar- 


Page  132 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figure  l.Perotrochus  maureri.  new  species.  Apertural.  lateral,  ventral  and  dorsal  views  of  holotype  (USNM  860320).  Off  Charleston, 
South  Carolina,  USA  (32°43'57"W,  78°05'41"W)  in  195-204  m.  Scale  bar  =  1.0  cm. 


gin  that  envelops  the  Filament-Tipped  Teeth  and  fa- 
cilitate,s  smooth  motion  of  the  radula  within  its  sheath 
on  the  rasping  stroke.  The  blunt  ends  of  the  Paddle- 
Shaped  Teeth  may  also  dislodge  entangled  sponge  tissue 
and  spicules  from  the  radula  and  free  them  to  pass  into 
the  esophagus. 

Type  locality:  90  nautical  miles  E  of  Charleston,  South 
Carolina,  USA  (.32°43'54"N,  78°06'00"W,  to  32°44'02"N, 
78°05'22"W),  in  195-213  m. 


Range:  Perotrochus  maureri  occurs  on  the  upper  con- 
tinental shelf  off  Charleston,  South  Carolina  and  Jack- 
sonville, Florida,  in  depths  of  193-366  m. 

Type  specimens:  Holotype,  USNM  860320;  Paratypes 
1-9,  USNM  875218;  Paratype  10,  AMS  C.  169400;  Para- 
type  11,  BM(NH);  Paratype  12,  MNHN,  Paratype  13, 
FMNH-  Paratype  14,  FSBC  I;  all  from  R/V  Nekton 
Delta  dive  561,  (32°43'57"N,  78°05'4r'W)  in  195-204 
m,  4  May  1987.  Paratypes  15-17,  LACM  2629;  Paratype 


M.  G.  Harasewvch  and  T.  M.  Askew,  1993 


Page  133 


Table  2.    (loniiiarison  cif  fornuilae  and  asviiiinetries  iii   plcunitiiniariiid  radula    Sources  of  data  footnoted 
character  could  not  be  inferred  from  cited  text  or  figure. 


indicates  that  trie 


A: 

Racl 
ian 

lid- 

Later; 

d  teeth  

Marginal  teetl: 

1  

15: 

Racl 

:,id- 

ian 

Central 

Lamellate 

Hooked 

Brush 

Flahelliform 

C: 

Racl 

lid- 

Inner 

Outer 

Filament- 

Paddle- 

Taxon 

N 

ian 

laterals 

laterals 

Side 

tipped 

shaped 

Skew* 

Mikadotrochtis  heijrichi^ 

■4? 

3 

20-21 

17-22 

63-65 

7-12 

R' 

Perot rochti.s  iiuoiianiis- 

1? 

3 

24 

13 

63 

6 

R9.10 

P.  aildcdoniciis^ 

1 

22 

14 

53 

7 

R" 

P   notialis' 

2 

29 

13 

63 

10 

? 

P.  amabilis'^ 

3 

24 

21 

63 

8 

R12 

P.  maureri'^ 

2 

24-26 

21 

61-63 

8-9 

L« 

P  midas' 

3 

26 

13 

63 

6 

L9.10 

P  lucai/a' 

3 

25 

13 

61 

6 

? 

P.  africana'* 

4 

25 

12 

50 

6 

? 

A  =  Standard  terminology  (e.g..  Fretter  &  Craham,   1962:169).  B  =  Terminolog\  of  Woodward,  1901,  and  other  authors.  C  = 

Terminolog)  of  Hickman,  1984. 

*  R  =  Right-skewed  asymmetry.  L  =  Left-skewed  as\  mmetr> . 

'  Tlie  radular  formula  given  by  Woodward  (1901:252)  and  subsec|uentl\  cited  by  other  authors,  is  an  incorrect  summation  of  the 

data  contained  in  his  paper.  His  report  that  there  are  223  teeth  per  row  (p.  247),  and  explicit  statement  that  there  are  21  lamellate 

teeth  (p.  249)  indicate  that  the  correct  radular  formula  is  (R-3-21-17-63-7),  at  least  for  the  three  specimens  that  he  examined. 

Bouvier  and  Fischer  (1902)  report  a  radular  formula  of  (R-3-20-22-65-12)  for  this  species. 

-  Bouvier  and  Fischer,  1899. 

'  Bouchet  and  Metivier,  1982:310. 

■■  Leme  and  Penna,  1969:227. 

5  Fretter,  1964:181. 

^  Herein 

"Fretter,  1966:608. 

"  Barnard,  1963:157.  Listed  as  approximate  counts. 

«  Hickman,  1981:190. 

'"  Hickman,  198430 

"  Bouchet  and  Metivier,  1982:fig.  2. 

'-  Herein,  n  =  2. 


18-20,  USNM  875221;  Paratype  21,  MCZ,  Paratvpe  22, 
ANSP,  Paratvpe  23,  AMNH,  R/V  Nekton  Delta  dive 
557  (32°43'54"N,  78°06'00"W)  in  198-213  m,  2  May 
1987.  Paratypes  24-26,  USNM  846900  (reported  in  Hara- 
sewvch et  al.  1988:94  as  Perotrochus  amabilis).  R/V 
Nekton  Delta  Dive  560,  (32°43'5S"N,  78°05'43"VV)  in 
198-210  m,  3  May  1987.  Paratypes  27-33,  USNM  860321, 
Paratypes  34-35,  FM  92.15.  1-2,  R/V  Nekton  Delta 
Dive  562  (32°44'02"N,  78°05'22"W)  in  204-213  m,  4  Mav 
1987.  Paratype  36,  HBOM  65:1988  Johnson-Sea-Link- 
1-1455,  about  80  n  miles  off  Charleston,  South  Carolina, 
USA  32°44.0'N,  78°05.6' W,  in  heavy  rubble  zone  of  rocks 
and  sand,  193  m.  September  6,  1983. 

Other  material  examined:  HBOM  65:2017  Off  Charles- 
ton, SC  circa  1982;  AMNH  226434,  AMNH  226435, 
AMNH  226436,  and  USNM  869531,  ail  from  Off  Jack- 
sonville, Florida,  in  366  m. 

Comparative  material  examined:  Perotrochus  amabilis 
(F.M.  Baver,  1963):  Holotype  USNM  635625,  S.E.  of 
Sombrero  Light,  Florida  ('24°29'N,  80°53'W-24°30'N, 
80°50'W),  trawled  in  220  m,  R/V  Gerda  Cruise  6333, 
haul  G-135;  USNM  801707,  100  miles  NNE  of  Sagua  La 
Grande,  Cuba  (23°35'N,  79°34'W),  in  18"3-238  m,  R/V 


Silver  Bay  sta.  2460;  USNM  846648,  W  of  Tampa,  Flor- 
ida, rubble  bottom,  210  m;  USNM  858215,  Green  Can- 
yon, 100  miles  S  of  Atchafalava  Bay,  Louisiana 
(27°44'35"N,  91°07'54"W),  in  170  m',  Johnson-Sea-Link- 
1-2385,  17.6°C;  AMNH  183151,  WNW  of  Ft.  Mvers, 
Florida,  in  220  m;  AMNH  244316,  Dry  Tortugas,  Florida 
(no  depth);  FSBC  I  30812,  About  97'  nautical  miles  W. 
of  Mullet  Key,  Pinellas  County.  Florida  (27°39'N, 
84°33'W),  in  12'6.2-128.0  m.  R/V  Hernan  Cortez;  FSBC 
I  33146,  About  95  nautical  miles  W.  of  Anna  Maria 
Island,  Manatee  County,  Florida  (27°31'N,84°31'W)  in 
135.6-126.5  m.  R/V  Hernan  Cortez;  HBOM  65:02190, 
About  100  nautical  miles  SE  of  Galveston,  Texas,  in  268.2 
m.  Johnson-Sea-Link-II-933.  Perotrochus  atlanticus 
(Rios  &  Mathews,  1968):  USNM  846647,  Off  Rio  Grande, 
Rio  Grande  do  Sul,  Brazil,  in  260  m;  AMNH  181294, 
Off  Rio  Grande,  Rio  Grande  do  Sul,  Brazil,  in  164  m, 
sand  &  mud;  AMNH  244317,  Solidao,  Brazil. 

Ecology:  Specimens  were  observed  in  situ  at  the  tvpe 
locality,  an  area  of  rugged  terrain  consisting  of  steep, 
large  (3-30  m)  hill  crests  and  \alleNs.  Perotrochus  maur- 
eri  was  largely  confined  to  hard  substrates,  composed  of 
slabs  or  fragments  of  relithified  phosphorite,  that  lined 


Page  134 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figures  2-4.  Perotrochus  maureri,  new  species.  2.  Apical  and  3.  oblique  views  of  protoconch.  Scale  bars  =  250  ^ni.  4.  Oviter  and 
inner  views  of  operculum.  Scale  bar  =  1.0  mm. 


the  tops  and  sides  of  the  hill  crests  (see  Askew,  1988:90, 
fig.  2).  Water  temperature  was  9.6-9.8°C.  Perotrochus 
maureri  occurs  in  considerable  densities  at  the  type  lo- 
cality, with  distances  between  specimens  ranging  from 
10  to  30  meters.  The  diet  of  P.  maureri  [as  P.  amabilis] 
was  reported  by  Harasewych  et.  al.,  (1988)  to  consist  on 
sponges  of  the  orders  Poecilosclerida,  Choristida  or  Spi- 
rophorida,  and  the  genus  Strongylophora .  Most  speci- 


mens have  7-12  repaired  shell  breaks,  indicating  fre- 
quent, unsuccessful  predation,  probably  by  crustaceans. 

Etymology:  This  species  is  named  in  honor  of  Mr.  Rich- 
ard S.  Maurer  in  recognition  of  his  long  and  devoted 
interest  in,  and  support  of  Malacology. 

Discussion:  Perotrochus  maureri  is  most  closely  related 
to  P.  amabilis,  and  more  remotely  related  to  P.  atlan- 


Figures  5-10.  Radiilar  ribijon  of  Perolruchux  muurcri.  7i.  Dorsal  view  of  critical-point  dried  buccal  mass,  showing  alternating 
arrangement  of  booked  teeth.  Scale  bar  =  .500  ^ni.  6.  Anterior  view  of  extended,  critical-point  dried  radula  Radula  is  seen  to  be 
asymmetrical  and  left-skewed.  Scale  bar  =  500  ^m.  7.  Left  side  of  radula.  Scale  bar  =  500  ^m  8.  Rachidian,  inner  and  outer 
lateral  teeth  Teeth  to  the  left  of  the  rachidian  are  further  anterior  than  their  homologues  on  the  right.  Scale  bar  =  250  ^m  9. 
Filament-tipped  teeth  from  two  adjacent  rows.  Scale  bar  =  100  ^m.  10.  Paddle-shaped  teeth.  Transition  from  filament-tipped  to 
paddle  shaped  teeth  discernible  on  teeth  9-10  (arrows).  Scale  bar  =  250  iim. 


M.  G.  Harasewvch  and  T.  M.  Askew,  1993 


Page  135 


^^M 

EW^V//? 

ki     IfAA 

kI 

^WK^ 

^ 

s/  ^S^ 

V^Jn^ 

^M 

^jm^i  m^^. 

SL  '/-'  r  1 

■^^     M^^m 

A^H  ^UBI^^^Sii.lr/^^iy  > 

"^Ski    /      J^ 

I^J^ 

ssfl^  \vHv  vlkr^ 

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^>^^ 

sfe^^  /.  JF^^/zcK 

^Sflki 

^^^ 

l^V  iJv^^^Si 

V^^J 

^^mM        //^^^ 

3^  y^i^H 

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^K^JC 

^^^^^K  >               I^H;  ^^^^^^^^^^H^S^^I 

S^Ih 

Page  136 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figure  11.  Map  showing  distribution  of  Perotrochtis  amabilis 
(Bayer)  (dots)  and  P.  maureri  n.  sp.  (open  circles).  Solid  and 
open  stars  denote  respective  type  localities. 

ticus.  It  can  be  distinguished  from  both  these  species  by 
its  smaller  size,  lower  spire,  and  its  more  darkly  pig- 
inented  shell  with  fewer  spiral  cords  on  the  selenizone 
and  base  (Table  1).  The  radula  of  P.  maureri  differs  from 
that  of  P.  amabilis  in  being  left-skewed  rather  than  right- 
skewed,  and  in  having  one  fewer  Inner  lateral  Teeth,  an 
equal  or  greater  number  of  Outer  Lateral  Teeth,  an  equal 
or  lesser  number  of  Filament-tipped  Teeth  and  an  equal 
or  greater  number  of  Paddle-shaped  Teeth  (Table  2). 
The  utility  of  minor  differences  in  radular  formulae  for 
distinguishing  ta.\a  is  questionable,  as  most  reports  to  date 
are  based  on  a  single  radula  per  species.  Intraspecific 
variation  in  the  numbers  of  Outer  Lateral,  Sickle,  Fila- 
ment-tipped and  Paddle-shaped  teeth  has  been  encoun- 
tered when  multiple  radulae  have  been  examined  (Table 
2).  As  the  transitions  between  the  types  of  teeth  are 
gradual  (especialK'  in  the  Marginal  Teeth),  it  is  also  pos- 
sible that  different  investigators  may  have  assigned  one 
or  more  transitional  teeth  to  different  tooth  types,  thus 
further  obscuring  the  discriminating  value  of  radular 
formulae.  Radular  asymmetry,  however,  appears  to  be 
constant  within  a  species  (P.  maureri,  left-skewed,  n  = 
5;  P.  amabilis,  right-skewed,  n  =  2),  and  is  tentatively 
regarded  as  a  u.seful  distinguishing  criterion. 

Perotroclms  maureri  is  known  only  from  the  upper 
continental  slope  off  northeastern  Florida  and  the  Car- 
olinas,  where  it  occurs  below  the  10°C  thermocline.  Its 
sister  species,  P.  amabilis,  is  limited  to  the  Gulf  of  Mexico 
and  the  Straits  of  Florida  (Fig.  11).  A  single  temperature 
record  from  Louisiana  (17.6°C)  indicates  that  this  species 
lives  substantially  above  the  10°C  thermocline.  We  sur- 
mise that  the  emergence  of  peninsular  Florida  separated 
tlie  ancestral  population  during  the  Neogcne,  with  /'. 
amabilis  evolving  in  the  Gulf  of  Mexico,  while  P.  maureri 
was  isolated  on  limited  areas  of  hard  substrate  off  the 
Carolinas,  and  subsequently  adapted  to  colder  temper- 
atures. 


ACKNOWLEDGMENTS 

We  thank  Richard  Cooper  and  Peter  Auster  of  the  NOAA 
National  Undersea  Research  Program,  University  of 
Connecticut,  Avery  Point,  for  making  submarine  time 
available  at  the  Charleston  Lumps  site.  The  assistance  of 
Susann  Braden  with  the  scanning  electron  microscopy  is 
gratefully  acknowledged.  This  is  Harbor  Branch  Ocean- 
ographic  Contribution  No.  943,  and  Smithsonian  Marine 
Station  at  Link  Port  Contribution  No.  315. 


LITERATURE  CITED 

Askew,  T.  M  1988.  A  new  species  of  pleiirotoniariid  gastro- 
pod from  the  western  Atlantic.  The  Nautilus  102(3):89- 
91. 

Barnard,  K.  H.  1963.  Notes  on  the  animals  of  Gyrinagiganfea 
(Lam.)  and  Pleurotomaria  africana  Tomlin.  Proceedings 
of  the  Malacoiogical  Societ\  of  London  .35(4):  155-158. 

Bayer,  F.  M  1963,  .\  new  pleurotomariid  gastropod  trawled 
in  the  Straits  of  Florida  by  R/\'  Gerda,  Bulletin  of  Marine 
Science  of  the  Gulf  and  Caribbean  13(3):488-492. 

Bouchet,  P.  and  B.  Metivier.  1982.  Living  Pleurotomariidae 
(Moiiusca;  Gastropoda)  from  the  South  Pacific.  New  Zea- 
land Journal  of  Zoology  9:309-318. 

Bouvier,  E.  L.  &  H.  Fischer  1899  Etude  monographique 
des  Pleurotomaires  actuels.  Bulletin  of  the  Museum  of 
Comparative  Zoolog\  ,  Harvard  32: 193-249,  pis.  1-4;  also 
published  in  Journal  de  (^oncln  liologie  47(2):77-151,  pis, 
4-7. 

Bouvier,  E.  L.  and  H.  Fischer.  1902.  L'organisation  et  les 
affinites  des  Gasteropodes  primitifs  d'apres  I'Etude  ana- 
tomique  du  Pleurotomaria  Beyrichi.  Journal  de  Conchy- 
liologie50(2):117-272,  pls.2-6. 

Fretter,  V  1964.  Observations  on  the  anatomy  of  Mikado- 
trochua  amabilis  Baser.  Bulletin  of  Marine  Science  of  the 
Gulf  and  Caribbean  14(1  );172-1S4, 

Fretter,  V.  1966,  Biological  investigations  of  the  deep  sea,  16. 
Observations  on  the  anatomv  of  Perutrochus.  Bulletin  of 
Marine  Science  16(3):603-614. 

Fretter,  V.  and  A.  Graham.  1962.  British  Prosobranch  Mol- 
lusks.  Ra>  Society,  London.  755  p. 

Harasewych,  M.  G.,  S.  A.  Pomponi  and  T  M.  .\skew.  1988. 
Spongivory  in  pleurotomariid  gastropods.  The  Nautilus 
102(3):92-98. 

Hickman,  C,  S.  1981.  Evolution  and  function  of  asymmetry 
in  the  .\rcheogastropod  Radula.  The  V'eliger  23(3):189- 
194,  figs.  1-22. 

Hickman,  C.  S.  1984.  Form  and  function  of  the  radulae  of 
pleurotomariid  gastropods.  The  X'eliger  27(l):29-36. 

Leme,  J.  L.  M.  and  L.  Penna.  1969.  Ocorrencia  de  Mika- 
dotrochus  no  Brasil.  com  descri^ao  de  uma  nova  especie 
(Gastropoda:  Pleurotomariidae).  Papeis  Avulsos  de  Zool- 
ogia  22:225-230. 

Rios,  E.  de  C.  and  H.  R.  Mathews.  1968.  Nova  especie  de 
Pleurotomariidae  do  Brasil  (Moiiusca:  Gastropoda).  -■Krq. 
Estafao  de  Biologia  Marinha  I'niversidade  Federal  do 
Ceara  8(l):65-68. 

Woodward,  M.  F.  1901.  The  -dnntomy  oi  Pleurotomaria  beyr- 
ichii  Hilg.  The  QuarterK  Journal  of  Microscopical  Science 
44(2):215-268,  pis.  13-16, 


THE  NAUTILUS  106(4):137-146,  1998 


Page  137 


The  Genus  Fulgoraria  (Gastropoda:  Volutidae)  of  the  northeastern 
Kamchatka  Peninsula  and  Sakhahn  Island,  with  Notes  on  the 
Paleoecology  and  Distribution  of  the  Subfamily  Fulgorariinae  in 
the  Oligocene  of  the  northern  Pacific 


Anton  E.  Oleinik' 

Hussiaii  Acadfiin  nt  Sciences 
Geological  Institute 
Pyzhevsky  per.  7 
109017  Moscow,  Russia 


ABSTRACT 

Four  new  species  of  the  genus  Fulgoraria  Sclnunacher,  1817, 
(subgenus  Musashia  Ha\ashi.  I960)  are  described  from  the 
Oligocene  Alugian  Formation  of  the  llpinsk\  Peninsula,  north- 
eastern Kamchatka  Peninsula,  Russia.  Previously  described  ful- 
gorariine  gastropods  from  the  same  region,  and  from  the  Oli- 
gocene of  Sakhalin  and  Karaginsky  Islands,  are  also  figured. 
Oxvgen  isotopic  analyses  of  contemporaneous  Cyclocardia  shells, 
along  with  a  comparison  of  the  ecology  of  Recent  congeneric 
taxa  and  a  paleoecologic  anal)  sis  suggest  a  bath)  al  environment 
as  most  probable  for  these  Oligocene  Fulgorariinae.  .\  review 
of  the  Oligocene  biogeography  of  northern  fuigorariines  along 
the  northern  Pacific  margin,  including  the  western  coast  of 
North  America,  indicates  that  this  subfamily  had  a  much  broad- 
er distribution  during  late  Paleogene  time  than  today.  These 
data  point  to  more  favorable  climatic  conditions  (including 
lower  water  temperatures)  for  dispersal  of  fulgorariine  volutes 
during  the  Oligocene 

Key  words:  Fulgorariinae;  systematics;  distribution;  Oligo- 
cene; Paleogene;  Northern  Pacific. 


INTRODUCTION 

Volutid  gastropods  are  common  as  fossils  in  Cenozoic 
faunas  of  the  northern  Pacific  region.  However,  their 
fossil  record  from  some  parts  of  this  region,  particularly 
the  far  east  of  Russia,  is  still  very  incompletely  known. 
The  subfamily  Fulgorariinae  Pilsbry  and  Oisson,  1954, 
is  the  dominant  group  among  North  Pacific  Cenozoic 
volutes.  Recent  members  of  the  "northern  group"  (Shi- 
kama,  1967)  of  this  subfamily  are  restricted  to  Japan  and 
adjacent  seas.  Less  well-known  extinct  species  occurred 
along  both  eastern  and  western  margins  of  the  North 
Pacific  during  both  early  and  late  Cenozoic  time. 


'  Present  Address:  Department  of  Earth  and  Atmospheric  Sci- 
ences, Purdue  University,  West  Lafayette,  Indiana  47907,  USA 


The  subfamily  Fulgorariinae  comprises  a  group  of  car- 
nivorous gastropods  with  a  uniserial  radula  composed  of 
tricuspid  rachidian  teeth  (Cooke,  1922;  Habe,  1943;  Oku- 
tani,  1963;  Weaver  &  du  Pont,  1970;  Watanabe  &  Habe, 
1978).  The  higher  systematics  of  this  subfamily  is  still 
not  fully  resolved,  with  the  two  most  recent  revisions 
(Shikama,  1967;  Weaver  &  du  Pont,  1970)  differing  pri- 
marily in  the  ranking  of  supraspecific  taxa.  Shikama  ( 1967) 
recognizes  three  genera:  Fulgoraria  Schumacher,  1817, 
Musashia  Hayashi,  1960,  and  Saotomea  Habe,  1943,  as 
well  as  the  subgenera  Psephaea  Crosse,  1871,  Nippon- 
ornelon  Shikama,  1967,  Neopscphaea  Takeda,  1953,  and 
Miopleiona  Dall,  1907.  The  last  two  are  known  only  as 
fossils.  Shikama's  classification  is  based  exclusively  on 
shell  characters  such  as  the  number  and  shape  of  colu- 
mellar  plaits,  the  size  and  form  of  the  protoconch,  and 
features  of  the  external  shell  morphology.  All  of  these 
characters,  especially  the  number  of  columellar  plaits, 
may  vary  during  ontogeny.  Interpreting  this  highly  vari- 
able shell  morphology  is  further  complicated  when  work- 
ing with  fossil  specimens,  as  they  are  often  incompletely 
preserved. 

Based  on  shell  and  radular  characters,  Weaver  and  du 
Pont  (1970)  recognized  only  a  single  Recent  north  Pacific 
genus  Fulgoraria,  with  the  subgenera  Psephaea,  Volu- 
tipisrna  Rehder,  1969,  Musashia.  Kurodina  Rehder,  1969 
and  Saotomea.  These  authors  regard  Nipponornelon  as 
a  synonsm  of  Musashia.  and  do  not  discuss  the  taxonomic 
position  of  the  fossil  Neopsephaea  and  Miopleiona. 

The  present  paper  provisionally  follows  the  classifi- 
cation of  Weaver  and  du  Pont  (1970),  but  includes  Nip- 
ponornelon. Miopleiona .  and  Neopsephaea  as  subgenera 
of  Fulgoraria. 

Twenty-six  species  of  Fulgorariinae  from  the  latest 
Eocene  and  Oligocene  formations  of  the  north  Pacific 
have  been  figured  or  described.  Of  these,  eight  are  known 
only  from  North  America  [Poul  Creek  and  Narrow  Cape 
(of  Sitkinak  Island)  Formations  of  Alaska;  Blakeley,  Twin 
River,  and  Eugene  Formations  of  Oregon  and  Washing- 


Page  138 


THE  NAUTILUS,  Vol.  106,  No.  4 


ton],  and  have  been  referred  to  the  subgenera  Nippon- 
omelon,  Musashia.  Miopleiona.  and  Seopsephaea  (Dur- 
ham, 1944;  Addicott  et  ai,  1971;  Tegland,  1933;  Allison 
&  Marincovich,  1981;  Moore,  1984).  Nine  species,  re- 
ferred to  the  subgenera  Fulgoraria,  Psephaea,  Musashia, 
i^'ipponomelon.  and  Neopsephaea,  are  restricted  to  the 
Oligocene  of  Japan  (Ashiya,  Kishima.  Nishisonogi,  Ku- 
mano,  and  Chikokubo  Formations  of  K%  ushu  and  Hon- 
shu Islands;  Momijiyama  Formation  of  Hokkaido)  (Oya- 
ma  et  ai,  1964;  Shikama,  1967;  Masuda  &  Noda,  1976). 
An  additional  seven  species,  from  various  localities  along 
the  northwestern  coast  of  the  United  States  and  the  Kam- 
chatka Peninsula,  have  only  been  identified  to  the  sub- 
generic  level  (Allison  &  Marincovich,  1981;  Moore,  1984; 
Gladenkov.  Sinelnikova  &  Bratseva,  1987).  Only  one  spe- 
cies endemic  to  the  Kamchatka  Peninsula  and  the  Koryak 
Upland,  Fitlgoraria  {Musashia)  olutorskiensis  L.  Krish- 
tofovich,  1973,  has  been  described  to  date.  A  second 
Siberian  species,  Fulgoraria  (Nipponomelon)  tohunagai 
(Kanehara,  1937)  is  more  widespread  and  is  also  known 
from  the  Oligocene  of  Japan. 

MATERIALS  AND  METHODS 

All  specimens  in  this  stud\-  were  collected  during  field 
work  in  eastern  Kamchatka  and  Sakhalin  Island  between 
1965  and  1986.  They  were  taken  from  the  Alugian  For- 
mation of  northeastern  Kamchatka,  the  "Laternula" 
sandstones  of  Karaginsky  Island,  and  the  Matchigarian 
Formation  of  central  Sakhalin  (fig.  21).  The  preservation 
of  the  specimens  was  often  fragmentary,  with  the  figured 
specimens  representing  the  most  complete  material 

In  order  to  address  the  questions  of  the  ecolog\  of 
these  Oligocene  fulgorariines,  a  paleotemperature  anal- 
ysis of  shells  of  Cyclocardia  ilpinensis  Pronina,  1973, 
which  occurs  in  abundance  in  the  same  strata  as  the 
volutes,  was  conducted.  This  standard  analysis  was  based 
on  the  ratio  of  the  oxygen  isotopes  ('''O/  '*0).  The  values 
of  '''O  ""O  (PDB  standard,  mass  spectrograph)  were  ad- 
justed to  the  Standard  Mean  Ocean  Water  (SMOW)  stan- 
dard and  corrected  for  altered  isotopic  composition  at 
higher  latitudes.  Temperature  values  were  calculated  us- 
ing the  following  formula: 

T  =  16.5  -  4.3  (b  -  A)  +  0.14(b  -  A)^, 

where  T  is  the  temperature  in  °C,  b  is  the  instrumentally 
determined  difference  in  the  '^'O/  "'O  ratio  between  the 
sample  and  the  standard,  and  A  is  the  correction  for  the 
original  isotopic  composition  of  sea  water.  For  high  lat- 
itudes A  =  —  1.  Prior  to  analysis,  all  shells  were  examined 
using  X-ray  diffraction  analysis  to  exclude  specimens  in 
which  the  aragonitic  structure  had  recr\stallized. 


The  following  institutional  acronyms  are  used;  GI — 
Geological  Institute,  Russian  Academy  of  Sciences.  Mos- 
cow; CMG — Central  Museum  of  Geology,  St.  Petersburg, 
Russia;  USNM  National  Museum  of  Natural  History, 
Smithsonian  Institution,  Washington,  DC,  USA. 

SYSTEMATICS 

Family  Volutidae  Rafinesque,  1815 
SubfamiK  Fulgorariinae  Pilsbry  and  Olsson,  1954 
Genus  Fulgoraria  Schumacher,  1817 
Subgenus  Musashia  Hayashi,  1960 

Fulgoraria  {Musashia)  novoilpinica  new  species 
Figures  1,  2,  17 

Description:  Shell  fusiform,  slender,  with  4-5  postnu- 
clear  whorls.  Last  whorl  stout,  comprising  -3  of  shell 
height.  Suture  moderateK  impressed,  subsutural  band 
absent,  .\perture  elliptical  (Length  width  —  3).  Inner 
lip  with  one  narrow  columellar  fold  and  siphonal  fold. 
Siphonal  canal  wide.  Axial  sculpture  of  numerous  (10- 
12  on  penultimate  whorl)  thin  ribs,  more  pronounced  on 
earlier  whorls,  smoother  and  wider  on  bod\  whorl.  Spiral 
sculpture  of  fine,  raised  threads  covering  entire  surface. 

Material  examined:  Holot\pe — USXM  468649.  length 
(incomplete)  119.5  mm,  width  58.3  mm;  Paratype — GI 
4072,  length  (shell  strongly  decorticated)  124.7  mm.  width 
62.2  mm;  1  ju\enile  shell.  7  broken  shells  and  fragments, 
2  molds;  all  from  the  t\  pe  localit\ . 

Type  locality:  Northwestern  part  of  the  Ilpinsky  Pen- 
insula, eastern  Kamchatka,  Russia.  Upper  part  of  the 
.Alugian  Formation,  Oligocene. 

Stratigraphic  range:  Known  onl\  from  the  Upper  part 
of  the  Alugian  Formation  of  eastern  Kamchatka.  Oli- 
gocene. 

Comparalive  remarks:  .\lthough  similar  to  Fulgoraria 
shutoi  Shikama,  1967,  from  the  Kishima  Formation  (Oli- 
gocene, Japan),  this  new  species  differs  in  having  a  more 
elongated  last  whorl,  narrower  axial  folds,  nearly  smooth 
body  whorl,  and  only  a  single  fold  rather  than  several 
strong  ones. 

Fulgoraria  {Musashia)  olutorskiensis  (L.  Krishtofovich, 

1973) 

Figures  11,  12,  15,  16 

Miopleiona  oregonensis  Khomenko,  193325,  pi  6,  fig.20. 
Miopleiona  olutorskiensis  L.  Krishtofovich,  1973:77,  pi.  22, 

figs.  8,9, 
Musashia  olutorskiensis  Dewatiiova  &  X'oiobueva,  1981:128, 

pi.  :32,  fig.  2. 


Figures  1 .  2.  Fulgoraria  {Musashia)  novoilpinica,  new  species.  Holotype,  I  SNM  468649,  1 19.5  mm,  .\lugian  Formation,  Oligocene. 
Figures  3.  4.  Fulgoraria  (Musashia)  genuata,  new  species.  Holotype,  USNM  468650,  84.5  mm,  .■\lugian  Formation.  Oligocene. 
Figures  5.  6.  Fulgoraria  {Seopsephaea)  tenuis  (Shikama,  1967).  GI  1164,  49.2  mm,  Matschigarian  Formation,  Oligocene.  Figures 
7,  8.  Fulgoraria  (Musashia)  tilitschikensis,  new  species.  Holotype,  USNM  468652,  51.1  mm,  Alugian  Formation,  Oligocene. 


A.  E.  Oleinik,  1993 


Page  139 


Page  140 


THE  NAUTILUS,  Vol.  106,  No.  4 


Musashia  (Musashia)  sp.  Gladenkov,  Siiielniko\a  &  Bratseva, 
1987:57,  pi.  15,  fig.2. 

Description:  Shell  oblong,  fusiform,  relatively  thin,  with 
5-6  postnuclear  whorls.  Body  whorl  comprises  appro.x- 
imately  -3  of  shell  height.  Suture  slightU  depressed,  with 
narrow  subsutural  band.  Aperture  elliptical,  acute  pos- 
teriorly. Inner  lip  arched,  with  one  narrow  columellar 
fold.  Callus  thin,  narrow.  Shell  surface  nearly  smooth, 
\\  ith  thin  growth  striae  and  very  fine,  raised  spiral  threads. 

Material  examined:  Holotype— CMG  21/10285,  length 
130.4  mm,  width  48  mm,  Pachatchi  River,  Olutorsky 
Region,  eastern  Kamchatka,  Russia,  "Hpinian"  Forma- 
tion, Oligocene;  Paratype— 19/10285,  Gulf  of  Olutorsk, 
Govena  Peninsula,  eastern  Kamchatka,  Russia,  Oligo- 
cene; GI  1486,  Gl  4055 — both  from  Ilpinsky  Peninsula, 
northeastern  Kamchatka,  Russia,  Alugian  Formation, 
Oligocene;  9  incomplete  shells  and  fragments  and  7  molds. 

Type  locality:  Olutorsky  Region,  northeastern  Kam- 
chatka, Russia. 

Stratigraphic  range:  Known  from  the  Oligocene  for- 
mations of  northeastern  Kamchatka;  abundant  in  the 
Alugian  Formation  of  the  Ilpinsk\-  Peninsula. 

Comparative  remarks:  With  its  nearly  smooth  sculpture, 
this  species  most  closely  resembles  Fulgoraria  [Musashia) 
nagaoi  (Shikama,  1967)  from  the  Poronai  Formation  (Eo- 
cene-Oligocene  of  Hokkaido),  but  differs  in  being  more 
elongated,  in  having  fine  spiral  sculpture,  and  in  having 
a  distinct  subsutural  band. 

Fulgoraria  (Musashia)  genuata  new  species 
Figures  3,  4,  19 

Description:  Shell  oblong,  fusiform,  slender,  w  ith  three 
stout,  preserved  whorls,  and  estimated  5-6  postnuclear 
whorls  in  intact  specimens.  Body  whorl  comprises  <  % 
total  shell  length.  Suture  shallow,  slightly  impressed.  Ap- 
erture elliptical.  Columella  with  one  weak  columellar 
fold.  Sculpture  of  straight,  smooth,  widely  spaced  axial 
ribs  (4  per  whorl),  most  prominent  on  early  whorls.  Spiral 
sculpture  of  numerous,  fine,  slightly  raised  threads  cov- 
ering shell  surface. 

Material  examined:  Holotype — USNM  468650,  length 
84.5  mm,  width  40.0  mm;  Paratype— GI  40721,  length 
75.9  mm,  width  46.1  mm;  both  from  type  locality. 

Type  locality:  Northwestern  part  of  Ilpinsky  Peninsula, 
northeastern  Kamchatka,  Russia.  Alugian  Formation. 
Oligocene. 

Stratigraphic  range:  Known  only  from  the  Alugian  For- 
mation of  the  Ilpinsky  Peninsula.  Oligocene. 


Comparative  remarks:  Although  similar  to  Fulgoraria 
(Musashia)  fujimotoi  (Kanno,  1958)  from  the  Hikokubo 
Formation  (Oligocene-Miocene,  Japan),  this  new  species 
differs  in  having  stouter  whorls  and  a  lower  spire,  in 
having  one  instead  of  two  columellar  folds,  and  in  having 
finer  spiral  sculpture. 

Fulgoraria  (Musashia)  cordafa  new  species 
Figures  13,  14 

Description:  Shell  elongate,  fusiform,  slender,  with  four 
postnuclear  whorls,  short  siphonal  canal  and  narrowly 
channeled  suture.  Body  whorl  comprises  approximately 
¥4  of  shell  length.  Aperture  oblong,  with  narrow  posterior 
angle.  Columella  with  one  weak  columellar  fold,  .\xial 
sculpture  of  low,  rounded  ribs  (—  12  per  whorl  on  pen- 
ultimate whorl).  Axial  ribs  crossed  by  fine,  closely  spaced 
spiral  lines. 

Material  examined:  Holotype — USNM  468651,  length 
59.1  mm,  width  24.3  mm;  Paratype  1— GI  4063/2,  length 
50.5  mm,  width  26.5  mm;  Paratype  2 — GI  40531,  length 
62.8  mm,  width  26.5  mm;  two  molds,  three  fragments 
and  one  impression;  all  from  the  type  locality. 

Type  locality:  Northwestern  part  of  Ilpinskv  Peninsula, 
northeastern  Kamchatka,  Russia.  Upper  part  of  the  .alu- 
gian Formation.  Oligocene. 

Stratigraphic  range:  Known  only  from  the  type  locality. 

Comparative  remarks:  This  new  species  is  most  similar 
to  Fulgoraria  (Musashia)  weaveri  (Tegland,  1933)  of  the 
Blakeley  Formation  (Oligocene,  Washington,  USA),  but 
has  a  more  elongated  shell,  more  numerous  axial  ribs, 
and  much  stronger  spiral  threads. 


Fulgoraria  (Musashia)  tilitschikensis  new  species 

Figures  7,  8,  18 

Description:  Shell  globoseK  fusiform,  solid  Teleoconch 
of  about  5  whorls.  Suture  narrowly  pressed.  Body  whorl 
comprises  %  shell  length.  Aperture  semiovate,  with  thin, 
narrow  callus.  Inner  lip  w  ith  one  weak,  subvertical  col- 
umellar fold.  Axial  sculpture  of  rounded  arcuate  ribs  (13 
on  body  whorl).  Spiral  sculpture  lacking. 

Material  examined:  Holot\pe — USNM  468652,  length 
(incomplete)  51.1  mm,  width  29.5  mm,  from  the  type 
locality;  Paratype  GI  40724,  length  (incomplete)  65.0 
mm,  width  37.6  mm,  Ilpinsk>  Peninsula,  northeastern 
Kamchatka,  Russia.  .Alugian  Formation,  Oligocene. 

Type  locality:  Korf  Settlement.  Coal  River,  northeastern 
Kamchatka,  Russia.  Alugian  Formation.  Oligocene. 


Figures  9,  10.  Fulgoraria  (Nipponumclon)  cLtokunagai  (Kanehara,  UW7).  GI  3586  1804.  149  mm,  " Lalcrnula"  Sandstones, 
Karagin.sky  Island,  Oiigocene-Lower  Miocene  (?).  Figures  1 1,  12,  15,  16.  Fulgoraria  (Musashia)  olutorskiensis  (L.  Krishtofovich, 
1973).  1 1,12.  GI  1486,  61.8  mm,  15,  16.  GI  4055,  103.9  mm,  Alugian  Formation,  Oligocene.  Figures  13,  14.  Fulgoraria  {Musashia) 
cordata,  new  species.  Holotype,  USNM  468651,  59.1  mm,  Alugian  Formation,  Oligocene. 


A.  E.  Oleinik.  1993 


Page  141 


cm. 


Page  142 


THE  NAUTILUS,  Vol.  106,  No.  4 


A.  E.  Oleinik,  1993 


Page  143 


Recent  species  of  Fulgorarinae: 


DEPTH  IN  METERS 
250  500 


750 


Fulgoraria  (Miisashia)  cancellata  Kuroda  et  Habe 
Fulgoraha  (Musashia)  formosana  Azuma 
Fulgoraria  (Musashia)  clara  (Sowerby) 
Fulgoraria  (Nipponomelon)  kamakurerisis  (Otuka) 
Fulgoraria  (Nipponomelon)  elongata  Shikanm 
Fulgoraria  (Nipponomelon)  prevostiana  Crosse 
Fulgoraria  (Nipponomelon)  angulosa  Shikama 


Figure  20.  Bath>Tnetric  distributions  of  Recent  species  of  Fulgorariinae  of  Japan  and  adjacent  seas,  based  on  data  in  Shikama 
(1967),  and  Weaver  and  du  Pont  (1970).  Thickness  of  hnes  indicates  relative  abundance. 


Slratigraphic  range:  Known  only  from  the  Alugian  For- 
mation of  northeastern  Kamchatka,  Russia.  Oligocene. 

Comparative  remarks:  This  species  resembles  Fulgoraria 
(Musashia)  shikamai  Moore,  1984,  from  the  Poul  Creek 
Formation  (Alaska,  USA)  and  the  Clallam  and  Twin  Riv- 
er Formations  (Washington,  USA),  but  this  new  species 
has  more  sharpened  axial  ribs  and  finer  spiral  threads. 

Subgenus  Nipponomelon  Shikama,  1967 

Fulgoraria  (Nipponomelon)  of.  tokiinagai  (Kanehara, 

1937) 

Figures  9,  10 

Psephaea  tokunagai  Kanehara,  1937:16-18,  pl.2,  figs.  2-,5. 
Fulgoraria  (Psephaea)  prevostiana  Crosse,  Shikama,  1954:pi.6, 

fig.  26, 
Fulgoraria  tokunagai  Kamada,  1962:192,  pl21,  figs.  5-8. 
Musashia  (Nipponomelon)  tokunagai  Shikama,  1967:100-101, 

pi.  14,  fig.  4. 
Musashia  tokunagai  Devvatilova  &  Volobueva,  1981:128,  pi. 33, 

fig-  1- 
Original  Description:  "Shell  tall  and  high  spired,  apical 
angle  about  25  degrees.  Suture  slightK  impressed  with 
narrow  shelf  around  it.  Surface  ornamented  with  a.xial 
folds,  which  ma\  be  somewhat  accentuated  above  and 
tend  to  disappear  on  the  body  whorl.  Spirally  grooved 
regularly,  also  over  the  folds,  .Aperture  spindle  shaped, 
obliquely  notched  posteriorly,  with  narrower  anterior 
canal.  Inner  lip  provided  with  two  columellar  folds  and 


covered  b\-  a  thin  callus,  which  adheres  to  the  columella 
and  thickens  anteriorly," 

Material  examined:  One  specimen,  GI  3586/1804,  length 
149  mm,  width  62.2  mm,  from  the  "Laternula"  sand- 
stones, Karaginsky  Island,  Gnunvayam  River,  Russia. 
Oligocene-Lower  Miocene  (?). 

Type  locality:  Yanagaya  bed,  \agakura  coal  mine,  Yu- 
moto,  Iwaki  City,  Fukushima  Prefecture,  Honshu,  Japan. 
Lower  Miocene. 

Stratigraphic  range:  "Laternula"  sandstones,  Karagin- 
sk\  Island,  Gnunvayam  River,  Russia,  Oligocene-Lower 
Miocene  (?),  to  Yanagaya  bed,  Nagakura  coal  mine,  Yu- 
moto,  Iwaki  Cit\ ,  Fukushima  Prefecture,  Honshu,  Japan. 
Lower  Miocene. 

Comparative  remarks:  Fulgoraria  tokunagai  is  charac- 
terized by  a  large  size,  high  spire,  large  protoconch,  and 
numerous  axial  folds.  The  spiral  sculpture  on  the  single 
specimen  from  Karaginsky  Island  is  poorly  preserved, 
but  this  specimen  retains  visible  traces  of  the  axial  folds 
and  thin  spiral  grooves.  The  shape  and  size  of  this  spec- 
imen, especialK  the  features  of  the  aperture,  suture,  and 
sculpture,  are  the  same  as  in  typical  F.  tokunagai.  How- 
ever, the  Russian  specimen  differs  from  the  type  in  hav- 
ing a  single  columellar  fold  rather  than  two.  This  dif- 
ference, together  u  ith  the  poor  preserv  ation  of  this  shell 
puts  the  species  determination  of  this  specimen  in  some 
doubt. 


Figure  17.  Fulgoraria  (Musashia)  novoilpinica,  new  species.  Reconstructed  drawing  of  holotype.  Figure  18.  Fulgoraria  (Musashia) 
tilitschikensis.  new  species.  Reconstructed  drawing  of  holotype.  Figure  19.  Fulgoraria  (Musashia)  genuata.  new  species.  Recon- 
structed drawing  of  holotype.  Scale  bars  =10  mm. 


Page  144 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figure  21.  Fossil  localities  of  Oligocene  Fulgorarinae  in  the  North  Pacific  referred  to  in  the  text.  1 — Honshu  Island,  2 — Hokkaido 
Island,  3 — Sakhalin  Island,  -4 — Kamchatka  Peninsula,  5 — Karaginsky  Island,  6 — Sitkinak  Island.  7 — Kodiak  Island,  8 — Oregon,  9 — 
Washington,  10 — California. 


Subgenus  Neopsephaea  Takeda,  1953 

Fulgoraria  {Neopsephaea)  tenuis  Shikama,  1967 
Figures  5,6 

Musashia  (Neopsephaea)  /emiis  Shikama,  1967:116-117,  pi.  13, 
figs.  3,4. 

Original  description:  "Shell  small  in  size,  slender,  fusi- 
form and  with  low  spire.  Suture  shallow  and  there  is  no 
subsutural  band.  Last  whorl  vaulted  at  middle  and  ap- 
erture relatively  narrow.  Surface  of  whorl  smooth  and 
a.xial  ribs  relatively  few  and  weak,  about  11  in  penulti- 


mate whorl,  gradually  becoming  obsolete  anteriorly.  Tip 
of  columella  straight  and  narrow.  There  may  be  one 
columellar  plait". 

Supplemental  description:  Shell  is  rather  slender,  elon- 
gate, fusiform,  with  about  5  (when  restored)  whorls  sep- 
arated b\  a  shallowK  impressed  suture.  Bod>  whorl  and 
aperture  comprising  slightl)  more  than  %  of  total  shell 
length.  Aperture  elliptical,  forming  acute  angles  at  an- 
terior and  posterior  margins.  Siphonal  notch  relatively 
narrow.  Outer  lip  thin,  simple.  Columella  with  one  weak 
columellar  fold.  Callus  verv  thin.  Shell  surface  smooth, 


Japan    Sakhalin       Kamchatka  Alaska  U.S. 

E     130       140       150      160        170       180       170       160       150       140       130       120  W 


Fulgoraria 

Psephaea 

Musashia 

[Nipponomelon 

Neopsephaea 

Miopleiona 


Cumulative 

Number 
of  Species     0 


Figure  22.  Geographic  distribution  of  the  subgenera  ot  Fulgoraria.  and  (heir  species  diversity  during  ihc  Oligocene. 


A.  E.  Oleinik,  1993 


Page  145 


with  thin  growth  Unes.  Axial  sculpture  ot  low ,  rounded 
ribs  (11-12  on  body  whorl)  that  disappear  on  anterior 
portion  of  body  whorl. 

Material  examined:  One  specimen,  Gl  1 164,  length  49.2 
nmi,  width  22  mm,  near  Lake  Matschigar,  central  Sa- 
khalin Island,  Russia.  Matschigarian  Formation.  Oligo- 
cene. 

Type  locality:  Zochuku  Sandstone,  Kishima  Group,  Na- 
gasaki Prefecture,  K)  ushu,  Japan.  Oligocene. 

Stratigraphic  range:  Near  Lake  Matschigar,  central  Sa- 
khalin Island,  Russia,  Matschigarian  Formation,  Oligo- 
cene, to  Zochuku  Sandstone,  Kishima  Group,  Nagasaki 
Prefecture,  K\ushu,  Japan,  Oligocene. 

Coniparaiive  remarks:  This  species  is  closeK  allied  to 
Fulgoraria  (Mcopscphaea)  antiquior  Takeda,  but  may 
be  distinguished  b\  its  more  swollen  body  whorl,  nar- 
rower aperture  w  ith  distinctive  posterior  margin,  smooth 
surface,  and  a.xial  ribs  that  are  fewer  in  number  or  en- 
tirely obsolete. 


DISCUSSION 

The  lithology  of  the  Alugian  Formation  of  the  llpinsky 
Peninsula  in  eastern  Kamchatka  consists  of  gray  to  dark 
gray  claystones  and  clays.  This  formation  has  a  total 
thickness  of  about  950  meters,  and  is  continuous,  lacking 
hiatuses  or  internal  unconformities.  No  shallowing  or 
deepening  trends  are  e\ident  within  the  section.  The 
strata  contain  numerous  carbonate  concretions  ranging 
in  size  from  1  cm  to  several  meters.  Some  of  the  claystone 
beds  are  delicately  laminated.  These  types  of  sediments 
are  indicative  of  soft  bottom  environments. 

Molluscan  remains  are  quite  common,  but  are  not 
concentrated  in  particular  layers  or  lenses.  Fossils  occur 
sporadically  throughout  the  section,  both  in  concretions 
and  within  the  matrix.  The  molluscan  fauna  does  not 
exhibit  mans-  changes  from  the  bottom  to  the  top  of  the 
formation.  These  data  point  to  a  depositional  regime 
comprising  relatively  rapid  and  uninterrupted  sedimen- 
tation (Kidwell,  1988). 

Species  diversit\  in  the  molluscan  assemblage  of  the 
Alugian  Formation  is  low.  The  most  abundant  taxa  oc- 
curring together  with  fulgorariine  volutes  include  Trotn- 
inina  bicordata  (Hatai  &  Koike,  1957),  Trominina  ish- 
ihariensis  (Ha\asaka  &  Matsui,  1951),  Neptiinea  ezoana 
Takeda,  1953,  Bathybembix  sakhalinensis  korjakensis 
Volobueva,  1981,  Optotiirris  (?)sp.,  Turritellasp.,  Cryp- 
tonatica  spp.,  Acila  praedivaricata  Nagao  &  Huzioka, 
1941,  and  Cyclocardia  ilpinensis  Pronina,  1973.  All  these 
taxa  have  relativeK  broad  bath\  metric  ranges,  occurring 
from  lower  neritic  to  bath)al  depths  (Hall,  1960;  Noda, 
1975;  Scarlato,  1981;  Moore,  1984). 

Recent  species  of  Fulgorariinae  are  mainly  bathyal 
animals,  dredged  most  often  from  depths  of  250  to  700 
or  more  meters  (fig.  20).  Bath\rnetricalK ,  the  bath\al 
zone  extends  from  a  mean  depth  of  200  m  to  2000  m, 
and  thermally,  from  the  15°C  isotherm  in  low  latitudes 


down  to  the  — 3°C  isotherm  in  high  latitudes.  Substrates 
are  predominantly  fine  silts,  muds,  and  oozes  (Encyclo- 
pedia of  Oceanography,  1974;  Hickman,  1974).  Oxygen 
isotope  anaKses  of  five  Cyclocardia  ilpinensis  shells  col- 
lected from  the  bottom  through  the  top  of  the  Alugian 
Formation  allowed  the  estimation  of  paleotemperatures 
as  ranging  from  3°C  to  5°C.  These  data  indicate  that 
water  temperatures  remained  quite  uniform  during  the 
deposition  ot  this  formation,  and  support  the  hypothesis 
of  a  bathyal  environment.  When  compared  to  Recent 
bathyal  bottom  temperatures,  these  data  suggest  that  the 
climatic  conditions  during  Alugian  deposition  were  sim- 
ilar to  those  found  between  32°N  and  45°N,  and  ap- 
proximate the  temperatures  that  define  the  distribution 
of  Recent  species  of  Fulgoraria.  Paleotemperatures  es- 
timated for  the  Alugian  Formation  exhibit  some  corre- 
lations with  those  estimated  for  Oligocene  formations  of 
the  western  coast  of  North  .America.  Paleotemperatures 
of  the  Blakeley  Formation  (Echinophoria  apta  zone), 
which  contains  five  species  of  Fulgorariinae.  were  esti- 
mated as  to  be  5-8°C.  The  remains  of  Aturia  nautiloids 
in  great  abundance,  which  may  have  required  a  tem- 
perature of  at  least  16°C  could  be  explained  by  post- 
mortem transport  of  empty  shells,  which  is  fairly  com- 
mon with  Recent  NautUus.  Paleodepth  was  thought  to 
range  from  100  m  to  350  m  (Moore,  1984).  Similarly, 
the  Narrow  Cape  Formation  of  Sitkinak  Island  (Alaska), 
which  contains  three  species  of  Fulgorariinae  ,  was  de- 
posited in  the  outer  neritic  zone  of  the  continental  shelf 
(Allison,  1978).  A  comparison  of  the  Narrow  Cape  data 
with  that  of  Recent  Alaskan  mollusks  suggests  a  paleo- 
depth of  100-200  m.  and  water  temperatures  of  10-12°C 
in  summer  months  during  the  Oligocene  (Allison  &  Mar- 
incovich,  1981).  According  to  Hall  (1964),  this  type  of 
marine  climate  can  be  defined  as  cool-temperate.  The 
Recent  species  Fulgoraria  {Nipponomelon}  prevostiana 
(Crosse,  1878)  has  been  dredged  from  depths  of  110- 
732  m  off  western  Hokkaido,  Japan,  at  a  bottom  tem- 
perature of  11.8°C  (Weaver  &  du  Pont,  1970:45).  Water 
temperatures  for  the  Alugian  Formation  thus  seem  to  be 
colder  than  those  in  western  America.  These  data  more 
probably  suggest  a  deeper  depositional  environment  for 
the  Alugian  Formation,  perhaps  no  shallower  that  400- 
500  m,  and  therefore  colder  water. 

Oligocene  climatic  conditions,  therefore,  must  have 
been  quite  favorable  for  a  wider  distribution  of  fulgo- 
rariine volutes  that  is  seen  today.  This  is  clearK'  supported 
by  the  fossil  record  (fig.  21).  I5ased  on  the  stratigraphic 
distribution  and  analysis  of  total  species  diversity  of  Ce- 
nozoic  volutes,  it  has  been  deduced  that  the  Oligocene- 
Early  Miocene  was  the  first  period  of  wide  distribution 
and  high  species  diversity  of  northern  Pacific  Fulgora- 
riinae (Oleinik,  1990).  There  was,  however,  an  appre- 
ciable amount  of  endemism,  especially  at  the  subgeneric 
level.  Fulgoraria  s.s.  and  Psephaea,  for  example,  were 
restricted  to  the  western  Pacific,  while  Miopleiona  is 
known  onl\  from  the  northeastern  Pacific.  These  re- 
stricted geographic  ranges  ma\  have  been  caused  by 
different  centers  of  origin  of  the  North  Pacific  Fulgo- 


Page  146 


THE  NAUTILUS,  Vol.  106,  No.  4 


rariinae,  or  were  the  results  of  differing  dispersal  strat- 
egies. Other  subgenera,  such  as  Musashia  and  Nippon- 
omelon,  were  abundant  throughout  the  north  Pacific. 
Total  species  diversity  of  Fulgorariinae,  however,  did 
not  vary  appreciably  during  the  Oligocene  of  the  north 
Pacific  (fig.  22). 

ACKNOWLEDGMENTS 

I  thank  Dr.  Yuri  B.  Gladenkov  and  Dr.  V'alentina  N. 
Sineiniko\a,  of  the  laboratory  of  Phanaerozoic  Stratig- 
raphy of  the  Geological  Institute  of  the  Russian  Academ\' 
of  Sciences,  for  providing  material;  Dr.  Sergei  I.  Kiyash- 
ko,  of  the  Institute  of  Marine  Biology  in  Vladivostok  for 
the  oxygen  isotope  analyses.  Photographs  of  the  shells 
were  taken  by  Mr.  Andrew  A.  Okunev  of  the  Geological 
Institute,  Moscow.  My  special  thanks  to  Dr.  M.  G.  Har- 
asewych.  National  Museum  of  Natural  History,  Smith- 
sonian Institution,  Washington,  D.C.,  Dr.  E.  J.  Petuch, 
Florida  Atlantic  University,  Boca  Raton,  Florida,  and 
Dr.  L.  K.  Marincovich,  U.S.G.S.,  Menlo  Park,  California 
for  critical  review  of  the  manuscript. 


LITERATURE  CITED 

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THE  NAUTILUS  106(4):  147-151,  1993 


Page  147 


A  New  Species  of  Columbellid  Gastropod  from  the 
Old  World  Tropics 


William  K.  Emerson 

Department  of  Invertebrates 
American  Museum  of  Natural  History 
New  York,  NY  10024-5192,  USA 


ABSTRACT 

Cotonopsis  monfilsi  new  species,  is  described  from  off  Senegal, 
West  .Africa  in  230-500  m.  The  generic  assignment  is  made  on 
the  basis  of  shell,  opercular,  and  radular  characters.  The  radula 
is  typically  columbellid,  whereas  the  shell  morphology  lacks 
some  characteristics  of  the  "Strombina"  gastropods.  Therefore, 
the  placement  in  the  S/romfoiia-group  genus  Cotonopsis  is 
tentative,  Metula  dockeryi  new  name  is  proposed  to  replace 
Metula  inflata  Dockerv,  1984,  not  Metula  inflata  (Houbrick, 
1984). 

Key  Words:  Prosobranchia,  Columbellidae,  Cotonopsis,  Buc- 
cinidae,  Sdetula,  west  Africa,  new  species,  new  name. 


INTRODUCTION 

During  the  past  five  years,  we  have  received  from  several 
shell  dealers  specimens  of  an  apparently  new  species  of 
columbellid  gastropod  for  study  and  report.  The  taxo- 
nomic  placement  of  these  shells  could  not  be  made  with 
certainty  until  the  morphology  of  the  radula  was  deter- 
mined. .\  radula  was  eventually  found  and  proved  to  be 
typically  columbellid  in  form  As  a  result,  this  gastropod 
can  be  assigned  provisionally  to  the  Strombina-group 
genus  Cotonopsis. 

I  take  pleasure  in  describing  this  new  species  in  honor 
of  Paul  Monfils,  who  was  the  first  to  call  tliis  perplexing 
species  to  my  attention. 

A  review  of  the  literature  disclosed  that  a  replacement 
name  was  required  for  a  fossil  species  of  Metula.  A  new 
name  is  provided  for  Metula  inflata  Dockery,  1984,  not 
Metula  inflata  Houbrick,  1984. 

ABBREVIATIONS 

AMNH  =  American  Museum  of  Natural  History,  New 
York 

LACM  =  Los  Angeles  County  Museum  of  Natural  His- 
tory, California. 


SYSTEMATICS 

Superfamily;  Buccinoidea  Rafinesque,  1815 
Family  Columbellidae  Swainson,  1840 
Genus  Cotonopsis  Olsson,  1942 

Type  species  (by  original  designation):  Strombina  {Co- 
tonopsis) panacostariceus  Olsson,  1942,  Pliocene  of  the 
Burica  Peninsula,  Costa  Rica,  Charco  Azul  Formation 

Cotonopsis  monfilsi  new  species 
(figures  1-10) 

Description:  Shell  solid,  fusiform,  whorls  inflated  with- 
out a  sutural  ramp;  protoconch  of  3^2  smooth,  convex 
whorls;  teleoconch  of  IVz  convex  whorls,  sculptured  with 
numerous  fine  spiral  lirae,  numbering  about  18  on  the 
penultimate  whorl  and  about  32  on  the  body  w  horl.  Some 
spiral  lines  bifid;  lirae  not  consistently  evenly  spaced. 
Thickened  varix  formed  on  body  whorl  above  the  rim 
of  the  aperture.  Aperture  ovate,  inner  lip  terminally  thin, 
but  with  a  sublabial  ridge  with  10  Urate  teeth;  columella 
w ith  4-5  weak  folds;  siphonal  canal  open,  short,  narrow. 
Fasciole  indistinct.  Periostracum  brownish  tan.  Back- 
ground color  brownish  buff-tan  with  broken  whitish  bands 
below  the  suture  and  with  axial  blotches  interrupting  the 
tanish  coloration.  Aperture  white.  Operculum  thin,  cor- 
neous, lenticular  with  a  terminal  nucleus  at  edge.  Radular 
ribbon  (figs.  9,  10)  rachiglossate,  typically  columbellid 
(cf.  Radwin,  1977,  fig.  22b).  Rachidian  tooth  a  narrow, 
nearly  rectangular  plate.  Lateral  tooth  shaft-like  with 
barbed  base,  tip  of  shaft  with  two  hooked  cusps. 

Material  examined:  Holotype  (AMNH  232519)  and  3 
paratypes  (AMNH  232091)  from  the  t\pe  localitv;  5 
Paratypes  (AMNH  232092)  dredged  in  300-350  m',  on 
muddy  bottom,  off  St.  Louis,  Senegal,  November,  1987, 
ex-Northeast  Natural  History  Imports;  3  Paratypes 
(AMNH  232152)  dredged  in  450-500  m,  on  muddy-sand 
bottom,  off  St.  Louis,  Senegal,  ex-Northeast  Natural  His- 
tory Imports;  6  referred  specimens  (5  specimens,  AMNH 


Page  148 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figurrs  1-8.    Cotonopsis  monfilsi  n.  sp.  1-3,  holotype  (AMNH  232519)   4-6,  paratype  (AMNH  232091);  figs.  1-6  xl.5.  Figs. 
8,  paratype  (AMNH  232092),  spire  enlarged  to  show  detail  of  earl\  whorls;  figs.  7,  8,  5.0x. 


232176;  1  specimen,  LACM  146956)  dredged  on  silty- 
sand  bottom,  230-260  m,  off  Casamance  Province,  Sen- 
egal (12''50'N,15°0'W),  ex-Northeast  Natural  History  Im- 
ports; 2  referred  specimens  (AMNH  255056)  trawled  in 
deep  water  from  off  the  Dakar  harbor  entrance,  Senegal 
(14°40'N,  17°26'W),  ex-Mal  de  Mer  Enterprises.  See  Ta- 
ble 1  for  shell  measurements. 

Type  locality:  dredged  in  300  m  off  St  Louis,  Senegal, 
West  Africa,  d6°02'N,16°30'W)  by  Marcel  Pin,  1987,  ex- 
Abbey  Specimen  Shells. 

Distribution:  Known  only  from  oU  Senegal,  West  Africa 
in  230  to  500  meters. 


Remarks:  The  shells  of  the  present  specimens  resemble 
certain  ""S7roni/n>irt"-group  gastropods  (such  as  Coton- 
opsis), but  the\  are  also  in  some  respects  reminiscent  of 
certain  buccinid  gastropods  (eg.  Pisania.  Mctula,  etc.). 
Before  the  nature  of  the  radula  was  known,  I  asked 
several  colleagues  for  their  opinions  based  on  the  ex- 
amination of  photographs  or  the  study  of  the  specimens. 
Some  of  them  referred  the  specimens  to  the  buccinid 
genus  Pisania  Bivona-Bernardi,  1832,  in  the  subfamily 
Pisaniinae  Gray,  1857:13,  formerly  credited  to  Tryon, 
1881:98  (see  Ceriiohorskv,  1971:138;  1975:192,  Beu  and 
Maxwell,  1987:56;  and  Bouchet,  1988:149,  for  comments 
on  the  subfamilial  status  of  Pisaniinae  in  Buccinidae,  and 


W.  K.  Emerson,  1993 


Page  149 


Table  1.  Slu-ll  climensioiis  and  proportions  of  the  specimens 
of  Cotonopsis  monfilsi  n.  sp.  n  =  20  specimens.  Width  incfndes 
terminal  \arix  on  k>odv  whorl.  Measurements  in  mm 


Figures  9,  10.  Cotono|)>.i>.  monfiUi  n  sp  9,  holotype  u^MNH 
2325f9),  radula,  about  one-thirti  of  the  ribbon  10,  detail  of  the 
rachidian  teeth;  fig.  9,  scale  bar  =  100  ^ni,  fig  10,  scale  bar  = 
200  ixm.  courtesy  of  J.  H.  McLean 


its  rejection  without  an  explanation  at  tiie  subfamilial 
level  b\'  Ponder  and  Waren,  1988:305)  The  resemblance 
of  the  new  species  to  Recent  species  of  Charitodoron 
(Mitridae)  from  South  Africa  was  also  suggested  (see 
Lozouet,  1991:206,  figs.  26-30).  Other  colleagues  be- 
lieved the  Senegalese  specimens  to  be  an  undescribed 
columbellid  species  referable  to  Cotonopsis  (sensu  stric- 
to). 

With  the  recovery  of  a  radula  from  a  dried  body  of 
the  new  species,  the  placement  of  the  new  species  in  the 
family  Columbellidae  can  be  confirmed.  The  radular 
characters  are  t>pically  columbellid  in  form  (figs.  9,  10). 
As  in  Cotonopsis  argentea  (Houbrick,  1983:352,  fig.  2), 
the  radular  dentition  consists  of  rachidian  teeth  that  are 
thin,  narrow  and  form  a  rectangular  plate.  A  wide  space 
separates  the  rachidian  teeth  from  the  lateral  teeth,  which 
are  composed  of  a  shaft  v\'ith  an  enlarged  base  and  are 
tipped  with  two  hooked  cusps.  The  shells  of  the  new- 
species  possess  the  thickened  varices  of  Stromhina-grou'p 
gastropods,  but  the  apertural  dentition  of  the  inner  lip 
differs  in  having  evenly  spaced  lirate  denticles  of  equal 
size.  These  are  comparable  to  those  of  the  buccinid  genus 
Bartschia  Rehder  (1943:199,  pi.  20,  fig.  17;  Olsson  and 
Bayer,  1972:924,  fig.  14).  Furthermore,  Bartschia  sig- 
nificans.  the  type  species,  has  strongly  cancellate  sculp- 
ture, whereas  the  new  species  possesses  fine  spiral  lirae. 
The  radular  characters  of  Bartschia  are  unknown,  but 
on  shell  features,  Beu  and  Maxwell  (1987:62)  believe 
Bartschia,  together  with  Metida  H.  and  A.  Adams  (1853: 
84;  Emerson,  1986:27),  to  be  closely  related  to  the  buc- 
cinid genus  Colubraria  Schumacher,  1817.  Some  of  the 
western  Atlantic  species  of  Metula  [e.g.  the  West  Indian 
M.  (Agassitida)  agassizi  Clench  and  Aguayo,  1941:179, 
pi.  14,  fig.  4;  Olsson  and  Bayer,  1972:917,  fig.  11  and  the 
east  African  M.  (Kanamarua)  rehderi  (Kilburn,  1977: 
193,  fig.  21)]  superficialK-  resemble  the  new  species. 
Bouchet  (1988:150,  fig.  1)  illustrated  the  typically  buc- 


Width- 

Type  specimens 

Height 

Width 

height 

*AMNH  232519  (holotvpe) 

40.1 

14,9 

0.37 

AMNH  232091  (paratvpe  1) 

41,7 

1,5.2 

0.36 

AMNH  232091  (paratype  2) 

37.2 

13.7 

0.37 

AMNH  232091  (paratype  3) 

37.1 

13.6 

0.37 

AMNH  232092  (paratype  1) 

41.1 

14.4 

0.35 

AMNH  232092  (paratype  2) 

37.9 

13.4 

0.35 

AMNH  232092  (paratype  3) 

.37.1 

13.8 

0.35 

AMNH  232092  (paratype  4) 

37.5 

13.6 

0.36 

AMNH  232092  (paratype  5) 

36.2 

12.8 

0.35 

AMNH  2321.52  (paratvpe  1) 

404 

14.6 

0.36 

AMNH  232152  (paratvpe  2) 

36.9 

12.8 

0.35 

AMNH  232152  (paratype  3) 

34.5 

12.6 

0.37 

Referred  specimens 

AMNH  232176,  a 

39.7 

14.5 

0.37 

AMNH  232176,  b 

36.3 

13.8 

0.38 

AMNH  232176,  c 

36,1 

13.9 

0.39 

AMNH  232176,  d 

35.0 

14.0 

0.40 

AMNH  2.32176,  e 

34.9 

13.7 

0.40 

LACM  146956 

36.2 

13.9 

0.38 

AMNH  255056  a 

39.1 

14.1 

0.36 

AMNH  255056,  b 

38.6 

13.4 

0.35 

Mean 

37.1 

13.8 

0.36 

Range 

34.5- 

12.6- 

0.35- 

41.7 

15.2 

0  40 

*  Radula  extracted. 


cinid  radula  of  his  new  species,  Metida  africana,  and 
concluded  the  radular  morphology-  is  significantly  dif- 
ferent from  that  of  the  buccinid  genus  Pisania. 

The  placement  of  the  new  species  in  the  genus  Co- 
tonopsis based  on  shell  characters  is  tentative.  As  the 
genus  is  defined  by  Jung  (1989:158),  the  teleoconch  whorls 
are  smooth  or  axialK  sculptured.  In  the  new  species,  the 
teleoconch  sculpture  is  composed  of  fine  spiral  lines  and 
no  axial  sculpture  is  present.  Cotonopsis  is  represented 
in  the  tropical  western  Atlantic  by  one  species  (C.  ar- 
gentea). The  remainder  of  the  15  species  of  Cotonopsis 
recognized  by  Jung  (1989),  2  fossil  and  12  living  species, 
occur  in  the  west  American  tropical  zone.  Previously, 
the  Strombina-gToup  gastropods  were  believed  to  be  re- 
stricted to  the  New  World  tropics,  dating  from  the  Mio- 
cene to  the  present,  w  ith  only  3  of  the  35  living  species 
occurring  in  the  western  Atlantic  (Jung,  1989:4).  There- 
fore, the  presence  of  a  species  of  Cotonopsis  in  west 
African  waters  may  reflect  a  separate  lineage  of  these 
columbellids  that  may  eventually  be  determined  to  re- 
quire recognization  as  a  new  genus-group  taxon. 

REPLACEMENT  NAME  FOR  A  HOMONYM 

During  my  review  of  the  literature  for  this  paper,  I  noted 
a  homonymic  species-group  name  in  Metula  (Buccini- 
dae).  Metula  {Metula)  inflata  Dockery  in  MacNeil  and 


Page  150 


THE  NAUTILUS,  Vol.  106,  No.  4 


Dockery  (1984:331,  pi.  52,  fig.  3;  published  in  "Novem- 
ber, 1984",  teste  D.  T.  Dockery  III,  in  Hit..  Dec.  9,  1991) 
is  a  junior  secondary  homonym  of  Acamptochetus  [=Me- 
tiila]  inflatus  Houbrick  (1984:421,  fig.  1;  published  July 
6,  1984).  I  here  rename  Metula  inflata  Dockery,  1984 
(not  Metula  inflata  [Houbrick,  1984]),  as  Metula  dock- 
eriji  Emerson,  NEW  NAME.  Dockery  s  ta.xon  is  a  Pa- 
leogene  fossil  from  Mississippi  and  Houbrick  s  ta.xon  is  a 
Recent  Philippine  species. 

Acamptochetus  Cossmann  (1901:123;  type  species  by 
original  designation:  Murex  mitraeformis  Brocchi,  1814) 
is  a  junior  subjective  synonym  of  Metula  H.  and  A.  Ad- 
ams (1853:84;  type  species  by  subsequent  designation 
Kobelt,  1876:  Buccinum  clathratum  Adams  and  Reeve, 
1850,  [not  Kiener,  1834,  nor  Anton,  1839]  =  Metula 
atnosi  Vanatta,  1913:22),  as  noted  by  Emerson  (1986:27) 
and  Beu  and  Maxwell  (1987:62)  and  accepted  by  Bouchet 
(1988:149). 


ACKNOWLEDGEMENTS 

In  addition  to  Paul  Monfils  of  Northeast  Natural  History 
Imports,  Providence,  RI,  I  am  grateful  to  Robert  Foster 
of  Abbey  Specimen  Shells  of  Santa  Barbara,  CA  and 
Robert  and  Dorothy  Janowsky  of  Mai  De  Mer  Enter- 
prises, West  Hempstead,  NY  for  providing  specimens  for 
study.  Silvard  P.  Kool  of  the  Museum  of  Comparative 
Zoology,  Harvard  University,  kindly  lent  specimens  of 
Metula  for  comparisons.  The  following  colleagues  co- 
operatively offered  comments  on  the  taxonomic  place- 
ment of  the  new  species  before  the  nature  of  the  radula 
was  known:  Walter  O.  Cernohorsky,  M.  G.  Harasewych, 
Richard  S.  Houbrick,  Peter  Jung,  R.  N.  Kilburn,  Jorgen 
Knudsen,  Paula  M.  Mikkelsen,  Gary  Rosenberg,  and  Gra- 
ham Saunders.  James  H.  McLean  generously  mounted 
the  radula  and  provided  the  photographs  for  the  radula. 
I  am  indebted  to  my  AMNH  colleagues,  Walter  E.  Sage, 
III  for  technical  assistance,  Andrew  S.  Modell  for  pho- 
tographic services  and  Stephanie  Grooms  for  word-pro- 
cessing the  manuscript. 

LITERATURE  CITED 

Adams,  H.  and  A.  1853  [-1852].  The  Genera  of  recent  Mol- 
lusca,  arranged  according  to  their  organization,  London 
l(13);65-96,  pis.  9-12  (June,  1853). 

Anton,  H.  E.  1839.  Verzeichniss  der  Conchy  lien  vvelche  sich 
in  der  Sammlung  von  Hermann  Eduard  .^nton  befinden. 
Halle,  xvi  -t-  110  p. 

Beu,  A.  G.  and  P.  A.  Maxwell  1987.  A  revision  of  the  fossil 
and  living  gastropods,  related  to  Picsiotriton  Fischer,  1884 
(Family  Gancellariidae,  subfamily  Plesiotritoninae  n, 
subfam.)  with  an  Appendix:  Genera  of  Buccinidae  Pisa- 
niinae  related  to  Cotubraria  Schumacher,  1817.  New  Zea- 
land Geological  Survey  Paleontological  Bulletin  54:1-140. 

Bivona-Bernardi,  A.  1832,  Caratteri  dun  nuovo  genere  di 
conchiglie  della  famiglia  delle  columellarie  del  Signer  De 
Lamarck  Effemeridi  Scientifiche  e  Letterarie  per  la  Sicilia 
2:8-13,  1  pi. 

Bouchet,  P.     1988.    Two  new  species  of  Metula  (Gastropoda: 


Buccinidae)  with  a  description  of  the  radula  of  the  genus. 
The  Nautilus  102(4):149-153. 

Cernohorsky,  W  O.  1971.  Indo-Pacific  Pisaniinae  (Mollusca: 
Gastropoda)  and  related  buccinid  genera.  Records  of  the 
Auckland  Institute  and  Museum  8:1.37-167. 

Cernohorsky,  W.  O.  1975.  Supplementary  notes  on  the  tax- 
onomy of  buccinid  species  of  the  subfamily  Pisaniinae 
(Mollusca:  Gastropoda).  Records  of  the  Auckland  Institute 
and  Museum  12:175-211. 

Clench,  W.  J.  and  C.  G.  Aguayo.  1941.  Notes  and  descriptions 
of  new  deep-water  Mollusca  obtained  by  the  Harvard- 
Havana  Expedition  off  the  coast  of  Cuba.  Memorias  de  la 
Sociedad  Cubana  de  Historia  Natural.  Ha\ana  15(2):  177- 
180, 

Cossmann,  M  1901  Essais  de  Paleconchologie  Comparee,  pt 
4,  293  p, 

Emerson,  W.  K.  1986.  On  the  type  species  of  Metula  H  and 
A.  Adams,  1853:  Buccinum  clathratum  A.  .Adams  and 
Reeve,  1850  (Gastropoda:  Buccinidae),  The  Nautilus  100(1): 
27-30, 

Gray,  J.  E.  1857,  Guide  to  the  systematic  distribution  of 
Mollusca  in  the  British  Museum.  Part  1,  London,  230  p. 

Houbrick,  R.  S,  1983.  A  new  Strombina  species  (Gastropoda; 
Prosobranchia)  from  the  tropical  western  .Atlantic.  Pro- 
ceedings of  the  Biological  Society  of  Washington  96(3): 
349-3.54, 

Houbrick,  R,  S.  1984.  A  new  "Me/i/Za"  species  from  the  Indo- 
West  Pacific  (Prosobranchia  Buccinidae).  Proceedings  of 
the  Biological  Society  of  Washington  97(2):420-424  (pub- 
lished July  6,  1984).' 

Jung,  P.  1989  Revision  of  the  S(roni/>i;!a-Group  (Gastropoda: 
Columbellidae),  fossil  and  living.  Distribution,  biostratig- 
raphy,  svstematics.  Memoires  Suisses  de  Paleontologie  HI: 
1-298. 

Kiener,  L.  C.  1834  [-1841].  Species  general  et  iconographie 
des  coquilles  vivantes.  .  .  .  Paris,  9:1-112, 

Kilburn,  R,  N  1977,  Taxonomic  studies  on  the  marine  Mol- 
lusca of  southern  .Africa  and  Mozambique  Part  1,  .Annals 
of  the  Natal  Museum  23(1):17.3-214. 

Kobelt,  W.  1876  [-1881].  Illustriertes  Conchylienbuch  Nu- 
remberg, 1:1-143. 

Lozouet,  P.  1991.  Mollusca  Gastropoda:  Eumiira  recentes  de 
la  region  neo-caledonienne  et  Charitodoron  fossiles  de 
I  Oligocene  superieur  d  .AquitainelMitridae)  hi.  t>osnier, 
A.  and  P.  Bouchet  (eds. )  Resultats  des  Campagnes  Ml'- 
SORSTOM,  vol  7,  no,  8.  Memoires  de  la  Museum  National 
d'Histoire  Naturelle,  Ser   .A,  Zoologie  1.50:205-222 

MacNeil,  F.  S,  and  D.  T  Dockery.  Ill  1984  Lower  Oligocene 
Gastropoda.  Scaphopoda,  and  Cephalopoda  of  the  V'icks- 
burg  Group  in  Mississippi.  Mississippi  Department  of  Nat- 
ural Resources,  Bureau  of  Geology,  Bulletin  24:1-415 
(published  in  November,  1984). 

Olsson,  A.  A.  1942.  Tertiary  and  Quaternary  fossils  from 
Burica  Peninsula  of  Panama  and  Costa  Rica.  Bulletins  of 
American  Paleontology  27(106):153-258, 

Olsson,  A  A.  and  F  M  Bayer  1972  .American  metulas  (Gas- 
tropoda: Buccinidae).  Bulletin  of  Marine  Science  (Uni- 
versity Miami,  FL)  22(4):900-925, 

Ponder,  W.  F.  and  A.  Waren.  1988.  In.  Ponder,  W.  F.,  D.  J. 
Eernisse,  and  J.  H.  Waterhouse,  eds.  Prosobranch  Phylog- 
eny,  .Appendix,  Classification  of  the  Caenogastropoda  and 
Heterostropha — .A  list  of  the  family-group  names  and 
higher  taxa  Malacological  Re\ie\\ .  Supplement  4:288-326. 

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Atlantic.  The  N'eliger  19(4):403-417. 


W.  K.  Emerson,  1993  Page  151 


Rehder,  H.  A.    1943.    New  marine  mollusks  from  the  Antilleaii  3:1-310,  Academy  of  Natural  Sciences,  Pfiiladelpfiia  [p. 

region.  Proceedings  of  the  United  States  National  Museum  98,1881] 

93(3161):187-203,  pis.  19,  20.  Vanatta,  E.  G.     1913.    Descriptions  of  new  species  of  marine 

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and  systematic.  Series  1,  Tritonidae,  Fusidae,  Buccinidae,  Philadelphia  65(l):22-27,  3  text  figs.,  pi.  2. 


THE  NAUTILUS  106(4):152-154,  1993 


Page  152 


The  Association  between  the  Gastropod  Biiccinaiiops  cochlidium 
(Dillwyn,  1817)  and  the  Sea  Anemone  Phlyctenanthus  australis 
Carlgren,  1949  in  Patagonian  Shallow  Waters 


Guido  Pastorino 

Division  Paleozoologia  Invertebrados 

Museo  de  Ciencias  Naturales 

Paseo  del  Bosque  s/  n 

1900  La  Plata,  Buenos  Aires, 

Argentina 


ABSTRACT 

The  association  between  Buccinanops  cochlidium  (Dillwyn) 
and  Phlyctenanthus  australis  Carlgren  is  described.  Both  spe- 
cies inhabit  in  the  sandy  infralittoral  of  Puerto  Piramide,  Chu- 
but  Province,  Argentina,  The  densities  and  percentages  of  gas- 
tropods with  anemones  attached  are  reported,  and  the  possible 
benefits  to  the  sea  anemone  and  the  gastropod  are  discussed. 

Key  ivords:  Symbiosis,  Ecology,  Anemones,  Buccinanops. 


INTRODUCTION 

The  association  between  actinians  and  gastropod  shells 
occupied  bv  hermit  crabs  has  been  frequently  described 
(e.  g.  Faurot,  1910,  1932;  Brunelli,  1913;  Ross^  1960;  Ross 
&  Sutton,  1961a,  1961b;  McLean  and  Mariscal,  1973; 
Balasch  et  a/.,  1977;  Brooks  and  Mariscal,  1986).  How- 
ever few  papers  have  been  published  describing  the  re- 
lationship between  actinians  and  living  gastropods  in  the 
South  Atlantic  (Bellisio  et  ai  1972;  Arnaud,  1978). 

This  paper  reports  on  the  association  between  Bucci- 
nanops cochlidium  (Dillwyn,  1817)  and  Phhjctenanthiis 
australis  Carlgren,  1949,  from  northern  Patagonia. 

MATERIALS  AND  METHODS 

Specimens  of  Buccinanops  cochlidium  and  B.  globulos- 
um  were  collected  during  November  1989  and  October 
1990  while  SCUBA  diving  at  depths  of  5-11  m  in  the 
sandy  infralittoral  zone  off  Puerto  Piramide  (42°35'S, 
64°17'W),  Chubut  Province,  Argentina.  In  February  1992, 
18  samples  were  collected  from  various  tlepths.  The  in- 
formation obtained  from  each  .sample  included  depth, 
area,  number  of  individuals  of  each  species  of  gastropod, 
and  number  of  anemones  attached  per  gastropod.  Data 
from  samples  were  combined  to  give  a  general  pattern 
for  Puerto  Piramide.  DejJth  was  determined  using  a  cap- 
illary depth  gauge  (Dacor  model  lA'.G  &  MCXi). 

The  material  was  first  relaxed  at  low  temperatures  and 


then  preserved  in  70  %  formalin.  The  specimens  are 
housed  at  the  Division  Zoologia  Invertebrados,  Museo  de 
Ciencias  Naturales,  La  Plata,  Buenos  Aires,  Argentina. 

RESULTS 

The  anemone  Phlyctenanthus  australis  commonly  cov- 
ers the  entire  dorsal  surface  of  the  mollusk  shell  with  its 
pedal  disc,  leaving  only  the  aperture  free  (fig.  4).  In  some 
cases  as  many  as  two  specimens  of  P.  australis  were 
attached  to  a  single  gastropod  (fig.  5).  Usually  only  a 
single  anemone  is  attached  either  to  the  spire  or  the  body 
whorl.  When  in  situ  specimens  are  viewed  from  above, 
only  the  completely  extended  oral  disc  of  the  anemone 
is  visible,  with  the  tip  of  the  siphon  of  B.  cochlidium 
extending  from  beneath  its  edge.  The  gastropods  usually 
remain  completely  buried.  Occasionally,  crawling  spec- 
imens with  more  than  one  attached  anemone  were  ob- 
served. 


-42-S 


Dib  C.  Tremouilles 


Figure  I.  Map  of  Peninsula  Valdes  area  showing  the  location 
of  the  study  area. 


G.  Pastorino,  1993 


Page  153 


Figures  2-5.  Buccinanops  cochlidium  (DilKvyn)  with  Phhjctenanthus  australis  Carlgren  attached.  1.  Oblique  apertural  view  of 
the  gastropod  showing  egg-capsules  attached  to  the  columellar  callus.  2.  Dorsal  view  of  the  same  specimen.  3.  P.  australis  covering 
the  entire  shell  of  the  gastropod.  4.  B.  cochlidium  with  two  anemones  attached  (arrowheads).  All  x   1 

Table  I.  Halli\  metric  distriljuliDii  oi  liuiiinaniipn  glohi»iiint.  B  cochliiliuiu.  ami  Flihiitcnantlnis  australis  at  Puerto  Piriimide, 
\ri;eiitina 


l)c|)lh  range 

(ni) 


5-5.9 


(i-B.9 


8-8.9 


9-9.9 


10-10  9 


No.  samples 

Species 

Gastropods  nr- 

SD 

%  with  one  anemone 

%  with  two  anemones 

'"c  wilhuul  ancinuiic^ 


B. 

C0( 

■hlid 

f. 

28.3 

0.60 

0.51 

0.68 

049 

13.8 

0.31 

0.35 

0.16 

0  10 

— 

82.35 

73.4 

67.74 

66.6 

— 

— 

13.3 

22.58 

20.0 

100 

17  (i5 

1 .  ■) .  ■) 

9(iS 

13  4 

Page  154 


THE  NAUTILUS,  Vol.  106,  No.  4 


As  samples  were  collected  just  after  the  hatching  pe- 
riod of  B.  globiilosum .  their  densit\  was  probably  over- 
estimated. However,  B.  globulosum  is  alwa\s  far  more 
abundant  than  B.  cochlidium.  Of  the  total  gastropods 
sampled  72.53%  had  one  anemone  attached,  13.97%  had 
two.  Anemone  free  specimens  comprised  only  13.50%  of 
the  sample.  No  correlation  between  depth  and  anemone/ 
gastropod  density  was  found. 

DISCUSSION 

Two  species  of  Buccinanops.  B.  glolmlosum  (Kiener, 
1834)  and  B.  cochlidium  (Dillwyn,  1817),  inhabit  the 
sublittoral  zone  near  Puerto  Piramide.  Buccinanops  glob- 
ulosum ranges  from  the  lower  midlittoral  into  the  sub- 
littoral to  a  depth  of  6  m  below  low  tide.  At  depths  greater 
than  6  m  it  is  replaced  b>  B.  cochlidium  (see  Table  1). 
Buccinanops  cochlidium  is  the  only  gastropod  that  par- 
ticipates in  the  gastropod-anemone  association,  although 
Phlyctenanthus  australis  was  also  found  attached  to  rocks 
in  the  intertidal  zone.  Phlyctenanthus  australis  was  not 
previously  known  from  south  of  Mar  del  Plata.  At  this 
locality  it  may  be  attached  to  the  decapod  Libinia  spi- 
nosa  Milne-Edwards,  1834  as  noted  by  Zamponi  (1977). 
.\mong  the  other  species  of  gastropods  living  in  the  area 
onK  Odontocymbiola  magellanica  (Gmelin,  1791)  is  large 
enough  to  be  able  to  carry  attached  anemones,  but  P. 
australis  is  not  known  to  attach  itself  to  the  shell  of  this 
volutid. 

Ross  (1974)  suggested  that  one  of  the  possible  benefits 
to  the  anemone  was  increased  mobility,  enabling  it  to 
obtain  greater  amounts  of  food  and  also  to  avoid  adverse 
physiological  conditions.  The  advantages  for  the  gastro- 
pod, on  the  other  hand,  are  not  as  clear.  McLean  and 
Mariscal  (1973)  and  Brooks  (1980)  suggested  that,  in  the 
case  of  the  hermit  crabs,  protection  provided  by  the 
nematocysts  of  the  anemone  would  be  a  benefit.  This 
ma>'  also  be  applicable  to  B.  cochlidium .  The  gastropod 
apparently  is  not  adversely  affected.  This  may  be  in- 
ferred from  the  normal  reproductive  activity,  indicated 
by  the  egg-capsules  attached  to  the  columellar  callus  of 
the  females  (figs.  1,  3). 

ACKNOWLEDGMENTS 

I  wish  to  acknowledge  M.  L.  Pastorino  who  supported 
part  of  the  travel  expenses.  J.  Muzon  and  T.  Simanauskas 
gave  valuable  comments.  I  am  grateful  M.  Griffin's  help 
with  the  English  version  of  the  manuscript.  The  manu- 
script benefited  from  reviews  by  G.  Vermeij  and  an  anon- 
\mous  reviser.  This  work  was  carried  out  during  the 
tenure  of  a  scholarship  granted  by  the  Consejo  Nacional 
de  Investigaciones  Cientificas  y  Tecnicas  (CONICET), 
Argentina. 


LITERATURE  CITED 

.\rnaud,  P.  M.  1978.  Observations  ecologiques  et  biologiques 
surle  Voliitidae  antarctique  Harpovohila  charcoti  (Lamy, 
1910)  (Gastropoda  prosobranchia).  Haliotis  7:44-46 

Balasch,  J.,  J.  Cuadras  and  G,  Alonso  1977,  Distribution  of 
Calliactis  parasitica  on  gastropod  shells  inhabited  by  Dar- 
danus  arrosor.  Marine  Behavior  and  Physiology  5(1):37- 
44. 

Bellisio,  N.  B.,  R.  B.  Lopez  and  A.  Tomo.  1972.  Distribucion 
vertical  de  la  fauna  bentonica  en  tres  localidades  antar- 
ticas;  Bahia  Esperanza,  Isla  Peterman  y  ,\rchipielago  Mel- 
chior.  Contribucion  del  Instituto  Antartico  .'Krgentinol42: 
1-87, 

Brooks,  W,  R.  1988,  The  influence  of  the  location  and  abun- 
dance of  the  sea  anemone  Calliactis  tricolor  (LeSueur)  in 
protecting  hermit  crabs  from  octopus  predators.  Journal 
of  E.\perimental  Marine  Biology  and  Ecology  116:15-21, 

Brooks,  W.  R.  and  R.  N.  Mariscal.  1986.  Population  variation 
and  behavioral  changes  in  two  pagurids  in  association  with 
the  sea  anemone  Calliactis  tricolor  (LeSueur).  Journal  of 
Experimental  Marine  Biology  and  Ecology  103:275-289. 

Brunelli,  G  1913,  Ricerche  etologiche.  Osservazioni  edes- 
perienze  sulla  simbiosi  dei  Paguridi  e  delle  .Attinie  Zool- 
ogische  Jahrbucher,  ,'\bteilung  fur  allgemeine  Zoologieund 
Physiologic  der  Tiere,  Jena  34:1-26. 

Faurot,  L.  1910.  Etude  sur  les  associations  entre  les  Pagures 
et  les  Actinies;  Eupagurus  Prideauxi  Heller  et  Adamsia 
palliata  Forbes,  Pagurus  striatus  Latreille  et  Sagartia  par- 
asitica Gosse.  .Archives  de  Zoologie  experimentale  et  ge- 
nerale  5:421-86. 

Faurot,  L.  1932.  .'Actinies  et  Pagures.  Etude  de  ps\chologie 
animale.  Archives  de  Zoologie  experimentale  et  generale 
74:139-54. 

McLean,  R.  B.  and  R.  N.  Mariscal.  1973.  Protection  of  a 
hermit  crab  by  its  symbiotic  sea  anemone  Calliactis  tri- 
color. Experientia  29:128-130. 

Ross,  D.  M,  1960.  The  association  between  the  hermit  crab 
Eupagurus  hernhardus  (L)  and  the  sea  anemone  Cal- 
liactis parasitica  (Couch),  Proceedings  of  the  Zoological 
Society  of  London  1.34(1  ):4.3-.57, 

Ross,  D.  M.  1974,  Behavior  patterns  in  associations  and  in- 
teractions with  other  animals.  In:  L.  Muscatine  and  H.  M. 
Lenhoff  (eds.)  Coelenterate  Biology,  Academic  Press,  New 
York,  p.  163-212. 

Ross,  D.  M.  and  L.  Sutton,  1961a,  The  association  between 
the  hermit  crab  Dardanus  arrosor  (Herbst)  and  the  sea 
anemone  Calliactis  parasitica  (Couch).  Proceedings  of  the 
Royal  Societ)  of  London  155:282-291. 

Ross,  D.  M.  and  L.  Sutton,  1961b  The  response  of  the  sea 
anemone  Calliactis  parasitica  to  shells  of  the  hermit  crab 
Pagurus  hernhardus.  Proceedings  of  the  Roval  Society  of 
London  155:266-281. 

Zamponi.  M  O  1977.  La  anemonofauna  de  Mar  del  Plata  y 
localidades  vecinas.  1.  Las  anemonas  Boloceroidaria  y  En- 
domvaria  Coelenterata,  .Actiniaria  Neotropica  23(70):  133- 
136.' 


THE  NAUTILUS  106{4):155-173.  1993 


Page  155 


Patterns  of  Diversity  and  Extinction  in  Transmarian 
Muricacean,  Buccinacean,  and  Conacean  Gastropods 


Edward  J.  Peluch 

Deparlnieiit  of  Cifology 
Florida  Atlantic  I'niversity 
Boca  Raton,  FL  33-131,  USA 


ABSTRACT 

.•\n  analysis  of  patterns  of  species-richness  in  ten  muricacean, 
buccinacean,  and  conacean  families  found  in  the  Maryland 
Miocene  formations  has  uncovered  a  previously-undetected 
Serravallian  Stage  extinction  event  that  decimated  the  fauna 
of  the  Calvertiaii  Subprovince  of  the  Transmarian  Province 
Prior  to  the  extinction  event,  the  molluscan  fauna  of  the  Langhi- 
an  Stage  (Calvert  Formation)  contained  man\  tropical  elements 
(here  referred  to  as  "caloosagenic  taxa  )  that  were  derivatives 
of  the  Miocene  Caloosahatchian  Province.  At  the  Langhian- 
Serravallian  boundary  and  the  initiation  of  Choptank  deposi- 
tion, most  of  the  earlier  caloosagenic  taxa  suffered  a  regional 
extinction.  Breakdown  of  local  ecosystems,  probably  due  to  a 
cooling  marine  climate,  increased  toward  the  end  of  Choptank 
time,  culminating  in  the  extinction  of  prominent  endemic  thai- 
did  and  turriil  genera  During  the  latest  Serravallian  Stage  and 
the  beginning  of  Tortonian  time  (St.  Mar\  s  Formation),  the 
marine  climate  returned  to  Langhian  conditions  and  there  was 
an  acconipan\ing  return  to  pre-Choptank  levels  of  diversity 
The  post-Choptank  molluscan  assemblages  differed,  however, 
in  containing  new  caloosagenic  taxa,  such  as  Conus,  and  had 
an  extremely  reduced  thaidid  fauna  Several  biogeographically 
important  new  taxa  are  also  proposed  and  these  include  three 
new  genera,  Patuxentrophon  n.gen  (Muricidae:  Trophoninae), 
Calvcrturris  n  gen.  (Turridae:  Mangeliinael,  and  Transma- 
riaturris  n  gen  (Turridae:  Mangeliinae),  and  six  new  species, 
Ecphorostjcon  lindajoijceac  n  sp.  (Thaididae:  Ecphorinae), 
Biiccinoftisus  paluxentcnsis  n.sp.  (Fasciolariidae),  Busycnty- 
pus  choptankciuis  n.sp.,  Turrifulgur  miinjiandicus  n.sp.,  and 
Turrtfulgur  prunicola  n.sp.  (all  Melongenidae:  Busyconinae), 
and  Calvcrturris  .schniidti  (Turridae:  Mangeliinae). 

Keywords:  Extinction,  Maryland,  Miocene,  Transmarian  Prov- 
ince. 


INTRODUCTION 

As  a  biogeographical  entity,  the  Transmarian  Molluscan 
Province  of  the  northern  and  central  Atlantic  Coastal 
Plain  has  been  relatively  unstudied.  This  early  Neogene 
paratropical  province,  whose  fauna  evolved  in  the  Oli- 
gocene  and  became  extinct  during  the  Messinian  Mio- 
cene, has  recently  been  shown  to  contain  numerous  en- 
demic gastropod  taxa  (Petuch,  I988a,b,c).  Primary  among 
these  are  a  plethora  of  distinctive  genera  in  the  super- 


families  Muricacea,  Buccinacea,  and  Conacea,  many  of 
which  are  indicators  of  the  provincial  boundaries. 

At  its  height,  the  Transmarian  province  extended  from 
the  present-day  Cape  Fear,  North  Carolina  area  north- 
ward to  at  least  Nova  Scotia  (figure  46),  and  encompassed 
three  distinct  subprovinces;  the  southern  Pungoian  Sub- 
province,  the  central  Calvertian  Subprovince,  and  the 
northern  Sankatian  Subprovince  (Petuch,  1988b).  The 
Pungoian  Subprovince  was  centered  on  the  Miocene  Al- 
bemarle Embayment  of  North  Carolina,  while  the  Cal- 
vertian Subprovince  had  its  focus  on  the  diverse  biotopes 
of  the  Salisbury  Embayment  of  Chesapeake  Bay,  south- 
ern New  Jersey ,  Delaware,  Maryland,  and  northern  Vir- 
ginia. Since  the  best-preserved  and  most  accessible  Trans- 
marian faunas  are  located  along  the  Patuxent  River,  the 
St.  Mary's  River  and  the  western  shore  of  Chesapeake 
Bay  in  Maryland,  and  since  these  were  the  principal 
faunas  used  in  my  provincial  analysis,  1  will  deal  only 
with  Calvertian  taxa  in  this  paper. 

In  Maryland,  the  Calvertian  Transmarian  molluscan 
fauna  is  contained  within  several  extremely  fossiliferous 
members  of  three  classic  Miocene  formations  and  a  pos- 
sible new  formation,  the  Calvert  (contemporaneous  with 
the  well-known  Kirkwood  Formation  of  New  Jersey) 
(Langhian  Stage),  the  Choptank  (early  Serravallian  Stage), 
the  Little  Cove  Point  Unit  (late  Serravallian  Stage)  (the 
status  of  this  unit  is  still  in  debate  among  authorities;  I 
here  follow  the  nomenclature  of  Ward  and  Blackwelder, 
1980:D4),  and  the  St.  Mary's  (latest  Servallian-early  Tor- 
tonian Stages).  Within  these  formations,  a  molluscan  ex- 
tinction event,  interspersed  between  two  periods  of  spe- 
ciation  and  diversity  augmentation,  can  be  recognized 
within  the  stratigraphic  record  and  these  give  insight 
into  possible  middle  Miocene  catastrophic  climatic  fluc- 
tuations along  the  eastern  coast  of  North  America.  In 
this  paper,  I  also  hope  to  shed  some  light  on  the  higher 
order  diversity  patterns  of  the  Transmarian  muricacean, 
buccinacean,  and  conacean  gastropod  families.  Several 
important  new  taxa  are  also  described  here  in  order  to 
fill  some  nomenciatural  gaps  in  the  Calvertian  fossil  rec- 
ord. These  include  three  new  genera,  Patuxentrophon 
(Muricidae:  Trophoninae),  Calverturris  (Turridae),  and 
Transmariaturris  (Turridae),  and  new  species  of  Ec- 


Page  156 


THE  NAUTILUS,  Vol.  106,  No.  4 


phorosycon  (Thaididae:  Ecphorinae),  Buccinofusus 
(Fasciolariidae).  Busycotypus  (Melongenidae:  Biis\con- 
inae),  Turrifulgur  (Melongenidae:  Busyconinae),  and 
Calverturris  (Turridae).  The  three  superfamilies  studied, 
which  include  ten  families  and  fort>  -one  genera  (listed 
here  in  Appendi.v  1 ),  are  excellent  indicators  of  ecosystem 
collapse  during  times  of  e.xtrenie  climatological  stress. 

THE  TRANSMARIAN  GASTROPOD  FAUNA 

As  demonstrated  in  my  earlier  work  (Petuch,  1988b:  11), 
the  Transmarian  molluscan  fauna  represents  one  of  the 
strangest  mixtures  of  gastropods  that  can  be  found  an\- 
where  in  the  fossil  record.  Unlike  any  known  Recent 
malacofauna,  the  Transmarian  assemblages  contain  sym- 
patric  suites  of  both  boreal,  high  arctic  gastropod  genera 
such  as  Boreotrophon  (Muricidae),  Admete  (Cancellari- 
idae),  Oenopota  (Turridae),  and  Euspira  (Natici- 
dae),  and  eutropical  genera  such  as  Conits  (Leptoconus) 
(Conidae).  Striotercbriim  ^Terebridae),  Laevityphis 
(Muricidae),  and  Cymatosyrinx  (Turridae).  These  arctic 
and  tropical  elements  coexisted  with  a  large  compliment 
of  endemic  Transmarian  genera.  Although  the  Trans- 
marian Province  contained  taxa  from  several  tropical 
families,  the  fauna  lacked  most  of  the  classic  eutropical 
index  groups,  such  as  the  Strombidae,  Turbinellidae,  Cy- 
praeidae,  Ovulidae,  Cerithiidae,  Potamididae,  and  Ly- 
riinae  (N'olutidae)  (Petuch,  1988b).  Based  on  the  absence 
of  these  eutropical  elements,  the  Transmarian  province 
can  only  be  considered  to  have  been  a  paratropical  faunal 
region. 

\s  determined  by  Gibson  (1967),  using  foraminiferal 
assemblages,  the  marine  climate  of  the  Transmarian 
Province  ranged  from  cool-temperate  to  temperate,  with 
only  a  small  seasonal  fluctuation  in  water  temperature. 
This  climatic  stabilit\  allowed  the  evolution  of  a  physi- 
ologicalK  stenothermal  temperate  molluscan  fauna,  with 
the  endemic  genera  being  unable  to  live  in  both  the 
colder  boreal  conditions  to  the  north  and  in  the  warmer 
tropical  conditions  of  the  Miocene  Caloosahatchian  Pro\- 
ince  to  the  south  (Petuch,  1988b)  (figure  46).  The  Trans- 
marian arctic  and  tropical  elements  represent  physiolog- 
ically eurythermal  "invaders"  from  the  Miocene  Boreal 
and  Caloosahatchian  Provinces.  These  opportunistic  spe- 
cies appear  to  have  evolved  physiological  tolerations  for 
the  warmer  (for  the  boreal  taxa)  or  colder  (for  the  tropical 
taxa)  water  conditions  of  the  Transmarian  region,  and 
evolved  their  own  endemic  Transmarian  temperate  spe- 
cies complexes. 

Transmarian  faunal  elements  that  were  derived  from 
the  tropical  Miocene  Caloosahatchian  Province  are  here 
referred  to  as  "caloosagenic".  The  caloosagenic  influence 
varied  through  time  in  the  Calvertian  Subprovince,  with 
some  stages,  such  as  the  early  Tortonian,  containing  many 
tropical  taxa  and  other  stages,  such  as  the  early  Serra- 
vallian,  containing  fewer  taxa.  Some  caloosagens  were 
present  only  during  the  Langhian  Stage  and  disappeared 
from  Mar\  land  during  the  Serravallian  Stage,  being  found 
subsequently  in  the  fossil  beds  of  Virginia  and  Florida. 


Primary  examples  of  these  early  Calvertian  caloosagens 
include  Phyllonolus  [P.  millvillcnsis  (Richards  and  Har- 
bison, 1942)),  Sfurcxiclla  (M.  shilohensis  (Heilprin, 
1887)),  Oliva  (O.  harrisi  Martin,  1904),  Amaea  (A.  re- 
ticulata Martin,  1904  and  A.  prunicola  Martin,  1904), 
Niso  (N.  lineata  (Conrad,  1841)),  Ficus  (F.  harrisi  (Mar- 
tin, 1904)),  and  Architectonica  (A.  trilineatum  (Conrad, 
1841)).  Other  caloosagens  make  their  first  appearance  in 
the  Calvertian  Subprovince  at  the  very  end  of  Salisbury 
deposition,  in  the  late  Serravallian  and  early  Tortonian 
Stages.  Some  of  these  late  Calvertian  caloosagenic  taxa 
include  the  previously-mentioned  conid  genus  Conus 
(Leptoconus  species  complex)  and  the  volutacean  genus 
Mitra  (A/,  mariana  Martin,  1904).  Still  other  caloosa- 
genic groups,  such  as  the  buss  conine  genera  Busycutypus 
and  Turrifulgur,  the  xolutid  Volutifusus,  and  the  tur- 
ritellid  Torcula,  existed  within  the  Calvertian  area 
throughout  the  Langhian,  Serra\allian,  and  Tortonian 
Stages. 

Transmarian  faunal  elements  that  were  derived  from 
the  arctic  Boreal  Province  are  here  referred  to  as  "boreo- 
gens".  As  in  the  case  of  the  caloosagens,  the  boreogenic 
influence  also  fluctuated  through  time,  being  more  prev- 
alent in  some  molluscan  assemblages  than  in  others.  Un- 
like the  caloosagens,  however,  very  few^  boreogens  were 
present  in  the  Calvertian  Subprovince  during  the  Lan- 
ghian stage.  Of  these,  only  a  single  genus  (the  naticid 
Euspira)  is  known  to  have  ranged  throughout  the  entire 
Maryland  Miocene.  The  Tortonian  Stage,  on  the  other 
hand,  saw  the  appearance  of  many  new  boreogenic  taxa 
that  previously  had  not  existed  within  the  Salisbury  Em- 
bavment.  Examples  ot  these  later  boreal  "invaders'  in- 
clude the  previously-mentioned  genera  Admete  and 
Oenopota. 


DYNAMICS  OF  TRANSMARIAN 
DIVERSITY 

By  recognizing  that  three  separate  groups  of  faunal  el- 
ements coexisted  within  the  Transmarian  Province,  the 
caloosagens,  the  boreogens,  and  the  Transmarian  endem- 
ics, it  is  now  possible  to  undertake  a  fine-resolution  anal- 
ysis of  the  diversit)'  patterns  of  C-alvertian  Transmarian 
muricacean,  buccinacean,  and  conacean  gastropods. 
These  are  discussed  in  the  following  sections.  A  listing 
of  all  known  Calvertian  species  in  these  three  superfam- 
ilies is  given  in  Appendix  2. 

Muricacean  diversity  patterns:  .Although  comprising  two 
families  and  fourteen  genera,  the  Transmarian  murica- 
cean taxa  were  not  distributed  equally  over  time.  Only 
two  muricid  genera,  Laevityphis  and  Chesatrophon  (fig- 
ure 32),  and  one  thaidid  genus,  Ecphora  s.s.,  are  tem- 
poralK  distributeil  throughout  the  Langhian,  Serraval- 
lian, and  Tortonian  Stages.  During  Langhian  time,  in 
the  Calvert  and  Kirkwood  (New  Jersey)  Formations,  only 
two  other  muricid  groups,  the  caloosagens  Murexiella, 
and  Phyllonutns.  are  present.  The  Langhian  ecphorine 
thaidids,  on  the  other  hand,  underwent  a  large  species 


E.  J.  Petuch,  1993 


Page  157 


radiation,  evolving  at  least  four  species  of  Ecphora  s.s., 
at  least  five  species  of  Trisecphora  (figure  38),  two  species 
of  Ecphorosycon.  and  two  species  of  Chesathais  (figure 
34)  (Petuch,  1989).  (Note:  The  genus  Chesathais,  al- 
though possibly  arising  from  a  Caloosahatchian  stem  spe- 
cies (C.  biconiciis),  was  represented  in  Maryland  by  the 
C.  ecclesiastictis  species  complex  and  can  be  considered 
to  be  an  endemic  Transmariaii  radiation. )  The  large  ec- 
phorines  apparently  dominated  the  Langhian  molluscan 
assemblages,  as  their  shells  are  abundant  in  most  units 
of  the  Calvert  Formation.  The  small  Calvert  muricids  of 
the  genera  Laevitijphis.  Chcsatrophon,  Miirexiella.  and 
Patuxentrophon  (figures  1,  2),  however,  are  rare  in  most 
assemblages  and  appear  to  have  occupied  the  ecological 
"fringe"  areas. 

During  Serravallian  time,  the  Maryland  Mtirexiella 
species  disappeared,  (and  presumabK  the  New  Jersey 
Kirkwood  Phyllonotus  species),  while  the  caloosagen 
Urosalpinx  first  made  an  appearance.  This  ubiquitous 
Caloosahatchian  ocenebrine  ta.xon  survived  to  become 
an  abundant  component  of  the  late  Serravallian-Tortoni- 
an  assemblages  of  the  St.  Marys  Formation,  and  is  also 
extant  in  the  Recent.  Also  appearing  in  the  early  Ser- 
ravallian was  the  small  muricid  Stephanosalpinx  (figure 
31).  This  distinctive  muricacean  disappeared  by  the  late 
Serravallian  and  is  confined  to  the  Choptank  Formation. 

As  during  the  Langhian,  the  predominant  muricacean 
radiation  during  the  early  and  middle  Serravallian  time 
was  a  complex  of  ecphorine  thaidids.  During  this  time, 
the  genus  Ecphora  s.s.  was  abundantly  represented  by 
at  least  four  species  and  four  stratigraphic  subspecies. 
The  subgenus  Trisecphora,  although  not  as  species-rich 
as  during  Langhian  time,  was  also  abundantly  repre- 
sented by  two  species  and  at  least  one  stratigraphic  sub- 
species. Likewise,  the  morphologically-conservative  ge- 
nus Chesathais  was  also  present  in  the  Choptank 
Formation,  in  this  case  as  a  species  and  two  stratigraphic 
subspecies.  Although  the  primitive  genus  Ecphorosijcon 
was  originally  thought  to  have  become  extinct  at  the  end 
of  Langhian  time  (Petuch,  1989:43),  the  recent  discovery 
of  a  rare  Choptank  species  (£.  lindajoyceae  n.sp.,  de- 
scribed here)  demonstrates  that  the  genus  survived  into 
earliest  Serravallian  time  and  was  part  of  the  rich  Chop- 
tank  ecphorine  fauna. 

Although  ecphorine  species  diversity  declined  at  the 
end  of  Choptank  time,  muricid  diversity  increased  dra- 
matically at  the  beginning  of  Little  Cove  Point-St.  Mary's 
time  (late  Serravallian-Tortonian).  The  abundant  pres- 
ence of  the  trophonine  genera  Chcsatrophon  (figure  32), 
Boreotrophon,  (figure  29,  30)  Scalaspira  (figure  4),  Li- 
rosoma  (figure  3),  and  the  ocenebrine  genera  Urosalpinx 
and  Mariasalpinx  (figure  33),  particularly  underscore 
this  shift  toward  muricid  dominance.  This  predominance 
is  emphasized  even  further  by  the  fact  that  only  a  single 
species  of  Ecphora  occurs  sympatrically  with  the  rich 
trophonine  and  ocenebrine  faunas  found  within  the  var- 
ious members  of  the  St.  Mary's  Formation.  Although  only 
a  single  Ecphora  species  is  found  within  any  bed  of  the 
St.  Mary's  Formation,  these  species  are  much  larger  than 


any  ancestral  species  from  the  Calvert  and  Choptank 
Formations,  and  are,  in  fact,  the  largest  muricaceans 
found  in  the  entire  Maryland  Miocene  {Ecphora  gard- 
nerae  Wilson,  1987  often  exceeds  lengths  of  110  mm). 
This  gigantism  gave  the  post-Choptank  ecphorine  sur- 
vivors an  ecological  advantage  over  their  more  numer- 
ous, but  much  smaller,  muricid  cousins — allowing  them 
to  utilize  large  prey  items,  such  as  adult  specimens  of 
the  bivalves  Mercenaria  and  Dosinia,  which  were  in- 
accessible to  the  contemporaneous  muricids  (Petuch,  1989: 
15).  Interestingly  enough,  of  all  the  late  Transmarian 
endemic  muricid  genera,  only  Scalaspira,  Lirosoma,  and 
Boreotrophon  (Transmarian  complex)  survived  into  the 
early  Pliocene  (Zanclian  Stage).  The  last  species  of  these 
groups,  S.  strumosa  (Conrad,  1832),  L.  sulcosa  (Conrad, 
1830),  and  B.  tetricus  (Conrad,  1832),  respectively,  can 
be  found  in  the  Yorktown  Formation. 

Buccinacean  diversity  patterns:  Comprising  five  fam- 
ilies and  twelve  genera,  the  Transmarian  buccinacean 
gastropods,  unlike  the  muricaceans,  were  more  equitably 
distributed  over  time.  Eight  of  the  twelve  genera  are 
found  in  all  three  Maryland  Miocene  formations  and  are 
represented  by  continuous  species  lineages.  Only  during 
St.  Mary's  time  do  additional  genera  appear,  and  these 
include  the  Transmarian  endemics  Mariafusus  (figure 
39),  Pseudaptyxis  (figure  35),  and  Bulliopsis.  No  boreo- 
genic  buccinacean  taxa  are  known  from  the  Pungoian, 
Calvertian,  or  Sankatian  Subprovinces. 

Of  the  five  Transmarian  buccinacean  families,  only 
the  Fasciolariidae  showed  any  appreciable  augmentation 
through  time.  During  Calvert  and  Choptank  time,  the 
family  Fasciolariidae  was  represented  by  only  a  single 
genus,  Buccinoftisus  (with  two  of  the  species  shown  here 
in  figures  7,  13,  14).  By  Little  Cove  Point-St.  Mary's  time, 
however,  the  number  of  fasciolariid  genera  had  jumped 
to  three,  with  the  Transmarian  endemics  Mariafusus  and 
Pseudaptyxis  possibly  representing  offshoots  that  now 
coexisted  with  their  Buccinofusus  ancestral  stock  (Pe- 
tuch, 1988b).  The  primitive  genus  Buccinofusus,  in  par- 
ticular, underwent  rapid  evolution,  producing  three  spe- 
cies in  the  Calvert  Formation  and  one  species,  each,  in 
the  Choptank  and  St.  Mary's  Formation  and  Little  Cove 
Point  Unit.  LInlike  the  other  Transmarian  buccinacean 
families,  all  the  Transmarian  fasciolariid  genera  were 
endemic  to  the  province,  with  most  taxa  being  confined 
to  the  Salisbury  Embayment  of  the  Calvertian  Subprov- 
ince. 

As  the  complete  opposite  of  the  Fasciolariidae,  the 
families  Buccinidae  and  Columbellidae  were  represent- 
ed in  the  Transmarian  Province  only  by  caloosagenic 
genera.  The  species  within  these  tropically-derived 
groups,  however,  represented  endemic  Transmarian  spe- 
cies radiations  that  paralleled  those  of  their  congeners  to 
the  south.  The  genera  Celatoconus,  Ptychosalpinx,  and 
Solenosteira  were  all  common  and  prominent  compo- 
nents of  the  Calvert,  Choptank,  and  St.  Mary's  faunas. 
In  the  case  of  Ptychosalpinx,  the  genus  had  radiated  into 
an  endemic  Transmarian  species  complex  with  at  least 


Page  158 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figure.  1-14.  Muricid,  thaidid.  and  fasciolariid  gastropods  from  the  Tran.sn.arian  Province.  1,  2.  Patuxentrophonpatuxentenm 
(Marfn  1904),  dorsal  and  ventral  views  of  18  tnm  specimen.  Zone  17,  Choptank  Formation,  Drum  Cliff,  St.  Mary  s  County, 
Maryland.  3.  Lirosoma  mariana  Petuch,  1988,  holotype,  length  27  mm.  St.  Mary's  Formation,  St.  Marys  River,  Maryland.  4. 
Scalasvira  harasewychi  Petuch,  1988,  holotvpe,  length  13  mm,  St.  Marys  Formation,  St  Mary  s  River,  \  ar>land  5  Ecphorosycon 
lindaiouceae  new  species,  holotvpe,  length  ,55  mm  (incomplete).  Zone  16,  Choptank  Formation,  Sandgates,  St.  Mary  s  County. 
Maryland   UF  21466.  6.  Ecphorosycon  pamlico  (Wilson,  1987).  68  mm  specimen  from  Zone  10,  Calvert  tormation.  Plum  1  omt. 


E.  J.  Petuch,  1993 


Page  159 


two  sympatric  species  during  St.  Mary  s  time.  The  Col- 
umbellidae,  which  normally  e.xhibits  a  high  degree  of 
species-richness  in  eutropical  areas,  was  represented  in 
the  paratropical  Transmarian  Province  by  only  a  single 
genus,  Mitrella.  Although  common  in  all  the  Maryland 
Miocene  exposures,  Mitrella  never  underwent  any  spe- 
cies radiation  and  was  represented  b\  only  two  sympatric 
species  in  the  Calvert  Formation  and  one  species,  each, 
in  the  Choptank  and  St.  Mary's  Formations  and  in  the 
Little  Cove  Point  Unit. 

Considering  that  the  overwhelming  majority  of  Trans- 
marian biotopes,  as  in  the  Recent  Carolinian  Province, 
were  composed  of  soft-bottom  sand  or  mud  areas,  the 
family  Nassariidae  could  be  expected  to  be  present  in 
the  Calvertian  region  as  a  large  species  radiation.  Al- 
though comprising  only  two  genera,  the  caloosagenic 
Ilyanassa  and  the  endemic  Bulliopsis,  the  Nassariidae 
dominated  many  facies  of  the  Calvert  and  St.  Mary  s 
Formations  and  were  the  predominant  group  of  small 
buccinaceans  in  the  Transmarian  Province.  Ilyanassa  is 
present  in  all  Maryland  Miocene  formations  as  a  series 
of  species  swarms,  with  at  least  two  species  in  the  Calvert 
Formation,  four  in  the  Choptank,  and  at  least  three  in 
the  St.  Mary's.  This  species-richness  is  comparable  to  that 
seen  in  the  Pliocene,  in  such  formations  as  the  Duplin 
and  Yorktown.  The  endemic  Transmarian  genus  Bul- 
liopsis, on  the  other  hand,  is  restricted  to  the  Calvertian 
Subprovince  and  appears  only  at  the  end  of  Salisbury 
deposition  during  Little  Cove  Point  and  St.  Mary's  time. 

The  most  extensive  macrobuccinacean  radiation  with- 
in the  Transmarian  Province  is  seen  in  the  busyconine 
melongenids.  These  large  and  abundant  predatory  gas- 
tropods were  represented  in  the  Calvertian  Subprovince 
by  species  swarms  in  three  genera;  Busycotypus  (ex- 
amples shown  here  in  figures  15-20),  Sycopsis  (figures 
36,  37),  and  Tiirrifulgur  (examples  shown  here  in  figures 
21-23,  26,  27).  These  groups  are  present  in  all  the  Mary- 
land Miocene  formations  and  exhibit  only  gradual  mor- 
phological shifts  through  time.  Sycopsis.  the  least  com- 
monly encountered  busyconine,  was  present  in  the 
Transmarian  Province  as  only  a  single  species  per  for- 
mation. The  caloosagenic  genera  Tiirrifulgur  and  Bu- 
sycotypus, on  the  other  hand,  were  often  present  as  sym- 
patric species  pairs,  and  in  some  cases,  such  as  Busycotypus 
during  St.  Mary's  time,  was  present  as  a  complex  of  four 
species.  The  species  radiations  of  both  of  these  genera 
will  be  discussed  in  greater  detail  under  the  description 
of  new  Busycotypus  and  Turrifulgur  species  in  the  sys- 
tematic section  at  the  end  of  this  paper. 


Conacean  diversity  patterns:  Of  the  three  Transmarian 
conacean  families,  only  two,  the  Turridae  and  the  Tereb- 
ridae,  are  temporally  distributed  throughout  the  Mary- 
land Miocene  and  are  found  in  all  three  formations.  The 
third  family,  the  Conidae,  is  only  known  from  the  St. 
Mary  s  Formation  and  makes  its  appearance  in  the  Cal- 
vertian Subprovince  only  at  the  end  of  Salisburv  depo- 
sition. Although  an  abundant  and  diverse  family  in  the 
Miocene  Caloosahatchian  Province,  the  Conidae  was 
represented  in  the  Transmarian  Province  by  only  a  single 
subgenus  of  Conus.  This  group,  Leptoconus  (figures  40, 
41),  had  evolved  into  a  complex  of  three  endemic  sym- 
patric species,  C.  diluvianus  Green,  1830,  C.  sanctae- 
mariae  Petuch,  1988,  and  C.  asheri  Petuch,  1989,  by  the 
end  of  St.  Mary's  time. 

The  Terebridae,  although  a  common  component  of  all 
Transmarian  molluscan  assemblages,  was  present  as  only 
two  genera,  the  caloosagenic  Strioterebrum  and  the  en- 
demic genus  Laevihastula.  The  absence  of  core  tropical 
terebrids  such  as  Myurella,  Hastula.  and  Paraterebra, 
which  are  all  found  in  the  Miocene  Caloosahatchian  and 
Gatunian  Provinces  to  the  south,  points  to  the  marginal 
paratropical  nature  of  the  Transmarian  Province,  and 
the  Calvertian  Subprovince  in  particular.  Strioterebrum 
shows  an  interesting  bimodal  evolutionary  pattern,  with 
two  spurts  of  speciation,  one  during  Calvert  time  and 
one  at  the  end  of  Salisbury  deposition,  during  St.  Mary's 
time.  The  decline  in  Strioterebrum  species-richness  dur- 
ing Choptank  time  may  be  linked  to  lower  water  tem- 
peratures during  the  early  Serravallian  stage.  L'nlike  Stri- 
oterebrum. Laevihastula  is  absent  in  the  Calvert 
Formation  but  underwent  a  large  species  radiation  dur- 
ing St.  Mary's  time,  evolving  at  least  four  sympatric 
species. 

The  third  Transmarian  conacean  family,  the  Turridae, 
is  present  in  the  Calvertian  Subprovince  as  four  subfam- 
ilies, twelve  genera,  and  over  thirty-five  species,  making 
it  the  single  largest  toxoglossate  radiation  in  the  Maryland 
Miocene.  Of  the  twelve  turrid  genera,  nine  were  endemic 
to  the  Transmarian  Province,  two  were  caloosagenic,  and 
one  was  boreogenic.  The  most  species-rich  subfamily  in 
the  Calvertian  Subprovince,  the  Clavinae,  comprised 
three  genera;  the  Transmarian  endemics  Chesaclava  (fig- 
ure 42)  and  Sedilopsis.  and  the  caloosagen  Cymatosy- 
rinx.  The  largest  number  of  species  of  these  genera,  par- 
ticularly Cymatosyrinx,  is  found  in  the  St.  Mary's 
Formation,  and  this  species-richness  probably  reflects  a 
response  to  the  warmer  marine  climate  during  the  early 
Tortonian. 


Calvert  County,  Marviand.  7.  Buccinojusus  devexus  (Conrad.  1843).  60  mm  specimen  from  Zone  10,  C^aivert  Formation.  Plum 
Point,  Calvert  County,  Maryland  8.  Ecphorosycon  kalyx  (Petucli,  1988).  23  mm  specimen  from  Zone  12,  Calvert  Formation, 
Scientist  s  Cliffs,  Calvert  County,  Maryland  9.  Ecphorosycon  lindajoyccae  new  species,  reconstructed  paratvpe,  length  69  mm. 
Zone  16,  Choptank  Formation.  Sandgates,  St.  Mar\ 's  Count),  Maryland  10.  Ecphorosycon  pandico  (Wilson,  1987),  juvenile 
specimen,  length  32  mm.  Zone  10,  Calvert  Formation.  Plum  Point,  Calvert  Count),  Marvland  11.  Buccinofusus  parilis  (Conrad, 
1832),  106  mm  specimen  from  St.  Mary  s  Formation,  St.  Mary's  River,  Maryland.  12.  Buccinofusus  chesapeakensis  Petuch,  1988, 
holotype,  length  93  mm.  Little  Cove  Point  Unit,  Little  Cove  Point,  Calvert  County,  Maryland.  13,  14.  Buccinofusus  patuxentensis 
new  species,  dorsal  and  ventral  views  of  holotvpe,  length  76  mm.  Zone  17,  Choptank  Formation.  Drum  Cliff,  St.  Marv's  County, 
Maryland,  UF  21499 


Page  160 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figures  15-28.  Busvconine  meloiigeiml  and  ti.rrid  gastropods  from  the  Transmarian  Province.  15,  16.  Busycotypuschoptankensis 
new  species,  dorsal  and  ventral  views  of  h.ilotNpe,  length  192  mm.  Zone  19,  Choptank  Formation,  Drum  C:liff,  St.  Mary  s  County, 
Maryland  UF  2.'3798.  17.  Rusycotypu.s  ralvcrtcnsis  I'etuch,  1988,  dorsal  view  of  holot>pe,  44  nun  (mcomplete).  Zone  10,  C,aivert 
Formation,  Plum  Point,  Calvert  County.  Maryland  18.  Unsyrolypus  clwsapcahcnsis  Petuch,  1988,  dorsal  view  of  holotype  length 
83  mm  Little  Cove  Point  Unit,  Little  Cove  Point,  Calvert  County,  Maryland  i9.  Biisycotypm  coronatum  (Conrad,  1840),  dorsa 
view  of  59  mm  specimen,  St.  Mary's  Formation,  St.  Marys  River,  Maryland.  20.  Busycotypus  rugosum  (Conrad,  1843)  ventra 
view  of  64  mm  specimen,  St.  Mary's  Formation,  St.  Mary's  River.  21,  26.  Turrifulgur  marylandicus  new  species,  dorsal  and  ventral 


E.  J.  Petuch,  1993 


Page  161 


In  the  Maryland  Miocene,  the  subtamily  Mangeliinae 
was  represented  by  the  most  number  of  genera,  with  the 
Transmarian  endetnics  Calvert urris,  MariadriUia.  and 
Transnmriatnrris,  tlie  caloosagen  Ghjphustuniti.  and  the 
boreogen  Oenopota.  L'nlike  the  C^lavinae,  the  mangeliine 
lurrids  were  not  equally  distributed  through  time,  but 
were  divided  into  two  groups:  one  that  was  confined  to 
the  CaKert  and  C^lioptank  Formations  and  the  other  that 
appeared  onl>  during  St.  Mar\  s  time.  The  former  group 
contains  Calverturris  (figures  24,  25),  Transmariat urris 
(figure  28),  and  Clyphostoma.  ail  of  which  disappeared 
during  the  mid-Serravallian.  The  latter  group  contains 
only  MariadriUia,  which  is  common  in  the  St.  Mary's 
Formation. 

The  subfamily  Turriculinae  contained  the  largest- 
known  Calvertian  turrid  species  radiation,  that  of  the 
endemic  genus  Mariatiirricida  (figure  45).  This  group  of 
large,  fusiform  conaceans  had  evolved  over  seven  species 
during  the  Maryland  Miocene,  and  these  were  probably 
the  primary  predators  on  large  infaunal  polychaetes.  The 
last-known  species,  Mariatiirricida  hiscatenaria  (Con- 
rad, 1834),  may  have  competed  with  the  caloosagenic 
Leptoconus  species  complex  of  the  St.  Mary's  Formation. 
Another  endemic  Transmarian  turriculine  group,  Ches- 
asyrinx  (figure  43),  is  rare  in  the  Calvert  and  Choptank 
Formations  but  is  abundant  in  the  St.  Mary's  Formation, 
where  it  evolved  into  at  least  two  sympatric  species.  This 
surge  of  Chesasyrinx  evolution  may  have  been  in  re- 
sponse to  the  warmer  marine  climate  during  St.  Mary's 
time.  Coinciding  with  the  early  Tortonian  speciation  acme 
of  Chesasyrinx  is  the  appearance  of  a  new  turrid  group, 
Nodisurculina  (figure  44),  which  is  restricted  to  the  St. 
Mary's  Formation. 

The  subfamily  Turrinae,  which  is  characterized  by 
large  species  radiations  and  complexes  of  genera  in  most 
Neogene  tropical  provinces,  was  poorly  represented  in 
the  Transmarian  Province.  Only  a  single  genus,  Herni- 
pleurotoma,  was  present,  but  was  equally  distributed 
throughout  the  entire  Maryland  Miocene  and  is  found 
in  all  three  formations.  This  endemic  Transmarian  genus 
is  morphologically  very  conservative,  with  all  known  spe- 
cies closely  resembling  each  other.  Throughout  its  tem- 
poral range,  Hemipleurotoma  probably  occupied  the 
ecological  "fringe  "  area,  having  had  to  compete  with 
the  extensive  vermivorous  radiations  of  the  subfamilies 
Clavinae,  Mangeliinae,  and  Turriculinae.  I'nderscoring 
this  possible  competitive  exclusion  is  the  fact  that  onl\ 
a  single  Hemipleurotoma  species  is  found  in  each  for- 
mation, as  opposed  to  the  contemporaneous  sympatric 


species  swarms  of  such  successful  vermivores  as  ^edi- 
liopsis,  Cymatosyrinx.  and  Mariaturricula. 

DYNAMICS  OF  TRANSMARIAN  EXTINCTION 

Based  on  both  the  presence  and  absence,  through  time, 
of  caloosagenic  and  endemic  taxa  in  the  Calvertian  area, 
an  early  Serravallian  extinction  event  can  be  delineated 
within  the  temporal  sequence  of  the  Maryland  Miocene. 
This  extinction  interval  began  at  the  initiation  of  Chop- 
tank  deposition,  at  the  boundary  of  Planktonic  Fora- 
minifer  Zones  NlO-Nll  and  N12'(Gibson,  1983:38),  and 
reached  a  peak  at  the  end  of  Choptank  deposition,  at 
the  boundary  of  Planktonic  Foraminifer  Zones  N12  and 
N13.  During  this  time,  which  ranges  from  13  million 
years  B.P.  to  approximately  12.3  million  years  B.P.  (Gib- 
son, 1983:  fig.  2),  several  dominant  gastropod  groups 
disappeared  from  the  Transmarian  ecosystems,  resulting 
in  post-Choptank  molluscan  assemblages  that  differed  in 
appearance  from  those  of  Langhian  age. 

Since  eutropical  caloosagenic  taxa  such  as  Clyphos- 
toma, Oliva,  Niso,  Ficus.  and  Architectonica  disap- 
peared at  the  end  of  Calvert  time,  but  boreogenic  taxa 
such  as  Oenopota  and  Euspira  persisted  throughout 
Choptank  time,  it  can  be  assumed  that  a  climatic  cooling 
event  was  the  primary  culprit  in  the  extinction  of  the 
dominant  Langhian  Transmarian  gastropod  taxa.  As 
pointed  out  by  Stanley  (1986),  a  "refrigeration  "  event, 
such  as  the  Serravallian  cooling  episode,  is  most  probably 
the  cause  of  both  regional  and  mass  extinctions  of  neritic 
faunas  throughout  the  Phanerozoic.  Since  many  of  the 
typical  Transmarian  endemic  genera,  such  as  Chesatro- 
phon,  Buccinofusus,  Sycopsis,  Chesaclava,  and  Maria- 
turricula. survived  the  Serravallian  extinction  and  are 
present  in  the  subsequent  St.  Mary's  Formation,  the 
Choptank-aged  refrigeration  event  was  not  as  severe  as 
that  of  the  mass  extinction  episode  during  the  late  Tor- 
tonian and  Messinian  Stages  (the  "Transmarian  Extinc- 
tion" of  Petuch,  1988b:  12).  During  that  late  Miocene 
time,  over  95%  of  the  Transmarian  gastropod  species 
lineages  disappeared.  The  Serravallian  extinction  epi- 
sode, then,  may  have  been  a  precursor  to  the  catastrophic 
Transmarian  Extinction  of  two  million  years  later.  The 
reduction  of  muricacean,  buccinacean,  and  conacean 
species-richness  during  Choptank  time  is  shown,  graph- 
ically, in  figure  47. 

Although  not  totally  devastating  to  Transmarian  gas- 
tropod assemblages,  the  Serravallian  extinction  was  ap- 
parently severe  enough  to  cause  the  extinction  of  tem- 


views  of  holotype,  length  (incomplete)  60  mm.  Zone  10,  Calvert  Formation,  Plum  Point,  Calvert  C;oiinty,  Maryland,  I'F  21467. 
22.  Turrifidfiur  lurriculus  Petuch,  1988,  holot>  pe,  length  32  mm,  .St.  Mary's  Formation.  .St,  Mar\  's  Hivt-r,  2.'$.  Turrifidgur  fusiforme 
(Conrad,  1840).  (i9  mm  specimen,  St.  Mary's  Formation,  St.  Mary's  River.  24.  Calverturris  hcllacrcnata  (Conrad,  1841),  28  mm 
specimen.  Zone  10,  Calvert  Formation,  Plum  Point,  (Calvert  County,  Maryland  25.  Calverturris  schmidti  new  species,  dorsal  view 
of  incomplete  holotype  (outline  reconstructed),  length  2.3  mm.  Zone  17,  Choptank  Formation  at  Drum  Cliff,  St.  Mary's  County, 
Maryland,  VF  21.500.  27.  Turrifulgur  prunicola  new  species,  dorsal  view  of  holotype,  length  3.5  mm.  Zone  10,  Calvert  Formation, 
Plum  Point,  Calvert  County,  Maryland,  UF  21468  28.  Transmariatiirris  calvertensis  (Martin,  1904),  dorsal  view  of  40  mm  specimen. 
Zone  10,  Calvert  Formation,  Plum  Point,  Calvert  County,  Maryland. 


Page  162 


THE  NAUTILUS,  Vol.  106,  No.  4 


Figures  29-45.  Representative  species  of  endemic  Traiismarian  gastropod  genera  and  endemic  species  complexes.  29.  Boreotro- 
phon  lindae  Petucli,  1988,  length  13  mm,  St  Mary's  Formation  (Boreotrophon  Fischer,  1884,  Transmarian  species  complex).  30. 
Boreotrophon  harasewychi  Petuch,  1988,  length  16  mm,  St  Mar>  's  Formation  {Boreotruphcm  Fischer,  1884,  Transmarian  species 
complex)  .31.  St ephanusalpinx  candelabra  Petuch,  1988,  length  18  mm,  Choptank  Formation,  lateral  view  showing  small  labial 
tooth  (Stephanosalpinx  Petuch,  1988).  32.  Chesatwpliun  chesapeaheanus  (Martin,  1904),  length  11  mm,  St.  Marys  Formation 
(Chesatrophon  Petuch,  1988).  33.  Mariasalpinx  emilyae  Petuch,  1988,  length  28  mm,  St  Mary's  Formation  {Mariasalpinx  Petuch, 


E.  J.  Petuch,  1993 


Page  163 


Figure  46.  Miocene  eastern  North  America,  showing  conti- 
nental configuration  (stippled  area)  and  molluscan  faunal  prov- 
inces, superimposed  upon  the  outline  of  Recent  North  America 
(dotted  line)  C  =  Miocene  Caloosahatchian  Molluscan  Prov- 
ince, T  =  Transmarian  Province,  with  its  three  subprovinces. 
the  Pungoian  U),  the  Calvertian  (2),  and  the  Sankatian  (3); 
thick  fence-line  represents  the  boundarv  between  the  Trans- 
marian and  Miocene  Caloosahatchian  Provinces;  S  =  Salisbury 
Embayment;  A  =  .Albemarle  Embayment 

perature-sensitive  organisms  such  as  the  Transmarian 
Nautilus  species  (Martin.  1904:130).  the  Transmarian 
archeocetian  whales  of  the  genera  Priscodelphinus  Cope, 
1868,  Ixacanthus  Cope,  1890,  Cetophis  Cope,  1868,  and 
Delphinodon  Leidy .  1869.  and  the  porpoise  genus  Oryc- 


30t 


§25 

CO 

m 

3) 

2201 


w 

TJ 

m 
0  15 

m 


10 


C  CH  L  S 

FORMATION 
Figure  47.  Species-richness  of  Calvertian  Transmarian  muri- 
cacean,  buccinacean.  and  conacean  gastropods  during  the  mid- 
dle Miocene,  show  ing  drop  in  diversity  during  Choptank-Little 
Cover  Point  time.  Dotted  line  =  number  of  known  murica- 
ceans;  dashed  line  =  number  of  know  n  buccinaceans;  solid  line 
=  number  of  know  n  conaceans  Calvertian  formations  include 
the  Calvert  (C),  Choptank  (CH),  Little  Cove  Point  Unit  (L), 
and  St  Mary's  (S).  Numbers  of  species  per  superfamily  are 
taken  from  the  data  listed  in  .Appendi.\  2. 


terocetus  Cope.  1867  (Case.  1904).  These  last  mentioned 
cetacean  genera  disappeared  abruptly  at  the  end  of  Cal- 
vert deposition,  contemporaneously  with  the  disappear- 
ance of  the  caloosagenic  gastropod  genera  Glyphostoma, 
Niso,  Oliva,  and  Ficus.  Besides  the  endemic  cetaeans,  a 
large  fauna  of  other  Transmarian  vertebrates  died  out 
at  the  Langhian-Serrav  allian  boundary .  Included  are  the 
giant  endemic  stingray  Raja  dux  Cope.  1867.  the  Myl- 
iobatus  gigas  Cope,  1867 — M.  pachydon  Cope,  1867 
comple.x  of  rays,  the  giant  carchariid  sharks  Carcharias 
collata  Case.  1904.  C.  laevissimiis  (Cope,  1867),  C.  mag- 


1988).  34.  Chesathais  lindae  Petuch.  1988,  length  50  mm,  Choptank  Formation  [Chesathais  Petuch,  1988 1.  35.  Pseuduptyxis 
sanctaemarine  Petuch,  1988,  length  26  mm,  St  Mary  s  Formation  [Pseudaptyxis  Petuch,  1988).  36.  Sycopsis  lindae  Petuch,  1988, 
length  65  mm,  Choptank  Formation  ySycopsis  Conrad,  1867).  37.  Sycopsis  tuberculatum  (Conrad,  1840),  length  52  mm,  St.  Mary's 
Formation  (Sycopsis  Conrad,  1867)  38.  Ecphora  [Trisecphora)  eccentrica  Petuch,  1989.  length  58  mm,  Calvert  Formation  [Tri- 
secphora  Petuch,  1988,  subgenus  of  Ecphora  Conrad,  1843).  39.  Mariafusus  marylandicus  (Martin.  1904),  length  60  mm,  St.  Mary's 
Formation  (Mariafusus  Petuch,  1988).  40.  Conus  {Leptoconus)  asheri  Petuch,  1988,  length  42  mm.  St.  Mary's  Formation  (Lep- 
toconus  Swainson.  1840,  subgenus  of  Conus  Linnaeus.  1758.  Transmarian  species  comple.x).  41.  Conus  (Leptoconnus)  diluvianus 
Green,  1830,  length  63  mm.  St  Mary  s  Formation  {Leptoconus  Swainson,  1840,  subgenus  of  Conus  Linnaeus,  1758,  Transmarian 
species  complex)  42.  Chesaclava  quarlesi  Petuch,  1988,  length  14  mm,  St.  Mary's  Formation  (Chesaclava  Petuch,  1988).  43. 
Chesasyrinx  rotijera  (Conrad,  1830),  length  24  mm,  St  Mar\  s  Formation  (Conrad,  1862),  length  14  mm,  St.  Mary's  Formation 
(Nodisurculina  Petuch,  1988).  45.  Mariaturricula  biscatenaria  (Conrad,  1834),  length  :35  mm,  St.  Mars  s  Formation  (Mariaturricula 
Petuch,  1988). 


Page  164 


THE  NAUTILUS,  Vol.  106,  No.  4 


na  (Cope,  1867),  and  C.  incidens  Case,  1904,  and  the 
giant  manatee  Trichcchus  giganteus  (DeKay,  l^i42) 
(Case,  1904).  It  seems  noteworthy  that  the  Calvert-Chop- 
tank  molluscan  extinction  correlates  with  the  impressive 
mammalian  and  elasmobranch  extinctions,  indicating  that 
there  was  a  province-wide  catastrophic  event  that  af- 
fected many  animal  groups. 

After  the  initial  Langhian-Serravallian  boundary  re- 
frigeration event,  there  was  a  gradual  and  further  cli- 
matic deterioration  during  Choptank  time.  This  gradu- 
alistic  pattern  is  demonstrated  by  the  sequential  extinction 
of  several  taxa.  Victims  of  the  initial  Langhian-Serra- 
vallian boundary  event  were  the  caloosagenic,  eutropicai 
muricid  genera  Murexiella  and  PhijUonotus,  which  dis- 
appeared at  the  end  of  Calvert-Kirkwood  time.  These 
thermophilic  genera,  which  are  widespread  in  the  Recent 
eutropicai  Caribbean  Province  (Petuch,  1987),  would  have 
been  particularly  sensitive  to  lowered  water  temperature. 
Another  caloosagenic  muricid  genus,  Laevityphis,  was 
extremely  rare  in  the  Choptank  Formation,  possibly  in- 
dicating a  sensitivity  to  cooler  temperatures  and  a  mar- 
ginal survival  ability.  Likewise,  the  ecphorine  thaidid 
subgenus  Trisecphora,  which  had  previously  undergone 
a  large  species  radiation  in  the  Calvert  Formation,  was 
present  during  Serravallian  time  only  as  a  single  species 
per  stratigraphic  unit.  This  drop  in  Trisecphora  species 
richness  may  also  have  been  in  response  to  lower  water 
temperatures. 

Of  particular  interest  is  the  thaidid  genus  Ecphuro- 
sycon,  which  appeared  to  have  its  center  of  speciation 
in  the  warmer  southern  Pungoian  Subprovince  (Petuch, 
1989).  This  distinctive  Transmarian  endemic  survived 
into  the  very  beginning  of  Choptank  deposition  (Zone 
16)  as  a  single  Calvertian  species,  but  became  extinct 
before  the  deposition  of  the  upper  bed  of  the  Drum  Cliff 
Member  (Zone  17).  This  ephemeral  existence  of  the  ther- 
mophilic genus  Ecphorosijcon  also  points  to  a  cooling 
event  during  earliest  Serravallian  time.  The  possibly  more 
cool-tolerant  genus  Ecphora  s.s.,  however,  proliferated 
into  several  species  complexes  during  Choptank  time  and 
this  accounts  for  the  lag  in  muricacean  extinction  seen 
in  figure  47.  Climatic  conditions  apparently  deteriorated 
so  greatly  by  the  end  of  Serravallian  time,  however,  that 
even  less  thermophilic  genera  such  as  Ecphora  s.s.,  Ches- 
athais,  and  Trisecphora  declined  in  species  richness.  At 
the  end  of  (>hoptank  deposition,  the  latter  two  thaidid 
genera  and  all  but  one  complex  of  Ecphora  s.s.  became 
extinct,  underscoring  the  ecological  impact  of  the  cooling 
event.  In  the  overlying  Little  Cove  Point  Unit  (late  Ser- 
ravallian Stage),  only  one,  or  possibly  two,  species  of 
Ecphora  s.s.  are  known  to  exist;  a  feeble  "ghost"  of  the 
large  Calvert  and  Choptank  ecphorine  radiations. 

The  turrid  genera  Calvcrturris  and  Transmariaturris 
and  the  trophonine  genus  Patuxcntrophon  also  disap- 
peared during  late  C;hoptaiik  time,  along  with  Ecpho- 
rosycon,  Chesathais  and  Trisecphora.  The  simultaneous 
loss  of  these  stenothermal  temperate  endemics  could  only 
have  been  produced  by  a  refrigeration  catastrophe.  In- 
terestingly enough,  the  large  C^alvertiaii  trophonine  ra- 


diation, which  appears  in  the  subprovince  only  at  the 
end  of  the  Serravallian  refrigeration  event,  may  have 
been  a  left-over  of  a  boreal  invasion  during  the  coldest 
time,  at  the  Choptank-Little  Cove  Point  Unit  boundary. 
This  strong  post-Choptank  boreogenic  influence,  which 
includes  Boreotrophon  and  Scalaspira,  may  have  rep- 
resented physiological  adaptations  to  warmer  water  con- 
ditions during  Little  Cove  Point-St.  Mary's  time. 

SYSTEMATICS 

The  type  material  of  the  following  new  species  is  de- 
posited in  the  invertebrate  paleontology  collection  of  the 
Florida  Museum  of  Natural  History,  University  of  Flor- 
ida, Gainesville,  Florida,  and  bears  UF  numbers. 

Order  Caenogastropoda 

Superfamily  Muricacea 

Family  Thaididae 

Subfamily  Ecphorinae  Petuch,  1988 

Genus  Ecphorosijcon  Petuch,  1988 

Ecphorosycon  lindajoyceae  new  species 
(figures  5,  9) 

Material  examined:  HOLOTYPE — length  55  mm  (in- 
complete), in  basal  bed  (Zone  16)  of  Choptank  Forma- 
tion, along  waterline  of  Patuxent  River  at  Sandgates,  St. 
Mary's  County,  Maryland,  UF21466;  PARATYPE— 
length  69  mm  (reconstructed  with  plasticine  clay),  same 
locality  as  holotype,  collection  of  author. 

Description:  Shell  inflated,  w  ith  globose  body  whorl  and 
distinctly  pyriform  outline;  spire  protracted,  scalariform; 
shoulder  sloping;  subsutural  area  flattened,  producing 
stepped  spire  whorls;  body  whorl  ornamented  v\ith  3 
rounded,  thin,  low,  adherent  cords;  cord  along  shoulder 
of  body  whorl  projecting  upward  (posteriorward),  be- 
coming progressivel)  more  bladelike  on  earlier  whorls; 
entire  body  whorl,  spire  whorls,  and  siphonal  canal  sculp- 
tured with  spiral  threads;  spiral  threads  give  entire  shell 
silky  texture;  spire  whorls  ornamented  w  ith  2  thin  cords, 
with  cord  along  shoulder  being  bladelike  in  form;  si- 
phonal  canal  proportionally  short  and  broad  for  genus; 
umbilicus  wide,  flaring,  well  developed. 

Etymology:  Named  for  my  wife,  Linda  Joyce  Petuch, 
who  assisted  me  with  my  collecting  along  the  St.  Mary's 
River  and  C-hesapeake  Ba\ . 

Discussion:  Previously  (Petuch,  1989:43)  I  had  stated 
that  the  genus  Ecphorosycon  had  become  extinct  at  the 
end  of  Langhian  time,  in  the  upper  beds  of  the  Calvert 
Formation.  The  discovery  of  an  Ecphorosycon  species  in 
the  lowest  bed  of  the  Choptank  Formation  (Zone  16), 
however,  demonstrates  that  the  genus  survived  into  early 
Serraxallian  time.  Within  Zone  16,  the  last  surviving 
Ecphorosycon,  E.  lindajoyceae,  is  rare,  with  only  a  few 
fragmentary  specimens  having  ever  been  collected.  The 
new  species  has  never,  to  my  knowledge,  been  collected 
in  the  upper  C^hoptank  beds  (Zones  17,  18,  and  19),  whose 
faunas  ha\e  been  relativelv  well-studied.  The  absence  of 


E.  J.  Petuch,  1993 


Page  165 


this  large  and  distinctive  ecphorine  from  Zone  17  indi- 
cates that  Ecphowsycon  became  extinct  in  earliest  Ser- 
ravalliaii  time,  making  it  an  excellent  index  fossil  for 
Zone  16. 

Ecphowsycon  lindajoyceae  is  most  similar  to  £.  pam- 
Uco  (Wilson,  1987)  from  Zone  10  of  the  Plum  Point 
Member  of  the  Calvert  Formation  and  from  the  Bon- 
nerton  Member  of  the  Pungo  River  Formation  of  North 
Carolina.  The  new  species  differs  from  £.  pamlico  (figure 
6,  10)  in  being  a  much  wider,  much  more  inflated  shell 
with  a  wider,  stumpier,  less  elongated  siphonal  canal. 
The  spire  of  £.  pamlico  is  higher  and  much  more  pro- 
tracted than  the  spire  of  E.  lindajoyceae,  which  is  dis- 
tinctly lower  and  depressed.  The  spiral  threads  on  the 
spire,  body  whorl,  and  siphonal  canal  of  the  new  species 
are  also  much  finer  than  those  of  the  coarsely-sculptured 
£.  pamlico.  The  new  species  is  also  quite  different  from 
E.  kalyx  (Petuch,  1988)  from  Zones  12  and  14  of  the 
Plum  Point  Member  of  the  Calvert  Formation  (figure 
8),  and  differs  in  being  a  much  more  inflated,  globose 
shell,  in  having  reduced,  less  developed  cords,  and  in 
having  a  sculpture  pattern  of  fine  spiral  threads. 

The  preservation  of  shells  within  Zone  16  is  poor,  with 
almost  every  specimen  being  collected  in  a  fragmentary 
state.  The  paratype  of  £.  lindajoyceae  (figure  9)  was 
actually  flattened  by  the  pressure  of  sedimentary  com- 
paction and  was  collected,  piecemeal,  as  a  handful  of 
fragments.  These  were  later  reassembled  over  a  template 
of  plasticine  clay.  The  holotype  is  the  best,  most  com- 
plete, specimen  found  to  date.  Although  fragmentary, 
the  holotype  exhibits  enough  salient  characteristics  to 
allow  for  easy  separation  from  the  older  £.  pamlico  and 
£.  kalyx.  At  the  type  locality,  Sandgates  on  the  Patuxent 
River  in  Zone  16  of  the  Choptank  Formation,  Ecpho- 
wsycon lindajoyceae  co-occurs  with  a  large  ecphorine 
fauna,  including  Erp/iora  (Ecphora)  me  ganae  sand  gate - 
sensis  Petuch,  1989,  £.  (Ecphora)  riheri  harasewychi 
Petuch,  1989,  £.  (Ecphora)  choptankensis  vokesi  Petuch, 
1989,  Ecphora  (Trisecphora)  smithac  Petuch,  1988,  and 
Chesathais  lindae  donaldasheri  Petuch,  1989. 

Family  Muricidae 

Subfamily  Trophoninae  Cossmann,  1903 

Genus  Patuxent rophon  new  genus 

Diagnosis:  Shell  vase-shaped,  with  sharply  angled,  car- 
inated  shoulder  and  high,  distinctly  scalariform  spire; 
body  whorl  wide  and  inflated,  tapering  rapidly  anteriorly 
into  siphonal  canal;  siphonal  canal  long,  slender,  straight, 
equal  in  length  to  body  whorl;  bod\  whorl  ornamented 
with  10  large,  rounded  cords;  smaller,  secondary  cords 
often  present  between  larger,  primary  cords;  shoulder 
cord  largest;  spire  whorls  ornamented  with  5  cords;  si- 
phonal canal  ornamented  with  15-20  large,  rounded 
cords;  aperture  oval. 

Type  species:  Chrysodomus  patuxentensis  Martin,  1904, 
C;hoptank  Formation,  Serravallian  Miocene  of  Maryland 
(figures  1,  2). 


Other  species  in  Patuxentrophon:  Patuxentrophon 
unnamed  species.  Zone  10  ot  the  Plum  Point  Member 
of  the  (Jalvert  Formation. 

Etymology:  A  combination  of  "Patuxent",  for  the  Pa- 
tuxent River  of  Maryland,  and  the  muricid  genus  Tro- 
phon. 

Discussion:  Martin  (1904:184)  referred  this  small  muri- 
cid group  to  the  archaic  neptuniid  genus  Chrysodomus 
Swainson,  1852,  primarily  on  the  basis  of  a  similarity  in 
ribbed  sculpture  patterns.  Patuxentrophon,  however,  dif- 
fers from  neptuniids  in  being  a  much  tinier  shell  with  a 
proportionally  much  longer  and  much  better  developed 
siphonal  canal.  The  new  genus  is  distinctly  trophonine, 
having  the  same  general  size,  siphonal  canal  develop- 
ment, and  spiral  sculpture  pattern  as  living  Boreotrophon 
species  such  as  the  Oregonian  B.  disparilis  (Dall,  1891) 
and  living  Trophonopsis  species  such  as  the  Alaskan  T. 
kamchatkanus  (Dall,  1902).  Patuxentrophon  also  shares 
many  characteristics  with  the  late  Serravallian,  Tortoni- 
an,  and  Messinian  trophonine  genera  Lirosoma  Conrad, 
1862  (figure  3),  and  Scalaspira  Conrad,  1862  (figure  4), 
having  the  sculpture  of  the  former  and  the  shape  of  the 
latter.  It  is  possible  that  the  Langhian-early  Serravallian 
Patuxentrophon  represents  the  stem-stock  from  which 
the  later  Miocene  Lirosoma  and  Scalaspira  evolved. 

Superfamily  Buccinacea 

Family  Fasciolariidae 

Subfamily  Fasciolariinae  Gray,  1853 

Genus  Buccinofusus  Conrad,  1868 

Buccinofusus  patuxentensis  new  species 

(figures  13,  14) 

Material  examined:  HOLOTYPE — Length  77  mm, 
from  Zone  17,  Drum  Cliff  Member  of  Choptank  For- 
mation, at  Drum  Cliff,  Jones  Wharf,  St.  Mary's  County. 
Maryland,  Patuxent  River,  UF  21499;  PARATYPE— 
length  54  mm,  same  locality  as  holotype,  collection  of 
author. 

Description:  Shell  elongately  fusiform,  with  very  ele- 
vated, protracted  spire;  subsutural  area  sloping,  shoulder 
rounded;  shoulder  and  middle  section  of  body  whorl 
ornamented  with  10-12  low,  axially-arranged  riblike 
knobs;  spire  whorls  with  8-12  riblike  knobs  bordering 
suture;  entire  shell,  in  turn,  sculptured  with  extremely 
numerous  fine  spiral  cords  and  threads;  cords  finest  along 
subsutural  area,  becoming  coarser  on  midbody,  and 
coarsest  on  siphonal  canal;  siphonal  canal  well  developed, 
short  in  proportion  to  length  of  spire. 

Etymology:  Named  for  the  Patuxent  River,  which  bor- 
ders the  type  locality. 

Discussion:  Buccinofusus  patuxentensis  occupies  the 
evolutionary  midpoint  between  the  primitive  B.  devexus 
(Conrad,  1843)  of  the  Calvert  Formation  (figure  7)  and 
the  more  advanced  B.  chesapeakensis  Petuch,  1988  of 
the  Little  Cove  Point  Unit  (figure  12)  and  B.  parilis 


Page  166 


THE  NAUTILUS,  Vol.  106,  No.  4 


(Conrad,  1832)  of  the  St.  Mary's  Formation  (figure  11). 
Morphologically,  B.  patuxentensis  resembles  the  slender 
and  proportionally  higher-spired  B.  devexus  in  shape, 
but  shares  the  coarser  spiral  sculpture  of  B.  chesapeak- 
ensis.  Of  the  known  Buccinofusus  species,  B.  patuxen- 
tensis is  closest  to  the  Little  Cove  Point  B.  chesapeak- 
ensis,  but  differs  in  having  finer  and  more  numerous 
spiral  cords  on  the  body  whorl  and  spire,  and  in  having 
stronger  and  more  numerous  knobs  on  the  spire  whorls. 
Through  time,  from  the  Langhian  to  the  Tortonian, 
the  genus  Buccinofusus  underwent  parallel  shifts  in  two 
morphological  characters;  shell  length-width  ratio  and 
degree  of  sculpture  coarseness.  The  oldest  species,  B. 
devexus,  is  the  most  slender  and  is  only  faintly  sculpted 
with  fine  spiral  threads.  The  Choptank  B.  patuxentensis 
retains  the  slender  shape  of  B.  devexus  but  presages  the 
heavier  sculptural  pattern  of  the  later  forms.  The  Little 
Cove  Point  B.  chesapeakensis  retains  the  sloping  shoulder 
and  fine  sculpturing  on  the  subsutural  area,  as  seen  in 
the  earlier  species,  but  has  the  inflated  body  whorl  and 
coarse,  widely-spaced  cords  around  the  midbody  and 
siphonal  canal  that  typify  the  advanced  forms.  This  char- 
acteristic Transmarian  lineage  culminates  in  the  St.  Mary's 
B.  parilis,  which  has  the  most  inflated  body  whorl,  lowest 
spire,  and  uniformly  coarse  sculpture  over  the  entire 
shell. 

Family  Melongenidae 

Subfamily  Busyconinae  Finlay  and  Marwick,  1937 

Genus  Busycotypus  Wenz,  1943 

Busycotypus  choptankensis  new  species 
(figures  15,  16) 

Material  examined:  HOLOTYPE— Length  192  mm 
(incomplete),  in  Zone  19  of  Choptank  Formation,  at  Drum 
Cliff,  Jones  Wharf,  St.  Mary's  County,  Maryland,  along 
Patu.xent  River,  UF  21498. 

Description:  Shell  vase-shaped,  very  inflated,  with 
ovately-cylindrical  body  whorl;  shoulder  very  broad, 
wider  than  length  of  body  whorl;  shoulder  sharply  an- 
gled, carinated,  ornamented  with  16-20  low,  undulating 
knobs  per  whorl;  spire  low,  stepped;  sutural  area  with 
wide,  deep,  flat-bottomed  sulcus,  producing  widely  can- 
aliculate spire  whorls;  periphery  of  sutural  sulcus  bor- 
dered with  large,  rounded,  undulating  cord;  body  whorl 
ornamented  with  longitudinal  growth  lines,  devoid  of 
spiral  sculpture;  spire  whorls  ornamented  with  6-8  faint 
spiral  threads;  siphonal  canal  broad,  ornamented  with 
10-12  low,  nearly  obsolete  spiral  cords;  aperture  e.\- 
tremely  wide,  open,  flaring,  with  smooth,  unornamented 
interior;  parietal  area  glazed. 

Etymology:  Named  for  the  Choptank  Formation,  for 
which  this  new  species  is  an  index  fossil. 

Discussion:  As  in  the  case  of  Buccinofusus  patuxen- 
tensis. Busycotypus  choptankensis  fills  in  a  gap  in  the 
evolutionary  record  of  the  Maryland  Miocene  Busyco- 
typus lineage.  The  new  species  is  closest  to  B.  chesa- 


peakensis Petuch,  1988  (figure  18)  of  the  younger  Little 
Cove  Point  Unit,  especially  in  being  a  smooth,  unorna- 
mented shell  with  low,  undulating  shoulder  coronations. 
The  Choptank  ancestor  differs  from  its  Little  Cove  Point 
descendant,  however,  in  being  a  much  more  inflated 
shell,  by  being  proportionally  thinner  and  more  fragile, 
and  in  having  less  developed,  lower  coronations  on  the 
shoulder  and  spire  whorls.  Of  the  two  related  St.  Mary's 
species,  B.  coronatum  (Conrad,  1840)  (figure  19)  and  B. 
rugosum  (Conrad,  1843)  (figure  20),  B.  choptankensis  is 
most  similar  to  the  latter,  but  differs  in  having  lower, 
less  developed  coronations  and  in  lacking  the  heavy  spiral 
corded  sculpture  that  characterizes  B.  rugosum.  From 
the  small,  ancestral  B.  calvertensis  Petuch,  1988  (figure 
17),  B.  choptankensis  differs  in  being  a  smooth,  unsculp- 
tured  shell  with  a  much  better  developed,  wider  cana- 
liculate suture. 

Through  time,  the  Transmarian  coronated  Busyco- 
typus species  exhibited  three  morphological  trends;  be- 
coming proportionally  less  inflated,  developing  wider  and 
deeper  sutural  sulci,  and  developing  larger  and  more 
prominent  shoulder  and  spire  coronations.  The  primitive 
B.  calvertensis  is  a  very  wide,  inflated  shell  with  small 
coronations  and  a  tiny,  almost  nonexistent  sutural  canal. 
Busycotypus  choptankensis  retains  the  inflated  shell  shape 
of  the  ancestral  B.  calvertensis,  but  has  large,  well  de- 
fined shoulder  coronations,  and  has  a  very  well  devel- 
oped, wide  sutural  canal.  The  next  youngest  in  the  se- 
quence, B.  chesapeakensis,  has  a  wide  sutural  canal  like 
B.  choptankensis  but  has  even  larger,  better  defined 
shoulder  coronations.  Unlike  the  primitive  species  B.  cal- 
vertensis and  B.  choptankensis,  however,  B.  chesapeak- 
ensis has  the  slender,  more  attenuated  body  form  of  the 
advanced  species.  The  two  St.  Mary's  species,  B.  corona- 
tum and  B.  rugosum,  both  appear  to  have  evolved  from 
the  Little  Cove  Point  B.  chesapeakensis,  and  share  the 
same  attenuated  body  form.  Busycotypus  coronatum  re- 
tains the  smooth,  unsculptured  bod\  whorl  of  6.  chop- 
tankensis and  B.  chesapeakensis,  but  has  the  largest,  most 
pronounced  shoulder  coronations  of  the  whole  complex. 
The  contemporaneous  and  sympatric  B.  rugosum.  on  the 
other  hand,  has  a  heavily  sculptured,  corded  bod>  whorl 
and  large,  rounded,  knoblike  shoulder  coronations.  The 
noncoronated  Bustjcotypus  species  from  the  St.  Mary's 
Formation,  B.  asheri  Petuch,  1988,  and  B.  alveatum 
(Conrad,  1863),  appear  to  belong  to  a  separate  species 
complex  and  are  not  closely  related  to  the  coronated 
forms. 


Genus  Turrifulgtir  Petuch,  1988 

Turrifulgur  marylandicus  new  species 

(figures  21,  26) 

Material  examined:  HOLOTYPE — Length  (incom- 
plete) 60  mm,  in  Zone  10,  Plum  Point  Member  of  Calvert 
Formation,  at  Plum  Point,  Calvert  County,  Maryland, 
UF21467. 

Description:    Shell  elongated,  cylindrical  in  form,  with 


E.  J.  Petuch,  1993 


Page  167 


high,  protracted  spire;  shoulder  sharply  angled,  low  on 
body  whorl,  below  wide,  ver\  sloping  subsutural  area; 
shoulder  ornamented  with  12  evenly-spaced,  small, 
sharply-pointed  knobs;  spire  whorl  ornamented  with  12 
knobs  per  whorl;  siphonal  canal  (missing  on  holotype) 
elongated  and  slender  (extrapolated  from  holotype);  body 
whorl  sculptured  with  numerous  fine  spiral  threads,  which 
become  nearl>  obsolete  around  mid-bod\;  sloping  sub- 
sutural area  sculptured  with  12  large  spiral  threads;  si- 
phonal  canal  sculptured  with  numerous  large,  evenly- 
spaced  spiral  cords;  aperture  proportionally  narrow. 

Etymology:    Named  for  the  State  of  Maryland, 

Discussion:  The  elongate,  high-spired  Turrifulgur 
marijlandicus  stands  out  as  unique  among  the  known 
Transmarian  Turrifulgur  species.  The  extremely  atten- 
uated bodv  form  is  reminiscent  of  T.  atraktoides  (Gard- 
ner, 1944)  from  the  late  Burdigalian  Oak  Grove  For- 
mation of  northern  Florida,  and  the  Langhian  T. 
marylandicus  appears  to  belong  to  the  same  species  com- 
plex. The  new  Maryland  busyconine,  however,  differs 
from  the  older  Oak  Grove  species  in  being  an  even  more- 
elongated  shell  with  a  more  drawn-out  body  whorl,  in 
having  a  more  protracted,  scalariform  spire,  and  in  hav- 
ing a  much  more  sloping  shoulder  and  subsutural  area. 
Turrifulgur  nmnjlandicus  also  has  fewer  knobs  per  whorl 
on  the  spire  than  does  its  more  finely-ornamented  Ca- 
loosahatchian  relative. 

This  new  protracted  Transmarian  Turrifulgur  is  sym- 
patric  with  T.  prunicola  n.sp.  (described  in  the  next 
section)  at  Plum  Point.  Turrifulgur  marylandicus  differs 
from  this  congener  in  being  a  larger,  more  elongated 
shell  with  a  much  higher  spire.  The  presence  of  two 
sympatric  species  of  Turrifulgur  in  the  Calvert  For- 
mation reflects  a  similar  pattern  seen  in  the  early  Langhi- 
an Shoal  River  Formation  of  northern  Florida,  where 
two  species,  T.  aldrichi  (Gardner,  1944)  and  T.  dasum 
(Gardner,  1944),  also  co-occur. 


Turrifulgur  prunicola  new  species 

(figure  27) 

Material  examined:  HOLOTYPE— Length  35  mm,  in 
Zone  10,  Plum  Point  Member  of  Calvert  Formation,  at 
Plum  Point  Calvert  County,  Maryland,  UF21468. 

Description:  Shell  vase-shaped,  fusiform,  with  sharply- 
angled  shoulder;  spire  slightly  stepped,  comparatively 
low  and  flattened  for  genus;  subsutural  area  only  slightly 
flattened,  subplanar;  shoulder  of  body  whorl  ornamented 
with  14  small,  low,  evenly-spaced  knobs;  spire  whorls 
ornamented  with  14  small  knobs  per  whorl;  body  whorl 
pinching-in  abruptly  to  base  of  siphonal  canal;  siphonal 
canal  slender,  narrow;  body  whorl,  siphonal  canal,  spire 
whorls  and  subsutural  area  sculptured  with  numerous 
very  fine  spiral  threads,  giving  shell  silky  appearance; 
aperture  oval. 

Etymology:  Named  for  Plum  Point.  Calvert  County, 
Maryland,  the  type  locality. 


Discussion:  Although  sympatric  with  Turrifulgur 
marylandicus  in  Zone  10,  T.  prunicola  belongs  to  a  sep- 
arate species  lineage.  The  narrow  T.  marylandicus  ap- 
pears to  be  a  member  of  the  T.  atraktoides  (Gardner, 
1944)-r.  dasum  (Gardner,  1944)  species  complex,  while 
T.  prunicola  probably  belongs  to  the  T.  aldrichi  (Gard- 
ner, 1944)  species  complex.  Both  Calvert  species  are  ca- 
loosagenic  offshoots  of  these  contemporaneous  complex- 
es. Although  the  T.  prunicola  lineage  survived  until  the 
Tortonian  Stage  (as  T.  fusiforme  (Conrad,  1840)  and  T. 
turriculus  Petuch,  1988),  the  T.  marylandicus  lineage 
died  off  during  the  Langhian  Stage,  and  is  not  found  in 
the  upper  beds  of  the  Calvert  Formation  nor  in  the 
Choptank  Formation. 

Turrifulgur  prunicola  is  closest  in  general  shell  mor- 
phology to  T.  turriculus  Petuch,  1988  (figure  22)  from 
the  Windmill  Point  Member  of  the  St.  Mary's  Formation. 
The  new  species  differs  from  its  St.  Mary's  descendant 
in  having  a  much  lower  spire,  in  having  fewer  and  small- 
er knobs  on  the  shoulder  and  spire  whorls,  and  in  having 
finer  and  more  delicate  spiral  threaded  sculpture.  Tur- 
rifulgur prunicola  is  also  similar  to  T.  fusiforme  (Con- 
rad, 1840)  (figure  23)  from  Zone  24  of  the  St.  Mary's 
Formation,  but  differs  in  being  a  smaller  shell  with  a 
more  sharply  angled  shoulder  and  proportionally  lower 
spire. 

Superfamily  Conacea 

Family  Turridae 

Subfamily  Mangeliinae  Fischer,  1887 

Genus  Calverturris  new  genus 

Diagnosis:  Shells  elongately  fusiform,  with  high,  pro- 
tracted spires  and  extended,  slender  siphonal  canals; 
shoulders  subcarinated,  ornamented  with  nodulose  cord 
below  greatly  sloping  subsutural  area;  nodulose  cord  bor- 
ders suture  on  spire  whorls;  body  whorls  ornamented 
with  6-8  large,  evenly-spaced  spiral  cords;  fine  spiral 
threads  present  between  cords  of  some  species;  siphonal 
canals  ornamented  with  numerous  spiral  threads;  anal 
notch  shallow,  with  greatest  indentation  corresponding 
to  nodulose  shoulder  cord;  protoconchs  proportionally 
small,  rounded,  domelike,  composed  of  one  and  one-half 
whorls;  apertures  narrow. 

Type  species:  Pleurotoma  bellacrenata  Conrad,  1841, 
Calvert  Formation,  Langhian  Miocene  of  Marvland  (fig- 
ure 24). 

Other  species  in  Calverturris:  Calverturris  schmidti 
n.sp.,  Choptank  Formation,  Serravallian  Miocene  of 
Maryland,  described  here. 

Etymology:  A  composite  of  "Calvert",  for  the  Calvert 
Cliffs  of  Maryland,  and  "turris",   "tower". 

Discussion:  This  small  group  of  Transmarian  turrids  is 
one  of  the  most  morphologically-distinctive  of  the  known 
Chesapeake  Miocene  conacean  gastropods.  The  promi- 
nent nodulose  carina  around  the  shoulders  of  Calver- 
turris species,  along  with  the  large,  evenly-spaced  cords 


Page  168 


THE  NAUTILUS,  Vol.  106,  No.  4 


on  the  body  whorls,  sets  this  genus  aside  from  all  other 
Transmarian  mangeliine  taxa.  In  general  body  form  and 
sculpture  pattern,  Calverturris  is  closest  to  Sediliopsis 
Petuch,  1988,  but  differs  in  having  larger  and  more  in- 
flated shells  and  in  having  the  characteristic  nodulose 
shoulder  carina. 

As  Calverturris  is  only  known  from  the  Calvert  and 
Choptank  Formations,  the  genus  appears  to  have  become 
extinct  during  mid-Serravallian  time. 

Calverturris  schniidti  new  species 

(figure  25) 

Material  examined:  HOLOTYPE — Length  (fragmen- 
tary) 23  mm,  in  Zone  17,  Drum  Cliff  Member  of  Chop- 
tank  Formation,  at  Drum  Cliff,  Jones  Wharf,  St.  Mary's 
County,  Maryland,  along  Patuxent  River,  UF  21500. 

Description:  Shell  shape  and  form  as  outlined  in  genus 
description;  shoulder  carina  with  evenly-spaced,  large 
nodules;  single  large  cord  present  on  sloping  subsutural 
area,  just  posterior  to  nodulose  carina;  body  whorl  or- 
namented with  67  large  spiral  cords  (extrapolated  from 
fragmentary  holotype);  fine  threads  present  between  large 
spiral  cords. 

Etymology:  Named  for  Mr.  Robert  Schmidt  of  Calvert 
Countv,  Maryland,  who  collected  the  holotvpe  at  Jones 
Wharf. 

Discussion:  The  Choptank  Calverturris  schmidti  is  sim- 
ilar to  the  Calvert  C.  bellacrenata,  the  type  of  the  genus, 
but  differs  in  having  larger  and  less  numerous  shoulder 
knobs  and  in  having  fine  spiral  threads  between  the  large 
cords  on  the  body  whorl.  The  steeply-sloping  subsutural 
area  of  C.  bellacrenata  is  also  smooth  and  devoid  of  spiral 
sculpture,  while  that  of  C.  schmidti  is  sculptured  with  a 
single  large  cord  and  several  faint  spiral  threads. 

While  Calverturris  bellacrenata  is  a  common  species 
in  Zone  10  at  Plum  Point,  C.  schmidti  is  a  much  rarer 
shell,  with  only  a  few  fragmentary  specimens  having 
ever  been  collected  along  the  Patuxent  River.  Appar- 
ently, the  genus  was  already  dying  out  by  Choptank  time. 

Genus  Transmariaturris  new  genus 

Diagnosis:  Shells  elongately  terebriform,  with  rounded 
shoulders  and  greatly  protracted,  elevated  spires;  si- 
phonal  canals  proportionally  short,  stubby;  subsutural 
area  of  all  species  slightly  depressed,  producing  faintly 
indented  sutural  band;  last  whorls  with  smooth,  unor- 
namented  shoulders;  spire  whorls  of  some  species  orna- 
mented with  rows  of  low,  riblike  knobs  along  suture 
margin;  spire  whorls  of  some  species  smooth,  unorna- 
mented;  body  and  spire  whorls  sculptured  with  only  very 
faint,  delicate  spiral  thread,  giving  shells  silky  texture; 
siphonal  canals  ornamented  with  10-12  thin  spiral  cords; 
apertures  proportionalK  small,  oval;  protoconchs  un- 
known; anal  notch  shallow,  with  indentation  correspond- 
ing to  depressed  subsutural  band. 


Type  species:  Pleurotoma  (Hemipleurotoma)  calver- 
tensis  Martin,  1904,  Calvert  Formation,  Langhian  Mio- 
cene of  Maryland  (figure  28). 

Other  species  in  Transmariaturris:  "Pleurotoma 
{Hemipleurotoma)"  choptankensis  Martin,  1904,  Chop- 
tank  Formation,  Serravallian  Miocene  of  Maryland. 

Etymology:  A  combination  of  "Transmaria" ,  for  the 
Transmarian  Province,  and  "turris". 

Discussion:  Based  on  general  shell  shape,  I  had  origi- 
nally placed  this  compact  group  of  elongated,  terebri- 
form turrids  in  the  clavine  genus  Chesaclava  Petuch, 
1988  (Petuch,  1988:35).  Further  study,  however  has  shown 
that  these  two  groups  of  turrids  differ  in  a  number  of 
ways  and  actually  belong  in  separate  subfamilies.  Trans- 
mariaturris, although  convergent  on  the  protracted  spire 
torm  of  Chesaclava,  differs  from  that  genus  in  having 
larger,  more  inflated  shells  with  proportionally  larger 
body  whorls,  by  having  longer  and  better  developed 
siphonal  canals,  in  having  more  indented  anal  notches 
that  produce  solenozones,  and  in  having  spriral  sculp- 
turing. The  small,  smooth  Chesaclava  species  closely  re- 
semble miniature  Cymatosyrinx  Dall,  1889  species  and 
develop  a  varix-like  adult  lip  and  stromboid  notch.  These 
two  characters  are  not  seen  in  Transmariaturris. 

Although  the  diagnostic  protoconchs  were  missing  on 
all  Transmariaturris  specimens  examined,  I  have  placed 
this  new  genus  in  the  subfamily  Mangeliinae,  primarily 
on  its  similarity  to  the  genus  Calverturris.  Both  genera 
are  of  the  same  general  shape  and  size,  and  share  the 
same  type  of  anal  notch  and  outer  lip  structure.  Trans- 
mariaturris differs  from  Calverturris,  however,  in  lack- 
ing the  large,  prominent  spiral  cords  on  the  body  whorl 
and  also  in  lacking  the  distinctive  nodulose  shoulder  ca- 
rina. The  subsutural  area  of  Calverturris  is  planar,  while 
that  of  Transmariaturris  is  slightly  depressed. 


ACKNOWLEDGEMENTS 

For  help  in  collecting  research  material  along  the  St. 
Mary's  and  Patuxent  Rivers  and  along  the  Calvert  Cliffs, 
I  thank  Mr.  Donald  F.  Asher,  Mechanicsville,  Maryland, 
Mr.  Robert  Schmidt.  Calvert  Countv',  Maryland,  and  my 
wife,  Linda  J.  Petuch. 


LITERATURE  CITED 

Case,  E.  C  1904.  Systematic  paleontology,  Miocene,  Mam- 
malia, Aves,  and  Reptilia.  In:  Clark,  W.  B.,  G.  B.  Shattuck, 
and  W.  H.  Dall  (eds).  The  Miocene  deposits  of  Maryland. 
Maryland  Geological  Survey,  Miocene,  p.  3-93. 

Gibson,  T  G.  1967  Stratigraphy  and  paleoenvironment  of 
the  Phosphatic  Miocene  Strata  of  North  C^arolina.  Geolog- 
ical Society  ol  ,\nierica  Bulletin  78:631-650 

Gibson,  T.  G.  1983  Straligra|)li\  of  Miocene  through  I.ower 
Pleistocene  Strata  of  the  L^nited  States  Central  .Atlantic 
Goastal  I'lain  In.  Ray,  C.  E.  (ed).  Geology  and  paleon- 
tology of  the  l^ee  Creek  Mine,  North  Carolina  I.  Smith- 
sonian Contributions  to  Paleobiology  53:35-80. 


E.  J.  Petuch,  1993 


Page  169 


Martin,  G.  C.  1904.  Systematic  Paleontology,  Miocene,  Mol- 
lusca.  Gastropoda.  In:  Clark,  W.  B.,  G.  B.  Shattuck,  and 
W.  H.  Dall  (eds  )  The  Miocene  deposits  of  Maryland 
Maryland  Geological  Survey,  Miocene,  p.  131-270 

Petuch,  E.  J.  1987.  New  Caribbean  molluscan  faunas.  The 
C'oastal  Education  and  Research  Foundation,  ('harlottes- 
\ille,  Virginia,  154  p 

Petuch,  E.  J.  1988a.  New  species  of  Ecphora  and  Ecphorine 
Thaidids  from  the  Miocene  of  Chesapeake  Bay,  Maryland. 
U.S.A.  Bulletin  of  Paleomalacology  1(1):1-16. 

Petuch,  E.  J.  1988b.  Neogene  history  of  tropical  American 
mollusks.  The  Coastal  Education  and  Research  Founda- 
tion, Charlottesville,  Virginia,  217  p. 

Petuch,  E.  J.  1988c.  New  gastropods  from  the  Maryland  Mio- 
cene. Bulletin  of  Paleomalacology  l(4):69-80. 

Petuch,  E  J  1989  Field  guide  to  the  Ecphoras  The  Coastal 
Education  and  Research  Foundation,  Charlottesville,  Vir- 
ginia, 140  p. 

Stanley,  S.  M.  1986.  Anatomy  of  a  regional  mass  extinction: 
Plio-Pleistocene  decimation  of  the  Western  Atlantic  Bi- 
valve fauna.  Palaios  1:17-36. 

Ward,  L.  W.  and  B.  W.  Blackvvelder  1980  Stratigraphic 
re\ision  of  I'pper  Miocene  and  Lower  Pliocene  Beds  of 
the  Chesapeake  Group,  Middle  Atlantic  Coastal  Plain.  U.S. 
Geological  Survey  Bulletin  1482-D  61  pp  .5  pis 


Appendix   i.    (Inntiiiued 


T       C       B 


Appendix  1.  Muricacean,  Buccinacean,  and  Conacean  Higher 
Taxa  from  the  Calvertian  Subprovince  of  the  Transmarian 
Province.  T  =  Transmarian  endemic  genus,  C  =  C'aloosagenic 
genus,  B  =  Boreogenic  genus 

T       C       B 

Superfamily  Muricacea 

Family  Muricidae 

Subfamily  Muricinae  da  Costa,  1776 
Phyllonotus  Swainson,  1833  * 

SubfamiK  Muricopsinae  Radwin  and 
D'Attilio,  1971 
Miircxiclla  Clench  and  Farfante,  1945  * 

SubfamiK  Ocenebrinae  Gray,  1847 
Mariawlpinx  Petuch,  1988  * 

Sephanosalpinx  Petuch,  1988  * 

Vrosalpinx  Stimpson,  1865  * 

Subfamily  Trophoninae  Cossmann,  1903 

Boreotrophon  Fischer,  1884  * 

Chesafrophon  Petuch,  1988  * 

Lirosoma  Conrad,  1862  * 

PatuxerUropbon  Petuch,  1992  * 

Scalaspira  C'onrad,  1862  * 

SubfamiK  Txphinae  Cossmann,  1903 
Laevitijphis  Cossmann,  1903  * 

Eamil)  Thaididae 

Subfamily  Ecphorinae  Petuch,  1988 

Chesathais  Petuch,  1988  * 

Ecphora  Conrad,  1843  * 

Ecphorosycon  Petuch,  1988  * 

Trisecphora  Petuch,  1988  * 

SuperfamiK  Buccinacea 

FamiK  Fasciolariidae 


Subfamily  Fasciolariinae  Gray,  1853 

Buccinofnstis  Conrad,  1868 

Marinfusus  Petuch,  1988 
SubfamiK   Fusininae  Swainson,  1840 

Pseudaptyxis  Petuch,  1988 
FamiK  Melongenidae 
Subfamily  Busyconinae  Finlay  and 
Mar  wick  1937 

Busycotypus  Wenz,  1943 

Sycopsis  Conrad,  1867 

Turrifiilgur  Petuch,  1988 
Fatnilv  Buccinidae 

CU'latocontts  Conrad,  1862 

Ptychosalpinx  Gill,  1867 

Solenosteira  Dall,  1890 
Familv  Nassariidae 

Bulliopsis  C^onrad,  1862 

Ilyanassa  Stimpson,  1865 
Eamil)  C^olutnbellidae 

Mitrella  Risso,  1826 
Superfamily  Conacea 
Eamil)  Conidae 

Leptoconus  Swainson,  1840 
Eamil)  Terebridae 

Lacvihastiila  Petuch,  1988 

Striotercbrum  Sacco,  1891 
Family  Turridae 
Subfamily  Clavinae  Powell,  1942 

Chcsaclava  Petuch,  1988 

Cymatosyrinx  Dall,  1889 

Sediliopsis  Petuch,  1988 
Subfamil)  Mangeliinae  P'ischer,  1887 

Calvcrturm  Petuch,  1992  T 

Glyphostonia  Gabb,  1872 

Ocnopota  Morch,  1852 

Mariadrillia  Petuch,  1988 

Transmariaturris  Petuch,  1992 
Subfamily  Turriculinae  Powell,  1942 

Chcsasyrinx  Petuch,  1988 

Mariaturricula  Petuch,  1988 

Nodisurculina  Petuch,  1988 
Subfamil)  Turrinae  Swainson,  1840 

Uemipleurotorua  Co.ssman,  1903 


Page  170 


THE  NAUTILUS,  Vol.  106,  No.  4 


Appendix  2.    Muricacean,  Buccinacean,  and  Conacean  Species  from  the  Calvertian  Subprovince  of  the  Transmarian  Province.  C 
=  Calvert  Formation,  Ch  =  Choptank  Formation,  L  =  Little  Cove  Point  Unit,  S  =  St.  Mary  s  Formation 


C 


CH 


L 


S 


Muricidae 

Muricinae 

'Phi/llonotus  millvillensis  (Richards  and  Harbison,  1942) 

Muric-opsinae 
Murexiella  shiluhcnsis  (Heilprin,  1887) 

Ocenebrinae 

Mariasalpinx  emilyae  Petuch,  1988 
Mariasalpinx  new  species 
Stephanosalpinx  candelabra  Petuch,  1988 
Urosalpinx  riisticus  (Conrad,  1830) 

Trophoninae 

Borcotrophon  hara.sewychi  Petuch,  1988 
Boreotrophon  laevis  (Martin,  1904) 
Borcotrophon  lindae  Petuch,  1988 
Che.satrophon  chesapeakeanus  (Martin,  1904) 
Chcsatrophon  new  species  a. 
Chesatrophon  new  species  b. 
Lirosoma  mariana  Petuch,  1988 
Patuxentrophon  patuxenicnsis  (Martin,  1904) 
Patuxcntrophon  new  species 
Scalaspira  harasewychi  Petuch,  1988 
Scalaspira  vohesae  Petuch,  1988 

Typinae 
Laevityphis  acuticosta  (Conrad,  1830) 
Laevityphis  new  species  a. 
Laevityphis  new  species  b. 

Thaididae 

Ecphorinae 
Chesathais  ecclesiasticus  (Dall,  1915) 
Chcsathai.s  lindae  lindae  Petuch,  1988 
Chcsathaix  lindae  dunaldasheri  Petuch,  1989 
Chesathais  lindae  drumcliffensis  Petuch.  1989 
Chcsathais  whitfieldi  Petuch,  1989 
Eephora  asheri  Petuch,  1988 
Eephora  ealvertensis  Petuch,  1988 
Eephora  ealvertensis  subspecies 
Eephora  chesapeakensis  Petuch,  1992 
Eephora  ehoptankensis  ehoptankensis  Petuch,  1988 
Eephora  ehoptankensis  delieata  Petuch,  1989 
Eephora  choptaiikensis  vokesi  Petuch,  1989 
Eephora  gardnerae  gardnerae  Wilson,  1987 
Eephora  gardnerae  angusticostata  Petuch,  1989 
Eephora  germonae  Ward  and  Gilinsky,  1988 
Eephora  mcganae  meganae  Ward  and  Gilinsky,  1988 
Eephora  meganae  sandgatescnsis  Petuch,  1989 
Eephora  rikeri  riken  Petuch,  1988 
Eephora  rikeri  harasewychi  Petuch,  1988 
Eephora  lurneri  Petuch,  1992 
Eephora  wardi  Petuch,  1989 
Eephora  tvardi  subspecies 
Eephora  leilliamsi  Ward  and  Gilinsky,  1988 
Eephora  (Trisecphora)  eeeentriea  Petuch,  1988 
Eephora  (Triseephoru)  martini  I'ctuch,  1988 
Eephora  (Trisecphora)  patiixentia  patuxentia  Petuch,  1988 
Eephora  (Trisec])}iora)  patuxentia  shaltucki  Pctuth,  1989 
Eephora  (Trisecphora)  ])runieolu  Pelucli,  1988 
Eephora  (Trisecphora)  scientistensis  Petuch,  1992 
Eephora  (Trisecphora)  schmidti  Petuch,  1989 
Eephora  (Trisecphora)  srtiithae  Petuch,  1988 
Eephora  (Trisecphora)  smithae  subspecies 


E.  J.  Petuch,  1993 


Page  171 


Appendix  2.    Continued 


CH 


Ecphora  (Trisecphora)  tricostata  Martin,  1904 
Ecphorosycon  kahjx  (Petuch,  1988) 
Ecphowsycun  lindajoijcvae  Petuch,  1992 
Ecphorosycon  paiulicu  (Wilson,  1987) 
Ecphorosycon  new  species 

Fasciolariidae 
Buccinofusus  (?)  calvertanus  (Martin,  1904) 
Buccinoftisus  chcsapeakensis  Petuch,  1988 
Buccinofusus  devexus  (Conrad,  1843) 
Buccinofusus  migrans  (Conrad,  1843) 
Buccinofusus  parilis  (Conrad,  1832) 
Buccinofusus  patuxentcnsis  Petuch,  1992 
Muriafusus  marylandicus  (Martin,  1904) 
Pscudaplyxis  sanclacrnuruic  Petuch,  1988 

Meiorjgenidae 

Busyconinae 
Busycotypus  alveatum  (Conrad,  1863) 
Busycotypus  asheri  Petuch,  1988 
Busycotypus  calvertensis  Petuch,  1988 
Busycotypus  chcsapeakensis  Petuch,  1988 
Busycotypus  choptankensis  Petuch,  1992 
Busycotypus  coronatum  (Conrad,  1840) 
Busycotypus  rugosum  (Conrad,  1843) 
Sycopsis  lindac  Petuch,  1988 
Sycopsis  tuberculatum  (Conrad,  1840) 
Syncopsis  new  species  a 
Sycopsis  new  species  b. 
Turrifulgur  fusiforme  (Conrad,  1840) 
Turrifulgur  marylandicus  Petuch,  1992 
Turrifulgur  prunicola  Petuch,  1992 
Turrifulgur  turriculus  Petuch,  1988 
Turrifulgur  new  species  a. 
Turrifulgur  nev\  species  b. 

Buccinidae 
Celatoconus  asheri  Petuch,  1988 
Celatoconus  protractus  (Conrad,  1843) 
Ptychosalpinx  lienosa  (Conrad,  1843) 
Ptychosalpinx  lindae  Petuch,  1988 
Ptychosalpinx  pustulosus  Petuch,  1988 
Ptychosalpinx  new  species 
Ptychosalpinx  new  species 
Solenostcira  cundierlandiana  (Gabb,  1860) 
Solenosteira  new  species 

Nassariidae 
Bulliopsis  integra  (Conrad,  1842) 
Bulliopsis  marylandica  (Conrad,  1862) 
Bulliopsis  ovata  (Conrad,  1862) 
Bulliopsis  quadrata  (Conrad,  1830) 
Bulliopsis  subcylindrica  (Conrad,  1862) 
Ilyanassa  calvertensis  (Martin,  1904) 
Ilyanassa  clongata  (Whitfield.  1894) 
Ilyanassa  greensboroensis  (Martin,  1904) 
Ilyanassa  gubernatoria  (Martin,  1904) 
Ilyanassa  marylandica  (Martin,  1904) 
Ilyanassa  pcralta  (Conrad,  1868) 
Ilyanassa  peraltoides  (Martin,  1904) 
Ilyanassa  trivittatoides  (Whitfield,  1894) 
Ilyanassa  new  species 

Columbellidae 
Mitrclla  calvertensis  (Martin,  1904) 
Mitrclla  communis  (Conrad,  1862) 


* 
* 

* 
* 
* 
* 

*  * 
«                     *                     * 

*  * 

* 
* 

* 

* 

* 
* 

*  * 

*  * 
* 

* 


Page  172 


THE  NAUTILUS,  Vol.  106,  No.  4 


.Appendix  2.    C^oiitinucd 


Cll 


Mitrella  new  species  a. 
Milrella  new  species  b. 
Mitrella  new  species  c. 
('onidae 
Contis  (Leptorimtis)  asheri  Petiicli,  1988 
Contis  (Leptoconus)  diluvianiis  Green,  1830 
Contis  (Lcptoco7itts)  sanctaemariae  Petucli,  1988 
Terebridae 
Laevihasliila  inornata  (Whitfield,  1894) 
Laevihastula  marylandica  Petucli,  1988 
Lacvihastiila  paluxentia  (Martin,  1904) 
Laevihastula  simplex  (Conrad,  18.30) 
Laevihastula  sublirata  (C^onrad,  1863) 
Strioterebrum  ealvertensis  (Martin,  1904) 
Strioterchrum  eurvilineata  (Whitfield,  1894) 
Strioterebrum  eurvilirata  (Conrad,  1843) 
Strioterebrum  dalli  (Martin,  1904) 
Strioterebrum  sineera  (Dall,  1895) 
Strioterebrum  ichitficldi  (Martin,  1904) 
Tiirridae 
(^la\  inae 
Chesaelava  dissimilis  (Conrad,  1830) 
Chesaclava  pseudeburnea  (Whitheld,  1894) 
Chesaelava  quarlesi  Petuch,  1988 
Chesaclava  whitfieldi  (Martin,  1904) 
Chesaclava  new^  species 
Cymatosijrinx  limatula  (Clonrad,  1830) 
Cynuitosyrinx  mariana  Petuch,  1988 
Cymatosyrinx  pyramidalis  (Martin,  1904) 
Cymntosyrinx  new  species  a, 

Cymatosyrinx  new  species  li. 

Scdiliopsis  angulata  (Martin,  1904) 

Sediliupsis  ealvertensis  (Martin,  1904) 

Sediliopsis  distans  (Conrad,  1862) 

Sediliopsis  gracilis  (Conrad,  1830) 

Sediliopsis  incilifera  (Conrad,  1830) 

Sediliopsis  paluxentia  (Martin,  1904) 
Mangehinae 

Calverturris  bellacrenata  (C^onrad,  1841) 

Calverturris  schmidti  Petuch,  1992 

Clyphosloma  obtusa  (Martin,  1904) 

Oenopota  cornelliana  (Martin,  1904) 

Ocnopota  marylandica  (Petuch,  1988) 

Oenopota  parva  (Conrad,  18.30) 

MariadriUia  parvoidea  (Martin,  1904) 

Transmariaturris  ealvertensis  (Martin,  1904) 

Trausmariaturris  choplankrnsis  (Martin,  1904) 
Turricuhnae 

Chesasyrinx  mariana  (Martin,  1904) 

Chesasyrinx  rotifera  (Conrad,  18.30) 

Chesasyrinx  new  species 

Mariaturricula  marylandica  (Conrad,  1834) 

Mariaturricula  biscalenaria  (Conrad,  1841) 

Mariaturricula  rugata  (Conrad,  1862) 

Mariaturricula  new  species  a, 

Mariaturricula  new  species  b. 

Mariaturricula  new  species  c. 

Mariaturricula  new  species  d. 

Nodisurculina  engonata  (Conrad,  1862) 
Turrinae 

Ilemipleurotoma  communis  (Conrad,  1830) 


E.  J.  Petuch,  1993  Page  173 


Appendix  2.    Continued 

c:  CM  1. 

Hcmiplcurotoma  protocommiinis  (Martin,  1904)  * 

Ilcmiplcurcitoma  new  species  * 

'  Known  from  the  Cal\ert-equi\alent  kirkwdod  Formation  of  New  Jersey;  possibi\  occurring  in  the  Calvert  Formation. 


THE  NAUTILUS  106(4):174-180,  1993 


Page  174 


Publication  Dates  of  The  Nautilus 


Eugene  V.  Coan 

Department  of  Invertebrate  Zoology 
California  Academy  of  Sciences 
Golden  Gate  Park 
San  Francisco,  CA  94118  USA 


M.  G.  Harasewych 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC  20560  USA 


As  the  publication  dates  of  earlier  issues  of  The  Nautihis 
are  nowhere  readily  accessible,  the  purpose  of  this  note 
is  to  provide  this  information  in  a  single  comprehensive 
listing. 

Although  six  indices  have  been  published  for  The  Nau- 
tilus and  for  its  predecessor.  The  Conchologists  Ex- 
change (Pilsbry  &  Johnson,  1895;  Henderson,  1927;  La 
Rocque,  1951,  1963,  1972;  Abbott,  1979),  none  provide 
a  listing  of  publication  dates.  The  dates  listed  below  are 
compiled  from  a  variety  of  sources. 

Dates  for  issues  published  during  Henry  A.  Pilsbry  s 
tenure  as  Editor  (volume  3,  number  1,  to  volume  71, 
number  3)  were  determined  largely  from  his  published 
bibliographies  (American  Malacological  Union,  1940; 
Baker^  1958;  Clench  &  Turner,  1962),  and  are  identified 
by  an  asterisk  (•)  following  the  date.  As  noted  in  the  first 
of  these  bibliographies  (American  Malacological  Union, 
1940;4),  the  dates  for  volumes  13-45  inclusive  were  de- 
rived from  ledgers  maintained  by  Charles  W.  Johnson, 
then  Managing  Editor. 

Publication  dates  of  issues  prior  to  volume  13,  as  well 
as  of  volume  46,  number  1,  had  been  extrapolated  based 
on  the  dates  of  receipt  of  these  issues  by  several  libraries 
(American  Malacological  Union,  1940:4).  These  dates, 
identified  by  a  dagger  (t),  are  estimates  and  may  in  some 
cases  prove  to  be  one  or  more  days  later  than  the  actual 
date  of  publication. 

During  H.  Burrington  Baker's  association  with  The 
Nautilus,  first  as  Business  Manager  (volume  46,  number 
2,  to  volume  71,  number  3)  then  as  Editor  and  Editor 
Emeritus  (volume  71,  number  4,  to  volume  84,  number 
1),  publication  dates  for  a  volume  were  published  in  the 
"Notes  and  News"  section  of  the  first  or  second  issue  of 
the  subsequent  volume.  This  practice  was  continued  by 
the  succeeding  editor  R.  Tucker  Abbott  (volume  83, 
number  1,  to  volume  100,  number  4)  for  volume  85. 
Dates  determined  from  these  listings  are  marked  by  a 
double  dagger  (|)  following  the  date. 

Exact  dates  for  volumes  86-89  have  not  previously 
been  published.  Mailing  dates  for  these  issues  were  de- 
termined from  records  on  file  in  the  editorial  offices  of 
the  journal  (PS  Forms  3541  and  3542),  are  identified  by 
a  section  mark  (§). 

Beginning  with  volume  90,  dates  of  publication  were 
printed  on  the  cover  and/or  title  page  of  every  issue, 
and  are  not  included  here. 

Dates  derived  from  the  bibliography  of  W.  H.  Dall 
(Bartsch  et  a!..  1946),  identified  by  a  number  sign  (#), 


supplement  and/or  corroborate  the  dating  of  the  earliest 
issues.  For  issues  that  could  not  be  dated  in  any  other 
manner,  the  date  of  the  library  stamp  of  the  Library  of 
Congress  (identified  as  LC),  the  National  Museum  of 
Natural  History  (identified  as  USNM)  the  Museum  of 
Comparative  Zoology,  Harvard  Ihiiversity  (identified  as 
MCZ),  or  the  Boston  Society  of  Natural  Historv  (iden- 
tified as  BSNH,  now  in  the  library  of  R.  Tucker  Abbott) 
is  provided. 


The  Conchologists'  Exchange 

1(1);  July  1886 

1(2);  August  1886 

1(;3);  27  September  1886  LC 

1(4);  October  1886 

1(5);  6  December  1886  t 

1(6);  after  28  December  1886  f 

1(7);  January  1887  [February  25  LC] 

1(8);  Februarv  1887  [March  9  LC] 

1(9/10);  30  April  1887  USNM 

1(11);  May  1887 

1(12);  28  June  1887  LC 

2(1);  20  July  1887  MCZ  [10  August  #,  12  August  LC 

2(2);  August  1887  [15  September  LC:] 

2(;3);  September  1887  [13  October  LC] 

2(4);  October  1887  [5  November  LC,  USNM] 

2(5);  November  1887  [15  December  LC] 

2(6):  December  1887  [23  January  1888  LC] 

2(7):  January  1888  [20  February  LC] 

2(8):  Februarv  1888  [26  March  LC] 

2(9);  2  May,  1888  t  [7  May  LC] 

[Publicalion  suspended.] 


The  Nautilus 


3(1);  5  Mav  1889  f,  # 
3(2):  9  June  1889  t 
3(3);  14  July  1889  f,  # 
3(4):  11  August  1889  t 
3(5):  1  October  1889  t 
3(6):  18  October  1889  t 
3(7);  13  November  1889  t 
3(8);  7  Januarv  1890  f 
3(9);  11  Februarv  1890  f.  # 
3(10);  12  March  1890  t 
3(11);  15  April  1890  t 
3(12):  7  May  1890  f 

4(1):  9  June  1890  t 
4(2):  27  June  1890  t 


E.  V.  Coan  and  M  G.  Harasewvch,  1993 


Page  175 


4(3):  6  July  1890  t 
4(4):  5  August  1890  t 
4(5):  1  October  1890  t 
4(6):  14  October  1890  f 
4(7):  7  November  1890  t 
4(8):  22  December  1890  f.  # 
4(9):  11  January  1891  t 
4(10):  1  February  1891  f 
4(11):  4  March  1891  f 
4(12):  5  April  1891  t 

5(1):  19  Mav  1891  f,  # 
5(2):  17  June  1891  f.  # 
5(3):  10  July  1891  t,  # 
5(4):  19  August  1891  t.  # 
5(5):  17  September  1891  f 
5(6):  18  October  1891  t 
5(7):  18  November  1891  f 
5(8):  9  December  1891  f 
5(9):  14  Januarv  1892  f.  # 
5(10):  5  Februarv  1892  t 
5(11):  25  March  1892  f.  # 
5(12):  17  April  1892  f 

6(1):  15  Mav  1892  t 

6(2):  1  June  1892  f 

6(3):  6  July  1892  f 

6(4):  [?]  August  1892  t 

6(5):  11  September  1892  t 

6(6):  4  October  1892  t 

6(7):  8  November  1892  f 

6(8):  12  December  1892  t 

6(9):  12  Januarv  1893  t 

6(10):  10  February  1893  #  [12  Februarv  f] 

6(11):  [?]  March  1893  t 

6(12):  9?  April  1893  t  [10  April  MCZ] 

7(1):  7  June  1893  t 
7(2):  19  June  1893  f 
7(3):  17  July  1893  f 
7(4):  15  August  1893  t 
7(5):  3  September  1893  f.  # 
7(6):  13  October  1983  t 
7(7):  5  November  1893  t 
7(8):  13  December  1893  f,  # 
7(9):  5  January  1894  t 
7(10):  30  Januarv  1894  f 
7(11):  1  March  1894  f 
7(12):  2  April  1894  t.  # 

8(1):  2  Mav  1894  t.  # 
8(2):  4  June  1894  f 
8(3):  8  July  1894  t.  # 
8(4):  3  August  1894  f.  # 
8(5):  7  September  1894  f 
8(6):  5  October  1894  t 
8(7):  1  November  1894  f.  # 
8(8):  3  December  1894  f,  # 
8(9):  2  January  1895  t 
8(10):  3  Februarv  1895  t,  # 
8(11):  4  March  1895  f,  # 
8(12):  1  April  1895  f 

9(1):  2  Mav  1895  t.  # 
9(2):  2  June  1895  t 
9(3):  8  Julv  1895  f.  # 


9(4):  7  August  1895  t 
9(5):  3  September  1895  f.  # 
9(6):  1  October  1895  t 
9(7);  4  November  1895  t,  # 
9(8):  5  December  1895  t 
9(9):  2  Januarv  1896  t.  # 
9(10):  3  Februarv  1896  t.  # 
9(11):  10  March  1896  t 
9(12):  12  April  1896  t 

10(1):  3  May  1896  t.  # 
10(2):  2  June  1896  f,  # 
10(3):  2  Julv  1896  t,  # 
10(4):  9  August  1896  f 
10(5):  1  September  1896  t.  # 
10(6):  9  October  1896  t.  # 
10(7):  3  November  1896  f 
10(8):  3  December  1896  f 
10(9):  31  December  1896  t 
10(10):  2  Februarv  1897  f 
10(11):  7  March  1897  t,  # 
10(12):  2  April  1897  t 

11(1):  6  Mav  1897  t,  # 
11(2):  1  June  1897  t 
11(3):  29  June  1897  f.  # 
11(4):  5  August  1897  f,  # 
11(5):  1  September  1897  f 
11(6):  4  October  1897  f.  # 
11(7):  1  November  1897  f.  # 
11(8):  6  December  1897  t,  # 
11(9):  3  January  1898  f.  # 
11(10):  1  Februarv  1898  t 
11(11):  4  March  1898  f 
11(12):  3  April  1898  t.  # 

12(1):  1  May  1898  f.  # 
12(2):  9  June  1898  t 
12(3):  30  June  1898  t.  # 
12(4):  4  August  1898  t,  # 
12(5);  1  September  1898  t 
12(6):  3  October  1898  t 
12(7);  7  November  1898  f.  # 
12(8):  2  December  1898  f 
12(9):  9  January  1899  t 
12(10);  1  February  1899  t 
12(11):  5  March  1899  t.  # 
12(12):  3  April  1899  f,  # 

13(1):  6  May  1899  ■ 
13(2):  3  June  1899  • 
13(3):  29  June  1899  • 
13(4):  31  Julv  1899  • 
13(5):  31  August  1899  • 
13(6);  2  October  1899  • 
13(7):  3  November  1899  •,  # 
13(8);  7  December  1899  •,  # 
13(9):  2  Januarv  1900  v  # 
13(10):  8  Februarv  1900  • 
13(11):  1  March  1900  •,  # 
13(12);  2  April  1900  • 

14(1):  2  May  1900  • 
14(2):  2  June  1900  •,  # 
14(3):  2  July  1900  - 
14(4):  1  August  1900  v  # 


Page  176 


THE  NAUTILUS,  Vol.  106,  No.  4 


14(5):  4  September  1900  • 
14(6):  2  October  1900  -,  # 
14(7):  3  November  1900  • 
14(8):  6  December  1900  •,  # 
14(9):  30  December  1900  • 
14(10):  1  Febniarv  1901  v  # 
14(11):  1  March  1901  v  # 
14(12):  6  .\pril  1901  •,  # 

15(1):  3  Mav  1901  •.  # 
15(2):  4  June  1901  • 
15(3):  1  July  1901  • 
15(4):  30  July  1901  •,  # 
15(5):  3  September  1901  •.  # 
15(6):  3  October  1901  • 
15(7):  4  November  1901  > 
15(8):  7  December  1901  v  # 
15(9):  8  January  1902  v  # 
15(10):  5  February  1902  • 
15(11):  6  March  1902  •,  # 
15(12):  5  April  1902  • 

16(1):  5  May  1902  • 
16(2):  2  June  1902  •.  # 
16(3);  5  July  1902  • 
16(4):  2  August  1902  •,  # 
16(5):  8  September  1902  • 
16(6):  6  October  1902  • 
16(7):  3  November  1902  •,  # 
16(8):  3  December  1902  •,  # 
16(9):  5  January  1903  •,  # 
16(10):  10  February  1903  • 
16(11):  28  February  1903  • 
16(12):  11  April  1903  •,  # 

17(1):  20  May  1903  • 
17(2):  11  June  1903  • 
17(3):  3  July  1903  • 
17(4):  12  August  1903  •,  # 
17(5):  4  September  1903  •,  # 
17(6):  7  October  1903  •,  # 
17(7):  6  November  1903  •,  # 
17(8):  7  December  1903  • 
17(9):  8  January  1904  •,  # 
17(10):  6  February  1904  •,  # 
17(11):  5  March  1904  •.  # 
17(12):  2  April  1904  • 

18(1):  5  May  1904  v  # 
18(2):  6  June  1904  • 
18(3):  8  July  1904  • 
18(4):  4  August  1904  • 
18(5):  6  September  1904  • 
18(6):  5  October  1904  • 
18(7):  7  November  1904  •,  # 
18(8);  17  December  1904  • 
18(9);  12  January  1905  • 
18(10):  11  February  1905  •,  * 
18(11):  6  March  1905  •,  # 
18(12):  10  April  1905  •,  # 

19(1):  9  May  1905  • 
19(2):  5  June  1905  •,  # 
19(3):  10  July  1905  •,  # 
19(4):  5  August  1905  • 
19(5):  9  September  1905  • 


19(6);  13  October  1905  • 
19(7):  4  November  1905  • 
19(8):  8  December  1905  v  # 
19(9):  6  January  1906  •,  # 
19(10);  8  February  1906  • 
19(11):  9  March  1906  -.  # 
19(12);  5  April  1906  -.  # 

20(1);  14  May  1906  • 
20(2);  11  June  1906  • 
20(3);  2  July  1906  • 
20(4):  18  August  1906  •,  # 
20(5):  22  September  1906  • 
20(6);  13  October  1906  • 
20(7);  5  November  1906  • 
20(8):  10  December  1906  • 
20(9):  11  January  1907  • 
20(10);  12  February  1907  v  # 
20(11);  4  March  1907  •,  # 
20(12);  12  April  1907  v  # 

21(1);  13  May  1907  • 

21(2);  12  June  1907  v  # 

21(3);  6  July  1907  • 

21(4):  16  August  1907  • 

21(5);  18  September  1907  • 

21(6):  10  October  1907  • 

21(7):  7  November  1907  • 

21(8):  9  December  1907  •,  #  [issue  mislabeled  as  vol.  22] 

21(9):  3  January  1908  •.  #  [issue  mislabeled  as  vol.  22] 

21(10):  3  February  1908  •  [issue  mislabeled  as  vol.  22] 

21(11):  7  March  1908  v  # 

21(12):  4  April  1908  •,  # 

22(1);  9  May  1908  •,  # 
22(2):  12  June  1908  • 
22(3):  16  July  1908  v  # 
22(4/5);  5  September  1908  • 
22(6);  12  October  1908  • 
22(7):  14  November  1908  •.  # 
22(8);  11  December  1908  •.  # 
22(9);  11  January  1909  • 
22(10);  13  February  1909  • 
22(11);  11  March  1909  •,  # 
22(12);  14  April  1909  •,  # 

23(1):  17  Mav  1909  • 
23(2):  4  June  1909  • 
23(3):  9  July  1909  • 
23(4):  4  September  1909  • 
23(5);  2  October  1909  v  # 
23(6):  13  November  1909  • 
23(7);  9  December  1909  • 
23(8);  17  January  1910  • 
23(9);  17  February  1910  • 
23(10):  8  March  1910  • 
23(11);  15  April  1910  •.  # 
23(12):  -  never  issued 

24(1);  19  Mav  1910  •,  # 
24(2);  17  June  1910  • 
24(3):  6  Jul\    1910  •,  # 
24(4):  2  August  1910  •,  # 
24(5):  2  September  1910  • 
24(6):  1  October  1910  • 
24(7);  14  November  1910  • 


E.  V.  Coan  and  M.  G.  Harasewvch,  1993 


Page  177 


24(8):  12  December  1910  •,  # 
24(9);  16  January  1911   • 
24(10):  4  February  1911  •,  # 
24(11):  6  March  1911  •,  # 
24(12):  10  April  1911  • 

25(1):  19Mav  1911  • 
25(2):  16  June  1911  •.  # 
25(3):  5  July  1911  •,  # 
25(4):  18  August  1911  • 
25(5):  11  September  1911  • 
25(6):  19  October  1911  •,  # 
25(7):  11  November  1911  • 
25(8):  13  December  1911  •,  # 
25(9):  15  January  1912  • 
25(10):  15  February  1912  • 
25(11):  8  March  1912  v  # 
25(12):  13  April  1912  • 


26(1):  11  May  1912  ' 
26(2):  1  June  1912  • 
26(3):  12  July  1912  • 
26(4):  5  August  1912  • 
26(5):  6  September  1912 
26(6):  10  October  1912  • 
26(7):  11  November  1912  • 
26(8):  12  December  1912  • 
26(9):  4  Januarv  1913  •,  # 
26(10);  5  February  1913  • 
26(11);  4  March  1913  •.  # 
26(12):  2  April  1913  v  # 

27(1):  8Mav  1913  • 
27(2):  21  June  1913  • 
27(3);  18  July  1913  • 
27(4);  7  August  1913  • 
27(5);  9  September  1913  ' 
27(6);  9  October  1913  • 
27(7);  6  November  1913  • 
27(8);  10  December  1913  •. 
27(9);  2  Januarv  1914  •,  # 
27(10):  9  February  1914  • 
27(11);  19  March  1914  -,  # 
27(12);  16  April  1914  • 


#  [issue  mislabeled  as  no,  4] 


28(1):  16  Mav  1914  -.  # 
28(2):  13  June  1914  •,  # 
28(3);  10  July  1914  - 
28(4);  17  August  1914  • 
28(5);  22  September  1914 
28(6);  15  October  1914  v 
28(7):  20  November  1914 
28(8);  21  December  1914  • 
28(9):  20  Januarv  1915  • 
28(10);  18  February  1915  • 
28(11):  10  March  1915  •,  # 
28(12):  16  April  1915  •,  # 

29(1);  19  Mav  1915  • 
29(2);  3  June  1915  • 
29(3);  6  July  1915  • 
29(4):  4  August  1915  • 
29(5);  4  September  1915  •. 
29(6);  11  October  1915  • 
29(7):  10  November  1915  • 
29(8);  17  December  1915  • 


#  [issue  mislabeled  as  no.  5] 


29(9);  7  January  1916  • 
29(10):  1  February  1916  v 
29(11):  6  March  1916  • 
29(12);  8  April  1916  • 

30(1):  19  May  1916  v  # 
30(2);  5  June  1916  • 
30(3):  14  July  1916  v  # 
30(4):  19  August  1916  • 
30(5);  28  September  1916  ■ 
30(6);  6  November  1916  • 
30(7):  4  December  1916  • 
30(8):  30  December  1917  • 
30(9);  20  January  1917  • 
30(10);  6  March  1917  • 
30(11);  29  March  1917  • 
30(12):  7  May  1917  • 


[THE  NAUTILUS  changes  to  a  quarterly  publication 
schedule.] 


31(1) 
31(2) 
31(3) 

14  July  1917  •,  # 
18  October  1917  • 
14  January  1918  • 

31(4) 

5  May  1918  • 

32(1) 

20  July  1918  V  # 

32(2) 

22  October  1918  • 

32(3) 

17  January  1919  •, 

# 

32(4) 

5  May  1919  • 

33(1) 

16  July  1919  •,  # 

33(2) 

6  November  1919 

33(3) 

22  January  1920  •, 

# 

33(4) 

20  April  1920  • 

34(1);  19  July  1920  •,  # 
34(2):  6  November  1920  • 
34(3);  11  January  1921  •,  # 
34(4):  5  May  1921  -,  # 


35(1) 
35(2) 
35(3) 
35(4) 

36(1) 
36(2) 
36(3) 

36(4) 


26  July  1921  •,  # 

5  December  1921  •.  # 

23  January  1922  -,  # 

24  April  1922  • 

24  July  1922  •,  #  [issue  mislabeled  as  vol.  35] 
7  October  1922  •,  # 
22  January  1923  • 
14  April  1923  • 


37(1):  23  July  1923  •,  # 
37(2):  11  October  1923  v  # 
37(3);  15  Januarv  1924  •,  # 
37(4):  24  April  1924  •,  # 


38(1) 

14  July  1924  •,  # 

38(2);  22  October  1924  • 

38(3);  19  January  1925  •.  # 

38(4);  2  May  1925  v  # 

39(1);  8  July  1925  •,  # 

39(2) 

31  October  1925  • 

39(3) 

11  January  1926  •,  # 

39(4) 

9  April  1926  ■ 

Page  178 


THE  NAUTILUS,  Vol.  106,  No.  4 


40(1):  12  July  1926  • 
40(2):  22  October  1926  •,  # 
40(3):  1  February  1927  • 
40(4):  29  .April  1927  •,  # 

41(1):  23  July  1927  • 
41(2):  27  October  1927  • 
41(3):  17  January  1928  • 
41(4):  25  .\pril  1928  • 

42(1):  14  Julv  1928  • 
42(2):  25  October  1928  • 
42(3):  15  January  1929  • 
42(4):  11  April  1929  • 

43(1):  11  July  1929  • 
43(2):  17  October  1929  • 
43(3):  15  January  1930  • 
43(4):  24  April  1930  • 

44(1):  17  July  1930  • 
44(2):  21  October  1930  • 
44(3):  27  January  1931  • 
44(4):  27  April  1931  • 

45(1):  13  July  1931  • 
45(2):  14  October  1931  • 
45(3):  9  January  1932  • 
45(4):  9  April  1932  • 

46(1):  20  July  1932  BSNH  [23  July 
46(2):  22  October  1932  t 
46(3):  25  January  1933  i 
46(4):  25  April  1933  i 

47(1):  16  June  1933  t 
47(2):  1  November  1933  t 
47(3):  26  January  1934  t 
47(4):  4  May  1934  t 

48(1):  10  July  1934  t 
48(2):  15  October  1934  t 
48(3):  19  January  1935  i 
48(4):  24  April  1935  i 

49(1);  22  July  1935  t 
49(2):  8  November  1935  :j: 
49(3):  30  January  1936  t 
49(4):  1  May  1936  t 

50(1):  14  July  1936$ 
50(2):  29  October  1936  t 
50(3):  29  January  1937  i 
50(4):  4  May  1937  i 

51(1):  3  July  1937  t 
51(2):  22  October  1937  i: 
51(3):  18  January  1938  i 
51(4):  21  April  1938  i: 

52(1):  22  July  1938  t 
52(2):  28  October  1938  t 
52(3):  23  January  1939  i 
52(4):  26  April  1939  t 

53(1):  21  July  1939:): 
53(2):  20  October  1939  t 


t] 


53(3):  28  January  1940  t 
53(4):  29  April  1940  t 

54(1):  23  July  1940  t 
54(2):  2  November  1940  i 
54(3):  4  February  1941  $ 
54(4):  5  May  1941  :j: 

55(1):  11  July  1941  t 
55(2):  24  October  1941  t 
55(3):  12  January  1942  i 
55(4):  7  May  1942  t 

56(1):  23  July  1942  1: 
56(2):  14  October  1942  t 
56(3):  15  February  1943  t 
56(4):  19  April  1943  t 

57(1):  23  July  1943  :j: 
57(2):  30  October  1943  t 
57(3):  9  February  1944  i 
57(4):  15  May  1944  t 

58(1):  17  August  1944  t 
58(2):  24  November  1944  % 
58(3):  19  February  1945  t 
58(4):  20  June  1945  t 

59(1):  6  September  1945  t 
59(2):  27  December  1945  t 
59(3):  9  February  1946  t 
59(4):  27  June  1946  t 

60(1):  30  August  1946  t 
60(2):  18  December  1946  t 
60(3):  11  March  1947  t 
60(4):  2  June  1947  t 

61(1):  14  Julv  1947  t 
61(2):  18  December  1947  $ 
61(3):  2  March  1948  $ 
61(4):  24  May  1948  t 

62(1):  22  July  1948  t 
62(2):  8  December  1948  t 
62(3):  18  March  1949  i 
62(4):  8  June  1949  :t: 

63(1):  19  September  1949  t 
63(2):  1  November  1949  t 
63(3):  13  February  1950  t 
63(4):  4  April  1950  t 

64(1):  5  July  1950  t 
64(2):  27  October  1950  t 
64(3):  15  February  1951  t 
64(4):  7  May  1951  t 

65(1):  27  August  1951  t 
65(2):  9  November  1951  t 
65(3):  25  February  1952  t 
65(4):  22  May  1952  i 

66(1):  25  July  1952  t 
66(2);  17  November  1952  t 
66(3):  2  February  1953  :|: 
66(4):  8  June  1953  t 


E.  V.  Coaii  and  M.  G.  Harasewych.  1993 


Page  179 


67(1) 
67(2) 

67(3): 
67(4): 


24  July  1953  t 
11  November  1953  t 
18  February  1954  $ 
17  May  1954  t 


80(3):  24  January  1967  t 
80(4):  24  April  1967  :j: 


68(1),  24  July  1954  t 


68(2) 
68(3) 
68(4) 


15  November  1954  t 
11  February  1955  t 
28  April  1955  t 


69(1):  1  August  1955  t 
69(2):  5  November  1955  $ 
69(3):  11  February  1956  •,  t 
69(4):  10  May  1956  •,  ij: 

70(1):  13  August  1956  t 
70(2):  12  November  1956  t 
70(3):  11  February  1957  •.  t 
70(4):  29  April  1957  •.  i 


71(1) 
71(2) 
71(3) 
71(4) 


16  August  1957  i 
4  November  1957 
4  March  1958  ij: 
24  April  1958  i 


72(1);  21  July  1958  t 
72(2):  1  October  1958  if. 
72(3):  15  January  1959  t 
72(4):  2  April  1959  t 

73(1):  20  July  1959:1: 
73(2):  3  October  1959  t 
73(3);  25  January  1960  ^ 
73(4):  4  April  1960  t 


74(1) 
74(2) 
74(3) 
74(4) 

75(1) 
75(2) 
75(3) 
75(4) 

76(1) 
76(2) 
76(3) 

76(4) 

77(1) 
77(2): 
77(3): 

77(4): 

78(1) 
78(2) 
78(3) 

78(4) 


I  July  1960  t 

5  October  1960  $ 

II  Januar\   1961  t 

6  April  1961  $ 

3  July  1961  t 

4  October  1961  ^r 
9  January  1962  i 
9  April  1962  $ 

5  July  1962  $ 

14  November  1962  t 
17  January  1963  t 

19  April  1963  t 

6  July  1963  t 

5  October  1963  i 

7  January  1964  i 
14  April  1964  t 

6  July  1964  t 

11  October  1964  t 
25  January  1965  i 

20  April  1965  t 


79(1);  9  July  1965  t 
79(2);  15  October  1965  t 
79(3);  25  January  1966  i 
79(4);  25  April  1966  t 

80(1);  6Julv  1966  t 
80(2);  11  October  1966  t 


81(1) 
81(2): 
81(3): 

81(4): 

82(1): 
82(2): 
82(3): 
82(4): 

83(1): 
83(2): 
83(3): 
83(4): 

84(1) 

84(2): 
84(3): 

84(4) 

85(1) 

85(2) 
85(3) 
85(4) 


6  July  1967  t 
16  October  1967  t 
25  January  1968  i 
18  April  1968  :|: 

24  July  1968  t- 

28  October  1968  t 
31  January  1969  i 
28  April  1969  t 

28  July  1969  t 
31  October  1969  t 
23  January  1970  i 
30  April  1970  t 

16  July  1970  t 
5  October  1970  i 

25  January  1971  i: 

26  April  1971  t 

20  July  1971  t 
4  October  1911  t 

27  January  1972  ij: 

28  April  1972  t 


86(1):  27  July  1972  § 
86(2-4):  1  December  1972  § 


[Publication  changes  to  one  volume  per  calendar  year] 


87(1) 
87(2) 
87(3) 
87(4) 

88(1) 
88(2) 
88(3) 
88(4) 

89(1) 
89(2) 
89(3) 
89(4) 


24  January  1973  § 
27  April  1973  § 
30  July  1973  § 

4  October  1973  § 

29  January  1974  § 

30  April  1974  § 
22  Julv  1974  § 

25  October  1974  § 

31  January  1975  § 
30  April  1975  § 
18  July  1975  § 

29  October  1975  § 


[Subsequent  issues  have  dates  of  publication  printed  on 
the  title  page  and  generally  also  on  the  cover.] 

LITERATURE  CITED 

Abbott,  R.  T.  (compiled  by  L,  B,  Hastings  and  M.  C.  Teskey). 
1979,  Indexes  to  The  Nautilus:  Geographical  (volumes 
1-90)  and  scientific  names  (volumes  61-90).  Melbourne, 
Florida  (American  Malacologists)  iv  +  238  pp. 

American  Malacological  Union.  1940.  Scientific  contributions 
made  from  1882  to  1939  b\  Henry  A.  Pilsbry,  Sc.  D.  The 
American  Malacological  Union.  63  pp 

Baker,  H  B.  1958.  Scientific  contributions  made  from  1940 
to  1957  by  Henry  A.  Pilsbry.  The  Nautilus  71(3);104-112. 

Bartsch.  P.,  h'  A.  Rehder,  and  B.  E.  Shields.  1946.  A  bibli- 
ography and  short  biographical  sketch  of  William  Healey 
Dal]  Smithsonian  Miscellaneous  Collections  104(15):l-96, 
Ipl. 


Page  180 


THE  NAUTILUS,  Vol.  106,  No.  4 


Clench,  V\'.  J.  and  R  D  Turner.  1962  New  names  introduced 
by  H  .\  Pilsbry  in  the  Mollusca  and  Crustacea  -Acadenn 
of  Natural  Sciences  of  Philadelphia,  Special  Publication  4: 
218  pp. 

Henderson,  J.  B.,  Jr.  (assisted  by  VI  U'.  Poole;  edited  by  W 
H.  Dall).  1927.  Index  to  The  Sautilus  volumes  3-34, 
1889-1921.  Michigan  (Clapp  it  Walker)  434  pp 

La  Rocque,  A.  (assisted  by  G.  Smithe  &  H.  W.  Harry).  1951. 
Index  to  The  Sautilus  volumes  35-60,  1921-1947  Ann 
Arbor,  Michigan  (University  of  Michigan).  332  pp 


La  Rocque,  A.  1963.  .Author  index  to  The  Nautilus  volumes 
3-75  and  its  predecessor  The  Conchologists'  Exchange, 
volumes  1  and  2.  Columbus,  Ohio,  279  pp 

La  Rocque,  .\.  1972  .Author  index  to  The  Saitliluf^  \olumes 
76-85.  Sterkiana  47:21-38. 

Pilsbry,  H  A.  and  C.  VV.  Johnson  1895.  Index  to  the  "Con- 
chologists Exchange."  The  Nautilus  9(4):43-48;  (5):57-60. 


INSTRUCTIONS  TO  AUTHORS 


THE  NAUTILUS  publishes  papers  on  all  aspects  of  the 
biology  and  systematics  of  mollusks.  Manuscripts  de- 
scribing original,  unpublished  research  as  well  as  review 
articles  will  be  considered.  Brief  articles,  not  exceeding 
1000  words,  will  be  published  as  notes  and  do  not  re- 
quire an  abstract.  Notices  of  meetings  and  other  items 
of  interest  to  malacologists  will  appear  in  a  news  and 
notices  section. 

Manuscripts:  Each  original  manuscript  and  accompa- 
nying illustrations  should  be  submitted  in  triplicate.  Text 
must  be  typed  on  one  side  of  8V2  x  H  inch  white  paper, 
double  spaced  throughout  (including  literature  cited,  ta- 
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on  all  sides.  All  pages  must  be  numbered  consecutively. 
If  printed  on  a  word  processor,  the  right  margin  should 
be  ragged  rather  than  justified.  Authors  should  follow 
the  recommendations  of  the  Council  of  Biology  Editors 
Style  Manual,  which  is  available  from  the  Council  of 
Biology  Editors,  Inc.,  9650  Rockville  Pike,  Bethesda,  MD 
20814,  U.S.A.  The  first  mention  of  a  scientific  name  in 
the  text  should  be  accompanied  by  the  taxonomic  au- 
thority, including  year.  Latin  names  and  words  to  be 
printed  in  italics  must  be  underlined;  leave  other  indi- 
cations to  the  editor.  Metric  and  Celsius  units  are  to  be 
used. 

The  sequence  of  sections  should  be:  title  page,  abstract 
page,  introduction,  materials  and  methods,  results,  dis- 
cussion, acknowledgements,  literature  cited,  tables,  fig- 
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title,  author's  name(s)  and  address(es).  The  abstract  page 
.should  contain  the  title  and  abstract,  which  should  sum- 
marize in  250  words  or  less  the  scope,  main  results  and 
conclusions  of  the  paper.  The  abstract  may  be  followed 
by  a  maximum  of  8  key  words.  All  references  cited  in 
the  text  must  appear  in  the  literature  cited  section  and 
vice  versa.  In  the  literature  cited  section,  all  authors 
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a  recent  issue  of  THE  NAUTILUS  for  bibliographic  style, 
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mation on  plates  and  figures  should  be  cited  only  if  not 
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and  each  placed  on  a  separate  sheet.  A  brief  legend  must 
accompany  each  table.  Captions  for  each  group  of  illus- 
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All  line  drawings  must  be  in  black,  high  quality  ink, 
clearly  detailed  and  completely  labeled.  Photographs 
must  be  on  glossy,  high  contrast  paper.  All  figures  are 
to  be  consecutively  numbered  (figs.  1,  2,  3,  ...  ,  NOT 
figs.  la.  lb,  Ic,  .  .  .  NOR  plate  1,  fig.  1  .  .  .).  Illustrations 
must  be  arranged  in  proportions  that  will  conform  with 
the  width  of  a  page  (6^4  inches  or  171  mm)  or  a  column 
(5Vi  inches  or  82  mm).  The  maximum  size  of  a  printed 
figure  is  6%  by  9  inches  or  171  by  228  mm.  All  illus- 
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backing,  numbered,  labeled  and  camera  ready.  The  au- 
thor's name,  paper  title  and  figure  rmmber(s)  should  ap- 
pear on  the  back.  Original  illustrations  must  be  between 
one  and  two  times  the  desired  final  size.  It  is  the  author's 
responsibility  that  the  line  weight  and  lettering  are  ap- 
propriate for  the  desired  reduction.  Original  illustrations 
will  be  returned  to  the  author  if  requested.  Color  illus- 
trations can  be  included  at  extra  cost  to  the  author. 

Voucher  Material:  Deposition  of  type  material  in  a 
recognized  public  museum  is  a  requirement  for  publi- 
cation of  papers  in  which  new  species  are  described. 
Deposition  of  representative  voucher  specimens  in  such 
institutions  is  strongly  encouraged  for  all  other  types  of 
research  papers. 

Processing  of  Manuscripts:  Upon  receipt,  every  manu- 
script is  acknowledged  and  sent  for  critical  review  by  at 
least  two  referees.  These  reviews  serve  as  the  basis  for 
acceptance  or  rejection.  Accepted  manuscripts  are  re- 
turned to  the  author  for  consideration  of  the  reviewers' 
comments.  A  finalized  version  of  the  manuscript  is  re- 
turned to  the  editor  and  sent  to  press.  Two  sets  of  proofs 
are  sent  to  the  author  for  correction.  Changes  other  than 
typesetting  errors  will  be  charged  to  the  author  at  cost. 
One  set  of  corrected  proofs  should  be  sent  to  the  editor 
as  soon  as  possible.  Authors  with  institutional,  grant  or 
other  research  support  will  be  billed  for  page  charges  at 
the  rate  of  $60.00  per  printed  page. 

An  order  form  for  reprints  will  accompany  the  proofs. 
Reprints  may  be  ordered  through  the  editor. 

Manuscripts,  corrected  proofs  and  correspondence  re- 
garding editorial  matters  should  be  sent  to:  Dr.  M.G. 
Harasewych,  Editor,  Division  of  Mollusks,  NHB  stop  118, 
National  Museum  of  Natural  History,  Smithsonian  In- 
stitution, Washington,  DC  20560,  USA. 


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