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THE  NAUTILUS 


Volume  114,  Number  1 
March  27,  2000 
ISSN  0028-1344 

A  cjuarteHij  devoted 
to  malacology. 


EDITOR-IN-CHIEF 

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The  Bailey- Matthews  Shell  Museum 
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National  Museum  of 
Natural  Historv' 
Smithsonian  Institution 
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Field  Museum  of 
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Laboratoire  de  Biologic  des 
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Museum  of  Comparative  Zoology 
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The  Academy  of  Natural  Sciences 
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Museum  of  Comparative  Zoology 
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TH  E€9NAUTI  LUS 


CONTENTS 


Volume  114,  Number  1 
March  27,  2000 
ISSN  0028-1344 


Adriaan  Gittenberger  Epitoniinii  (Gastropoda:  Epitoniidae)  associated  with 

Jeroen  Goud  iinislirooni  corals  (Scleractinia:  Fun<fiidae)  from  Sulawesi, 

Edmund  Gittenberger  Indonesia,  with  the  description  ot  four  new  species 1 

Marta  J.  deMaintenon  A  new  species  ot  CohtmbcUa  (Neogastropoda: 

Columbellidae)  from  the  Caribbean  Neogene 14 

Bruce  A.  Marshall  The  New  Zealand  Recent  species  of  Muricopsis  Bucquoy, 

Kevin  W.  Burch  Dautzenberg  and  Dollfus,  1882  (Gastropoda:  Muricidae) 18 

Notices 3{) 


APRl 


THE  NAUTILUS  114(1):1-13,  2000 


Page  1 


Epitoniiun  (Gastropoda:  Epitoniidae)  associated  with  mushroom 
corals  (Scleractinia:  Fungiidae)  from  Sulawesi,  Indonesia,  with 
the  description  of  four  new  species 


Adriaan  Gittenberger 

Institute  of  Evolutionan-  and 

Ecological  Sciences 
Universitv  of  Leiden 
P.O.  Box'9516 
NL  2300  RA  Leiden 
THE  NETHERLANDS 


Jeroen  Coud 

National  Museum  of  Natural  Histon' 

RO.  Box  9.517 

NL  2.300  RA  Leiden 

THE  NETHERLANDS 


Edmund  Gittenberger' 

Institute  of  Evolutionary  and 

Ecological  Sciences 
National  Museum  of  Natural  Historv 
RO.  Box  9517 
NL  2300  RA  Leiden 
THE  NETHERLANDS 


ABSTRACT 

At  least  six  species  of  the  genus  Epitoniiim  scnsii  lata  are  found 
associated  witli  mushroom  corals  (Fungiidae)  off  Ujinig  Pan- 
dang,  Sulawesi.  Indonesia.  Revised  descriptions  of  £.  costiila- 
tiim  (Kiener,  1838)  and  E.  iilu  Pilsbry,  1921  based  on  t\pe 
specimens  and  additional  material  are  given.  Four  new  species 
are  described:  E.  hoeksemai.  E.  ingridae,  E.  lochi,  and  E.  tivi- 
lae.  The  true  identity  of  £.  bullntum  (Sowerbw  1844),  a  species 
not  associated  with  corals  and  not  found  in  Sulawesi  and  near- 
by areas,  is  clarified.  Examination  of  tyjie  sjiecimens  made  pos- 
sible the  characterization  of  nominal  species  that  appear  to  be 
either  identical  with  or  closely  related  to  the  fungiid-associated 
epitoniids  found  off  Sulawesi. 

Ki-ij  words:  Indo-Pacific,  parasites,  coral  reefs,  coral/mollusk 
association,  egg  capsules. 


INTRODUCTION 

Several  epitoniid  species  are  known  to  live  in  association 
with  sea  anemones  (phvluni  Cnidaria,  order  Actiniaria) 
(Robertson,  1963;  1983a,  b;  1993;  Vecchi(j,  1964;  Salo, 
1977;  Perron,  1978;  Kay,  1979;  Schimek.  1986;  Hartog, 
1987;  Dushane,  1988a-<';  Yamashiro,  1990;  Nakayama, 
1991;  Mienis,  1994).  Less  commonly,  epitoniids  are  found 
associated  with  stony  corals  (piiylum  Cnidaria,  order 
Scleractinia),  in  particular  with  species  of  the  free-living 
Fungiidae  or  mushroom  corals  (Robertson,  1963;  1970; 
Bosch,  1965;  Hadfield,  1976;  Kav,  1979;  Bratcher,  1982; 
Loch,  1982;  Sabelh  and  Taviani,  1984;  Bell,  1985;  Dusha- 
ne, 1988a-c;  Loo  and  Chou,  1988;  Page  and  Willan,  1988; 
Hoeksema,  1988;  1989;  Yamashiro,  1990;  Mienis,  1994; 
Oliverio  ct  ai,  1997).  OnK'  three  Epitoiiitim  species  are 
usualK'  mentioned  in  the  literature  in  association  with 
fungiids;  in  one  case  (Lcjch,  1982)  a  fourth  species  is  re- 
ported but  not  named.  This  paper  deals  mainK'  widi  die 
taxonomy  of  die  surprisingly  high  number  of  species  of 

'  Author  for  correspondence;  gittenberger@naturalis.nnni  nl 


Epitonium  found  associated  widi  mushroom  corals  during 
a  survey  in  a  relatively  restricted  area  in  Indonesia,  off 
Ujung  Pandang  (Sulawesi).  Four  of  these  species  proved 
to  be  new  to  science,  although  at  least  one  of  them  had 
frequenth'  been  cited  and  illustrated  under  an  incorrect 
name.  The  shells  of  diese  species  are  ver)'  fragile,  which 
might  explain  why  diey  are  mosdy  poorly  represented  or 
not  represented  at  all  in  most  institutional  collections. 
These  species  are  only  kiiouni  from  live-collected  speci- 
mens: it  is  very  unlikely  diat  emptv  shells  will  be  found 
washed  ashore  without  being  seriously  damaged  or  un- 
recognizable. A  more  elaborate  analysis  of  the  ecological 
data  collected  during  the  project  is  being  prepared  (Git- 
tenberger, A.,  unpul)lished  data). 

The  systematic  and  evolutionary  importance  of  vari- 
able characters  such  as  egg  capsules  (Figures  36-38), 
eggs  (Figure  41)  and  mucous  threads  (Figures  43-47), 
which  can  be  either  straight  or  hoisted,  is  still  poorlv 
known.  We  observed,  however,  that  populations  of  the 
different  species  may  differ  in  these  characters.  With 
exception  of  one  article  by  Oliverio  et  al.  (1997).  the 
literature  is  scantx-  in  respect  to  these  characters.  Oliv- 
erio ct  nl.,  while  discussing  the  coral-associated  epitoniid 
Epitonium  billccanuni  Dushane  and  Bratcher,  1965,  fig- 
ured the  egg  capsules,  eggs,  mucous  threads  (of  the 
tswsted  type)  and  shells  of  veliger  larvae.  We  did  not 
obseiA'c  a  difference  in  sculpture  or  well-defined  tran- 
sition between  the  protoconch  1,  formed  by  the  shell 
gland  of  the  larva  inside  the  egg  capsule,  and  protoconch 
2,  secreted  by  the  velum  of  the  swimming  veliger  be- 
tween hatching  and  settling.  The  protoconchs  (Figures 
16,  25-29,  42,  48)  turned  out  to  be  verv  uniform  among 
the  various  species  studied  here,  all  of  which  apparently 
have  planktotrophic  development.  Sclerites  of  at  least 
one  species  of  soft  coral,  probabh'  of  the  genus  Sinularia 
May,  1898  (subclass  Octocorallia,  order  Alcyonacea,  fam- 
ily AlcNoniidae)  (L.  P.  van  Ofvvegen,  NNM)  were  found 
associated  with  the  egg  capsules  of  some  species  (Fig- 
ures 39-40). 


Page  2 


THE  NAUTILUS.  Vol.  114.  No.  1 


Figure  1.  Sun'e\ed  area  off  Ujung  Paiidang.  S.  Sulawesi.  Indonesia.  The  coral  reefs  investigated  in  particular  are:  1.  W  (Pulau) 
Lae-Lae;  2,  W  Bone  Baku;  3,  E  (Pulau)  Samalona;  4,  W  (Pulau)  Samalona;  5,  E  and  ESE  (Pulau)  Kudingareng  Keke;  6.  W  (Pulau) 
Kudingareng  Keke:  7.  NW  Lan[g]kai;  8.  Pulau  Badi:  9.  Bone  Lola:  10.  (Pulau)  Bone  Tanibung. 


In  a  monograph  on  Epitoniidae  from  southern  Africa 
and  Mozambique,  Kilbum  (1985:  240)  stated  that  "epi- 
toniid  taxonomy  remains  in  a  chaotic  state,  particularK^ 
above  the  species  level."  Kilbum  observed  that  the  clas- 
sification of  the  genus  Epitonium  is  (p.  280)  "vers'  ten- 
tative and  is  aimed  soieK  at  grouping  together  similar 
species  for  convenience  sake."  Clench  and  Turner  (1951) 
and  Bouchet  and  Waren  (1986)  followed  a  similar  ap- 
proach in  their  revision  of  eastern  Atlantic  Epitoniidae. 
Because  we  could  not  imequivocalK'  classify-  all  Indo- 
nesian species  within  one  or  more  of  the  19  subgenera 
used  by  Kilbum  (1985)  or  the  39  subgenera  listed  bv 
Wenz  (1940)  under  "Scala"  (=  Epitonium).  we  decided 
to  refrain  from  following  anv  subgeneric  classification. 

The  epitoniid  species  described  in  this  studv  live  as- 
sociated with  mushroom  corals  and  at  least  some  of 
them  are  so  similar  that  they  seem  to  be  closelv  related 
phvlogenedcalK-.  Thev  point  to  possible  adaptive  radia- 
tion within  a  single  clade.  Adequate  ph\logenetic  anal- 
yses including  other  species  of  Epitonium  co-occurring 
with  different  hosts  in  the  same  general  area  could  help 
clarify-  whether  these  species  found  in  association  with 
fungiid  corals  form  a  monophsletic  group. 

MATERIALS  AND  METHODS 

Samples  were  collected  off  the  coast  of  Ujung  Pandang, 
Sulawesi,  Indonesia.  The  fungiid  fauna  of  the  area  (Fig- 
ure 1)  is  relatively  well  known;  see  Hoeksema  (1989)  for 
details.  During  the  period  April-June,  1997,  9  coral  reefs 


were  inspected,  to  a  depth  of  18  (rarelv  24)  meters.  Ap- 
proximately 10'  mushroom  corals,  belonging  to  about  30 
species  of  free-living  Fungiidae,  were  searched  for  epi- 
toniids  and  their  eggs.  Beneath  these  corals,  about  10' 
specimens  of  Epitonium.  some  of  which  with  egg  cap- 
sules, were  collected.  The  identifications  of  the  coral 
species  were  made  or  at  least  checked  b)'  Dr.  B.  W. 
Hoeksema.  The  snails  were  classified  into  morphological 
categories  according  to  characters  of  shell,  egg  capsules, 
and  proboscis.  Secondarilv,  the  species  thus  disdn- 
guished  were  anaKzed  ecologicallv  for  preferential 
depths,  hosts,  and  substrates.  The  ecologiciU  data  will  be 
discussed  in  a  future  article  (Gittenberger,  A.,  unpub- 
lished data). 

The  various  Epitonium  species  recorded  during  the 
surve\'  were  identified  bv  review  of  the  literature,  con- 
sultation with  specialists,  and  by  comparison  with  ma- 
terial deposited  in  several  collections;  this  includes  com- 
parison with  hpe  specimens  of  conchologicallv  similar 
taxa.  These  tvpes  are  mentioned  in  the  systematic  treat- 
ment of  each  species. 

From  the  about  10'  specimens  collected,  only  shells 
with  more  than  4  mm  length  were  measured.  The  num- 
ber of  specimens  (n)  measured  in  the  calculation  of 
mean  values  is  mentioned  at  the  beginning  of  the  de- 
scriptions. Means  are  indicated  between  the  e.xtremes 
(minimum-mean-maximum).  To  allow  for  better  com- 
parisons, shell  sculpture  is  described  for  both  the  fifth 
teleoconch  whorl  and  where  the  teleoconch  is  5  mm  in 
width,  a  part  of  die  spire  that  is  in  part  independent  of 


Gittenberger  et  ai,  2000 


Page  3 


Figure  2.  Maps  of  the  Indo-Pacific  Region,  from  the  Red  Sea  to  the  Hawaiian  Archipelago,  showing  records  of  the  Epitonium 
species  in  this  study  known  from  more  than  tvvo  localities:  Epitonium  costulatum  (Kiener,  1838)  (stars),  E.  ingridae  new  species 
(triangles),  E.  twilae  new  species  (squares)  and  E.  iilu  Pilsbry,  1921  (circles). 


the  actual  whorl  number.  The  term  protoconch  refers  to 
the  protoconchs  1+2.  The  maximum  diameter  of  pro- 
toconch 1  was  measured  in  two  shells  for  each  species 
(except  tor  E.  lochi  because  of  insufficient  material),  us- 
ing SEM  photographs  of  specimens  prepared  from  egg 
capsules  (Figures  30-31);  because  ver\'  similar  values 
were  consistently  found,  no  more  measurements  were 
taken.  Shells  of  Epitonium  species  cannot  be  recognized 
as  fulK'  grown  or  not.  Comparative  informal  observations 
indicate  that  when  the  snails  start  laving  eggs  thev  have 
not  yet  reached  maximum  size.  No  minimum  values  are 
included  in  the  descriptions  but  only  the  largest  speci- 
men and  the  largest  number  f)f  whorls.  After  removal 
from  70%  ethanol,  egg  capsules  without  embedded  sand 
(juicklv  collapsed;  these  could  not  be  photographed.  Un- 
less stated  otherwise,  all  descriptions  refer  to  material 
from  off  Ujung  Pandang. 

The  following  institutional  abbreviations  are  used: 
AMS,  Australian  Museum,  Sydney;  ANSP,  Academy  of 
Natural  Sciences,  Philadelphia;  BMNH,  The  Natural 
Histon'  Museum,  London;  LACM,  Natural  Histoiv  Mu- 


seum of  Los  Angeles  County,  Los  Angeles;  MHNG,  Mu- 
seum d'Histoire  Naturelle,  Geneve;  MNHN,  Museum 
national  d'Histoire  naturelle,  Paris;  MZB,  Museum  Zool- 
ogicum  Bogoriense,  Bogor,  Indonesia;  NNM,  National 
Museum  of  Natural  Historv,  Leiden.  Numbers  following 
a  slash  sign  after  collection  numbers  refer  to  number  of 
shells  in  relevant  lots. 

SYSTEMATICS 

Family  Epitoniidae  Berrv,  1910 
Genus  Epitonium  Boding,  1798 

Epitonium  costulatum  (Kiener,  1838) 
(Figures  2-6,  22,  25,  38-41,  47) 

Scalaha  costulatum  Kiener,  1838:  pi.  2,  fig.  4;  1838:  5. 
Epitonium  costulatum. — Robertson,    1963:   57,   pi.   5,   fig.   4; 

1970:  45;  Loch.  1982:  4,  1  fig.;  Dushane,  1988a;  30,  figs. 

1,  2. 

Description:  Shell  (Figures  3-6,  22,  25)  (n  =  7):  Frag- 
ile (large  specimens)  to  verv  fragile,  moderately  elongate- 


Page  4 


THE  NAUTILUS,  Vol.  114,  No,  1 


Figures  3 — 1.     Epitonium  costiilatum  (Kiener,  1838),  holotype 
(MHNG  1152/16).  Shell  length  3.3  cm. 


conical,  cream\  white,  reacliing  32  mm  in  length,  with  at 
least  1  damaged  specimen  (from  Ambon)  measuring  41.2 
mm.  Lengtli/vvidth  1.6-1.9-2.2.  Protoconch  whorls  3  .3/8; 
maximum  protoconch  1  diameter  0.14  mm  (n  =  2).  Pro- 
toconch with  numerous  fine,  incised,  axial  lines.  Teleo- 
conch  whorls  up  to  10,  separated  bv  vers'  deep  (fenes- 
trate) suture.  Successive  whorls  are  almost  detached.  Te- 
leoconch  with  e\enK-  spaced,  orthocline,  thin  costae,  dam- 
aged in  all  examined  specimens.  Over  most  of  their 
lengdi,  costae  appear  to  be  curved  abaperturally  at  outer 
margin.  Costae  adapicallv  relativelv  high  and  erect,  not 
coronate,  becoming  short  toward  columella.  Costae  most- 
K'  continuous,  but  touching  onK'  slightK'  diose  of  adjoining 
whorls.  Ver\-  weak  spiral  lines  present.  Fifth  teleoconcli 
whorl  (widtli  4.9  mm)  with  16-18.4^26  costae.  Five  mm 
width  whorl  (whorl  4,  5,  or  6)  widi  16-17.4-20  costae. 
Aperture  subcircular.  Apertural  lengtli/shell  length  0.3. 
Umbilicus  moderately  wide. 

Egg  capsules  (Fif^iires  38,  3.9,  40,  47):  Embedded  with 
sand  and  closel)-  connected  along  a  straight,  longitudi- 
nally striated,  mucous  diread  (Figure  47).  Capsules 
asymmetrical,  somewhat  conical  with  a  circular  widest 
part.  Capsules  3. 0-3. .3-3. 5  mm  in  length  and  1.5-1.6- 
2.0  mm  in  width  (n  =  8).  One  egg  capsule  contains  70- 
17.5-335  eggs  (n  =  5). 

Proboscis:  With  some  irreguIarK'  interrupted,  longitvi- 
dinal,  white  zones,  which  are  as  wide  as  transparent  in- 
terspaces. 

Type  material  (Figures  3-4): 

1152/16. 


Holotype  MHNG 


Type  locality:     Unknown. 

Other   material   examined:     NNM,   Indonesia,   Am- 
bon, Hitu,  outer  part  of  Ambon  Bay,  E  and  W  sides  of 


Laha,  A.  Fortuin  and  J.  C.  den  Hartog  leg.;  LACM 
124.505,  Thailand.  Phuket  Island. 

Records  in  the  Uterature:  Australia:  Queensland, 
Thetford  Reef  off  Caims  (Loch,  1982:  4).  Philippines; 
Bongao  Channel,  Sanga  Sanga  (Robertson,  1963;  57-.58, 
pi.  5,  fig.  4).  Thailand;"Rava  Island  (Dushane,  1988a;  32). 
Maldives,  Little  Hiva  (Dushane,  1988a:  32).  Red  Sea: 
Straits  of  Tiran  (Dushane,  1988a:  .30-31);  Sinai,  Thomas 
Reef  27°.59'N,  34°27'E  (Dushane,  1988a;  .32). 

Distribution  (Figure  2):  Australia  (Queensland),  In- 
donesia (Sulawesi),  Philippines,  Thailand,  and  Red  Sea. 

Habitat:  Snails  were  recorded  at  6-12  m  depth.  Coral 
hosts  were  Ctenactis  cchinata  (Pallas,  1766),  and  Her- 
poUtlia  Umax  (Esper,  1797).  Groups  of  one  to  four  snails 
were  found  in  the  sand  (sometimes  buried)  under  a  sin- 
gle coral;  sometimes  close  to  a  few  hundred  egg  cap- 
sules. 

Remarks:  The  data  provided  bv  Sherbom  and  Wood- 
ward (1901)  are  insufficient  to  indicate  the  exact  vear  of 
publication  of  the  new  taxa  in  Kiener's  monograph  on 
the  Genre  Scalaire'.  We  follow  Troschel  (18.39),  who 
Listed  Kiener's  undated  work,  with  the  new  species  in  it, 
in  his  'Report  on  the  achievements  in  zoology  during  the 
year  1838.  V.  MoUusca'  [in  German].  The  names  are 
printed  botli  on  the  plates  and  in  the  main  text  of  Kie- 
ner's work.  The  severely  damaged  holotspe  of  this  spe- 
cies (Figures  .'3-4)  is  a  relatively  elongate  shell.  Shells  of 
this  species  are  most  similar  to  those  of  Epitonium  pal- 
lasii  (Kiener,  1838),  a  species  originally  described  from 
an  unknown  locality,  but  now  known  from  the  Indo-West 
Pacific  (Kaicher,  1980:  2382;  Eisenberg,  1981;  pi.  37,  fig. 
9;  Wilson,  1993:  278,  pi.  44,  fig.  6a-b).  According  to 
Kiener  (1838)  and  in  agreement  with  Wilson's  descrip- 
tion ("about  ten  costae  on  the  last  whorl")  and  the  fig- 
ures in  the  litterature,  E.  paUasii  differs  from  E.  costii- 
latum h\  the  stronger  shells  with  thicker  costae,  which 
are  more  widely  spaced.  Dushane  (19S8a:  .'30,  fig.  2)  fig- 
ured very  similar  egg  capsules  of  this  species,  reporting 
two  connecting  threads  for  material  from  the  Red  Sea. 

Epitonium  hoeksemai  A.  Gittenberger  and  Goud  new  species 
(Figures  9-10,  18,  20,  26,  43) 

Description:  Shell  (Figures  9-10,  18,  20,  26)  (n  =  9): 
Ven-  fragile,  elongate-conical,  creamy  white,  reaching 
19  mm  in  length.  Length/width  1.6-1.9-2.4.  Protoconch 
whorls  3.  Maximum  protoconch  1  diameter  0.13  mm  (n 
=  2).  Protoconch  with  numerous  very  fine,  incised,  axial 
lines.  Teleoconch  whorls  up  to  9  1/8,  separated  b)'  a 
moderately  deep  suture.  Teleoconch  sculpture  (Figures 
IS,  20)  of  somewhat  unevenly  spaced,  orthocline,  rela- 
tively low  costae,  and  low  spiral  threads  that  become 
conspicuously  more  numerous  and  variable  on  the  ab- 
apical  whorls.  Costae  on  entire  teleoconch  more  prom- 
inent than  spiral  sculpture.  Third  teleoconch  whorl  has 
ca.  12  spiral  threads,  fifth  ca.  25.  Costae  not  alwa\s  con- 
tinuous, touching  the  adjoining  whorls,  where  thev  are 
curved  adaperturallv.  Fiftli  teleoconch  whorl  (width  2.1 


Gittenberger  et  al,  2000 


Page  5 


Figures  5-14.  Species  of  Epitonium  associated  with  mushrooni  corals  off  Ujung  Pandang.  5-6.  E.  costulatuin  (Kiener,  1838), 
length  2.8  cm.  7-8.  E.  ingridae  new  species,  holotype,  NNM  59088,  length  2.0  cm.  9-10.  £.  hoeksemai  new  species,  holotype, 
NNM  59074,  length  1.3  cm.  11-12.  E.  tihi  Pilsbry,  1921,  length  1.6  cm  and  1.0  cm,  respectively.  13-14.  E.  lochi  new  species, 
holot>pe,  NNM  .59094,  length  0.9  cm. 


mm)  with  24—27-29  costae.  Five  mm  width  whorl  (whorl 
8,  9  or  10)  with  32-35-38  costae.  Aperture  subcircular. 
Apertural  length/shell  length  0.28-0.29-0..30.  Umbilicus 
very  narrow. 


Egg  capsules  (Figure  43):  Sub-.spherical,  white,  tran.s- 
parent.  with  protuberances,  no  noticeable  embedded 
sand.  Capsules  closelv  connected  to  each  other  along  a 
twisted  mucous  thread. 


Page  6 


THE  NAUTILUS,  Vol.  114,  No.  1 


Figures  15-21.  SEM  micrographs  of  species  of  Epitonitim  associated  with  uiuslinKim  corals  off  Ujung  Pandang.  15-16.  E.  lochi 
new  species  15.  Shell.  Scale  line  =  1  mm.  16.  Protoconch.  Scale  line  =  0.1  mm.  17.  £.  tihi  Pilsbrw  1921,  shell.  Scale  line  =  1 
mm.  18.  E.  hock.scmai  new  species,  apical  whorls.  Scale  line  =  1  mm.  19.  E.  twilac  new  species,  teleoconch  whorl  sculpture.  Scale 
line  =  0.1  mm.  20.  E.  hoeksemai  new  species,  teleoconch  sculpture.  Scale  line  =  0.1  mm.  21.  E.  iilii  Pilsbr\',  1921,  teleoconch 
whorl  sculpture.  Scale  line  =  0.1  mm. 


Habitat:  This  species  was  recorded  between  5-15  me- 
ters depth.  Coral  hosts  were  Heliofungia  actinifonnis 
(Quoy  and  Gaimard,  1833)  and  Fungia  ftiugitcs  (Lin- 
naeus, 1758).  One  to  5  specimens  were  found  attached 
by  mucous  threads  to  the  underside  of  a  coral,  near  a 
few  hundred  egg  capsules. 


Type  material:  Holohpe  NNM  59074,  from  t)pe  lo- 
caliU-.  Parat\pes:  NNM  59081/1,  Indonesia,  Sulawesi,  off 
Ujung  Pandang,  W.  Lae-Lae,  9  m;  NNM  59079/1,  MZB/ 
1,  W'  Bone  Baku,  6  m:  NNM  59080/1,  59082/1,  9  ni; 
NNM  59086/2,  t\pe  localit\-:  NNM  59076/1,  6  m;  NNM 
59077/5,  12  m:  NNM  59083/1,  E  Kudingareng  Keke,  3 


Gittenberger  et  al,  2000 


Page  7 


m:  NNM  59078/1.  W.  Kudingareng  Keke,  12  m;  NNM 
59075/1,  14  m;  NNM  59084/1,  NW.  Lankai,  6  m;  NNM 
59087/2,  Bone  Lola.  15  m;  NNM  59085/1.  Bone  Tam- 
biing.  6  ni. 

Type  locality:  Indonesia,  Sulawesi,  off  Ujimg  Pan- 
dang.  W  Samalona.  5  m  tlepth. 

Distribution:  Onlv  known  from  Indone.sia,  off  Sula- 
wesi. 

Etymology:  This  species  is  named  after  Dr.  B.  W. 
Hoeksema,  who  supervised  the  field  portion  of  this  pro- 
ject. 

Remark.s:  Shells  of  this  .species  resemble  those  of  Ep- 
ituniuin  iilu,  but  differ  b)-  a  lengtli/width  ratio  of  ca.  1.9 
instead  of  ca.  2.6.  Because  most  examined  specimens  are 
damaged,  the  fine  structure  of  the  costae  could  not  be 
observed.  The  teleoconch  sculpture  appears  alwavs  ob- 
solete to  the  naked  eye  and  the  number  of  spiral  threads 
increases  more  evidently  in  E.  huehscmai,  with  ca.  13 
spiral  threads  added  between  the  third  and  the  whorl. 
In  £.  lochi  new  species  (see  below),  on  the  other  hand, 
there  is  a  more  clearly  reticulate  sculpture  on  the  early 
teleoconch  whorls. 

Epitonium  ingridae  A.  Gittenberger  and  Govid  new  species 

(Figures  2.  7-8.  23-24.  27.  .30.  36.  46) 

Description:  Shell  (Figures  7-8,  23-24,  27,  30)  (n  = 
5):  Verv  fragile,  moderatelv  slender  conical,  creamy 
white;  reaching  20.8  in  length.  Lengtli/width  2.0-2.2- 
2.3.  Protoconch  whorls  ca.  3.  Protoconch  with  three 
whorls,  with  numerous  fine,  incised,  axial  lines.  Maxi- 
mum diameter  of  protoconch  1.  0.14-0.15  mm  (n  =  2) 
(Figure  30).  Teleoconch  whorls  up  to  10,  separated  by 
deep  suture.  Teleoconch  sculpture  of  evenly  spaced,  or- 
thocline,  thin,  lamellate  costae,  and  numerous  very  fine 
spiral  threads  (>100  on  the  9th  whorl),  superimposed 
on  somewhat  coarser  spiral  cordlets  (ca.  15  on  fifth  te- 
leoconch whorl).  Initial  whorls  with  multiple,  lamellate 
costae,  fused  together  to  form  thicker  ones  (Figures  23, 
24).  Coarser  spiral  cordlets  are  most  prominent  on  initial 
teleoconch  whorls,  where  thev  are  superimposed  on  cos- 
tae (Figure  24);  coarser  cordlets  become  obsolete  on 
most  abapical  whorls.  Costae  are  more  or  less  damaged 
in  all  specimens;  better  preserved  costae  coronate.  Par- 
ticularly below  the  periphery',  costae  somewhat  curved 
abaperturally  at  their  free  margins,  whereas  adapically 
more  erect  and  shghtly  curved  abaperturally  or  adaper- 
turally  near  suture,  depending  on  position  of  costa  on 
adjoining  whorl.  Costae  mostly  continuous,  but  hardly 
touching  each  other.  Fifth  teleoconch  whorl  (width  3.8 
mm)  with  20-24-31  costae.  Five  mm  width  whorl  (whorl 
6  or  7)  with  23-30-33  costae.  Aperture  subcircular.  Ap- 
ertural  lengtli/shell  length  0.3.  Umbilicus  very  narrow. 

Egg  capsules  (Figures  36,  46)  (n  =  8):  Oval  (Figure 
36),  embedded  with  sand  and  closelv  C(jnnected  along 
straight,  longitudinalK  striated,  mucous  thread  (Figure 
46).  Capsules  3,0-3.3-3.5  mm  length  and  1.5-1.6-2.0 
mm  in  width.  Capsules  cont;un  93-120-173  white  eggs. 


ProlM 


)scis: 


Whitish 


Type  material:  lloiotvpe  NNM  59088,  from  t\pe  lo- 
calih.  Paratopes:  NNM  59089/1.  Indonesia,  Sulawesi,  off 
Ujung  Pandang,  W.  Kudingareng  Keke,  12  m;  NNM 
59090/2.  59092/1.  E  Samaknia.  9^m:  NNM  59091/1,  24 
m;  NNM  59093/1,  Bone  Tambung,  7  m. 

Type  locality:  Indonesia,  Sulawesi,  off  Ujung  Pan- 
dang, ESE.  Kudingareng  Keke,  15  m. 

Other  material  examined:  AMS  329657,  Australia, 
Queensland,  off  Macgillivray  Bav,  Lizard  Island, 
14°39'S,  145°29'E.  10  m,  I.  Loch  leg.;  NNM  unnum- 
bered, 1  shell,  Indonesia,  Ambon,  llitu.  outer  part  of 
Ambon  Bav,  eastern  Laha,  J.  C.  den  Hartog  leg. 

Distribution  (Figure  2):  Australia,  Queensland;  In- 
donesia, Ambon,  and  Sulawesi. 

Habitat:  Specimens  of  this  species  were  found  be- 
tween 7-24  meters  depth.  Coral  hosts  were  Fungia  con- 
cinna  Verrill,  1864,  F  fungites,  Hcliofungia  actinifonnis, 
Herpolitha  limax  and  Foh/pht/llia  talpina  (Lamarck, 
1801).  Specimens  were  found  attached  by  mucous 
threads  to  the  underside  of  a  coral;  one  or  two  speci- 
mens were  found  associated  with  up  to  a  few  hundred 
egg  capsules. 

Etymology:  This  species  is  named  after  Ms.  Ingrid 
van  der  Loo.  Leiden. 

Remarks:  Conchologically  this  species  resembles  the 
■probaiile  hoIot)pe"  (Kaicher,  1981:  3036)  of  Epitonium 
clubium  Sowerby  1844  (BMNH  1981234)  from  the  Phil- 
ippines, which  is  an  imperfect  shell  with  a  broken  ap- 
erture and  several  apical  wliorls  missing.  Its  length  could 
have  been  ca.  20  mm.  Costae  of  adjacent  whorls  are 
continuous,  slightly  curved  toward  aperture  adapically 
and  away  from  aperture  abapically,  not  projecting  over 
suture.  The  holotvpe  of  Epitonium  duhium  most  clearlv 
differs  from  E.  ingridae  bv  its  less  prominent  teleoconch 
sculpture  and  thicker,  not  lamellate  costae.  The  speci- 
men figured  bv  De  Boury  (1912:  pi.  7,  fig.  4,  Scala  du- 
hia),  which  might  represent  E.  duhium  (of.  Kilbum, 
1985:  327)  has  more  oblique  costae.  The  identih'  of  Sca- 
laria  graiji  Nvst.  1871,  (nomen  novum  for  Scalaria  stri- 
ata Gray,  1847,  not  Defrance,  1827)  is  unclear;  Tryon 
(1887:  60,  as  S.  striata)  and  De  Bour\-  (1912:  95,  as  S. 
striata  and  S.  graiji)  considered  this  nominal  ta.xon  a  syn- 
onym of  S.  duhia.  Kilburn  (1985:  327)  questioned  this 
synonym)'.  The  shell  of  S.  graiji  figured  by  Tryon  (1887: 
pi.  12,  fig.  68,  as  S.  striata)  has  a  more  shallow  suture 
and  relatively  larger  aperture  when  compared  to  E.  in- 
gridae. The  new  species  also  resembles  Epitonium  fria- 
hilis  (Sowerby  1844)  from  Western  Australia.  Swan  Riv- 
er The  holotvpe  (BMNH  1966653).  figured  b\  Kaicher 
(1980:  2329).'  is  16  mm  in  length  ancf  7  mm  in  width, 
with  ten  whorls.  It  differs  most  conspicuousK'  from  E. 
ingridae  bv  its  closed  lunbilicus  and  absence  of  spiral 
threads.  The  species  described  and  illustrated  from  Svd- 
ney  Harbour  as  Foliaceiscala  barLssa  by  Iredale  (1936: 


Page  8 


THE  NAUTILUS,  Vol.  114,  No.  1 


Figures  22-31.  SEM  micrographs  of  species  of  Epitoniiiin  associated  with  mushroom  corals  off  Ujimg  Pandang  (unless  stated 
otherwise).  22.  E.  costulatum  (Kiener,  1838),  Indonesia,  Ambon,  Hitu,  outer  part  of  Ambon  Bay,  W  Lalia,  apical  whorls.  Scale  line 
=  0.5  mm.  23-24.  E.  in^ridae  new  species  23.  Apical  whorls.  Scale  line  =  0.5  mm.  24.  Teleoconch  whorl  sculpture.  Scale  line  = 
0.1  mm.  25-29.  Protoconch.  25.  E.  costulatum  (Kiener,  1838)  (same  shells  as  figure  22).  Scale  hne  =  0.1  mm.  26.  E.  hoeksemai 
new  species.  27.  £.  inffidae  new  species.  28.  E.  twilae  new  species.  29.  E.  uhi  Pilsbr\',  1921.  Scale  hne  =  0.1  mm.  30-31. 
Protoconch  1.  30.  £  ingridae  new  species.  31.  E.  twilae  new  species.  Scale  line  =  0.02  mm. 


Gittenberger  el  ai.  2()()() 


Page  9 


Figures  32-35.  Species  of  Epitoniiim  often  confused  in  the 
literature.  32-33.  E.  tivilae  new  species,  holotype,  NNM 
59104,  length  1.5  cm.  34-35.  £.  buUntitm  (Sowerby,  1844), 
holotype,  BMNH  198136,  length  2.0  cm. 


300,  pi.  22,  fig.  15)  seems  to  be  similar  in  shape  and 
size,  but  the  costae  are  described  as  "of  different 
strength,  some  fine,  others  large  and  recurved,  while  still 
others  approach  varices  in  size." 

Epitoniiim  lochi  A.  Gittenberger  and  Goud  new  species 
(Figures  13-16,  37,  45) 

?Epitonium  species  4:  Loch,   1982:  4-5,   1   fig.  (see  remarks 
below). 

Description:  Shell  (Fig^urc.s  13-16)  (n  =  4):  Very 
fragile,  elongate-conical,  creamy  white,  reaching  8.5  mm 
in  length.  Length/width  2. 0-2. .3-2. 7.  Protoconch  whorls 
3.25.  Maximum  diameter  of  protoconch  1  diameter  0.12 
mm  (n  =  1).  Protoconch  with  numerous  fine,  incised, 
axial  lines.  Teleoconch  whorls  up  to  8,  separated  by  a 
very  deep  suture.  Teleoconch  with  evenly  spaced,  or- 
thocline,  lamellate  costae,  crossing  low  spiral  threads 
that  are  approximateK'  a  half  to  a  fifth  as  wide  as  inter- 
spaces (Figure  15).  Reticulate  pattern  present  on  most 
adapical   whorls,   replaced   by  spiral   threads   on   later 


whorls.  Third  whorl  with  ca.  13  spiral  threads,  fifth  one 
with  ca.  15;  spiral  threads  equally  prominent  throughout 
whorl.  C^ostae  usualK  not  continuous,  lamellar  hut  rather 
low,  barely  touching  preceding  whorl.  Due  to  damage  in 
most  specimens,  fine  structure  of  costae  could  not  be 
examined.  Fifth  teleoconch  whorl  (width  1.3  mm)  with 
24-25-26  costae  and  12-13-15  spiral  threads.  Aperture 
subcircular  Apertural  length/shell  length  0.22-0.23. 
Umbihcus  absent. 

Egg  capsules  {Figures  37,  45):  The  roundish,  white, 
egg  capsules  (Figure  37)  are  mixed  with  sand,  and  close- 
ly connected  to  each  other  along  a  straight  mucous 
thread  without  well-defined  sculpture  (Figure  45). 

Type  material:      Holotv'pe  NNM  59094,  from  type  lo- 

calit)'.  Paratopes:  NNM  59095/2,  59096/1,  Indonesia,  Su- 
lawesi, off  Ujung  Pandang,  MZB/1,  tyi^e  locality;  16  m; 
NNM  59098/1,  18  m;  NNM  59099/1,  E  Kudingareng 
Keke,  3  m;  59100/1,  12  m;  59102/1,  18  m;  NNM  59101/ 
2,  ESE  Kudingareng  Keke,  15  m;  59103/1,  Pulau  Badi, 
24  m.  See  also  Remarks. 

Type  localitj':  Indonesia,  Sulawesi,  off  Ujung  Pan- 
dang, W  Kudingareng  Keke,  12  m. 

Other  material  examined:  AMS  329687/2,  Australia, 
Queensland,  Lizard  Island,  Watsons  Bav,  14°40'S, 
145°27'E,  24  m,  I.  Loch  leg.;  AMS  329688/1,  329689/1, 
Granite  Bluff,  14°39'S,  145°27'E,  23  m,  I.  Loch  leg.  (see 
Remarks  below). 

Distribution:     Indonesia  and  probably  Australia. 

Habitat:  The  snails  were  found  between  3-24  m. 
Fuitgia  costulata  Ortmann,  1889,  and  F.  tenuis  Dana, 
1846,  were  coral  hosts.  If  the  Australian  record  really 
proves  to  be  this  new  species,  Fungia  ci/clolitcs  La- 
marck, 1816,  should  be  included  as  an  additional  host 
(Loch,  1982:  4).  One  to  4  specimens  were  found  at- 
tached by  a  straight  mucous  thread  (Figure  45)  to  the 
underside  of  a  coral  accompanied  by  up  to  a  few  hun- 
dred egg  capsules. 

Etymology:  This  species  is  named  after  Mr  Ian  Loch, 
who  described  this  or  a  very  similar  species  from  Aus- 
tralia, without  naming  it. 

Remarks:  Loch  (1982)  referred  to  and  figured  an  un- 
named species  4'  from  Australia,  distinguishing  it  from 
E.  ulii.  We  were  able  to  compare  that  material  with  the 
specimens  collected  off  Ujung  Pandang.  In  the  Austra- 
lian specimens,  the  spiral  threads  are  somewhat  more 
prominent,  which  could  represent  some  degree  of  intra- 
specific  variation.  The  limited  amount  of  material  does 
not  allow  for  conclusions  on  the  identity'  of  the  Austra- 
lian specimens;  this  prevented  their  inclusion  as  para- 
types. 

Conchologically  this  species  is  most  similar  to  Epiton- 
ium  zat  rephes  MeUiW.  1910  (holotype  BMNH 
191281683),  from  the  Mekran  coast.  This  shell  is  figured 
by  Kaicher  (1980;  2377);  it  differs  bv  having  continuous 


Paee  10 


THE  NAUTILUS,  Vol.  114,  No.  1 


Figures  36 — 48.  .Spt'cii-N  ol  Epiloniiim  associated  with  imislirooiu  corals  oii  L'luiig  Pandang.  36-38.  Egg  capsules.  36.  E.  ingridae. 
Scale  line  =  1  mm.  37.  E.  lochi  new  species.  Scale  line  =  1  mm.  38.  E.  costiilatum  (Kiener.  LS38).  Scale  line  =  1  mm.  39.  Sclerite 
of  alcvoniid  octocoral  found  in  association  with  egg  capsules  ot  E.  costiilatiim.  Scale  line  =  0.1  mm.  40.  Detail  of  figure  39.  Scale 
hue  =  0.1  mm.  41.  E  costiilatum.  Eggs  within  a  capsule.  42.  E  iilii  Egg  capsule  with  protoconchs  1.  Scale  line  =  0.1  mm.  43- 
47.  Mucous  threads.  Scale  line,  with  figure  43,  =  0.01  mm.  43.  E.  hoekscinai.  44.  E  iilii.  45.  E.  lochi.  46.  E.  ingridac.  47.  £. 
costulatiiin.  48.  E.  twilac,  lar\'al  shells  (=  protoconchs  1).  Scale  line  =  0.1  mm. 


costae  and  b\-  the  more  narro\\l\  spaced  spiral  threads, 
which  are  about  as  wide  as  their  interspaces. 

The  holotv'pe  of  Epitoniuin  obliqita  (Sowerbw  1844) 
[Scalaria]  (BMNH  19812.31)  also  resembles  E.  lochi.  but 
differs  in  having  a  clearh'  open,  though  narrow,  umbili- 
cus, and  continuous  costae. 

The  holotvpe  of  Epitoniuin  dcficrsi  (Jousseaume, 
1911)  [Tenuiscala]  (MNHN  De  Bou'r\-2706)  from  Aden, 
which  has  a  broken  aperture  and  missing  apical  whorls, 
can  notwithstanding  be  distinguished  from  £.  loclii  bv 
the  relatively  small  shell  lengtli/width  ratio  (only  ca.  1.9) 
and  the  presence  of  some  varices.  This  holot\pe  is  also 
figured  by  Kaicher  (1981:  3116). 

Epitoniuin  twilae  A.  Gittenberger  and  Goud  new  species 
(Figures  2,  19,  28,  32-33,  48) 

Epitonium  bullatum  (Sowerbv,  1S44):  Dushane.  1988;  .30,  figs. 
5,  6.  Yamashiro,  1990:  299,  figs.  1-6.  Not  Scahiha  hulla- 
tum  Sowerby,  1844. 


Epitonium  species  2:  Loch,  1982:  3^,  1  fig. 
Epitonium  sp.:  Bratcher,  1982:  3,  fig.  1. 

DescripHon:  Slull  (Fifiures  19,  28,  31,  32^3,  48)  (n 
=  20):  Very  fragile,  broad-conical,  creamy  white, 
reaching  17  mm  in  length.  Lengtli/width  1.2-1.-1-1.6. 
Protoconch  whorls  3.  Protoconch  widi  numerous  verv 
fine,  incised,  axial  lines.  Ma.\imum  protoconch  1  diam- 
eter 0.12-0.13  mm  (n  =  2).  Teleoconch  whorls  up  to  9, 
straight-sided,  separated  h\  a  shallow  suture.  Teleoconch 
sculpture  (Figure  19)  of  unevenly  spaced,  fine,  ortho- 
cline,  more  or  less  obsolete  costae,  not  or  onK'  in  part 
lamellate,  and  numerous  low  spiral  threads  that  are  sep- 
arated bv  interspaces  that  varv  from  as  wide  as  the  spiral 
threads  to  three  times  as  wide.  Costae  not  continuous, 
not  curved,  barely  touching  adjacent  whorls.  Fifth  teleo- 
conch whorl  (width  =  2. .5  mm)  with  19-24.4-30  costae. 
Five  mm  width  whorl  (whorl  6  or  7)  widi  24-31-36  cos- 


Gittenberger  et  ai,  2()()() 


Page  11 


tae.  Aperture  subcircular.  Apertural  length/shell  lengtli 
0.37-0.44-{).52.  Umbilicus  ven  uanou. 

E<i^  capsules  (ti  =  10):  Egg  capsules  roundisli,  white, 
transparent,  with  protuberances,  without  embedded 
sand,  closely  connected  along  straiglit  mucous  thread. 
Egg  capsules  1.2-1.4—1.6  mm  diameter  with  342^25- 
532  white  eggs  per  capsule. 

Type  material:  Holotvpe  NNM  59104,  from  type  lo- 
calit\.  Paratopes:  NNM  59149/1,  Indonesia,  Sulawesi,  off 
Ujung  Pandang:  W  Lae-Lae,  7  m:  NNM  59148/2,  9  m; 
NNM  59145/L  t^pe  !ocalit^■,  3  m;  NNM  59105/1.  591.38/ 
3,  6  m:  NNM  '59127/1,  59129/1,  .59141/10,  59147/1, 
59150/2,  9  m;  NNM  59126/1,  59142/1,  59143/1,  12  m; 
NNM  59146,  15  ni;  NNM  .59139,  ESE  Samalona,  5  m; 
NNM  59131/4,  12  ni;  NNM  59140/5,  13  ni;  NNM 
59116/1,  59117/1,  591.32/3,  591.3.3/2,  59134/1,  .591.35/1, 
W  Samalona,  9  m;  NNM  .59121/1,  59122/1,  12  m:  NNM 
59118/1,  15  m;  NNM  .59151/1,  E  Kudingareng  Keke,  9 
m:  NNM  59106/1,  .59107/1,  W  Kudingareng  Keke,  9  ni; 
NNM  .5911.5/3,  10  m;  NNM  .5911.3/1,  .59114/1,  12  ni; 
NNM  59108/1,  .59109/1,  .59110/1,  .59111/2,  .59112/1,  15 
m:  NNM  .5912.3/1,  .59124/1,  18  m;  NNM  .591.52/1, 
.591.5.3/1,  24  m;  NNM  .591.37/1,  NW  Lankai.  12  ni;  NNM 
.59159/5,  Pulau  Badi,  25  m;  NNM  59161/2,  Bone  Lola, 
8  m:  NNM  5959160/1,  9  m;  NNM  59154/2,  591.5.5/2, 
.59156/2,  5916,3/1,  Bone  Tambung,  5  m;  NNM  59157/1, 
22  m.  Onh'  the  specimens  from  off  Ujung  Pandang  are 
considered  the  t\pe  series  (see  Distribution  below). 

Type  locality:  Indonesia,  Sulawesi,  off  Ujung  Pan- 
dang, W.  Bone  Baku,  6  m. 

Other  material  examined:  AMS  3296.5.3/1,  Austraha, 
Queensland  (see  also  Loch,  1982:  3,  4,  1  fig.):  No.  5 
Sandbank  Reef,  1.3°45'S,  144°16'E,  9  m,  I.  Loch  leg.; 
AMS  09980.3/2,  0998041,  09980.5/1,  099806/1,  100188/ 
14,  .329680/1,  .32968.3/2,  off  Lizard  Island,  14°39'- 
14°42'S,  14,5°23'-145°28'E,  2-11  m,  P  H.  Colman,  1. 
Loch  and  W.  F.  Ponder  leg.;  AMS  329672/1,  Opal  Reef, 
N.  of  Cairns,  16°15'S,  14.5°.50'E,  9  m,  I.  Loch  leg.;  AMS 
09657.5/2,  101238/2,  147334/2,  329676/4,  329679/3, 
329670/1,  E-NE  of  Townsville.  18°46'-18°57'S, 
147°31'-147°44'E,  9-18  m,  I.  Loch  leg.  NNM  unnum- 
bered, Indonesia,  Ambon,  Hitu,  outer  part  of  Ambon 
Bay,  W  Laha,  J.  C.  den  Hartog  leg.;  NNM  unnumbered, 
Sulawesi,  off  Ujung  Pandang. 

Records  in  the  literature:     Papua  New  Guinea,  Na- 

gada  (16  km  N  of  Madang)  (Bratcher,  1982:  3,  1  fig.). 
Japan:  Sesoko  Island,  Okinawa  (Yamashiro,  1990;  299- 
305,  figs.  1-6).  Red  Sea,  Sinai,  Thomas  Reef,  27°59'N, 
.34°27'E  (Dushane,  19SSa:  31,  figs.  5,  6). 

Distribution  (Figure  2):  Australia  (Queensland), 
Papua  New  Guinea,  Indonesia,  Japan,  and  Red  Sea. 

Habitat:  This  species  was  found  from  3  m  to  the  max- 
imum diving  depth  of  24  meters.  In  the  literature  that 
might  refer  to  this  species  a  depth  of  45  meters  was 
mentioned  (Loch,  19S2).  The  following  coral  host  spe- 


cies were  recorded:  Ctcnactis  echinata,  Heiyolillui  li- 
max,  Sandtilolitlui  dcntata  Quelch,  1884,  S'.  rolnista 
(Quelch,  1886)  and  Zoopiltis  echinatus  Dana,  1846. 
Clung  with  mucous  threads  to  the  underside  of  a  coral, 
one  to  fourteen  specimens  were  found  accompanied  by 
up  to  a  few  hundreds  of  egg  capsules. 

Etymology:  This  species  is  named  after  Mrs.  Twila 
Bratcher,  of  Los  Angeles,  California,  USA,  vylio  first  dif- 
ferentiated the  new  taxon  from  E.  Imllotiint. 

Remarks:  This  species  has  been  misidentified  by  var- 
ious authors  (Dushane,  1988a;  Yamashiro,  1990;  Mienis, 
1994,  conditionally)  as  Epitonium  hullatum  (Sowerbv, 
1844),  a  species  associated  with  sea  anemones  (Kilburn 
and  Rippey,  1982;  Kilbum.  1985;  Mienis,  1994).  The 
badly  damaged  holot)pe  of  E.  buHaturn  (Figtires  34,  .35) 
has  a  more  globular,  far  less  fragile  shell  with  convex 
whorls,  costae  occasionally  forming  a  varix,  and  only 
about  5  teleoconch  whorls  at  a  length  of  19  mm  (several 
apical  whorls  are  missing).  The  specimens  illustrated  by 
Jousseaume  (1921:  pi.  3,  fig.  2),  Azuma  (1962:  fig.  2,  as 
Globiscala  koshiwajimcnsis),  Kilbum  and  Rippev-  (1982; 
pi.  11,  fig.  15),  Kilbum  (1985:  .3.30,  figures  160-163)  and 
Wilson  (1993:  pi.  44,  fig.  9)  exemplifs-  the  variability-  of 
E.  hiillnfiiin.  Although  £.  tuihic  differs  conspicuousK- in 
shape  from  the  other  Epitonium  species  in  this  studv;  its 
protoconch  (Figures  28,  31,  48)  cannot  be  distinguished 
from  that  of  these  other  species. 

Yamashiro  (1990)  published  various  data  on  the  life 
history  of  £.  twilac  (as  E.  huUatum).  That  author  de- 
scribed the  egg  capsules  as  elliptical,  0.88  mm  in  length 
and  0.75  mm  in  width,  containing  38-98  eggs  each. 
These  data  chffer  from  our  results.  Based  on  very  similar 
shell  morphologies,  however,  we  consider  his  specimens 
and  the  ones  examined  in  this  section  to  be  conspecific. 

Despite  the  fact  that  E.  twilac  differs  markedly  in 
shell  morphology  from  E  ulu,  the  protoconchs  of  these 
species  are  yer\'  similar 

Epitonium  uhi  Pilsbrv,  1921 
(Figures  2,  11-12,  17,  21,  29,  44) 

Epitonium  ulu  Pilsbiy,  1921:  376,  fig.  llc;  Bosch,  196.5;  267, 
fig.  1:  Robertson,  1970:  4.5;  Hadfield,  1976:  1.35,  Table  1; 
Tavlor,  1977:  253,  2.58,  fig.  7;  Kav,  1979:  1.56,  fig.  5.3a,  b; 
Loch,  1982:  3,  1  fig.;  Bell,  1985:  '1.59,  figs.  1-6;  Dushane, 
1988a:  31,  figs.  3,  4;  1988c:  9,  1  fig.;  Wilson,  1993:  273. 

Description:  Shell  (Fipircs  11-12,  17,  21,  29)  (n  = 
20):  Very  fragile,  elongate-conical,  creamy  white; 
reaching  16  mm  in  length.  Lengtli/width  2. .3-2. 6-3. 6. 
Protoconch  whorls  3.  Maximiun  protoconch  1  diameter 
0.13  mm  (n  =  2).  Protoconch  with  numerous,  ver\-  fine, 
incised,  axial  lines.  Teleoconch  whorls  up  to  12,  sepa- 
rated by  moderately  deep  suture.  Teleoconch  sculpture 
varxing  in  intensit)'  from  well-defined  to  obsolete.  Cos- 
tae unevenly  spaced,  orthocline,  more  or  less  lamellate 
or  obsolete,  not  continuous,  relatively  prominent  adapi- 
cally  and  clearK'  encroaching  on  adjacent  whorl,  curxed 
in  adapertural  direction  (Figure  21).  Spiral  threads  vary 
in  strength  on  a  single  whorl;  spiral  threads  only  slightly 


Page  12 


THE  NAUTILUS,  Vol.  114,  No.  1 


increasing  in  number  on  later  whorls.  Fifth  teleoconch 
whorl  (width  2.0  mm)  with  15-23-28  costae  and  9-11- 
15  spiral  threads.  Five  mm  width  whorl  (between  whorl 
8  and  11)  with  19-28-33  costae  and  10-14-25  spiral 
direads.  Aperture  circular  to  somewhat  oval.  Apertural 
length/shell  length  0.20-0.26-0.29.  Umbilicus  very  nar- 
row to  closed. 

Egg  capsules  (Figure  44):  Egg  capsules  roundish  to 
oval,  white,  granulated,  sometimes  embedded  with  sand. 
Egg  capsules  closely  connected  along  a  twisted  mucous 
diread  (Figure  44)';  Dushane  (19S8a;  32)  reported  3 
t\\isted  threads.  Capsule  diameter  0.8-1.3-1.7  mm  (n  = 
5).  One  capsule  contains  67-225-405  eggs.  Dushane 
(1988a:  32)  reported  400-600  eggs  within  a  capsule, 
which  she  described  as  papillose,  with  softly  rounded 
papillae.  Kay  (1979:  fig.  53B)  figures  the  egg  capsules  as 
elliptical,  1.1  mm  in  width  and  ca.  1.6  mm  in  length.  See 
Bell  (1982;  1985)  and  Dushane  (1988a)  for  further  data 
on  egg  capsules,  lite  history  and  other  relevant  data. 

Proboscis:  Whitish,  with  some  transversal,  transparent 
bands. 

Type  material:  Holotype  ANSP  127818,  from  t\pe  lo- 
caliU'. 

Type  locality:     USA,  Hawaii,  Hilo. 

Other  material  examined:  ANSP  127818.  USA,  Ha- 
waii, Big  Island,  Hilo;  AMS  138321/1,  Australia,  Queens- 
land (see  also  Loch,  1982:  3,  1  fig.).  Eel  Reef,  12°24'S, 
143°22'E,  4-8  m,  I.  Loch  leg.;  AMS  329660/1,  Long 
Sandy  Reef  12°29'S,  143°46'E,  10  m,  I.  L<xh,  leg.;  AMS 
099801/3,  099802/2,  100821/1,  329656/3,  near  Lizard  Is- 
land, 14°40'-14°42'S,  145°23'-145°28'E,  1.5-14  m,  P 
H.  Colman  and  I.  Loch  leg.;  AMS  138320/1,  S  Escape 
Reef  15°53'S,  145°49'E,  18  m.  I.  Loch,  leg.;  AMS 
096573/7,  329649/2,  329650/1,  329651/1,  329652/1, 
329655/2,  329658/7,  E-NE  of  Townsville,  18°46'- 
18°57'S,  147°31'-147°44'E,  6-15  m,  I.  Loch  leg.;  NNM 
unnumbered,  Indonesia:  Ambon,  Hitu,  outer  part  of 
Ambon  Bay,  E  and  W  Laha,  A.  Fortuin  and  J.  C.  den 
Hartog  leg.;  LACM  86-163.  Java,  off  Jakarta,  Kepulauan 
Seribu  (=  Thousand  Islands),  Pulau  Pelangi  and  Pulau 
Putri;  AMS  138318/1,  Malaysia:  Pulau  Singa  Besar  Pu- 
lau Langkawi,  6°14'S,  99°44'E,  I  m,  I.  Loch,  leg. 

Records  in  the  literature:  USA,  Hawaii,  Oaliu,  Ka- 
neoke  Bay  (Bell  1985:  159-164,  figs.  1-6);  Papua  New 
Guinea  (Dushane.  1988a:  32;  Maldives  (Dushane, 
1988a:  32);  Red  Sea,  Straits  of  Tiran,  Tiran  Island  and 
Sinafir  Island  (Dushane,  1988a:  31,  32,  figs.  3,  4);  Sinai, 
Thomas  Reef  27°59'N,  34°27'E  (Dushane,  1988a:  31, 
figs.  5,  6). 

Distribution  (Figure  2):  Hawaii,  Australia  (Queens- 
land), Indonesia,  Malaysia,  Maldives,  and  Red  Sea. 

Habitat:  Tliis  species  was  recorded  between  3-24  me- 
ters depth.  Coral  hosts  were  Fungia  spinifcr  Claere- 
boudt  and  Hoeksema,  1987,  F.  scabra  Doderlein,  1901, 
F.  concinna,  F.  horrkla  Dana,  1846,  F.  scniposa  Klun- 


zinger,  1879,  F  fungites,  F  granulosa  Klunzinger,  1879, 
F.  scutaria  Lamarck,  1801,  F.  moluccensis  Van  der  Horst, 
1919,  F.  gravis  Nemenzo,  1955,  and  F.  paumotensis 
Stutchbury,  1833.  One  to  11  specimens,  free  or  accom- 
panied by  up  to  a  few  hundreds  of  egg  capsules,  were 
observed  on  the  individual  corals,  attached  witli  mucous 
threads  to  the  underside  or  on  the  substrate  of  these 
hosts. 

Remarks:  Shells  of  this  species  yar\'  considerably  in 
lengtli/width  ratio,  intensit\'  of  teleoconch  sculpture  and 
number  of  costae.  They  differ  from  E.  hoeksemai  b)'  a 
lengtli/width  ratio  of  ca.  2.6  instead  of  ca.  1.9  and  b\-  the 
presence  of  less  than  20  spiral  threads  on  the  fifth  te- 
leoconch whorl.  It  is  the  most  common  epitoniid  species 
associated  with  Fungiidae  in  the  study  area. 

ACKNOWLEDGMENTS 

We  are  particularK-  grateful  to  Dr  B.  W.  Hoeksema  for 
his  most  valuable  assistance  in  collecting  and  identifying 
the  fungiid  corals  examined  in  this  article  and  for  con- 
tributing his  expertise  on  corals.  We  thank  all  who  al- 
lowed us  to  examine  material  in  the  molluscan  collec- 
tions under  their  care;  Dr  P.  Bouchet  (MNHN),  Mr  I. 
Loch  (AMS),  Dr  Y.  Finet  (MHNG),  Dr  J.  H.  McLean 
(LACM),  Dr  G.  Rosenberg  (ANSP).  and  Mrs.  K.  Way 
(BMNH).  We  also  thank' Mr  A.  Weil  and  Mrs.  L. 
Brown,  who  contributed  expertise  on  epitoniids,  and  Dr 
L.  P.  van  ORvegen  (NNM),  who  identified  the  sclerites 
associated  with  some  egg  capsules.  Some  photographs 
were  taken  b\-  A  "t  Hooft  (Universitv-  of  Leiden  Institute 
of  Evolutionary  and  Ecological  Sciences,  Leiden).  This 
project  was  subsidized  h\  the  Royal  Dutch  Academ\'  of 
Sciences,  the  SHchting  Jan  Joost  ter  Pelkwijk-fonds  and 
the  Leiden  Universits  Foundation. 


LITERATURE  CITED 

Azuma,  M.  1962.  Descriptions  of  five  new  species  of  Japanese 
Epitoniidae.  Venus  22:1.30-13.5. 

Bell,  J.  L.  1982.  Larval  development  and  metamorphosis  of  die 
prosobranch  mollusc  Epitonium  ulii.  associated  v\ith  a  sol- 
itary coral.  Pacific  Science  16:508. 

Bell,  J.  L.  1985.  Larval  growth  and  metamoqshosis  of  a  pros- 
obranch gastropod  associated  with  a  solitary  coral.  Pro- 
ceedings of  the  fifth  international  coral  reef  congress,  Ta- 
hiti 5:159-164. 

Bosch,  H.  F.  1965.  \  gastropod  parasite  of  solitan'  corals  in 
Haw;iii.  Pacific  Science  19:267-268. 

Bouchet,  P.  and  A.  Waren.  1986.  Revision  of  the  Nordieast 
Adantic  bathyal  and  abvssal  Aclididae,  Eulimidae,  Epito- 
niidae (Mollusca,  Gastropoda).  Bollettino  Malacologico, 
Supplemento  2:299-576. 

Bratcher  T.  1982.  The  Fungia  coral:  It's  full  of  snrjsrises.  Ha- 
waiian Shell  News  30{.3);3. 

Clench,  W.  J.  and  R.  D.  Tvmier  1951.  The  genus  Epiionium 
in  the  western  .\tlantic.  Part  I.  Jolnrsonia  2:249-.3.56. 

De  Boury,  E.  1912.  Description  de  Scalidae  nouseaux  ou  pen 
connus.  Journal  de  Conchvliologie  60:87-107. 

Dushane,  H.  1988a.  Geographical  distribution  of  some  Epito- 


Gittenherger  et  «/.,  2000 


Page  13 


niidae  (Mollusca:  Gastropoda)  associated  with  liint!;iid  cor- 
als. The  Nautilus  102:30-35. 

Dushane,  H.  1988b.  Hawaiian  Ejiitoiiiidac.  Hawaiian  .Shell 
News  36(4):6-7. 

Dushane,  H.  1988c.  Hawaiian  Epitoniidae  (continued).  Ha- 
waiian Shell  News,  36(11):7,  9. 

Eisenberg,  J.  M.  1981.  \  collector's  guide  to  sea  shells  of  the 
world.  Bloonisburv,  Books,  London.  2.39  pp. 

Hadfield,  M.  G.  1976.  .Vlolhi.scs  as.sociated  with  li\ing  tropical 
corals.  Micronesica  12:133-148. 

Hartog,  J.  C.  den.  1987.  Observations  on  the  wentletrap  Epi- 
tonium  clathraUilum  (Kanmacher,  1797)  (Prosobranchia, 
Epitoniidae)  and  the  sea  anemone  Btinodosoma  hiscaij- 
ensis  (Fischer,  1874)  (Actinaria,  Actiniidae).  Basteria 
5I(4-6):9.5-108. 

Hoeksenia,  B.  W.  1988.  Mobilih'  of  free-living  fungiid  corals 
(Scleractinia),  a  dispersion  mechanism  and  sui"viv;J  strat- 
egy in  dviiamic  reef  habitats.  /(!,  Choat,  ].  H.,  D.  Barnes, 
M.  A.  Borowitzka,  J.  C.  Coll,  P.  ].  Davies,  P  Flood,  B.  G. 
Hatcher,  D.  Hopley,  P  A.  Hutchings,  D.  Kinsey,  G.  R. 
Orme,  M.  Pichon,  P.  F  Sale,  P.  Sammarco,  C.  C.  Wallace, 
G.  Wilkinson,  E.  Wolanski  and  O.  Bellwood  (eds.)  Pro- 
ceedings of  the  6th  International  Coral  Reef  S\inposiuni, 
Townsville,  Australia,  pp.  715-720. 

Hoeksenia,  B.  W.  1989.  Taxonomy,  plnlogenv  and  biogeogra- 
phy  of  mushroom  corals  (Scleractina:  Finigiidae).  Zoolo- 
gische  Verhandelingen  254:1-295. 

Iredale,  T.  1936.  Australian  niolluscan  notes.  No.  2.  Records 
of  the  Australian  Museum  I9(5):267-340. 

Jousseaume,  F.  1921.  Sur  qnelques  mollusques  de  la  Mer 
Rouge  nouveaux  ou  non  figures.  Memoires  de  la  Societe 
Zoologique  de  France  28:53-60,  pi.  3. 

Kaicher,  S.  D.  1980-1983.  Epitoniidae  I-IH.  Card  catalogue 
of  world-wide  shells.  Pack  23  [1980],  Pack  .30  [1981]  and 
Pack  35  [1983]. 

Kay,  E.  A.  1979.  Hawaiian  marine  shells.  Reef  and  shore  fauna 
of  Hawaii.  Section  4:  Mollusca.  Special  Publication  64(4). 
Beniice  P.  Bishop  Museum,  Honolulu,  653  pp. 

Kiener,  L.  1838.  Genre  Scalaire.  Species  general  et  iconogra- 
phie  des  coquilles  vivantes.  pis.  1-7,  1-22. 

Kilbuni,  R.  N.  1985.  The  family  Epitoniidae  (Mollusca:  Gas- 
tropoda) in  southern  Africa  and  Mozambique.  Annals  of 
the  Natal  Museum  27:239^337. 

Kilbuni,  R.  N.  and  E.  Rippev.  1982,  Sea  shells  of  southern 
Africa.  Macmillan  South  Africa,  Johannesburg,  249  pp. 

Loch,  1.  1982.  Queensland  epitoniids.  Australian  Shell  News 
39:3-6. 

Loo,  G.  K.  M.  and  L.  M.  Cliou.  1988,  Corals  of  the  genus 
Fimgia.  Nature  Malaysiana  13:26-29. 

Mienis,  H.  K.  1994.  CalUnctis  poh/piis:  a  new  host  of  Epiton- 
itim  buUatum.  Epinet  3(2):.5-6. 

Nakayama,  T  1991.  A  new  epitoniid  species  from  the  Pacific 
Coast  of  the  Kii  Peninsula,  Japan.  The  Veliger  .34(1):88- 
90. 

Ohverio,  M,,  M.  Taviani  and  R.  Clieiiiello.  1997.  A  coral-as- 
sociated epitoniid,  new  to  the  Red  Sea  (Prosobranchia, 
Ptenoglossa).  Argonauta  9(10-12):.3-10. 

Page,  A.  J.  and  R.  C.  Willan.  1988.  Ontogenetic  change  in  the 
radula  of  the  gastropod  Epitoniuiii  hillccniw  (Prosobran- 
chia: Epitoniidae).  The  X'eliger  .30:222-229. 


Perron,  F.  1978.  The  habitat  and  feeding  behavior  of  the  wen- 
tletrap Epitonium  grccnlandicum.  Malacologia  17:6.3-72. 

Pilsbrv-,  H.  A.  1921.  Marine  mollusks  of  Hawaii.  Proceedings 
of  the  Academy  of  Natural  Sciences  of  Philadelphia  72; 
360-382. 

Robertson,  R.  1963.  Wentletraps  (Epitoniidae)  feeding  on  sea 
anemones  and  corals.  Proceedings  of  the  Malacological 
Society  of  London  35:51-63. 

Robertson,  R.  1970.  Review  of  the  predators  and  parasites  of 
stony  corals,  with  special  reference  to  sviiibiotic  proso- 
branch  gastropods.  Pacific  Science  24:4.'3-54. 

Robertson,  R.  1983a.  Observations  on  the  life  history  of  the 
wentletrap  Epitonium  echinaticostum  in  tlie  Bahamas. 
The  Nautilus  97:98-103. 

Robertson,  R.  1983b.  Obseryations  on  the  life  history^  of  the 
wentletrap  Epitonium  alhidum  in  the  West  Indies.  Amer- 
ican Malacological  Bulletin  1:1-11. 

Robertson,  R.  1993.  Two  new  tropical  western  Atlantic  species 
of  Epitonium.  with  notes  on  similar  global  species  and 
natural  history.  The  Nautilus  107:81-93. 

Sabelli,  B.  and  M.  Taviani.  1984.  Red  Sea  record  of  a  Fungia- 
associated  epitonid.  BoUettino  Malacologico  20:91-94. 

Sale,  S.  1977.  Observations  on  feeding,  chemoreception  and 
toxins  in  two  species  of  Epitonium.  The  Veliger  20:168- 
172. 

Schimek,  R.  L.  1986.  A  diet  that  stings;  sea  anemones  as  food 
for  snails.  Shells  and  Sea  Life  18(11);I73-I75. 

Sherboni,  C.  D.  and  B.  B.  Woodward.  1901.  Notes  on  the 
dates  of  publication  of  part  of  Kiener's  "Species  general 
et  iconographie  des  coquilles  vivantes,  .  .  . '  etc.  Proceed- 
ings of  the  Malacological  Societv'  of  London  4:216-219. 

Taylor,  J.  B.  1977.  Growth  rates  in  juvenile  carnivorous  pros- 
obranchs  (Mollusca:  Gastropoda)  of  Kaneohe  Bay,  Oahu, 
(Hawaii).  In:  Taylor,  D.  L.  (ed.)  Proceedings:  Third  Inter- 
national Coral  Reef  Svanposium,  Biology  1,  Miami,  pp. 
253-259. 

Troschel,  F.  H.  1839.  V.  Mollusca.  7(i:  Erickson,  [initial(s)  not 
indicated],  F  H.  Troschel,  F.  Stein  and  "Herausgeber "  [A. 
F  A.  Wiegmann],  Bericht  iiberdie  Leistungen  im  Gebiete 
der  Zoologie  vviihrend  des  Jahres  1838.  Archiv  fiir  Natur- 
geschichte  5:201-241. 

Ti-von,  G.  W.,  Jr.  1887.  Manual  of  Conchologv;  stnictural  and 
systematic.  Series  1,  Volume  9.  Solariidae,  lanthinidae, 
Tricliotropidae,  Scalariidae,  Cerithiidae,  Rissoidae,  Litto- 
rinidae.  Author's  edition,  Philadelphia,  488  pp.,  71  pis. 

Vecchio,  H.  1964.  Henrietta's  tank  obserx'ations,  wentletraps 
and  anemones.  San  Diego  Shell  Club  5(6):D24. 

Wenz,  W.  1940.  Gastropoda.  Teil  I;  Allgemeiner  Teil  nnd  Pro- 
sobranchia. In:  Schindewolf  O.  H.  (ed.)  Handbuch  der 
Paliiozoologie  6(I[4]):721-960. 

Wilson,  B.  1993.  Australian  marine  shells.  Part  1.  Prosobranch 
gastropods.  Odvs.sey  Publishing,  Kallaroo,  408  pp. 

Yamashiro,  H.  1990.  A  wentletrap  Epitonium  bullntnm  asso- 
ciated with  a  coral  Siindalolithii  rohusta.  Venus  49:299- 
305. 

At  the  time  the  manuscript  was  being  sent  for  tvpesetting,  the 
authors  received  the  following  book: 

Weil,  A.,  L.  Brown  and  B.  Neville.  1999.  The  wentletrap  book. 
Guide  to  the  Recent  Epitoniidae  of  the  world.  Evolver  srl, 
Roma,  246  pp. 

This  work  apparently  does  not  contain  data  that  could  substan- 
tially add  to  or  change  the  contents  of  the  present  article. 


THE  NAUTILUS  114(1):14-17,  2000 


Page  14 


A  new  species  of  Columhella  (Neogastropoda:  Columbellidae) 
from  the  Caribbean  Neogene 


Maria  J.  deMaintenon 

Department  of  Marine  Science 
Universitv  of  Hawaii  at  Hilo 
200  W.  Kawili  Street 
Hilo.  HI  96720  USA 
deniainte@ha\vaii.edu 


ABSTRACT 

A  new  species  of  CohimbeUa  is  described  from  the  Neogene 
of  the  southwestern  Caribbean  and  compared  widi  other  Ca- 
ribbean CohimhcUa  species.  The  new  species  is  a  member  of 
a  species  pair  diitering  primarily  in  inferred  lar^'al  ecolog\-.  Pro- 
toconch  morphology  suggests  that  the  new  species  had  planktic 
development,  whereas  its  extant  analogue.  Columhella  mcrca- 
toria  (Linnaeus,  1758),  has  nonplanktic  development.  Though 
tlie  2  species  were  contemporaneous.  the\'  do  not  appear  to 
have  been  svmpatric. 

Kcif  words:  Colombia.  Costa  Rica,  lanal  ecolog\'.  protoconch. 


INTRODUCTION 

Evolutionary  divergence  in  larval  ecology  between 
planktic  and  nonplanktic  modes  i.s  a  common  pattern  in 
marine  molluscs.  Many  gastropod  genera  in  several  fam- 
ilies display  both  reproductive  modes.  Recent  evolution- 
arv  divergence  may  result  in  pairs  of  .species  differing 
primariK-  in  lar\al  shell  moq^hologx-.  Traditionally  such 
variation  was  considered  as  evidence  for  poecilogony  in- 
traspecific  variation  in  larval  developmental  modes.  Re- 
cent reviews  of  poecilogony  in  marine  invertebrates 
(Hoagland  and  Robertson,  1988;  Bouchet.  1989;  Knowl- 
ton,  1993;  Oli\erio,  1996)  however,  ha\e  found  httle  ev- 
idence to  definitiveK'  support  that  poecilogony  occurs. 
Hoagland  and  Robertson  (I98S)  reviewed  reported  cases 
of  poecilogon\'  in  marine  gastropods,  and  concluded  that 
the  pattern  of  larval  development  is  generalK  informa- 
Hve  in  species-level  invertebrate  swstematics.  The  pur- 
pose of  this  paper  is  to  describe  a  new  species  of  col- 
umbellid  gastropod,  CohimhcUa  mohiciisis,  that  differs 
from  another  Caribbean  CohimhcUo  species  in  its  in- 
ferred developmental  mode.  AdditionalK.  the  2  species 
appear  to  be  allopatric,  lending  further  support  to  their 
taxonomic  distinction. 

CohimhcUa  Lamarck,  1799  is  primariK-  a  tropical 
American  taxon,  consisting  of  15  Neogene  and  Recent 
American  species  and  2  species  in  the  eastern  Atlantic. 
Phylogenetic  analyses  of  representative  columbellid  ta.\a 


(deMaintenon.  1999)  supports  the  monoph\K  of  Col- 
umhcUa.  based  on  8  characters  of  anatom\-,  radular  mor- 
phology' and  shell  morpholog)-.  Recent  species  of  Col- 
iimhcUa  are  differentiated  primarily  on  the  basis  of  con- 
chological  characters.  The  extant  species  comprise  2 
morphological  groups,  one  of  which  occurs  in  the  Atlan- 
tic. The  Atlantic  CohimhcUa  species  are  very  similar,  and 
are  characterized  by  having  shells  with  spiral  cords  over 
die  entire  shell  surface.  The\'  differ  primariK'  in  shell 
coloration  and  in  the  number  and  strength  of  spiral 
cords.  Jung  (1994)  recently  reviewed  the  fossil  species 
of  the  Atlantic  group  in  a  discussion  of  the  CohimhcUa 
species  from  the  Neogene  of  the  Dominican  Republic. 
The  extant  members  of  the  Atlantic  group  include  2  spe- 
cies in  the  eastern  Atlantic  and  2  in  die  western  Atlantic. 
The  known  fossil  record  of  the  group  extends  back  to 
the  late  Miocene.  Of  the  living  and  fossil  species  of  Col- 
iimhcUa  in  the  Atlantic,  the  new  species  is  the  onK'  one 
considered  to  have  planktic  development. 

The  second  group  comprises  the  10  extant  Panamic 
CohimhcUa  species.  These  differ  from  the  Atlantic  group 
by  lacking  spiral  cords  on  the  shell  except  as  juveniles. 
All  members  of  the  Pacific  group  have  multiwhorled 
protoconchs,  and  are  inferred  to  have  planktic  devel- 
opment. The  fossil  record  of  this  group  is  unk-nown  be- 
fore the  Pliocene,  when  the  extant  species  appear  Al- 
lopatric species  pairs  differing  in  developmental  mode 
alread\'  have  been  reported  in  the  group  of  Atlantic  Col- 
umhella species.  Moolenbeek  and  Hoenselaar  (I99I)  dif- 
ferentiated 2  species  in  the  eastern  Atlantic:  CohimhcUa 
nistica  (Linnaeus.  1758)  with  nonplanktic  development, 
in  the  Mediterranean  and  eastern  Atlantic  coast,  and 
CohimhcUa  adansoni  (Menke,  1853)  with  planktic  de- 
velopment, endemic  to  the  offshore  islands  of  the  east- 
em  Atlantic.  Stud\'  of  alloz\me  data  from  these  2  species 
(Oliverio,  1995,  1996)  indicates  that  they  thverged  about 
2  million  years  ago.  The  new  species  described  herein 
has  a  similar  relationship  with  one  of  the  western  Atlan- 
tic species,  Columhella  mcrcatoria  (Linnaeus,  1758). 

The  following  institutional  abbrexiations  are  used:  UF, 
Universit\'  of  Florida,  Florida  Museum  of  Natural  His- 


M.  J.  deMaintenon,  2000 


Page  15 


torv';  SBMNH,  Santa  Barbara  Museum  of  Natural  His- 
tor\-:  UCMP,  Unhersih'  of  California  Museiun  of  Pale- 
ontologN ;  uses,  United  States  (Geological  Survev; 
USNM,  National  Museum  of  Natural  History-,  Smithson- 
ian Institution. 

The  micrographs  in  figures  1  and  2  were  taken  using 
the  Elettroscan  Environmental  Scanning  Electron  Mi- 
croscope at  the  Universit)'  of  California  Museum  ot  Pa- 
leontology. Specimens  were  scanned  uncoated,  at  15  k\' 
and  between  2  and  3  Torr  water  vapor. 

SYSTEMATICS 

Famil\-  Columbellidae  Swainson,  1840 
Genus  ColumheUa  Lamarck,  1799 


Columbclla  luoincnsis  new  species 
(Figures  2,  4-8,  Table  1) 

Description:  Shell  small,  stronibiform,  up  to  19  mm 
shell  length.  Aperture  length  slightK'  more  than  half 
shell  length.  Protoconch  conical,  multiwhorled,  smooth, 
widi  3  whorls.  Teleoconch  with  6-7.5  whorls,  with  spiral 
cords  over  all  whorls.  Bodv  whorl  with  12-18  cords 
counted  at  the  aperture  edge,  continuing  anteriorlv  to  a 
few  finer  cords  at  the  base  of  the  shell.  Each  spire  whorl 
with  4-6  cords  visible.  Usually  vvith  a  prominent,  slighth- 
nodulose  cord  at  the  shoulder,  which  is  most  often  the 
second  cord  below  the  suture.  First  2-3  teleoconch 
whorls  with  nodules  at  the  periphers'  and  weak  axial 
ridges,  which  fade  out  on  subsequent  whorls.  Aperture 
constricted  by  inrolled  labial  edge,  thickest  in  the  center. 
Labial  edge  with  10-12  denticles,  facing  inward.  Colu- 
mella with  2  weak  folds.  Anterior  parietal  wall  with  5- 
10  denticles  in  an  axial  row.  Short  posterior  parietal  cal- 
lus, continuous  with  aperture  edge. 

Type  material:  Holot>pe,  UCMP  39918,  12.8  mm 
long,  7.5  mm  wide,  Uq^e  iocalit>'.  Para^pe  UCMP  39919, 
t\-pe  localit)';  Paratopes  UCMP  39920  and  39921,  Upper 
Tubara  Group,  below  type  localit\',  Punta  Pua,  Bolivar, 
Colombia;  Paratype  USNM  501150,  Moi'n  Fm.  2  km 
west  of  Puerto  Limon,  Costa  Rica;  Paratvpe  USNM 
501151,  Mofn  Formation,  between  Puerto  Limon  and 
Pueblo  Nuevo,  Costa  Rica. 

Type  locality:  Upper  Tubara  Group,  Punta  Pua,  about 
15  miles  northeast  of  Cartagena,  Bolivar,  Colombia 
(UCMP  S-66). 

Other  material  examined:  USGS  localit\'  21037,  1 
specimen,  Moi'n  Formation,  outskirts  of  Puerto  Limon, 
Costa  Rica;  UCMP  S-66,  11  .specimens;  UCMP  S-65,  2 
specimens;  TU  954,  1  specimen;  TU  956,  1  specimen. 

Distribution:  The  k-nown  lots  of  Coliimbella  moinen- 
sis  are  from  the  Pliocene  to  Pleistocene  (?)  of  Costa  Rica 
and  Colombia.  The  material  from  Colombia  lacks  pre- 
cise stratigraphic  data  but  is  thought  to  be  from  the  Mio- 
cene to  Pliocene  Tubara  Group  (Yokes,  1990).  Yokes 
(1990)  reported  that  the  muricid  species  she  studied 


from  these  2  (^oloniiiian  localities  are  tspical  of  the 
Pleistocene  Moi'n  fauna  from  Costa  Rica.  .Although  the 
Moi'n  Formation  has  better  stratigraphic  definition  than 
the  above  mentioned  units,  some  controversy  exists 
about  the  age  of  the  formation.  Coates  et  al.  (1993)  state 
that  the  Moi'n  Formation  is  Late  Pliocene  in  age,  how- 
ever its  fainias  are  more  usuallv  considered  to  be  Pleis- 
tocene in  age  (Robinson,  1993,  and  references  therein). 
Thus  the  stratigraphic  range  of  this  species  is  currently 
imprecise. 

Etymology:  Columbclla  itwiucitsis  is  named  after  the 
Moi'n  Formation  of  Costa  Rica. 

Discussion:  Columbclla  uioincusis  is  verv'  similar  to 
Columbclla  mercatoiia  (Linnaeus,  1758)  (Figure  3).  a 
common  western  Atlantic  species  k-nown  from  the  Pli- 
ocene to  the  Recent.  The  new  species  can  be  differen- 
tiated from  C.  mercatoiia  by  its  multiwhorled  proto- 
conch. Columbclla  mercatoria  has  1.5  to  2  bulbous  pro- 
toconch whorls  (Figure  1),  with  at  least  one  specimen 
having  2.25  whorls.  The  new  species  has  a  conical  pro- 
toconch with  3  whorls  (Figure  2),  and  its  nuclear  whorl 
is  smaller  than  that  of  C.  mercatoria.  This  latter  species 
has  nonplanktic  development;  its  eggs  hatch  as  crawling 
juveniles  after  about  33  days,  during  which  the  larvae 
feed  on  nurse  eggs  in  the  capsules  (Bandel,  1974).  The 
multiwhorled  protoconch  of  C.  moinensis,  in  contrast,  is 
indicative  of  plank-tic  development. 

Columbclla  mercatoria  varies  greatly  in  adult  size  and 
strength  of  sculpturing,  as  do  many  other  Columbclla 
species.  In  the  case  of  C  mercatoria,  this  variation  may 
be  a  function  of  environment.  In  southern  Florida,  the 
species  is  commonlv  associated  with  both  reef  rock  and 
seagrass  habitats,  and  specimens  found  in  seagrasses 
tend  to  be  smaller  and  less  brightK  patterned  than  reef 
specimens.  The  k-nown  specimens  of  C.  moinensis  come 
from  only  6  lots,  and  most  of  the  material  is  similar  in 
size.  The  single  specimen  from  LISGS  localit\'  21037 
(USNM  501152,  Figure  8)  is  inarkedK'  larger  than  other 
specimens  (Table  1),  and  has  a  weaker  shoulder.  It  does 
have  a  multiwhorled  protoconch  (broken  off  though  still 
with  the  shell)  consistent  with  that  of  C.  moinensis,  so 
will  be  referred  to  the  new  species  in  spite  of  the  dif- 
ferences between  it  and  odier  specimens. 

Many  of  the  specimens  that  are  considered  to  repre- 
sent C  moinensis  have  eroded  protoconchs  and  thus 
cannot  be  diagnosed  with  certaintv.  However,  the\-  are 
comparable  in  shape  and  size  to  specimens  of  C  mo- 
inensis from  the  same  localities. 

Columbclla  tnercatoria  was  contemporaneous  with  C. 
nwinensis.  However,  the  2  species  have  not  so  far  been 
found  svmpatricallv.  Columbclla  mercatoiia  has  been 
collected  in  the  Pliocene  and  Pleistocene  Mare  and  Abi- 
sinia  Formations  of  Yenezuela  (Weisbord,  1962)  and  in 
the  Pliocene  Bowden  Formation  of  |amaica  (a  worn  and 
damaged  .specimen  incorrectK'  identified  as  Columbclla 
submercatoiia  is  illustrated  by  Woodring,  1928). 

Columbclla  submercatoria  Olsson,  1922  occurs  in  the 
Late  Miocene  of  the  Dominican  Republic  (Jung,  1994) 


Page  16 


THE  NAUTILUS,  Vol.  114,  No.  1 


Figure  1.  Recent  CoIuml>elln  mercntoria.  protoeonch.  UF  126820.  Colon  Iskuid.  Bocas  del  Tom  Province,  Panama.  Scale  line  =  2(X) 
jxni.  Figure  2.  Coliimhdln  moineim.s  new  species.  Protoconch  of  paratvpe,  UCMP  .39919.  Sc;ilc  line  =  2(X)  ]xm.  Figure  3.  Recent 
Columbdla  inercntuiia  (SBMNH  144H5I,  Los  Totiinios,  Venezuela).  Scale  line  =  5  mm.  Figures  4—5.  Colunibclla  iitoinciisLs  new 
species.  Holohpe,  UCMP  39918.  Scale  line  =  5  mm.  Figures  6-7.  Columbdla  inoinensi.s  new  species.  Paratvpe,  UCMP  39919.  Scale 
line  =  5  mm.  Figure  8.      Larsje  specimen  itf  (^ohnnlirlla  niciiiirii'ii-s  now  species.  USNM  .5011.52.  Sc;Je  line  =  5  mm. 


M.  J.  deMaintenon,  2000 


Page  17 


Table  1.  Lengths  and  widths  in  mm,  and  number  of  cords  on 
the  bodv  whorl  of  t\pes  and  figured  specimens 


Spe 


Max. 
length 
(mm) 


Ma\.  width       No.  of 
(mm)       spiral  cords 


Holotvpe,  UCMP  39918 
Paratype,  UCMP  .39919 
Paratvpe,  UCMP  39920 
Paratvpe,  UCMP  .39921 
Parat\pe,  USNM  .5011.50 
Paratvpe,  USNM  501151 
USNM  .5011.52 


12.8 

7.5 

14 

11.4 

6.3 

14 

15.3 

8.7 

12 

15.8 

9.1 

17 

10.1 

5.5 

15 

12.5 

7.2 

16 

18.6 

10.4 

17 

and  in  the  Neogene  of  Costa  Rica.  Whether  this  is  a 
distinct  species  or  a  form  of  C.  mcrcatoiia  lias  been  de- 
bated and  should  still  be  regarded  as  uncertain.  The  pri- 
marv'  difference  between  them  is  the  greater  number  of 
spiral  cords  on  C.  submcrcatoiia  (22  to  25  per  whorl 
rather  than  15  to  20),  but  large  specimens  of  e.xtant  C. 
mcrcatoiia  from  some  areas  of  the  Caribbean  also  have 
a  greater  number  of  spiral  cords  than  usual.  The  type 
locahty  for  C.  submcrcatoria  is  Red  Cliff  Creek,  Costa 
Rica,  a  locality-  that  is  presentK'  imlocated  but  thought 
to  correspond  to  late  Miocene  or  earl\  Pliocene  beds  in 
the  Limon  Basin  (Jung,  1994).  Coliiinhclhi  submcrcato- 
ria has  a  paucispiral  protoconch  of  about  1.5  whorls 
(Jung,  1994),  similar  to  that  of  C.  rnercatoria. 

LITERATURE  CITED 

Bandel,  K.  1974.  Spawning  and  development  of  some  Col- 
umbellidae  from  the  Caribbean  Sea  of  Colombia  (South 
America).  The  Veliger  16:271-282, 

Bouchet,  P.  1989.  A  review  of  poecilogonv  in  gastropods.  Jour- 
nal of  Molluscan  Studies  55:67-78. 


Coates,  A.  C,  J.  B.  C.  Jackson,  L.  S.  Collins,  T.  M.  Cronin,  H. 
J.  Dowsett,  L.  M.  Bybell,  P  Jung,  and  J.  A.  Obando.  1992. 
Closure  of  the  Isthmus  of  Panama:  the  near-shore  marine 
record  of  Costa  Rica  and  western  Panama.  Bulletin  of  the 
Geological  SocieU-  of  America  104:814-828. 

Jung,  P.  1994.  Neogene  paleontolog\-  in  the  Northern  Domin- 
ican Republic  15.  The  genera  Coluinbdla.  Eiin/pi/rciie, 
Panniictdrin.  Coiwlla,  Nitidella,  and  Mctulella  (Gastro- 
poda: Columbellidae).  Bulletin  of  American  Paleontology 
106(. 344):  1-45. 

deMaintenon,  M.  1999.  The  phylogenetic  relationships  of  mod- 
em columbellid  taxa  (Neogastropoda:  Columbellidae),  and 
the  evolution  of  herbivorv  from  camivorv.  Invertebrate  Bi- 
ologv' 118:2.58-288. 

Hoagland.  K.  E.  and  R.  Robertson.  1988.  An  assessment  of 
poecilogonv  in  marine  invertebrates:  phenomenon  or  fan- 
tasy? Biolo'gical  Bulletin  174:109-125. 

Knowlton,  N.  1993.  Sibling  species  in  the  .sea.  Annual  Review 
of  Ecology'  and  Systematics  24:189-216. 

Moolenbeek,  R.  G.  and  H.  J.  Hoenselaar.  1991.  On  the  identity 
of  'Coliiinbi'llti  nistica'  from  West  Africa  and  the  Maca- 
ronesian  Islands.  Bulletin  Zoologisch  Museum  1.3:6.5-70. 

Oliverio,  M.  1995.  Larval  development  and  allozyme  variation 
in  East  Atlantic  Columhella  (Gastropoda:  Prosobranchia: 
Columbellidae).  Scientia  Marina  .52:77-86. 

Oliverio,  M.  1996.  Life-histories,  speciation,  and  biodiversih"  in 
Mediterranean  prosobranch  gastropods.  Vie  et  Milieu  46: 
16.3-169. 

Robinson,  D.  G.  1993.  The  zoogeographic  implications  of  the 
prosobranch  gastropods  ot  the  Moi'n  Formation  of  Costa 
Rica.  American  Malacological  Bulletin  10:251-255. 

Weisbord,  N.  E.  1962.  Late  Cenozoie  gastropods  from  North- 
ern Venezuela.  Bulletin  of  American  Paleontology 
42(193):  1-672. 

Woodring,  W.  P.  1928.  Miocene  mollusks  from  Bowden,  Ja- 
maica. Part  2:  gastropods  and  discussion  of  results.  Car- 
negie Institute  of  Wasliington.  564  pp.,  40  pis. 

Vokes,  E.  1990.  Cenozoie  Muricidae  of  the  West  Atlantic  re- 
gion Pt.  VIII — Miircx  S.S.,  Haiistclliim.  Cliicorciis  and 
Hcxnplcx,  additions  and  corrections.  Tulane  Studies  in  Ge- 
ology and  Paleontolog}'  23:1-96. 


THE  NAUTILUS  114(l):18-29,  2000 


Page  18 


The  New  Zealand  Recent  species  oi  Muricopsis  Bucquoy, 
Dautzenberg  and  Dollfus,  1882  (Gastropoda:  Muricidae) 


Bruce  A.  Marshall 

Museum  of  New  Zealand  Te  Papa 

Tongarewa 
PO.  Box  467 
Wellington 
NEW  ZEALAND 
bnieeni@tepapa.govt.nz 


Kevin  W.  Burch 

6  Oakland  Avenue 
Whangarei 
NEW  ZEALAND 


ABSTRACT 

Four  New  Zealand  Recent  Miiricopsis  species  are  recognized: 
M.  octogonus  (Quoy  and  Gainiard,  1833),  M.  mariae  (Finlay, 
1930)  and  M.  profunda  new  species,  which  belong  in  subgenus 
Murcxsul  Iredale,  1915,  and  M.  scotti  new  species,  which  is 
referred  to  Rolandiclla  new  subgenus  together  with  the  south- 
em  Australian  species  A/,  umhilicatus  (Tenison  Woods,  1876). 
Their  shells  and  radulae  are  illustrated  and  distributions  plot- 
ted. Mtiiicopsis  mariae  is  interpreted  as  a  species  rather  than 
a  chronosubspecies  of  Af.  espinosiis  (Hutton,  1886)  as  has  been 
traditional. 

Key  words:  New  Zealand,  Muricidae,  Muiicopsis,  Murcxsul, 
new  taxa. 


INTRODUCTION 

For  a  number  of  vears  a  lar2e,  locall\'  common,  sublit- 
tcjral  form  of  Muricopsis  from  rockv  ground  off  northern 
New  Zealand  was  suspected  to  be  an  imnamed  species 
distinct  from  M.  octogonus  (Quoy  and  Gaimard,  1833). 
Extensive  collecting  and  field  observ-ations  led  Scott 
(1989)  to  conclude  that  distinct  species  were  indeed  in- 
volved, and  he  reported  differences  in  shell  morphologv 
operculum  colour  and  habitat.  On  areas  of  rock\'  ground 
at  12-15  m  depth,  cut  by  channels  with  sand  floors,  Scott 
found  that  whereas  M.  octogonus  t\picalh'  lives  both  on 
horizontal  rockv  surfaces  with  dominant  kelp  (Eklonia 
radiata)  and  on  sand  at  the  channel  floors,  the  undes- 
cribed  species  typically  lives  on  the  channel  walls 
amongst  abundant  sponges  and  brachiopods.  The  pref- 
erence for  vertical  sublittoral  rock  surfaces  would  ac- 
count for  its  extreme  raritv-  on  beaches  and  in  dredge 
and  trawl  samples.  The  following  revision  was  initiated 
after  recent  discovery  of  well  preserved  juvenile  speci- 
mens (hitherto  unavailable  or  unrecognised)  of  the  un- 
described  species,  which  show  additional  differences 
from  M.  octogonus  that  in  turn  suggest  a  closer  relation- 
ship with  the  southern  Australian  species  M.  umbilicatus 
(Tenison  Woods,  1876).  Abbreviations  and  text  conven- 


tions are:  AIM,  Auckland  Institute  and  Museum; 
BMNH,  The  Natural  History  Museum,  London;  CM, 
Canterbury^  Museum,  Christchurch;  IS,  Ian  Scott  collec- 
tion, Auckland:  K\VB,  Kevin  W.  Burch  collection,  Whan- 
garei (includes  the  outstanding  collection  formed  bv  our 
late  friend  Dave  Gibbs);  NMNZ,  Museum  of  New  Zea- 
land Te  Papa  Tongarewa,  Weflington;  NZGS,  Institute 
of  Geological  and  Nuclear  Sciences,  Lower  Hutt.  Unless 
specified,  all  material  is  at  NMNZ  (registration  numbers 
preceded  bv  "M.").  Length  dimension  precedes  width  in 
all  cited  measurements. 

SYSTEMATICS 

Superfamilv  Muricoidea  Rafinesque,  1815 

Familv  Muricidae  Rafinesque,  1815 

SubfamiK  Muricidae  Rafinesque,  1815 

Genus  Muricopsis  Bucquoy,  Dautzenberg  and  Dollfus, 

1882 

Muricopsis  Bucquoy,  Dautzenberg  and  Dollfus,  1882:  19.  Type 
species  (by  original  designation):  Murcx  hlainviUii  Pa\Tau- 
deau,  1826:  Recent,  Mediterranean. 

Subgenus  Murcxsul  Iredale,  1915 

Murcxsul  Iredale,  1915:  471.  Tvpe  species  (by  original  desig- 
nation): Murex  octogonus  Quoy  and  Gainiard,  1833;  Re- 
cent, New  Zealand. 

Remarks:  Ponder  (1972)  considered  Murcxsul  to  be  a 
s\iion\Tn  of  Muricopsis.  though  Radwin  and  D'Attilio 
(1976),  Yokes  (1970,  1988)  and  Beu  and  Max-wefl  (1990) 
have  treated  them  as  cbstinct  genera.  We  agree  with  Vo- 
kes  (1988)  that  they  are  closelv  related,  but  Uke  Houart 
(1988,  1991,  1993),  prefer  to  interpret  Murcxsul  as  a 
subgenus  oi  Muricopsis. 

Muricopsis  (Murcxsul)  octogonus  (Quov  and  Gainiard,  1833) 
(Figures  1-15,  24,  25,  .33,  .37) 

Murcx  octogonus  Quov  and  Giiiniard,  1833:  531,  pi.  36,  fig.  8, 
9;  Kiener,  1843:  64,  pi.  15,  fig.  2;  Grav,  1843:  229;  Reeve, 
1845,  pi.  29,  fig.  1.34;  Kiister  and  Kobelt,  1869:  79,  pi.  28, 


B.  A.  Marshall  and  K.  \X.  Burch,  2000 


Page  19 


fig.  23;  Hutton,  1880:  47;  Poirk-r,  1883;  112;  Snter.  1901: 

61. 
Mnrex  penividiius  Sowerln,  1841a:  8,  lit;.  103;  Sovverhw  1841b: 

147;  \bkes,  1970;  .327.  Not  Miinx  pinniiiuiis  Lamarck. 

1816  (preoccupied). 
Miircx  (Phi/Uonotus)  octo^i^oiiiis. — Tr\on.  1880;  110,  pi.  .30.  fig. 

272.  273  onK'  I,  in  purt — fig.  274  =  Muricop.si.s  aispidatiis 

(Sowerbv,  1879));  Hutton,  1884;  218. 
Miirex  clipsactis. —Tnon.  1880;  110,  pi.  .30,  fig.  281  onl\-.  Not 

Broderip,  18.33. ' 
Miirex  octogoniis  var  iimhilicata. — Suter  1901;  61.  Not  Tcni- 

son  Woods,  1876. 
Murex  {Miirii-antlui)  octofionii.s. — Suter,  1913:  400.  pi,  48,  fig. 

1. 
Murc'x  (Miiricantha)  octog,oiiu.s  var  iiiiihilirdtii.s. — Suter  1913: 

401.  Not  Tenison  Woods,  1876. 
Hcxiifilcx  (Miircxsul)  octogonii.s. — Iredale,    1915;  471;  Wenz, 

1941;  1090,  text  fig.  3096. 
Hexaplcx  (Murexsul)  octogonus  var  umbilicntns. — Iredale, 

191.5;  471.  Not  Tenison  Woods,  1876. 
Murexsul  octogonus. — Finlav,  1926;  419;  Yokes,  1964;  13,  fig. 

20,  .50,  61;  Ponder  1968;  31.  fig.  1,  37-41;  Yokes,  1970; 

327;  Fiiir  1976;  63,  pi,  17.  fig.  239;  Radwin  and  DAttilio. 

1976;  163,  text  fig.  104,  pi.  26,  fig,  6,  7;  Abbott  and  Dance, 

1983;  143,  te.xt  fig.;  Scott.  1989;  6,  text  figs.;  Ben  and  Max- 
well. 1990:  .359,  pi.  481. 
Murexsul  cuvierensis  Finlav,  1926;  419  =  noiiun  nudum. 
Murexsul  cuvierensis  Finlav,  1927:  487,  pi.  24,  fig.  2. 
Murex  (Murex.'iul)  octogonus. — Thiele,  1929;  290;  Shikama  and 

Horikoshi,  1963;  69,  te.xt  fig.  104. 
Murex  {Murex.sul)  ednae  Smitli,"l940;  43,  pi.  2,  fig.  10. 
Muricopsis  octogonus  octogonus. — Ponder  1972:  237,  te.xt  fig. 

.3/24. 
Murcx.sut  octagunus  [sic]. — Kaicher  1978,  card  1608. 
Muricopsis  octogonus. — Powell,  1979;  170,  pi.  .35,  fig.  1;  Scott, 

1989;  6,  text  figs.;  Hart,  1993;  44,  text  fig.  (in  part— far 

right  figure  =  holotvpe  of  Af .  scotti  new  species). 

Type  material:  M.  octogonus:  syntypes  (2),  including 
the  originally  figured  specimen  (Figure  15)  MNHN,  Bay 
of  Islands,  New  Zealand;  M.  pcniiianus:  repositors'  of 
txpe  material  unknown  (not  located  at  BMNH.  K.M. 
Way,  pers.  comm.),  "Pacosma\o,  Peru"  =  New  Zealand; 
M.  cuvierensis:  holotxpe  AIM  70500,  off  Cuvier  Island, 
73  m;  M.  ednae:  repositorx-  of  txpe  material  unhiown, 
"Japan"  =  New  Zealand. 

Other  material  e.xamined:  About  1000  specimens  in 
NMNZ  (152  lots)  and  K.W.  Burch  (24  lots)  collections. 

Di.stribution  (Figure  37):  Pleist(K'ene  (Castlecliffian) 
to  Recent.  Three  Kings  Islands  and  North  Island,  New 
Zealand,  as  far  south  as  Kapiti  Island  (west  coast)  and 
Mahia  Peninsula  (east  coast),  0-508  m;  taken  alive  at  low 
tide  to  121  m. 

Remark.s:  Muricopsis  octogoitus  is  exceptionally  vari- 
able in  teleoconch  moq'jhologx',  and  there  is  complete 
intergradation  between  the  most  extreme  of  the  forms 
illustrated  here  (Figures  1-15).  Despite  great  variation 
in  sculpture  of  the  last  few  whorls  in  adults,  all  of  the 
specimens  are  identical  in  protoconch  and  earlv  teleo- 
conch morphologx'.  Moreover,  we  could  detect  no  cbf- 
ferences  between  the  most  extreme  forms  in  radular 


morphology  or  e.xternal  anatomy.  It  thus  seems  ilear  that 
all  specimens  under  consideration  here  represent  a  sin- 
gle higliK  variable  .species. 

Mature  specimens  range  from  29  to  92  (est.)  mm  in 
length.  Spines  may  be  entirely  absent,  or  short  to  long. 
The  secondary-  spiral  cords  ma\-  remain  weaker  than  the 
primaries  and  spineless  throughout,  or  the%  ma\-  enlarge 
to  resemble  the  primaries  before  the  last  adult  whorl, 
and  some  or  all  may  develop  spines  that  may  be  as  long 
as  those  on  the  primaries.  The  siphonal  canal  may  be 
short  and  broad  and  the  fasciole  rounded  with  low  ridg- 
es, or  the  canal  ma\'  be  long  and  narrow,  and  the  fasciole 
set  with  canals  from  earlier  stages  of  growth  that  encircle 
a  narrow  to  rather  large  false  umbilicus.  The  teleoconch 
may  be  reddisli  brown  with  black  spiral  cords,  or  white 
or  cream  with  vellovvish  or  (txpicalK )  reddish  brown  spi- 
ral cords.  Shells  with  dark,  extensive  pigmentation  pre- 
dominate in  the  littoral  and  locally  to  about  12  m.  Spec- 
imens with  short  spines,  and  with  secondary'  spirals  that 
enlarge  to  resemble  the  primaries  predominate  in  the 
littoral  and  localK'  to  about  50  m,  whereas  long-spined 
shells  on  which  the  secondarv  spirals  remain  weaker 
than  the  primaries  txpically  occur  deeper  than  about  30 
m,  thfjugh  localK-  as  shallow  as  about  20  m.  The  outer 
lip  of  mature  specimens  ma\'  be  smooth  or  (hpicalK  ) 
dentate,  and  the  inner  lip  rim  ma\'  be  free  and  projecting 
below  a  narrow  parietal  area,  or  almost  fully  adlierent. 
Unusually  large,  entirely  spineless  shells  occur  off  Spirits 
Bax',  where  thev'  intergrade  with  short-spined  specimens 
(Figures  9,  II). 

Muricopsis  octogonus  has  been  observed  on  a  number 
of  occasions  on  sandy  substrata,  boring  holes  in  and 
feeding  upon  the  shallowK'-burrowing  bivalve  Tawera 
spissa  (Desha\es,  18.35)  (I.  Scott,  pers.  comm.;  K.W.B., 
pers.  obs.).  What  it  eats  on  rock-\-  ground  remains  to  be 
established.  The  radula  is  illustrated  (Figure  33)  for 
comparison  with  those  of  M.  mariac,  M.  scotti  and  M. 
unibilicatus  (Figures  34-36). 

There  are  no  records  of  M.  octogonus  from  the  west 
coast  of  the  North  Island  between  Cape  Maria  van  Die- 
men  and  Cape  Egmont,  or  from  the  east  coast  south  of 
Mahia  Peninsula  (Figure  37),  and  it  would  seem  that  the 
present  distribution  off  the  northeastern  and  southwest- 
em  North  Island  is  relictual,  perhaps  dating  from  prior 
to  the  last  glacial  maximum. 

How  far  back  this  .species  extends  in  the  fossil  record 
is  unclear.  Specimens  from  a  Late  Pliocene  (Nukuma- 
ruan)  horizon  near  Waipukurau  (GS 10858,  former  cut- 
ting on  disused  section  of  Ashcott  Road,  coll.  A.C.  Beu, 
NZGS)  appear  to  be  indistinguishable  from  Recent  spec- 
imens in  teleoconch  moqihology  but  differ  in  having 
smoothly  rountleil  instead  of  angulate  protoconchs  as  in 
Recent  material  (Figure  30)  (see  below).  If  it  proves  to 
be  specifically  distinct  from  M.  octogonus,  it  mav-  be  as- 
signable to  M.  dihtcidus  Marwick,  1931  (Early  Pliocene, 
Gisborne  District).  Some  of  the  New  Zealand  Cenozoic 
ta.xa  are  scarceK'  distinguishable  from  M.  octogonus  or 
from  each  other,  and  it  is  likely  diat  M.  octogonus  is  a 
direct  descendent  of  one  or  other  of  them,  such  as  M. 


Page  20 


THE  NAUTILUS.  Vol.  114,  No.  1 


B.  A.  Marshall  and  K.  W.  Burch,  2000 


Pa^e  21 


proavitiis  (Laws,  1935),  M.  proi^enitor  (Laws,  1935),  or 
M.  manvicki  (Max^A'ell,  1971),  all  from  Middle  Miocene 
(Lilbumian)  beds. 

Muricopsis  (Murexsul)  mariae  (Finlav,  1930) 
(Figures  17,  19,  27,  28,  34,  38) 

Murexsul  inariac  Finlay,  1930:  237;  Kaiclicr,  197S,  card  1639; 

Ben  and  Maxwell,'  1990:  359. 
Murexsul  cspinosus  mariae. — Ponder,  1968:  32,  fig.  2,  32,  33, 

.34. 
Muricopsis  espinosus  marine. — Powell,  1979:  170,  pi.  37.  fig. 

5;  Scott,  1989:  6,  text  figs. 

Ty-pe  material:  HolotApe  AIM  70502,  Cape  Maria  van 
Dienien,  northern  New  Zealand. 

Other  material  examined:  Several  hundred  speci- 
mens in  NMNZ  (66  lots)  and  K.W.  Burch  (13  lots)  col- 
lections. 

Distribution  (Figure  38):  Three  Kings  Islands  and 
northeastern  North  Island,  New  Zealand,  as  far  south  as 
Anaura  Bav,  0-233  m;  taken  living  under  rocks  at  low 
tide  to  40  ni. 

Remarks:  Muricopsis  mariae  differs  principally  from 
M.  octogonus  in  consistentK-  lacking  spines,  in  being 
smaller  relative  to  the  number  of  whorls,  in  having  stron- 
ger denticles  within  the  adult  outer  lip,  in  attaining 
smaller  size  (ma.\imum  length  29  mm  as  against  ca.  9.3 
mm),  in  being  bluish  white  instead  of  tvpicalh'  white 
within  the  aperture,  and  in  tvpicalh'  lacking  a  distinct 
shoulder  angulation,  at  least  on  the  early  teleoconch 
whorls  (some  specimens  have  a  distinct  shoulder  angu- 
lation on  later  whorls).  In  a  sample  of  both  species  taken 
living  together  at  25  m  in  Whangaroa  Harbour  entrance 
(M. 1:37244,  M. 74899),  the  protoconch  ranges  from  800- 
1000  |j.m  wide  (mean  =  900  ixnx  SD  =  6.053,  n  =  21) 
in  M.  inariac,  and  650-1050  |jLm  wide  (mean  =  930  |jLm, 
SD  =  0.097,  n  =  17)  in  M.  octogonus.  The  largest  M. 
mariae  protoconch  seen  is  1250  |jLni  wide  (M. 133712). 
The  first  whorl  of  the  protoconch  has  a  distinct  angula- 
tion and  the  summit  is  more  or  less  flattened  in  both 
species.  Juveniles  of  A/,  mariae  and  M,  octogonus  mav 
be  difficult  to  distinguish,  but  in  M  mariae  the  primarv- 
spiral  cords  are  stronger  after  the  second  teleoconch 
whorl,  the  secondary  spirals  enlarge  more  rapidly  to  re- 
semble the  primaries,  and  the  Secondaries  on  the  sutural 
ramp  become  pigmented  early  on  the  third  whorl  in- 
stead of  after  the  third  or  fourth  whorl  (compare  Figures 
24,  25  with  27,  28).  Specimens  from  Cape  Maria  van 


Diemen  to  North  Cape  and  off  the  Three  Kings  Islands 
are  more  broadly  conical  than  most  specimens  from  fur- 
ther south  (Figures  17,  19)  but  are  otherwise  indistin- 
guishable. 

The  diet  of  this  species  is  unknown.  The  distinctive 
radiila  is  illustrated  (Figure  34)  for  comparison  with  that 
of  A/,  octogonus,  M.  scotti  and  M.  timhilicatus  (F'igures 
.33,  35,  .36). 

Ponder  (1968)  interpreted  M.  mariae  as  a  chronosub- 
species  of  A/,  espinosus  (Hutton,  1886),  though  Beu  and 
Maxwell  (1990)  suggested  that  they  might  well  be  dis- 
tinct species.  Compared  with  15  well  preserved  .speci- 
mens of  M.  espinosus  obtained  near  Waipukurau 
(GS10858,  map  ref.  U22/085.308,  former  cutting  on  dis- 
used section  of  Ashcott  Road,  WNW  of  Waipukurau, 
A.G.  Beu,  NZCS — lowest  Nukumaruan,  Late  Pliocene), 
which  are  indistinguishable  from  the  hpe  material  ob- 
tained nearln',  M.  mariae  differs  in  having  a  protoconch 
that  is  shorter  relative  to  its  width  with  a  shouldered 
(rarely  rounded)  instead  of  rounded,  more  or  less  bul- 
bous first  whorl  (Figure  27),  and  in  having  narrower  spi- 
ral interspaces,  especially  on  the  first  three  or  four  te- 
leoconch whorls.  In  most  specimens  of  M.  mariae  the 
secondary  spiral  cords  enlarge  to  resemble  the  primaries 
more  rapidlv  than  in  M.  espinosus.  InterestingK-  enough, 
the  sample  includes  well-preserved  specimens  of  a  larger 
species  that  also  lacks  a  shoulder  angulation  on  the  pro- 
toconch (Figure  30).  The  status  of  this  larger  fossil  spe- 
cies is  unclear,  but  it  is  certainly  more  closely  related  to 
M.  octogonus  than  to  M.  espinosus.  Muricopsis  espinosus 
or  a  very  similar  species  occurs  in  the  bath\a!  "coral 
thicket".  Lake  Ferry,  Palliser  Bay  (Mangapanian:  Late 
Phocene)  (Figure  31).  The  protoconch  has  thus  inde- 
pendently become  angulate  in  the  M.  octogonus  and  M. 
mariae  lineages  since  the  Pliocene.  The  differences  be- 
tween M.  espinosus  and  M.  mariae  in  protoconch  mor- 
phology and  similarities  in  teleoconch  morphology  sug- 
gest that  thev  are  distinct  species. 

The  original  description  o(  Murcx  espinosus  (Hutton, 
1886)  is  based  on  more  than  one  specimen,  the  larger 
of  which  provided  the  cited  dimensions  (30.0  X  14.5  mm, 
CM3294),  the  smaller  of  which  was  subsecjuently  illus- 
trated bv-  Hutton  (1893)  (28.0X13.3  mm,  CM3295).  Al- 
though these  specimens  each  have  two  labels  (none  orig- 
inal) staring  that  they  are  "holotype '  and  "paratype"  re- 
spectively, they  are  in  fact  synt\pes  because  Hutton  did 
not  use  these  unecjuivocal  terms.  Ponder  (1968,  figs  35, 
36)  effectively  designated  the  larger  specimen  as  lecto- 
type  by  stating  that  it  was  the  "holotype".  Incidentally, 


Figures  1-12.  Shells  of  adult  Muricopsis  (Murexsul)  octogonus  (Quoy  and  Gainiard).  1.  Off  Thompson  Point,  Wailieke  Island, 
22  m,  KWB  (length  .54  mm).  2.  Off  Cuvier  Island,  70  m,  M..5911  (length  78.5  mm).  3.  Off  Motiikalian  Island,  W  of  Paparoha, 
Coramandel,  18-20  m,  KWB  (length  45.5  mm).  4.  Scallop  bed  near  Matiatia  Bav  entrance,  Waiheke  Island,  13  m,  K\^'B  (length 
48  mm).  5.  Off  Three  Kings  Islands,  craxpot,  KWB  (length  90  mm,  est.  93  mm).  6.  Raiifurlv  Bank.  East  Cape,  31-47  m,  M. 75079 
(length  46  mm).  7.  Off  Mavor  Island,  .59-74  m,  M.64825  (length  63.5  mm).  8.  Off  Motnhoropapa  Island,  The  Noises,  25  m,  K'WB 
(length  42.5  mm).  9.  Off  Spirits  Bav,  48  m,  M.  137051  (length  54  mm).  10.  Hooper  Point,  Spirits  Bav  12  m,  M.  1.346.30  (length  29 
mm).  11.  Off  Tom  Bowling  Bay,  49  m,  M. 137090  (length  64  mm).  12.  Reotalii,  Whangarei,  low  tide,  KWB  (length  45.5  mm). 


Page  22 


THE  NAUTILUS,  Vol.  114,  No.  1 


Figures  13-23.  Shells  of  Muricopsis  species  {all  specimens  adult  except  IS).  13-15.  Muricopsis  (Murexsul)  octogonits  (Quoy  and 
Gaimard).  13.  Ranfurlv  Bank.  Ea.st  Cape,  89-94  m,  M.64812  (length  46  mm).  14.  Whangaroa  Harbour  entrance^  2,5  m,  M. 74899 
(length  .38  mm).  15.  originallv  figured  s\iit\pe,  MNHN  (length  .3.5. .5  mm).  16,  23.  Murexsul  (Muricop.sis)  profunda  new  species, 
holohpe,  off  Three  Kings  Islands,  91  m,  M. 70356.  17,  19.  Muricopsis  (Murexsul)  marine  (Finlav).  17.  Archvvav  Island.  Three  Kings 
Islands,  15  m.  M. 117096  (22.3X12.3  mm).  19.  Whangaroa  Harbour  entrance.  25  m.  M. 137244  (26x13  mm).  18,  20.  Murexsul 
(Rolniicliella)  umhilicatus  (Tenison  Woods),  Gulf  St.  Vincent,  South  Australia,  South  Australian  Museiun  (Figure  20,  length  .32  mm). 
21,  22.  Murexsul  (Rolandiella)  scotti  new  species.  21.  Holotvpe,  Mathesons  Bay,  Leigh,  6-9  m,  M. 138186  (length  44  mm).  22. 
Paratype,  Southeast  Bay,  Great  Island,  Three  Kings  Islands,  18  m,  KWB  (length  53.5  mm).  Scale  Una  =  1  mm. 


B.  A.  Marshall  and  K.  W.  Burcli,  2()()() 


Page  23 


Figures  24-32.  Early  whorls  of  Miiiicopsi.s  species.  24,  25.  Muiicop.sK  {Miircxsiil)  ociof^oiius  (Qiio\'  aiid  Gainiard),  VVhangaroa 
Harlioiir  entrance,  25  m,  M. 74899  (shell  length  7.05  mm).  26.  Miiirx.stil  (Roland icllti)  scotti  new  species,  submarine  cave  S  side  of 
Ro.seman'  Rock,  Princes  Islands,  Three  Kings  Islands,  20  m,  M. 117084  (length  6  nnn).  27,  28.  Muricopsi.s  (Murcx.sul)  niariac  (Finlay), 
Whangaroa  Harbour  entrance,  25  m,  M.  1.37244  (shell  length  4.75  mm).  29,  32.  A/i/rico/i.v/.v  (Miircx.wl)  profunda  new  species,  early 
whorls  (29)  and  teleoconch  (left)  and  protoconcli  (right)  microsculpture  (32),  NW  ol  Great  Island,  Three  Kings  Islands,  310  m, 
M. 93876.  30.  Mttricopsis  (Mtirexsul)  sp.  aff  octogonu.s  (Quoy  and  Gaimard),  GSI0858,  Ashcott  Road,  VVaipuknrau,  shallow  water. 
Late  Pliocene  (lowest  Nukvnnanian),  NZGS  TM8078.  31.  Muricopsi.s  (Murexsul)  sp.  cf  espinosus  (Hutton),  "coral  thicket".  Lake  Ferry, 
Palliser  Bay,  bathyal.  Late  Pliocene  (Mangapanian),  M. 40408.  Scale  line  for  figure  32  =  50  |xm,  other  scale  lines  =  500  |jini. 


Page  24 


THE  NAUTILUS,  Vol.  114,  No.  1 


Figures  33-36.  Radulae  of  Muricopsis  species.  33.  Muricopsis  (Murexsul)  octogonus  (Quoy  and  Gaimard),  off  Te  Arai  Point, 
Northland,  41  ni,  M.  137251  (shell  length  65  mm).  34.  Muricopsis  (Murexsul)  mariae  (Finlay),  Governors  Pass,  Great  Barrier  Island, 
11-29  m,  M. 21784  (shell  length  22  mm).  35.  Muricopsis  (Rolandiella)  scotti  new  species,  off  Cape  Rodney,  11-13  m,  M. 137416 
(shell  length  40  mm).  36.  Muricopsis  (Rolandiella)  umbilicatus  (Tenison  Woods),  Edithburgh,  South  Australia:  South  Australian 
Museum,  Adelaide,  unregistered  (shell  length  28  mm).  Scale  lines  =  50  jxm. 


this  t)-pe  material  was  mislaid  imdl  recentK',  and  omitted 
from  the  catalogue  of  tspe  and  figured  fossils  in  the  Ge- 
ology Department  of  the  Canterbury  Museum  (Brad- 
shaw  et  al,  1992). 

Muricopsis  (Murexsul)  profunda  new  species 
(Figures  16,  23,  29,  39) 

Description:  Shell  up  to  10.5  mm  high  (holotype 
adult?),  fusiform,  of  moderate  thickness;  protoconch  and 
teleoconch  pale  orange,  adapical  4  primary  spirals  and 
secondary  spirals  between  them  reddish  brown  on  last 
whorl.  Protoconch  830-870  |xm  wide  excluding  flared 
rim,  of  1.5  convex  whorls,  covered  with  minute  hemi- 
spherical granules,  summit  of  first  whorl  angulate,  last 
half  whorl  rather  e\enly  convex.  Teleoconch  of  4.5  reg- 
ularly expanding  whorls;  first  whorl  broadly  convex;  sub- 
sequent whorls  with  pronounced  supramedian  angula- 
tion, sutural  ramp  broad  and  flat,  side  broadl\'  convex, 
smoothly  confluent  with  siphonal  canal.  Sculpture  con- 
sisting of  axial  varices,  and  rounded  primarv'  and  sec- 


ondary spiral  cords.  Axial  varices  strongly  retracted  from 
adapical  insertion  throughout,  12  on  last  whorl  (holo- 
tspe),  narrow  on  1st  whorl,  sloping  adaxialh  so  that  bases 
are  overhung,  blade  like  and  strongly  adapicalK-  pro- 
duced above  protoconcli/teleoconch  suture  and  cement- 
ed to  side  of  last  whorl  of  protoconch,  e.xtending  beyond 
adapical  extremitv  of  last  protoconch  whorl  or  to  about 
mid  whorl  length.  .Axial  varices  on  subsecjuent  whorls 
becoming  diicker  and  rounded,  each  sumiounted  by  4 
or  5  fine,  collabral,  ada.xially  sloping,  axial  lamellae.  Axial 
interspaces  with  fine  collabral  growth  lines  and  a  few 
lamellar  growth  lines.  Primarv  spiral  cords  numbering  3 
on  spire  and  3  on  base;  spire  spirals  commencing  ini- 
niediatelw  similar  throughout,  adapical  spiral  at  shoulder 
angulation,  abapical  spiral  bordering  suture;  adapical  2 
basal  spirals  similar  to  spire  spirals,  abapical  spiral  stron- 
ger, widely  separated.  Secondary  spirals  numbering  com- 
mencing on  3rd  whorl,  remaining  weaker  than  primaries, 
2  on  ramp,  1  between  each  priniar\-  spiral,  3  between 
abapical  basal  primaries,  and  1  between  abapical  basal 
primarv  and  tip  of  canal.  Siphonal  canal  of  moderate 


B.  A.  Marshall  and  K.  W.  Burch,  2000 


Page  25 


Figures  37.  Distributions  of  New  Zealand  Recent  Muricop- 
sis  {Miircxstd)  octogonus  (Quov  and  Gaimard)  (500  and  1000 
m  isobaths  indicated). 


Figure  38.  Distributions  of  New  Zealand  Recent  Muricopsis 
(Miircxsiil)  marine  (Finlay)  (500  and  1000  m  isobaths  indicat- 
ed). 


length,  gently  flexed,  partly  enclosed  by  thin  extension 
of  inner  lip.  Aperture  p\riform;  outer  lip  rather  thin  at 
rim,  weakly  thickened  within,  simple;  inner  lip  fully  ad- 
herent adapically,  rim  free  at  abapical  extremity  and  ex- 
tending almost  to  canal  tip.  Animal  unktiown. 

Type  material:  Holotype  NMNZ  M. 70356,  34°11'S, 
172°10'E,  off  Three  Kings  Islands,  northern  New  Zea- 
land, 91  m,  19  Februar\'  1974,  r.v.  Acheron.  Paratx^jes: 
33°59.2'S,  172°13.6'E,  18  km  N  of  Great  Island,  Three 
Kings  Islands,  155  m,  23  June  1978,  r.y.  Tangaroa  (1 
juvenile,  M. 137247);  34°06.5'S,  172°04.7'E,  llkm  NW 
of  Great  Island,  310  m,  30  June  1978,  r.v.  Tam^awa  (1 
juvenile,  M.93876). 

Other  material  examined:  34°009'S,  171°44  7'E, 
Middlesex  Bank,  NW  of  Three  Kings  Islands,  201-216 
m,  31  January  1981,  r.v.  Tan^aroa  (2  juveniles, 
M.93302);  34°02.0'S,  171°44.0'E,  Middlesex  Bank,  246- 
291  m,  31  January  1981,  r.v.  Tan^aroa  (3  juveniles, 
M. 137245);  .34°05.9'S,  171°55.1'E,  24  km  NW  of  Great 
Island,  Three  Kings  Islands,  710  m,  27  June  1978,  r.v. 
Tangaroa  (4  juveniles,  M. 137246);  34°17.6'S, 
17r45.3'E,  39  km  SW  of  Great  Island,  427  m,  21  June 
1978,  r.v  Tangaroa  (3  juveniles,  M. 94338). 

Distribution:  Off  Three  Kings  Islands,  northern  New 
Zealand,  91-710  m  (shells  only). 

Remarks:  Muricopsis  profunda  differs  from  M.  octo- 
goniis  and  M.  mariac  principalK'  in  having  ;Lxial  varices 
that  ride  up  onto  the  protoconch  from  the  next  whorl, 
in  that  the  outer  lip  is  much  more  strongly  retracted 


from  the  suture,  and  in  that  the  spiral  cords  are  stronger 
at  equivalent  stages  of  growth  and  become  pigmented 
at  a  later  stage  of  growth.  The  size  attained  by  this  spe- 
cies is  unclear,  though  it  may  not  grow  much  larger  than 
the  holotvj^e  (length  10.5  mm). 

Etymology:     Deep  (Latin). 

Rolandiella  new  subgenus  (of  Muricopsis) 

Type  species:  Muricopsis  (Rolandiella)  scotti  new 
species;  Recent,  northern  New  Zealand. 

Diagnosis:  Sliell  similar  to  those  of  species  of  Muri- 
copsis (Murexsul),  but  with  median  shoulder  angulation 
and  weak  abapical  2  primary  spiral  cords  on  first  1.5 
teleoconch  whorls.  Inner  Lip  of  most  specimens  upstand- 
ing over  most  of  its  length. 

Description:  Shell  21-62  mm  high  at  maturity,  spire 
about  as  high  as  last  adult  whorl  or  slightly  shorter,  stout, 
of  moderate  thickness.  Protoconch  of  1.5-2.0  rovmded 
whorls.  Teleoconch  of  mature  specimens  widi  5.5-7.0 
convex,  shouldered  whorls,  shoulder  median  on  first  2 
whorls,  ascending  to  adapical  third  or  slightly  higher  on 
subsequent  whorls;  sutural  ramp  shallowly  concave;  side 
broadly  convex,  smoothly  confluent  with  siphonal  canal. 
Primary-  sculpture  consisting  of  strong,  axial  varices  and 
spiral  cords  with  spines  at  intersections.  Varices  travers- 
ing all  whorls,  angulate  in  section,  numbering  8-11  (7 
or  12  in  occasional  specimens)  on  adult  penultimate 
whorl.  Primary  spiral  cords  rounded,  numbering  3  on 
spire  and  3  on  base  in  adults.  Adapical  primary  spiral  at 


Page  26 


THE  NAUTILUS,  Vol.  114,  No.  1 


shoulder  angulation,  peripheral;  abapical  primary  bor- 
dering suture  throughout,  or  separating  as  insertion  of 
last  few  whorls  descends  to  ne.xt  priniarv  spiral:  adapical 
and  abapical  primaries  commencing  immediately;  inter- 
mediate primary*  commencing  at  about  mid  first  whorl, 
gradually  enlarging  to  resemble  other  primaries.  Abapi- 
cal basal  primary  spiral  wideK'  separated  from  adjacent 
primary,  interspaces  ot  other  primar)'  spirals  closer  and 
of  similar  width  to  each  other.  Secondary'  spirals  pro- 
gressively developing,  1-5  on  sutural  ramp,  1  each  in 
some  or  all  interspaces  of  primaiT  spirals,  0-.3  betyveen 
abapical  basal  priman'  spiral  and  tip  of  canal,  others  on 
summits  of  some  or  all  primary-  spirals,  1  or  more  en- 
larging to  resemble  primaries  on  last  adult  whorl.  Pri- 
mar\'  and  1  (abapical  shoulder  spiral)  or  more  secondarv 
spirals  procKiced  as  short  to  moderate-length  spines  at 
varices,  spines  on  primar\'  spirals  of  similar  length  or 
shoulder  spiral  longest.  Secondary  axial  sculpture  com- 
prising fine  collabral  growdi  lines  that  are  raised  as  thin, 
fragile  lamellae  on  ramp  against  suture  and  where  in- 
tersecting spiral  sculpture.  Aperture  pvriform.  Outer  lip 
thickened  within  at  maturity,  strongh'  so  in  particularly 
large  specimens,  which  may  also  develop  spirally  elon- 
gate denticles.  Inner  lip  thin,  fulK*  adherent  over  adap- 
ical third  or  quarter,  abapical  part  typically  free,  flared 
and  strongK'  projecting,  ajjapical  extremity  e.xtenchng 
well  beyond  flexure  of  siphonal  canal.  Siphonal  canal  of 
moderate  length,  almost  enclosed  by  infolding:  siphonal 
fasciole  strong,  forming  an  umbilicus  in  some  ver\'  large 
specimens.  Radula  and  operculum  similar  to  those  in 
Miiricop.sis  (s.  str)  and  Murcxsul. 

Remarks:  Rolandiclla  is  introduced  for  Muiicopsis 
scotti  new  species  from  northern  New  Zealand,  and  M. 
umbilicatiis  (Tenison  Woods,  1876)  from  southern  Aus- 
tralia, both  of  which  resemble  typical  Murcxsul  species 
in  gross  facies,  but  differ  in  that  the  shoulder  spiral  is 
situated  medially  on  the  early  spire  whorls  then  ascends 
adapicalK',  and  the  intermediate  primary  spiral  on  the 
spire  commences  later  than  the  others  and  develops 
slowly.  By  contrast,  in  Muiicopsis  species,  all  three  pri- 
maries are  strong  and  commence  immediately  on  die 
first  teleoconch  whorl,  and  the  shoulder  spiral  borders 
the  suture  then  descends  abapically  over  subsequent 
whorls. 

None  of  the  named  New  Zealand  Cenozoic  species 
referred  to  Murcxsul  by  Beu  and  Maxwell  (1990)  seem 
to  belong  in  Muiicopsis  (Rolandiclla).  though  the  rela- 
tionships of  the  Early  Miocene  species  Muiicopsis  cchin- 
ophorus  (Powell  and  Bartrum,  1929)  are  unclear  be- 
cause the  early  teleoconch  morphology  is  unknown. 

Both  M.  (Rolaiuliclla)  scotti  and  M.  (R.)  uinbilicatus 
have  large,  rounded,  paucispiral  protoconchs  indicating 
non-planktotrophic  development,  and  suggesting  that 
they  have  either  crawl-away  larvae  or  drifting  larval  stag- 
es of  short  duration.  We  suppose  that  the\'  are  derived 
from  a  common  ancestor  with  planktotrophic  larval  de- 
velopment that  had  a  trans-Tasman  distribution.  How  far 
back  in  time  diis  putative  common  ancestor  may  have 


@      . 


Figure  39.     Distributions  of  New  Zealand  Recent  Muricopsis 

(MtirexstiJ)  profunda  new  species  (circle)  and  Muricopsis  {Ro- 
landicUa)  scotti  new  species  (squares)  (500  and  1000  m  iso- 
baths indicated). 


lived  is  impossible  to  estimate  without  fossil  records  or 
Tiiolecular  data. 

Etymology:  After  Roland  Houart,  Belgium,  in  appre- 
ciation of  his  outstanding  contribution  to  world  muricid 
systematics. 

Muricopsis  (Rolandiclla)  scotti  new  species 
(Figures  21,  22,  26,  35.  39) 

Muricopsis  species  Scott,  1989:  6,  text  figs. 
Muricopsis  octogonus. — Hart,    1993:   4.5,   right  text  fig.   onlv 
(  =  holotype).  Not  Qnov  and  Gainiard,  18.3.3. 

Description:  Shell  up  to  62  mm  high,  fusiform,  spire 
about  as  high  as  last  adult  whorl  or  slightK'  shorter,  stout, 
of  moderate  diickness.  Protoconch  and  first  1  or  2  te- 
leoconch whorls  white,  subsequent  whorls  white  or  pale 
buff,  primary,  secondary  and  some  finer  spiral  cords  red- 
dish brown,  aperture  porcellaneous  white.  Protoconch 
sharply  delineated  b\'  low  varix,  of  1.5-2.0  smooth,  con- 
vex whorls.  Teleoconch  of  up  to  7  convex,  shouldered 
whorls,  shoulder  median  on  first  2  whorls,  ascending  to 
adapical  third  or  slightly  higher  on  subsequent  whorls; 
sutural  ramp  shallowK'  conca\e:  side  broadk'  convex, 
smoothly  confluent  with  siphonal  canal.  Primary  sculp- 
ture consisting  of  strong,  axial  varices  and  spiral  cords 
with  spines  at  intersections.  Varices  traversing  all  whorls, 
angulate  in  section,  numbering  8-11  (7  or  12  in  occa- 
sional specimens)  on  adult  penultimate  whorl.  Primary 
spiral  cords  rounded,  numbering  3  on  spire  and  3  on 
base  in  adults.  Adapical  primary  spiral  at  shoulder  an- 


B.  A.  Marshall  and  K.  W.  Burch,  2000 


Page  27 


gulatioii,  peripheral;  ahapical  spire  spiral  bordering  su- 
ture at  first,  separating  as  insertion  oi  last  few  whorls 
descends  to  next  (iornierK'  adapieal  basal)  priinar\' spiral; 
adapical  and  abapical  spire  spirals  commencing  imme- 
diatek":  intermediate  spire  spiriil  commencing  at  about 
mid  first  whorl,  slowlv  enlarging  to  resemble  abapical 
spire  spiral  and  basal  primary-  spirals.  Abapical  basal  pri- 
mar\  spiral  wideK  separated  from  adjacent  primarw  in- 
terspaces of  other  primars'  spirals  closer  and  of  similar 
width  to  each  other.  Secondar\'  spirals  appearing  pro- 
gressively, numbering  3-5  on  sutural  ramp,  5-7  in  zone 
comprising  summit  of  shoulder  spiral  space  behveen  it 
and  adjacent  primars,  usualK  3  or  4  on  sununits  of  other 
primary  spire  spirals,  3  or  4  between  abapical  2  basal 
primaries,  3  between  abapical  basal  priman*  spiral  and 
tip  of  canal,  and  generally  1  each  in  some  or  all  inter- 
spaces of  primar\'  spirals,  most  or  all  secondaries  re- 
maining weaker  than  primaries  throughout,  or  some  en- 
larging to  resemble  primaries  on  last  adult  whorl.  Pri- 
man  and  some  secondars'  spirals  produced  as  short  to 
moderate-length  spines  at  varices,  shoulder  spines  lon- 
ger than  others  on  spire,  spines  on  larger  spirals  tspicalK 
of  similar  length  on  last  adult  whorl.  Secondar\'  axial 
sculpture  comprising  fine  collabral  growth  lines  that  are 
raised  as  thin,  fragile  lamellae  on  ramp  against  suture 
and  where  intersecting  spiral  sculpture.  Aperture  p\Ti- 
form.  Outer  lip  thickened  within  at  maturity,  strongly  so 
in  particularK'  large  specimens,  which  ma\'  also  develop 
spiralK'  elongate  denticles.  Inner  lip  thin,  fidh"  adherent 
over  adapical  third  or  quarter,  abapical  part  hpicalK' 
free,  flared  and  strongly  projecting,  abapical  extremits* 
extending  well  beyond  flexure  of  siphonal  canal.  Siphon- 
al  canal  of  moderate  length,  almost  enclosed  bv  infold- 
ing; siphonal  fasciole  strong,  forming  an  umbilicus  in 
some  vers'  large  specimens.  Operculum  with  terminal 
nucleus,  yellowish  brown.  Radula  (figure  35):  Central 
teeth  each  with  strong,  conical  central  cusp;  smaller, 
conical  lateral  and  marginal  cusps,  and  2  \er\'  small  inner 
lateral  denticles  that  are  well  separated  from  lateral 
cusp.  Lateral  teeth  narrow. 

Type  material:  Holotype  NMNZ  M. 138186,  Mathe- 
sons  Bay,  Leigh,  northern  New  Zealand,  6-9  m,  Feb- 
niar\-March  1992,  D.W.  Gibbs  (ex  D.W.  Gibbs  and 
K.W.  Burch  collections).  Paratvpes:  34°09.5'S, 
172°08.8'E,  Southeast  Bay,  Great  Island,  Three  Kings 
Islands,  20-22  m,  4  March  1997,  K.W.  Burch  and  D.D. 
Crosby  (2  juveniles,  M. 134760);  Southeast  Bay,  alive,  18 
m,  ISMay  1982,  K.W.  Burch  (6,  KWB);  Princes  Islands, 
Three  Kings  Islands,  15  m,  29  November  1983,  G.S. 
Hardy  and  A.L.  Stewart  (1,  M. 75135);  S  side  of  South- 
west Island,  Three  Kings  Islands,  ;ilive,  27  m,  12  Feb- 
nian'  1986,  G.S.  Hardv  (2,  M.84267):  off  Cape  Maria 
van  Dienien,  craypot.  May  1991  (1,  KWB);  Whangaroa 
Harbour  entrance,  alive,  18  m,  November  1995,  K.W. 
Burch  and  D.W  Gibbs  (2,  KWB);  Rikoriko  Cave,  Poor 
Knights  Islands,  alive,  30-40  m,  1982,  M.II.B.  ONeill 
(1,  'm.75181);  Poor  Knights  Islands,  alive.  May  1979, 
A.S.W.  Penniket  (1,  M.  133691);  Poor  Knights  Islands, 


December  1980,  K.W.  Burch  (1,  M. 133692);  Poor 
Knights  Islands,  alive,  15-20  m,  1983,  K.W.  Burch  (4, 
KWB);  Poor  Knights  Islands,  alive,  1970,  1972,  W.  Doak 
(5,  M.  133690);  South  Cave,  Poor  Knights  Islands,  alive, 
17  m,  21  May  1969,  A.N.  Baker  (1,'  M.23617);  South 
Harbour,  Poor  Knights  Islands,  tilive,  25  Mav  1969,  A.N. 
Baker  (3,  M.23618);  off  Cape  Karikari,  30m,  20  June 
1981  (1,  M. 133694);  Urtjuarts  Bav  Whangarei  Heads, 
alive,  March  1965  (1,  M. 133695);  "The  Canyon",  N  side 
of  Burgess  Island,  Mokohinau  Islands,  alive,  15-21  m, 
Januari-  1985,  D.W.  Gibbs  (3,  KWB);  Maori  Island, 
Leigh,' 22  Mav  1975,  A.S.W.  Penniket  (1,  M. 133695); 
NE  of  Mathesons  Bay,  Leigh,  alive,  18  m,  Mav  1995, 
D.W.  Gibbs  (1,  M.127040);'Mathesons  Bay,  alive,  6-10 
m,  Februars-March  1992,  D.W.  Gibbs  (6,'  KWB);  S  of 
entrance  to  Omaha  Cove,  Leigh,  20  ni,  December  1967, 
I.  Scott  (2,  IS);  off  Little  Barrier  Island,  alive,  craypot, 
T.  Riley  (2,  M.  133693);  Cape  Rodney  alive,  15  m,'june 
1987,  D.W.  Gibbs  (7,  M.90125);  Cape  Rodney,  alive,  15 
m,  4  March  1989,  I.  Scott  (12,  M. 117277);  Cape  Rodnev, 
alive,  11-13  m,  I.  Scott,  Mav  1989  (53,  M. 137416);  Cape 
Rodney  alive,  18  m,  June  1977  (1,  M.  133698);  Waika- 
wau  Bay,  Coramandel,  alive,  16  m,  April  1989,  D.W. 
Gibbs  (2,  KWB);  off  Needle  Island,  Mercury  Bay,  alive, 
16  m,  January  1985,  D.W.  Gibbs  (3,  KWB).' 

Other  material  examined:  S  side  of  Rosemar\-  Rock, 
Princes  Islands,  Three  Kings  Islands,  20  ni,  18  January 
1985,  FJ.  Brook  (5,  M. 117084);  off  West  Island,  Three 
Kings  Islands,  Elinfiamitc  wreck,  alive,  W  Doak  (1, 
M. 133696);  .34°2().0'S,  173°06.6'E,  N  of  North  Cape, 
163-168  m,  27  Januar\'  1981,  r.v.  Tonj^aroa  (1, 
M. 137225);  Rarawa  Reef'  Great  E.xliibition  Bay  crav- 
pots,  alive,  40  ni,  August  1989,  I.  McMillan  (2, 
M.  100382);  Rarawa  Reef  cra\pots,  alive,  73  m,  28  Sep- 
tember 1988,  I.  McMillan  (6,  M.95234);  Rarawa  Reef 
cra\pots,  alive.  40  m,  16  December  1987,  I.  McMillan 
(3,  M. 95315);  off  Cape  Karikari,  alive,  cra\pots,  1988,  I. 
McMillan  (1,  M.95322);  Henr\-  Island, '  Whangaruni, 
alive,  13  m,  5  December  1971,  \.N.  Baker  (1,  M.2592()); 
off  Hen  and  Chickens  Islands,  alive  (1,  M.2686;  1, 
M.83882);  Mathesons  Bay,  Leigh,  alive,  15-20  m,  26 
March  1995,  A.  Spurgeon  (2,  MT26998). 

Distribution  (Figure  39):  Three  Kings  Islands  and 
northeastern  North  Island,  New  Zealand  as  far  south  as 
Little  Barrier  Island,  13-168  m;  taken  alive  at  13-40  m 
from  rocky  groirnd. 

Remarks:  Miiricopsis  ■•icotti  is  most  similar  to  the 
southern  Australian  species  M.  nmbilicatus  (Figures  18, 
20,  26)  in  early  shell  ontogeny  and  gross  adult  shell  fa- 
cies  than  to  any  New  Zealand  Recent  Murcxsiil  species. 
Muricopsis  scotfi  differs  from  M.  nmbilicatus  in  a  num- 
ber of  details  including  larger  adult  size  (shell  length  34— 
62  mm  versus  21-30  mm),  stronger  and  more  numerous 
secondary-  spirals,  insertion  point  of  the  last  few  whorls 
desceruling  froirr  the  abapical  spire  spiral  to  the  next 
(formerly  adapical  basal)  primary  spiral,  and  abapical 
secondary  spiral  on  the  sutural  ramp  never  becoming  as 


Page  28 


THE  NAUTILUS,  Vol.  114,  No.  1 


large  as  tlie  shoulder  spiral.  The  radula  of  M.  scotti  is 
similar  to  those  of  M.  umhihcatus  and  M.  octogomis 
(Figures  3,  35,  36). 

Muricopsis  scotti  resembles  M.  octof^onus  in  size,  but 
is  readily  distinguishable  bv  the  following  characteristics: 
orange  brown  instead  of  dark  reddish  brown  operculum, 
rounded  instead  of  angulate  protoconch,  generally  larger 
aperture,  shorter  siphonal  canal,  and  much  stronger  sec- 
ondary spiral  sculpture  on  the  primary  spiral  cords,  es- 
pecially on  the  shoulder  spiral  and  between  it  and  the 
adjacent  primary.  Unlike  M.  octogonus,  which  has  ex- 
tremeK'  variable  sculpture,  M.  scotti  consistently  has  a 
distinct  shoulder  angulation,  angular  yarices  and  short 
spines  that  are  not  curved  backwards,  or  at  most  only 
slightK'  so.  In  M.  scotti  the  shoulder  spiral  (adapical  pri- 
man)  is  much  stronger  than  the  others  on  the  first  two 
teleoconch  whorls  and  occupies  a  median  position,  then 
gradualK'  ascends  to  about  the  adapical  third  on  subse- 
quent whorls  (Figure  26).  By  contrast,  on  the  first  1.5 
teleoconch  whorls  in  M.  ocfogoiius,  the  three  primary' 
spire  spirals  are  all  strong  and  similar,  and  the  shoulder 
spiral  commences  beside  the  suture  and  descends  to 
about  the  adapical  third,  remaining  there  or  at  the  adap- 
ical quarter  on  subsequent  whorls  (Figure  24).  Although 
the  rim  of  the  inner  lip  may  be  upstanding  or  more  or 
less  fulK'  adherent  in  both  species,  that  in  M.  scotti  is 
typically  free  and  more  strongly  upstanding  abapically, 
and  extends  abapicallv  further  beyond  the  angulation  at 
the  top  of  the  siphonal  canal. 

Shells  of  living  M.  scotti  are  invariably  more  or  less 
covered  widi  living  Br\'ozoa,  barnacles  and  other  epi- 
bionts,  often  with  the  addition  of  Waltonia  inconspiciia 
(Sowerb\',  1846)  (Brachiopoda),  upon  which  it  has  been 
observed  to  feed  (Scott,  1989  and  pers.  comm;  K.W.B., 
pers.  obs.). 

Etjinology:     After  Ian  Scott,  Auckland. 


ACKNOWLEDGMENTS 

For  the  loan  of  type  material,  other  specimens,  and  in- 
formation, we  are  grateful  to  P.  Bouchet  and  V.  Heros 
(Museum  National  d'Histoire  Naturelle,  Paris),  N.  Hiller 
(Canterburv'  Museum,  Christchurch),  I.  Loch  (Austra- 
han  Museum,  Sydney),  I.  Scott  (Auckland),  A.  Spurgeon 
(Hamilton),  FG.  Thompson  (Florida  Museum  of  Natu- 
ral Historv',  Gainesville),  and  K.M.  Way  (The  Natural 
History  Museum,  London).  We  also  record  our  indebt- 
edness to  our  late  friends  D.W.  Gibbs,  PR.  Jamieson, 
and  J.R.  Penniket,  who  supplied  some  of  die  finest  ma- 
terial used  in  this  study;  and  Wellington  Shell  Club 
members,  who  provided  Mahia  Peninsula  records  of  M. 
octogomis.  Thanks  also  to  A.G.  Beu  (Institute  of  Geo- 
logical and  Nuclear  Sciences,  Lower  Hutt),  I.  Scott 
(Auckland),  and  anonymous  referees  for  comments  on 
the  manuscript,  N.  Heke  and  M.  Hall  (Museum  of  New 
Zealand)  for  pliotography  and  photographic  printing  re- 
spectively, and  to  W'.  St.  George  (Institute  of  Geological 


and  Nuclear  Sciences,  Lower  Hutt)  for  scanning  elec- 
tron microscope  facilities. 


LITERATURE  CITED 

Abbott,  R.  T.  and  S.  P.  Dance.  198.3.  Compendium  of  seashells. 
A  color  guide  to  more  than  4,200  of  the  world's  marine 
shells.  American  Malacologists,  Melbourne,  410  pp. 

Beu,  A.  G.  and  P.  A.  Maxwell.  1990.  Cenozoic  Molluscaof  New 
Zealand.  New  Zealand  Geological  Survey  Paleontological 
Bulletin  .58:1-518. 

Bradshaw,  M.  A.,  A.  J.  Lobb,  M.  R  Cave  and  K.  Watson.  1992. 
Catalogue  of  type  and  figured  fossils  in  the  Geology  De- 
partment Canterbury  Museum.  Canterbury  Museum  Bul- 
letin 8:1-11.3. 

Bucquoy,  E.,  P.  Dautzenberg  and  G.-F.  DoUfus.  1882.  Les 
mollusques  marins  du  Roussillon.  1.  Bailliere,  Paris,  570 
pp. 

Fair,  R.  H.  1976.  The  Murex  book:  an  illustrated  catalogue  of 
Recent  Muricidae  (Muricinae,  Muricopsinae,  Ocenebri- 
nae).  Audior's  edition,  Honolulu,  138  pp. 

Finlay,  H.J.  1926.  A  further  commentary  on  New  Zealand  mol- 
luscan  systematics.  Transactions  of  the  New  Zealand  In- 
stitute 57:320^85. 

Finlay.  H.  J.  1927.  Additions  to  the  Recent  molluscan  fauna  of 
New  Zealand.  No.  2.  Transactions  of  the  New  Zealand 
Institute  57:48.5-487. 

Finlav,  H.  J.  1930.  Additions  to  the  Recent  molluscan  fauna  of 
New  Zealand.  No.  3.  Transactions  of  the  New  Zealand 
InstihJte  61:222-247. 

Gray,  J.  E.  1843.  Catalogue  of  die  species  of  Mollusca  and  their 
shells,  which  have  hitherto  been  recorded  as  found  at  New 
Zealand,  with  the  description  of  some  lateK-  discovered 
species.  Appendix  4,  pp.  22^8-265.  In:  Dieffenbach,  E., 
Travels  in  New  Zealand:  with  contributions  to  the  geog- 
raphy, geology,  botany,  and  natiual  histor\'  of  that  country. 
2.  Murray,  London. 

Hart,  M.  1993.  New  Zealand  Muricidae.  World  Shells  6:4:3- 
45. 

Houart,  R.  1988.  Description  of  seven  new  species  of  Muri- 
cidae (Neogastropoda)  from  the  southwestern  Pacific 
Ocean.  Venus  47:185-196. 

Houart,  R.  1991.  Description  of  thirteen  new  species  of  Mur- 
icidae (Gastropoda)  from  Australia  and  the  New  Caledo- 
nia region,  with  range  extensions  to  Soudi  Africa.  Journal 
of  the  Malaeological  Society  of  Australia  12:3.5-55. 

Houart,  R.  1993.  Three  new  species  of  Muricinae  and  Muri- 
copsinae (Gastropoda:  Muricidae)  from  Somalia,  Christ- 
mas (Line  Islands)  and  the  Philippine  Islands.  Venus  52: 
41-46. 

Hutton,  F  W.  1880.  Manual  of  die  New  Zealand  Mollusca. 
Hughes,  Wellington,  224  pp. 

Hutton,  F  W.  1884.  Rexision  of  the  rachiglossate  Molluscaof 
New  Zealand.  Transactions  of  the  New  Zealand  Institute 
16:216-2.33. 

Hutton,  F  W  1886.  New  species  of  Tertiary  shells.  Transac- 
tions of  the  New  Ze;iland  Institute  18:33:3-335. 

Hutton,  F  W  1893.  The  Pliocene  Mollusca  of  New  Zealand. 
In:  J.  J.  Fletcher  (ed.).  The  MacLeay  Memorial  Voliniie. 
Linnean  Society  of  New  South  Wales,  Sydney,  3.5-92. 

Iredale,  T  1915.  A  commentar\'  on  Sviter's  "Manual  of  the  New 
Zealand  Mollusca".  Transactions  of  the  New  Zealand  In- 
stitute 47:417-i97. 


B.  A.  Marshall  and  K.  W.  Burch.  2000 


Page  29 


Kaicher,  S.  D.  1978.  Card  catalogue  of  world-wide  shells.  Pack 
16 — Muricidae  .3.  Kaicher,  St.  Petersburg,  Florida. 

Kiener,  L.  C.  1843.  Species  general  et  iconograpiiie  des  co- 
quilles  \i\autcs,  coiuprenant  la  collection  du  Museum 
d'Histoire  uaturelle  de  Paris,  la  collection  Lamarck,  celle 
du  Prince  Massena,  et  les  decouvertes  recentes  des  vova- 
geurs.  Genr(>  Rocher  {Miirfx,  Lin.).  Bailliere,  Paris,  1.30 
pp. 

Kuster,  H.  C.  and  W.  Kobelt.  18.39-1878.  Die  ge.schwanzten 
vmd  bevvehrten  Puqjurschnecken  (Adurex.  Rtmclla.  Triton- 
ium,  Trophoit.  Hindsia).  In:  Kiister,  H.  C.  and  W.  Kobelt 
(eds.),  Systemati.sches  Conchylien-Cabinet  von  Maituii 
und  Chemnitz.  .  .  3  (2)  193.  Bauer  and  Raspe,  Leipzig, 
336  pp. 

Poirier,  J.  1883.  Re\ision  des  Miirex  du  Museinn.  Nouvelles 
Archives  du  Museum  dHistoire  Naturelle  (2)  .5:1.3-128. 

Ponder,  W.  F.  1968.  Nomenclatural  notes  on  some  New  Zea- 
land rachiglossan  gastropods  with  descriptions  of  five  new 
species.  Records  of  the  Dominion  Museum  6:29-47. 

Ponder,  W.  F.  1972.  Notes  on  some  Australian  genera  and  spe- 
cies of  the  family  Muricidae  (Neogastropoda).  Journal  of 
the  Malacological  Societv  of  Australia  2:21.5-248. 

Powell,  A.  W.  B.  1979.  New' Zealand  Mollu.sca.  Marine,  land 
and  freshwater  shells.  Collins,  Auckland,  500  pp. 

Quoy,  J.  C.  R.  and  P.  Gaimard.  1833.  Voyage  de  decouverts  de 
I'Astrolabe,  execute  par  ordre  du  Roi  pendant  les  annees 
1826-1827-1828-1829,  sous  le  commandement  de  M.J. 
Dumont  d'Umlle.  Zoologie.  MoUusques  2  (2).  Tastu,  Par- 
is: .321-686. 

Radwin,  G.  E.  and  A.  DAttilio.  1976.  Miirex  shells  of  the 
world.  An  illustrated  guide  to  the  Muricidae.  Stanford 
University'  Press,  Stanford.  284  pp. 

Reeve,  L.  184.5-1846.  Monograph  of  the  genus  Miirex.  Con- 
chologia  iconica:  or  illustrations  of  the  shells  of  mollus- 
cous animals.  3.  Reeve,  London.  36  plates  and  captions. 


Scott,  1.  1989.  A  closer  look  at  Miiricopsis  octogonus.  Poirieria. 

Auckland  Museum  Concholog\'  Section  L5:6-8. 
Shlkama,  T.  and  M.  Horikoshi.   1963.  Selected  .shells  of  the 

world  illustrated  in  colours.  HokurvTi-kan,  Tokyo.  154  pp. 
Smith,  M.  1940.  Two  new  marine  molluscs  from  japan.  The 

Nantilus  .54:43. 
Sowerby,  G.  B.  1841a  (in  1832-1841).  The  conchological  illus- 
trations. Miirex.  Sowerby,  London,  9  pp,  23  plates. 
Sowerby,  G.  B.  1841b.  Descriptions  of  some  new  species  of 

Miirex.  principally  from  the  collection  of  H.  Cuming,  Esq. 

Proceedings  of  the  Zoological  Societv  of  London  8  (1840): 

137-147. 
Suter,    H.    1901.    Note   upon    Trophoii   iimhiUcatns,   Tenison 

Woods.  The  Journal  of  M;ilacology  8:61-63. 
Suter,  H.  1913.  Manual  of  the  New  Zealand  Mollusca.  With 

an  atlas  of  plates  [1915].   Mackav,  Government  Printer, 

Wellington,  1120  pp. 
Thiele,  J.  1929.  Handbuch  der  Systematischen  Weichtierk-unde 

1  (1).  Fischer,  Jena,  376  pp. 
Tryon,  G.  W  1880.  Manual  of  concholog)-:  structural  and  sys- 
tematic. With  illustrations  of  the  species.  2.  Muricinae, 

Puqjurinae.  Tryon,  Philadelphia,  289  pp. 
Yokes,  E.  H.   1964.  Supraspecific  groups  in  the  subfamilies 

Muricinae  and  Tritonaliinae  (Gastropoda:  Muricidae).  Ma- 

lacologia  2:1^1. 
Yokes,  E.  H.  1970.  The  West  American  species  of  Murexiella, 

(Gastropoda:  Muricidae)  including  hvo  new  species.  The 

Yeliger  12:325-329. 
Yokes,  E.  H.  1988:  Muricidae  (Mollusca:  Gastropoda)  of  the 

Esmeraldas  beds,  northwestern  Ecuador,  Tulane  Studies 

in  Geolog)-  and  Palaeontology  21:1-50. 
Wenz,  W.  1941.  Gastropoda.  Teil  1:  Allgemeiner  Teil  und  Pro- 

sobranchia  5  (7),  pp.  961-1200.  7».-  Schindewolf  O.  H. 

(ed.),  Handbuch  der  Palaozoologie  6.  Bomtraeger,  Berlin. 


THE  NAUTILUS  114(1):30,  2000 


Page  30 


Notices 


CONSTANCE  E. 
MALACOLOGY 


BOONE  GRANTS  TO 


The  newly  established  Constance  E.  Boone  Grants  to 
Malacologv'  proxddes  annually  up  to  $1000  to  qualified 
persons  undertaking  c(jllection-based  research  on  Re- 
cent mollusks  at  the  Houston  Museum  ot  Natural  Sci- 
ence, Houston,  Texas.  The  HMNS  malacology  collection 
contains  over  48,000  cataloged  lots  (45,000  in  database 
format),  comprising  approximately  L5  million  speci- 
mens. The  main  geographic  focus  of  the  HMNS  is  the 
Gulf  of  Mexico.  The  museum  also  has  one  of  the  finest 
malacological  research  fibraries  in  the  country.  Awards 
will  be  made  only  to  citizens  or  permanent  residents  of 
the  Americas  or  to  undergraduate  and  graduate  students 
attending  colleges/universities  in  the  United  States.  Al- 


though proposals  of  up  to  $1000  will  be  entertained, 
grants  generally  do  not  exceed  $.500.  Partial  funding  is 
not  unusual.  The  Houston  Conchology  Society  (HCS) 
Grants  Committee  judges  applications. 

Grant  recipients  are  encouraged  to  submit  a  short  paper 
concerning  their  research  to  the  editor  of  the  HCS  pub- 
lication, the  Texas  Conclwlogist. 

Postmark  deadline  for  applications  is  May  1,  2000. 
Awards  will  be  announced  by  June  1 ,  2000. 

For  more  information,  please  contact: 

Dr.  John  B.  Wise 

Houston  Museum  of  Natural  Science 

One  Hermann  Circle  Dr 

Houston,  TX  770.30-1799 

jwise@hmns.org 


Florida  Department  of  State 

Katherine  Harris 

Secretary  of  State 

Florida  Arts  Council 

Division  of  Cultural  Affairs 

This  program  is  sponsored 

in  part  by  the  State  of  Florida, 

Florida  Department  of  State, 

Division  of  Cultural  Affairs,  and 

the  Florida  Arts  Council 


INSTRUCTIONS  TO  AUTHORS 


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biology  and  systematics  of  mollusks.  Manuscripts  describing 
original,  unpuljlished  research  as  well  as  review  articles  will 
be  considered.  Brief  articles,  not  exceeding  1000  words,  will 
be  published  as  notes  and  do  not  require  an  abstract.  No- 
tices of  meetings  and  other  items  of  interest  to  malacolo- 
gists  will  appear  in  a  news  and  notices  section. 

Manuscripts:  Each  original  manuscript  and  accompauviiig 
illustrations  should  be  submitted  in  triplicate.  Text  must  be 
t)'ped  on  one  side  of  8V4  X  11  inch  white  paper,  double 
spaced  throughout  (including  literature  cited,  tables  and 
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Authors,  Editors,  and  Publishers,  which  is  available  from 
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cbe).  The  first  mention  of  a  scientific  name  in  the  text 
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TH  EfyNAUTI  LUS 


CONTENTS 


Valium'  114.  Number  2 

June  6,  2000 

ISSN  0028-1344 


Douglas  G.  Smith  Notes  on  the  ta\onoiny  of  introduced  Bcllainija 

(Gastropoda:  Viviparidae)  species  in  northeastern  North 

America 31 

Guido  Pastorino  A  revision  of  the  Patagonian  genus  Xijmeiiopsis  Powell, 

M.  G.  Harasewych  1951  (Gastropoda:  Muricidae) 38 

Jose  H.  Leal  Copiilabi/ssia  riosi,  a  new  deep-sea  limpet  (Gastropoda: 

Luiz  Rieardo  L.  Simone  Pseudococculinidae)  from  the  continental  slope  off  Brazil 

with  comments  on  the  svstematics  of  the  genus 59 

G.  Darrigran  Invasion  of  the  exotic  freshwater  mussel  Limnopcnw 

I.  Ezcurra  de  Drago  fortunei  (Dunker,  1857)  (Bivalvia:  Mytilidae)  in  South 

America 69 

Warton  Monteiro  Location  of  allospermatozoa  in  the  freshwater  gastropod 

Toshie  Kawano  Bioiuphalaiia  tcna^iophila  (d'Orbigny,  1835)  (Pulmonata: 

Planorbidae) 74 

Jay  Cordeiro  Status  of  the  tidewater  mucket,  Lcptodca  ochracea  (Say, 

1817)  (Bivalvia:  Unionidae),  in  Halfway  Pond, 
Massachusetts.  USA .' 80 

Notices   84 


THE  NAUTILUS  114(2):31-37,  2000 


Page  31 


Notes  on  the  tcixonomy  of  introduced  BeUaniya  (Gastropoda: 
Viviparidae)  species  in  northeastern  North  America 


Douglas  G.  Smith 

Department  i)t  Biologv'  and 
Graduate  Program  in  Organismic  anti 

E\olntionar\  BiologN 
Uni\ersit\'  ot  Massachusetts 
Amherst,' MA  01003-5810  USA 
dgsmith@bio.uaniss.edu 


ABSTRACT 

The  genus  BeUnmi/a  has  become  well  established  in  Nortli 
America  since  its  first  introduction  arovmd  the  turn  ot  the  20"' 
centurv.  Some  malacologists  have  recognized  two  species,  usu- 
alK'  placed  in  the  genus  Cipaiigopahidina.  C.  chinen.sis.  and  C. 
japonica.  Other  investigators,  however,  have  questioned  a  two 
species  concept  or  simply  considered  all  populations  to  be  xar- 
iants  of  a  single  species,  C.  chinensis.  Differences  observed  in 
the  juvenile  shell  and  male  reproductive  anatomy,  along  with 
size-specific  features  of  adult  shells,  enable  easv  separation  of 
the  hvo  taxa  and  support  the  two  species  concept.  Characters 
of  female  and  male  reproductive  anatomv  confirms  the  place- 
nient  ot  the  species  in  the  subfamik'  Bellanninae.  Anatomical 
features  which  ha\e  been  used  to  substantiate  Cipanaopahi- 
dina  are  proposed  to  be  of  subgeneric  value  only. 

Kei/  words:  Introduced  snails,  Cipangopaludina,  Idiopoma.  Vi- 
vipams.  Lecijthoconcha. 


INTRODUCTION 

Among  the  more  conspicuous  members  of  the  North 
American  freshwater  gastropod  fauna  are  the  large  ex- 
otic viviparid  snails  customarily  placed  in  the  genus  Ci- 
pan^opaJiidina.  The  North  American  history  of  Cipaii- 
gopahidina and  its  distribution  was  discussed  and  re- 
viewed by  Clench  and  Fuller  (1965)  (as  a  subgenus  of 
Vivipanis),  Dundee  (1974),  and  Joldnen  (1982).  The 
majority  of  North  American  populations  are  confined  to 
the  coasts,  with  scattered  records  from  the  interior  At 
present,  the  east  coast  appears  to  comprise  the  greatest 
number  of  occurrences  (Joldnen,  1982). 

Since  the  earliest  reports  of  Cipangopaludina  popu- 
lations in  North  America  around  1900  there  has  been 
much  debate  regarding  the  identification  and  \ali(lit\'  oi 
the  various  species  and  subspecies  reptjrted  and,  to  a 
lesser  extent,  the  status  and  position  of  the  taxon  Cipan- 
gopaludina in  the  Vixaparidae.  Hannibal  (1912)  proposed 
the  subgenus  Cipangopaludina  to  accomodate  Idiopoma 
niallcata  (Reeve,  1863),  leaving  /.  japonica  (von  Mar- 
tens, 1861)  in  Idiopoma  s.  s.  Subsequent  authors  (e.g.. 


Annandale,  1920;  Prashad,  1928)  regarded  Idiopoma 
Pilsbrv',  1901,  recognized  primarily  on  features  of  the 
operculum,  as  a  synonym  of  Vivipanis.  Annandale 
( 1920)  used  anatomical  characters  to  propose  a  new  ge- 
nus Lecijthoconcha  to  include  L.  cJiincncsis  and,  in  a 
later  paper  (Annandale,  1921),  to  include  L.  japonica  as 
well.  Rao  (1925)  maintained  use  oi  Lcci/thoconcha  and 
expanded  the  number  of  anatomical  characters  that  de- 
fine the  genus,  most  notablv  a  folded  and  thickened  gill 
filament,  a  thickened  muscular  mantle  edge,  and  the 
"strongly  developed  brain".  Prashad  (1928)  subsequently 
synonymized  Lecijthoconcha  with  Cipangopahidina  but 
retained  Lecijthoconcha  as  a  subgenus.  Furthermore,  he 
reduced  Idiopoma  maUeata  to  a  subspecies  of  V  chinen- 
sis (Gray,  1817).  Kuroda  (1929)  adopted  all  of  Prashads 
(1928)  revisions.  Rohrbach  (1937)  primarily  used  char- 
acters of  the  reproductive  anatomv  to  divide  the  vivi- 
parids  into  two  subfamilies,  the  Vi\iparinae  and  a  new 
subfamily  Bellamyinae.  Under  the  Bellamyinae  he  listed 
a  single  large  genus,  Bcllamija  Jousseaume.  Although 
considered  to  be  a  genus  largelv  limited  to  tropical  spe- 
cies, Rohrbach  (1937)  was  able  to  examine  specimens  of 
a  race  of  non-tropical  V.  chinensis  that  displa\ed  anatom- 
ical features  of  BcUaintja.  Yen  (1943)  extended  Rohr- 
bach s  (1937)  classification  to  Chinese  species  and  sub- 
species, restateil  the  close  relationship  between  Bella- 
mija  and  Cipangopahidina,  but  retiuned  Cipangopahi- 
dina at  the  genus  level. 

The  first  major  work  on  North  American  \i\iparids 
following  this  period  is  Clench  and  Fuller's  (1965)  study 
in  which,  for  unkTiown  reasons,  they  treat  Cipangopa- 
hidina as  a  subgenus  of  Vivipanis.  These  authors  res- 
urrected the  trinomen  V  chinensis  malleatus  for  all 
North  American  populations  of  this  species  and  listed  it 
as  a  species  separated  from  V'  (Cipangopahidina)  japon- 
iciis.  Most  subsequent  studies  on  viviparid  faunas  (eg. 
Pace,  1973:  Taylor,  1981;  Rao,  1989)  have  considered 
Cipangopahidina  a  genus  of  the  Bellamyinae. 

North  American  opinion  has  been  di\aded  concerning 
the  \'alidit\'  of  the  two  introduced  species  (see  Jokinen, 


Page  32 


THE  NAUTILUS.  Vol.  114.  No.  2 


1991,  for  review).  Many  North  American  faunal  studies 
including  these  viviparids  have  listed  all  large  introduced 
\iviparid  populations  with  unhanded  shells  as  Cipango- 
pahidina  chincnsis  (eg.  Robertson  and  Blakeslee,  1948; 
Jacobson  and  Emerson,  1961;  Dundee,  1974;  Clarke, 
197S).  Other  reports  have  recognized  the  two  species  C 
chincnsis  and  C.  japonica  (eg.  Wolfert  and  HUtunen, 
1968;  Burch,  1982;  Tavlor,  1981;  Jokinen,  1983,  1991; 
Smith,  1995). 

Certainly  one  of  the  problems  affecting  the  species 
issue  is  the  apparent  rant)-  of  the  ta.\on  C.  japonica. 
Clench  and  Fuller  (1965)  gave  only  four  North  Ameri- 
can records,  and  Taylor  (1981)  listed  only  one  in  Cali- 
fornia. Wolfert  and  Hiltunen  (1968)  listed  an  Ohio  re- 
cord and  Jokinen  (1984)  provided  an  additional  record 
in  Connecticut,  but  has  before  and  since  considered  the 
population  C.  chincnsis  or  doubtfully  C.  Japonica  (Joki- 
nen, 1983,  pers.  comm.).  Jokinen  (1991)  subsequently 
added  a  New  York  record  that  would  be  the  eighth  pub- 
hshed  North  American  record  for  C.  japonica. 

The  impetus  for  the  present  study  came  from  the  dis- 
covery of  two  wild,  previously  unreported  populations  of 
C.  japonica-morph  sn;iils  in  Connecticut  from  which  se\'- 
eral  variously  aged  specimens  and  reproductively  active 
females  were  collected.  The  present  study  provides  com- 
parative data  on  a  number  of  characters  of  each  species 
in  an  attempt  to  determine  whether  two  species  or  a 
single  variable  species  occurs  in  North  America.  Addi- 
tionally, using  data  from  reproductive  organs  and  other 
anatomical  characters  plus  observations  by  previous  au- 
thors on  viviparid  subgroups,  a  proposal  to  reduce  Ci- 
pangopahidina  to  a  subgenus  ot  BeUanu/a  is  presented. 
Hereafter  in  this  study  these  two  species  will  be  referred 
to  as  Bellamija  spp. 

MATERIALS  AND  METHODS 

Specimens  from  12  extant  or  extirpated  BcIIamt/a  pop- 
ulations were  examined.  The  original  assignment  ot 
specimens  to  either  specific  taxon  was  by  use  of  adult 
shell  characters.  The  distribution  of  specimens  is  listed 
below.  All  were  fixed  in  10%  formalin,  relaxed  or  unre- 
laxed,  and  stored  intact  in  50-60%  isopropyl  alcohol.  In- 
tact shells  of  deceased  animals  were  also  collected.  Each 
collection  has  been  catalogued  in  the  Invertebrate  Di- 
vision, Museum  of  Zoology,  University  of  Massachusetts, 
Amherst  (UMA).  Collection  numbers  and  other  details 
regarding  the  collections  can  be  furnished  upon  request. 
BcUami/a  chincnsis  s.l:  UMA  0668,  University  Pond, 
University  of  Massachusetts  Campus,  Amherst,  Hamp- 
shire County;  UMA  0885,  unnamed  pond  in  Medford, 
and  UMA  1705,  Lake  Winthrop,  Holhston,  both  Mid- 
dlesex County;  UMA  1635,  Forest  Park  Ponds,  Spring- 
field, Hampden  County;  UMA  1663,  Wachusett  Reser- 
voir Outlet,  Worcester  County;  UMA  1094,  Purchase 
Louisiana  Brook,  Northfield,  Frankhn  County;  UMA 
1330,  Pine  Tree  Brook,  Milton,  Norfolk  County;  UMA 
1805,  Snake  River,  Norton,  Bristol  Count>-  (all  Massa- 


chusetts); UMA  1755,  Black  Pond,  Middlefield,  Middle- 
sex County,  Connecticut. 

BcIIamija  japonica:  UMA  1754,  Lake  Mohegan, 
Fairfield,  F;iirfield  County;  UMA  1816,  Connecticut  Riv- 
er, Lviue,  New  London  County;  UMA  1755a,  Black 
Pond,  Middlefield,  Middlesex  County,  all  Connecticut. 
Also  examined  were  specimens  (UMA  1700)  purchased 
from  a  pet  store  in  Hadley,  Hampshire  County,  Massa- 
chusetts. 

Samples  ol  intra-uterine  juveniles  were  removed  from 
females.  The  animal  was  removed  from  the  shell  of  each 
juvenOe  and  the  shell  iiir  dried.  Radulae  were  also  re- 
moved from  adult  specimens,  cleaned  in  10%  potassium 
hydroxide,  subsequently  washed,  and  air  dried.  All  dried 
material  was  mounted  on  individual  Cambridge  stubs 
with  Duco  cement  and  sputter-coated  with  gold.  All  pre- 
pared specimens  were  then  examined  with  a  JEOL 
Model  JSM-5200  scanning  electron  microscope  (SEM). 

For  anatomiciil  study,  at  least  five  specimens  of  each 
sex  were  dissected  from  the  two  largest  collections  of 
each  species  as  determined  by  shell  characters.  There- 
after, dissections  were  made  on  at  least  one  adult  spec- 
imen of  each  sex  (if  available)  from  each  remaining  lo- 
cation. A  total  of  44  specimens  divided  almost  evenly 
among  both  species  were  analyzed.  Dissections  were 
performed  with  the  aid  of  a  Wild  M5  dissecting  micro- 
scope. 

RESULTS 

The  principal  conchological  characters  distinguishing  the 
tsvo  taxa  are  the  shape  of  the  spire  and  the  presence  of 
carination;  to  quote  from  Clench  and  Fuller  (1965)  re- 
garding B.  japonica:  ".  .  .can  be  distinguished  from  V 
mallcatiis  (=  C.  chinen.sis)  by  having  a  more  acute  spire 
and  by  having  fine  carina  or  carinae."  An  examination  of 
animals  from  presumed  B.  japonica  populations  reveal 
that  these  characteristics  are  most  evident  in  shells  35- 
45  mm  shell  length  (Figure  1).  At  this  size,  the  species 
can  be  separated  using  the  diagnosis  of  Clench  and  Ful- 
ler (1965).  Below  35  mm  shell  length  both  species  have 
a  carina  on  the  body  whorl.  Above  45  mm,  the  carina  of 
B.  japonica  becomes  nearly  obsolete,  and  only  a  sfightly 
more  acute  spire  in  B.  japonica  enables  identification. 

The  source  of  differentiation  in  shell  characters  in 
these  two  species  is  in  part  derived  from  an  allometric 
growth  pattern  first  described  by  Jokinen  (1982),  who 
showed  that  shell  width  in  B.  japonica  increases  at  a 
slower  rate  than  in  B.  chincnsis.  This  phenomenon 
would  account  for  the  more  "acute  spire"  in  B.  japonica 
described  by  Clench  and  Fuller  (1965).  The  regression 
slopes  for  each  species,  however,  are  only  slightly  differ- 
ent (see  Jokinen,  1982,  fig.  2).  Therefore,  adult  shell 
moi-phology  has  its  limitations  in  the  distinction  of  these 
two  species. 

A  characteristic  that  appears  to  be  reliable  in  separat- 
ing B.  japonica  from  B.  chincnsis  is  the  morphology  of 
the  intra-uterine  juvenile  shell.  The  juveniles  occurring 
in  the  uteiTis  or  brood  chamber,  considered  a  functional 


D.  G.  Smith,  2()()() 


Page  3.3 


Figure  1.  Comparison  of  similar  sized  shells  of  Bdlamija. 
Upper  and  lower  left,  B.  japonica  from  Lake  Mohegan,  Con- 
necticut. Upper  right,  B.  chinensis  from  Medford,  Mussachn- 
sets,  and  lower  right,  B  chinensis  from  Black  Pond,  Connect- 
icut. Arrow  denotes  carina.  Scale  line  =  10  mm. 


modification  of  what  is  otherwise  called  the  pallial  ovi- 
duct, demonstrate  clear  differences  in  the  position  of  the 
embryonic  whorl  (protoconch)  and  in  surface  sculpture. 
Most  viviparid  snails  in  their  juvenile  stages  have  specific 
features  of  the  periostracum,  including  the  presence  of 
periostracal  hairs  encircling  the  shell  v\'horls,  and  surface 
sculpture  (Fretter  and  Griiliam,  1962;  Joldnen,  19(S4). 

In  the  juvenile  shell  of  B.  chinensis,  the  spire  com- 
prises only  two  elevated  whorls  (Figures  2-A).  The  em- 
bryonic whorl  is  depressed  in  frontal  view  below  the  suc- 
cessive whorl  (first  teloconch  whorl),  characterized  by 
two  fines  ot  periostracal  hairs,  <3ne  clearly  evident  on  the 
whorl,  the  other  along  the  upper  border  (Figure  2,  ar- 
row). Such  was  observed  in  all  examined  populations.  In 
contrast,  the  embryonic  whorl  of  B.  japonica  (Figure  5) 
is  somewhat  elevated  above  the  first  teloconch  whorl, 
indicated  by  two  spiral  fines  of  periostracal  hairs,  which 
in  frontal  view  gives  the  shell  the  appearance  of  having 
four  whorls. 

Differences  in  periostracum  are  even  more  eNddent 
than  those  in  whorl  development.  The  periostracum  of 
the  body  whorl  of  juvenile  B.  chinensis  (Figures  6,  7)  is 


finely  sculptured  with  a  series  of  closely  set  striae,  about 
20/mm  near  the  midpoint  of  the  body  whorl,  interrupted 
by  peqiencficular  shallow  grooves  more  widelv  spaced 
than  tlie  striae.  The  spiral  lines  bearing  the  fine  hairs  are 
not  raised  as  spiral  carinae.  In  B.  japonica  (Figure  8), 
the  surface  of  the  periostracum  is  strongly  marked  by 
raised  carinae,  each  having  a  series  of  periostracal  hairs. 
Between  the  carinae  are  widely  spaced  spiral  striae, 
about  8/mm  near  the  middle  of  the  whorl.  Although  oc- 
casional interiTiptions  occur,  there  is  no  pattern  of  dis- 
tinct grooves  as  found  in  and  characteristic  of  B.  chinen- 
sis. 

Regarding  anatomical  characters,  the  radula  provides 
no  specific  or  cfistinctive  characters  for  interspecific  dif- 
ferentiation. Veiy  subtle  cfifferences  (Figures  9,  10)  oc- 
cur in  the  spacing  of  denticles  of  the  lateral  teeth  for 
example,  but  these  differences  are  probablv  subject  to 
variation  and  should  not  be  considered  as  refiable  taxo- 
nomic  characters.  The  gill  filament  of  each  species  con- 
sists of  an  elongate  structure,  measuring  up  to  5  mm 
length  in  the  largest  (>50  mm  shell  length)  individuals. 
The  filament  is  broadest  at  its  base  where  it  is  attached 
to  the  mantle,  narrowing  in  depth  cfistally.  A  cfistinct 
blood  vessel  courses  its  length  just  below  the  thickened 
superior  margin.  The  "fokfing"  of  the  filament  described 
by  Rao  (192.5)  in  a  few  species,  and  used  subsequently 
as  a  character  of  Cipangopahulina.  was  not  observed  in 
material  of  either  species. 

Major  features  of  both  the  male  and  female  repro- 
ducti\'e  svstems  demonstrate  an  overall  conservative 
structural  plan  clearly  characteristic  of  the  Bellamyinae 
as  defined  by  Rohrbach  (1937).  Dissection  of  the  female 
reproductive  system  revealed  no  consistent  differences 
between  B.  chinensis  and  B.  japonica.  Within  the  male 
reproductive  system,  the  \'as  deferens  of  each  species 
shows  distinct  differences  in  its  cfimensions  and  its  re- 
lationship with  both  the  columellar  muscle  and  the  tes- 
tes. In  B.  chinensis  (Figure  11),  the  vas  deferens  arises 
from  the  testes  as  an  unbranched  tube.  All  further 
branching  of  the  vas  deferens  occurs  internally  within 
the  testes.  Occasionally,  a  single  large  anterior  branch 
merging  with  the  vas  deferens  is  partially  visible.  The 
emerged  vas  deferens  widens  considerablv  as  a  complex 
duct  and  proceeds  a  short  cfistance,  suspended  by  mes- 
enteries, until  it  reaches  the  columellar  muscle.  At  this 
point,  the  remaining  portion  of  the  vas  deferens  bends 
shaqiely  downwards.  It  then  abruptly  and  without  en- 
largement enters  the  prostate.  The  downwardlv  cfirected 
portion  of  the  vas  deferens  remains  free  of  the  colu- 
mellar muscle  although  it  is  supported  throughout  by 
mesentaiy  tissue. 

The  vas  deferens  of  B,  japonica  departs  from  the  tes- 
tes multibranched,  with  at  least  three  and  usually  four 
visible  branches  converging  to  a  single  duct  (Figure  12). 
The  duct  then  enlarges,  but  not  to  the  degree  seen  in 
B.  chincn.'iis,  and  extends  a  greater  distance,  suspended 
by  mesentery  tissue,  until  meeting  with  the  columellar 
muscle  where,  as  in  B.  chinensis,  it  bends  downward 
and,  without  enlargement,  joins  the  prostate  gland.  Un- 


Page  34 


THE  NAUTILUS,  Vol.  114.  No.  2 


ij^gaiit«^Jgi*jfj 


BD 


m 


v^ 
m 


3  - 


■N  "1 


BD 


Figures  2-5.  Spire  of  juvenile  shells  ot  Bellnmya.  2-4.  Bellamtja  chinensis.  2.  Black  Pond,  Connecticut.  3.  Wachusett  Reservoir, 
Massachusetts.  4.  Campus  pond,  Amherst,  Massachusetts.  Figures  3,  4,  apical  whorls  only.  5.  Bcllami/a  jnponica.  Lake  Mohegan, 
Connecticut.  BD  =  hodv  whorl.  Scale  line  =  0.5  mm. 


hke  B.  chinensis,  however,  the  portion  of  the  vas  defer- 
ens pro>diiial  to  the  prostate  gland  is  invested  with  col- 
umellar  muscle  tissue.  Odier  aspects  of  gross  anatomy 
of  the  male  reproductive  system,  including  testes  size 
and  shape  and  prostate  size  and  shape,  are  similar  in  the 
two  species. 

The  characteristics  of  the  vas  deferens  in  each  species 
were  most  pronounced  in  the  larger  animals.  The  hvo 
patterns  were  concordant  with  shell  characters  in  all  but 
one  case,  the  Black  Pond  locality.  Both  species  occur  in 
this  pond  and  examined  males  show  features  of  shell  and 
reproductive  anatomy  consistent  with  above  observations 
or  variations  of  each.  Most  males  from  this  collection, 
however,  are  young  adults  (<.'35  mm  shell  length)  and, 
as  stated  above,  the  anatomical  characters  inchcative  of 
each  species  are  best  developed  in  larger  specimens. 
Nevertheless,  it  is  possible  that  hybridization  is  occur- 
ring. Previously,  in  North  America,  both  species  have 
been  reported  in  sympatiy  (Lake  Erie;  Wolfert  and  Hil- 


tunen,  196S),  but  not  in  svmtopy  so  it  is  not  known  if 
hydridization  is  a  usuiJ  outcome  of  contact. 


DISCUSSION 

Although  adult  shell  structure  as  presented  by  Clench 
and  Fuller  (1965)  does  not  in  and  of  itself  provide  con- 
\incing  evidence  for  the  recognition  of  two  distinct  spe- 
cies, the  characters  of  the  embrvonic  shell  and  male  re- 
proilucti\e  system  in  combination  with  shell  features  in- 
dicate that  at  least  Uvo  similar  but  distinct  species  of 
Bcllami/a  are  present  in  North  America.  The  question 
remains,  however,  regarding  which  names  to  assign 
these  species;  is  enough  information  available  to  warrant 
use  of  the  two  specific  epithets,  mallcata  and  japonica 
as  hsted  by  Hannibal  (1911)  soon  after  the  species  were 
first  reported  in  North  America?  If  so,  then  what  is  the 
the  status  of  the  name  B.  chinensis? 

Clench  and  Fuller  (1965)  gave  a  partial  history  of  the 


D.  G.  Smith,  2000 


Page  35 


Figures  6-8.  Penostracal  surface  of  body  whorl  ol  ju\eiiile  Bellami/a^  6.  B.  chinensis  from  Black  Pond,  Connecticwt.  7.  Same, 
Forest  Park  ponds,  Springfield,  Massachusetts.  8.  B.  japonica  from  Lake  Mohegan,  Connecticut.  Scale  line  =  0..5  mm.  Figures  9- 
10.     Radula  (central  and  lateral  teeth).  9.  Bilhiim/a  japonica.  10.  Bcllainija  chinensis.  Scale  line  =  0.05  mm. 


taxonomy  and  nomenclature  of  these  species.  As  can  be 
deduced  from  their  account,  the  original  descriptions  of 
these  species  were  essentially  brief  accounts  of  the  adult 
shell  with  very  little  geographical  data  provided.  Pace's 
(1973)  review  of  the  titxonomv  of  B.  chinensis  probably 
best  summarizes  the  nomenclatural  and  taxonomic  con- 
fusion that  accompanies  this  species.  Seemingly  every 
race  or  form  encountered  by  earlv  workers  was  estab- 
lished on  the  most  subtle  of  characters.  Differences  of 
opinion  on  the  validity  of  named  forms  and  generic  as- 


signment was  quite  likely  compounded  by  difficulties  in 
communication  between  Asian  and  European  investiga- 
tors. Unfortunately,  the  practice  of  naming  new  species 
of  Asiatic  viviparids  on  the  basis  of  a  few  qualitative  shell 
characters  continues  (Li,  1990;  Liu  ct  ai,  1994).  Until 
future  study  requires  otherwise,  I  follow  Pace  (1973)  and 
other  earlier  workers  who  assert  that  many  of  the  sub- 
sequentlv  named  forms  related  to  B.  chinensis.  including 
B.  mallcata,  are  at  most  subspecies.  Given  that  subspe- 
cific  status  is  normal!)-  maintained  through  geographic. 


Page  36 


THE  NAUTILUS,  Vol.  114,  No.  2 


CM 


VD 


CM 


Figures  11-12.  Anatomy  of  male  reproductive  structures. 
11.  Bellaimja  chinensis.  12.  Bellamya  japonica.  Scale  applies 
to  figures  11  and  12.  CM  =  coliimellar  muscle,  CT  =  con- 
nective tissue  (mesentery),  F  =  fibers  of  columellar  muscle,  P 
=  prostate,  T  =  testes,  VD  =  vas  deferens. 


(1912)  in  his  description,  the  foundation  of  Cipangppa- 
liidina  hes  chiefly  in  certain  anatomical  features  dis- 
cussed above.  Folding  of  the  gill  filament,  a  character  of 
Cipangopahidina  estabhshed  by  Rao  (1925),  was  not  ob- 
served in  either  species.  Folding  of  the  filament  mav  be 
caused  by  contraction  or  mav  represent  distortion  as  a 
result  of  preservation.  A  thick  mantle  edge  and  a  devel- 
oped sphincter  muscle  are  present  in  both  species,  but 
based  on  comparison  with  smaller  Vivipants  georgianus 
(I.  Lea,  1834)  thickness  and  muscle  size  can  be  related 
to  animal  size  (Smith,  pers.  observ.).  A  comparison  of 
mantle  types  illustrated  by  Annandale  (1920:  112)  and 
used  to  characterize  Lecijthoconcha  (=  Cipangopahidi- 
na) apart  from  other  genera  more  clearlv  demonstrates 
a  gradation  of  mantle  development  among  the  groups  he 
discussed.  In  relation  to  brain  comple.xity  (sensu  Rao, 
1925),  more  detailed  studies  are  necessary  for  the  prop- 
er evaluation  of  this  character.  Following  Rohrbach's 
(1937)  anatomical  svstem,  which  was  somewhat  extend- 
ed by  Pace  (1973),  B.  japonica  represents  a  group  within 
Bellamya  that  is  characterized  by  a  branched  vas  defer- 
ens proximal  to  the  testes;  Bellamya  chinensis  represents 
a  sister  group  that  has  an  unbranched  tree  portion  of 
the  vas  deferens.  Vail  (1977)  has  addressed  the  problems 
of  subgroups  within  Bellamya  in  a  more  general  fashion 
and  suggested  that  further  studies  are  necessary  to  es- 
tablish natural  groups  within  the  Bellamyinae.  It  is  thus 
proposed  that  Cipangopahidina  should  be  treated  as  a 
subgenus  of  Bellamya,  a  decision  with  some  precedent. 
Cipangopahidina  would  provisionally  embrace  the  large 
bellamvids  with  unhanded  shells  during  any  part  of  their 
life  historv  and  with  native  distributions  fimited  to  Asia. 

ACKNOWLEDGMENTS 

I  thank  Eileen  Jokinen  for  providing  useful  input  and 
comment  on  various  drafts  of  the  paper.  Kenneth  Boss 
provided  extensive  information  on  the  taxonomy  and  lit- 
erature of  Asian  bellamvids. 


not  genetic,  separation,  and  that  several  populations  of 
interbreeding  B.  chinensis  have  been  introduced  into 
North  America,  the  features  that  defined  any  particular 
subspecies  have  most  likelv  been  obliterated.  Further- 
more, in  that  most  opinion  leans  towards  a  Japanese  or- 
igin for  at  least  some  of  the  North  American  populations 
and  that  populations  of  each  species  have  been  reported 
in  Japan  (Kuroda,  1929),  I  believe  that  B.  japonica  s.  I. 
and  B   chinensis  s.  I.  are  the  species  involved. 

The  decision  to  place  both  species  with  Bellannia  re- 
sults in  part  from  Taylor's  (1981)  statement,  substanti- 
ated by  anatomical  data  pro\ided  in  this  studv,  that  B. 
japonica  belongs  to  the  Bellamvinae.  With  respect  to  the 
continued  use  of  the  name  Cipangopahidina  lor  6,  chi- 
nensis, those  characters  used  to  substantiate  the  genus 
are  either  variable  or  characteristic  of  animals  reaching 
a  relatively  large  size.  Notwithstanding  the  absence  of  a 
carina  on  the  adult  shell,  originally  used  by  Hannibal 


LITERATURE  CITED 

Annandale,  N.  1920.  Materials  for  a  generic  revision  of  the 
freshwater  gastropod  molluscs  of  the  Indian  Empire.  Re- 
cords of  the  Indian  Museum  19:107-115. 

Annandale,  N.  1921.  Zoological  results  of  a  tour  in  the  Far 
East.  The  viviparous  water-snail  of  Lake  Biwa,  Japan. 
Memoirs  of  the  Asiatic  Society,  Bengal  6:399—401. 

Burch,  J.  B.  1982.  The  freshwater  snails  (Mollusca:  Gastro- 
poda) of  North  America.  Environmental  Monitoring  and 
Support  Laboratory,  U.  S.  Environmental  Protection 
Agencv  (EPA -600/3-82-026),  Cincinnati,  294  pp. 

Clarke,  A.  H.  1978.  The  Asian  apple  snail,  Cipangopahidina 
chinensis  (Vixiparidae)  in  Oneida  Lake,  New  York.  The 
Nautilus  92:1.34. 

Clench.  W.  J.  and  S.  L.  H.  Fuller.  1965.  The  genus  Vivipams 
(Viviparidae)  in  North  America.  Occasional  Papers  on 
Mollusks  2:385-412. 

Dundee,  D.  S.  1974.  Catalog  of  introduced  mollusks  of  Eastern 
North  America  (North  of  Mexico).  Sterkiana  55;l-.36. 


D.  G.  Smith,  2()()() 


Page  37 


Fretter,  V.  and  A.  Graliam.  1962.  British  Prosohiainh  Mullusrs. 

Ra\-  SocietN'.  London.  755  pp. 
Hannihal,  H.  1911.  Further  notes  on  Asiatic  \i\iparas  in  Cah- 

toniia.  The  Nautilus  25:31-32. 
Hannihal.  H.  1912.  A  sviiopsis  of  the  Recent  and  Tertian' lanil 

and   fresh   water   Mollusca  of   the  Califoniian  Pro\ince, 

hased  on  an  ontogenetic  classification.  Proceedings  ot  the 

Malacological  Societv  of  London  10:112-211, 
Jacobson,  M.  K.  and  W.  K.  Emerson.  1961.  Shells  ot  the  New 

York  Citv  Area.  Argonaut  Books,  Larchniont,  142  pp. 
Jokinen,  E.  H.  1982.  Cipanoppaludina  chinensis  {Gastropoda: 

\'i\iparidae)  in  North  America.  The  Nautilus  96:89-95. 
Jokinen,  E.  H.  1983.  The  Freshwater  Snails  of  Connecticut. 

Bulletin  Connecticut  Geological  and  Natural  History  Sur- 
vey 109:1-83. 
Jokinen,  E.  H.  1984.  Periostracal  moqiholog)' of  \i\iparid  snail 

shells.  Transactions  of  the  American  Microscopical  Societv 

103:312-316. 
Jokinen,  E.  H.  1991.  The  Freshwater  Snails  (Mollusca:  Gas- 
tropoda) of  New  York  State.  Bulletin  of  the  New  York 

State  Museum  482:1-112. 
Kuroda,  T.  1929.  Notes  on  the  nomenclature  of  the  Japanese 

species  of  the  genus  Vivipanis.  The  Venus  1:98-102. 
Li.  Z.   1990.  A  new  species  of  genus  CApangopahuUna  from 

Dianchi  Lake  in  Yunnan  Province,  China.  Acta  Zootax- 

onomica  Sinica  15:2.5-27. 
Liu,  Y.  Z.,  W.  Zans,  Y.  Wane,  and  Y.  Duan.  1994.  Eight  new 

species  of  freshwater  molluscs  in  southeast  China.  Acta 

Zootaxonomica  Sinica  19:2.5-36. 
Pace,  G.  L.  1973.  The  freshwater  snails  of  Taiwan  (Formosa). 

Malacological  Review,  Supplement  1:1-118. 


Prashad,  B.  1928.  Recent  and  fossil  Viviparidae.  Stud\-  iu  dis- 
tribution, evolution  and  Paleogeography.  Memoirs  ol  the 
Indian  Museuui  8:153-252. 

Rao,  H.  S.  1925.  On  the  comparative  anatomv  of  Oriental  Vi- 
viparidae. Records  of  the  Indian  Museum  27:129-1.35. 

Rao.  N.  \.  S.  1989.  Handbook  Fresliwater  Molluscs  of  India. 
Zoological  Survey  ot  India,  Calcutta.  289  pp. 

Robertson,  I.  C.  andC.  L.  Blakeslee.  1948.  The  Mollusca  of 
the  Niagara  Frontier  Region.  Bulletin  of  the  Buffalo  So- 
ciety of 'Natural  Sciences  19(3):  1-191. 

Rohrbach,  F.  1937.  Oekologische  und  morphologische  Unter- 
suchungen  an  Vivipanis  (Bellarmja)  ca))illntHs  Franenfeld 
und  Vivipanis  {Brllamt/a)  unicolor  01i\er,  unter  Benick- 
sichtigung  anderer  tropischer  F'ormen  luid  im  Hinblick 
auf  phvletische  Beziehungen.  Archi\-  fur  Molluskenkunde 
69:177-218. 

Smith,  D.  G.  1995.  Keys  to  the  Freshwater  Macroinvertebrates 
of  Massachusetts.  Second  Edition.  University  of  Ma.ssa- 
chusetts,  Amherst,  243  pp. 

Taylor,  D.  W.  1981.  Freshwater  mollusks  of  California:  a  dis- 
tributional checklist.  California  Fish  and  Game  67:140- 
163. 

Vail,  V.  A.  1977.  Comparative  reproductive  anatomy  of  3  \ivi- 
parid  gastropods.  Malacologia  16:519-540. 

Wolfert,  D.  R.  and  J.  K.  Hiltunen.  1968.  Distribution  and 
abundance  of  the  Japanese  snail,  Vivipanis  japoniciis  and 
associated  macrobenthos  in  Sandusky  Bay,  Ohio.  Ohio 
Journal  of  Science  68:.32— 40 

Yen,  T.  1943.  A  preliminary  revision  ot  the  Recent  species  of 
Chinese  Viviparidae.  The  Nautilus  56:124-130. 


THE  NAUTILUS  114(2):38-58,  2000 


Page  38 


A  revision  of  the  Patagonian  genus  Xijryienopsis  Powell,  1951 
(Gastropoda:  Muricidae) 


Cuido  Pastorino 

Museo  Argentino  de  Ciencias  Natiirales 
Av.  Angel  G;illardo  470,  3°  piso,  lab.  57 
CUOsbjR  Bvienos  Aires 
ARGENTINA 
rvpastor@criba.edu .  ar 


M.  G.  Harasewych 

Department  of  Inxertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washmgton.  D.C.  20560-0118  USA 
Harasewych@nmnh.si.edu 


ABSTRACT 

The  genus  Xt/menopsi.s  Powell,  1951  is  re\ised  and  restricted 
to  4  Recent  species:  Xtjmenopsis  miiriciformis  (King  and  Brod- 
erip,  1832),  X.  comigatus  (Reeve,  1848),  X.  huccineus  (La- 
marck, 1816)  and  X.  suhnodosus  (Gray,  1839).  Of  the  34  nom- 
inal species  that  have  been  described  as  or  assigned  to  Xtjmen- 
opsis by  various  authors,  3  belong  to  the  buccinid  genus  Pa- 
reuthria  and  I  is  a  trophonine  nuiricid  not  closely  related  to 
Xymenopsis.  The  ta.xononiic  affinities  of  each  of  the  remaining 
.30  names  (one  is  a  replacement  name)  are  discussed.  Ele\en 
holot)pes  and  ten  lectotypes  (nine  newly  designated)  are  illus- 
trated, many  for  the  first  time.  Tvpes  of  .3  taxa  were  destroyed 
during  World  War  II  and  another  5  t)pes  are  lost.  Neotvpes 
are  designated  for  Fusus  huccineus  Lamarck,  1816,  the  oldest 
name  to  be  applied  to  any  species  of  Xymenopsis,  as  well  as 
for  Buccinum  subnodosa  Gray,  1839,  Trophon  brucei  Strebel, 
1904,  Trophon  fcnestratus  Strebel,  1904,  Trophon  paessleri  tur- 
rita  Strebel,  1904,  Trophon  ringci  Strebel,  1904  and  Trophon 
standcni  Strebel,  1904.  This  is  done  to  remo\e  anv  ambiguity 
regarding  the  identity  of  these  ta.\a,  and  thus  stabilize  the  no- 
menclature of  Recent  species  of  Xymenopsis.  The  gross  anat- 
omy and  moqjhology  of  the  radula,  egg  capsule,  protoconch, 
and  shell  ultrastructure  of  the  type  species  of  Xymenopsis  are 
described  and  illustrated.  Xymenopsis  is  compared  to,  and  dif- 
ferentiated from,  the  genera  Xynwne.  which  is  restricted  to 
New  Zealand,  and  Trophon.  with  which  it  co-occurs  in  the 
Magellanic  province. 

Key  words:  Neogastropoda,  Trophoninae,  Patagonia,  Magel- 
lanic Province. 


INTRODUCTION 

The  genus  Xymenopsis  encompasses  a  group  of  small, 
conchologicaUy  variable  muricid  gastropods  endemic  to 
the  Magellanic  Prox-ince.  These  shell-drilling  predators 
commonly  co-occur  wdth  species  of  Trophon  on  mussel 
beds,  and  range  from  the  intertidal  zone  to  depths  ex- 
ceeding 100  meters.  The  majority  of  the  published  Bt- 
erature  on  this  group  dates  from  the  early  era  of  Ant- 
arctic and  subAntarctic  exploration,  and  has  concentrat- 
ed on  describing  numerous,  minor  phenotypic  variants 


as  species.  The  biologv'  ot  these  animals  has  not  previ- 
ously been  stuched. 

Powell  (1951)  erected  the  genus  Xymenopsis  to  dis- 
tinguish Patagonian  muricids  with  a  conical,  multi- 
whorled  protoconch  and  rounded  ;mal  ribs  from  those 
with  a  rounded,  paucispiral  protoconch  and  numerous, 
lamellose  varices  that  belong  to  the  genus  Trophon.  He 
attributed  30  nominal  taxa  to  Xymenopsis,  but  recog- 
nized that  many  would  prove  to  be  synonyms,  stating 
that  "An  evaluation  of  these  names  is  not  possible  with- 
out recourse  to  the  tyj^e  material  which  is  in  European 
museums."  Imphcit  in  the  generic  name  Xymenopsis  is 
Powell's  recognition  of  the  close  affinity  of  this  Magel- 
lanic group  to  Xymenc  Iredale,  1915,  which  is  endemic 
to  New  Zealand.  Powell  (1951)  noted  that  the  shell 
sculpture  of  Xymenopsis  was  similar  to  that  of  Xyniene, 
but  that  its  protoconch  resembled  that  of  Zcatroplwn 
Finlay,  1927,  another  New  Zealand  genus. 

In  his  revision  of  New  Zealand  muricids  assigned  to 
the  genus  Trophon,  Finlay  (1926)  proposed  several  su- 
praspecific  taxa,  each  representing  what  he  regarded  to 
be  a  hneage  or  "natural  grouping."  He  divided  these  taxa 
into  2  groups,  based  primarily  on  protoconch  morphol- 
ogy. One  group,  defined  on  the  basis  of  having  a  sym- 
metrically conical  protoconch  of  >  2  whorls  and  a  small 
nucleus,  contained  the  genera  Xymenc  Iredale,  1915, 
XymcncUa  Finlav,  1926,  and  Zcatrophon  Finlaw  1927. 
The  second  group,  recognized  by  having  an  asymmetri- 
cal, rounded  protoconch  of  1-2  whorls,  with  a  large,  lat- 
eral nucleus,  included  Axymene  .scvi.si/  stricto  Finlay, 
1926,  Axymcnc  (Lcnitrophon)  Finlay,  1926,  Comptella 
Finlay,  1926,  Tcrcfundus  scnsit  stiicfo  Finlay,  1926,  and 
Terefiindus  iMinoiirophon)  Finlay,  1926. 

Ponder  (1972)  rexdewed  the  Recent  and  fossil  New 
Zealand  species  related  to  Xymenc.  Based  on  an  analysis 
of  gross  anatomy,  shell,  and  radular  moqihology,  he  con- 
cluded that  the  taxa  Axymene,  Lenitrophon,  XymeneUo, 
Zcafroplion,  and  probably  Vc.5flri(//fl  Finlay,  1926,  did  not 
merit  taxonomic  recognition  even  at  the  subgeneric  lev- 
el, and  reduced  all  but  Vesatuda  to  .svmonyms  oi Xymenc. 
However,  he  regarded  Xymenopsis  sufficiently  distinct  to 


G.  Pastorino  and  M.  G.  Harasewvch,  2()()0 


Page  39 


Table  1.  Tiixa  attributed  to  the  genus  Xijiiu'iiopsis  by  Powell  (1951)  and  Dell  (1971,  1972).  Taxa  listed  ni  bold  are  valid  species. 


Troplion  iiciiininntus  Strebel,  1904 

Trcphon  (dints  Strebel,  1904 

Fiisiis  cilhithis  Philippi,  1845 

Tniplioti  bnitti  Strebel,  1904 

Xymenopsis  buccineus  (Lamarck,  1816) 

Troplion  canccllarioidi's  Ree\e,  1847 

Xymenopsis  canccllinns  (Philippi,  1S45) 

Fiisiis  candidatiis  Rochebnnie  and  Mabille,  1889 

Xymenopsis  corrttgattts  (Reeve,  1S4S) 

Troplion  coiithoiii/i  Strebel,  1904 

Fiisiis  decolor  Philippi,  1845 

Fiisiis  dispar  Rochebnine  and  Mabille,  1889 

Troplion  elcfians  Strebel,  1904 

Troplion  elongatiis  Strebel,  1904 

Trojihon  fnlklandiciis  Strebel,  1908 

Troplion  fenestratiis  Strebel,  1904 

Troplion  hoijlei  Strebel,  1904 

Fiisns  jarqiiinoti  Philippi,  18.55 

Fiisiis  lebnini  Mabille  and  RochebiTnie,  1889 

Fiisiis  liratiis  Gould,  1849 

Fiisiis  loebbeekei  Kobelt,  1878 

Xymenopsis  muriciformis  (King  and  Broderip,  1832) 

Troplion  obesiis  Strebel,  1904 

Troplion  ornatiis  Strebel,  1904 

Trophon  paessleri  Strebel,  1904 

Troplion  paessleri  turrita  Strebel,  1904 

Fiisiis  pliimbeiis  Gould,  19.52 

Trophon  pseiidoelonoatiis  Strebel,  1904 

Trophon  ringei  Strebel,  1904 

Fiisiis  roseiis  Honibron  and  Jacquinot,  1854 

Trophon  standeni  Strebel,  1904 

Xymenopsis  subnodosus  (Gray,  1839) 

Fiisiis  textdiosiis  Honibron  and  Jacquinot,  1854 

Fiisiis  violaceiis  Mabille  and  Rochebnnie,  1889 


(syiiony-m  of  Xipnenopsis  nuiricifoniiis  < 
(synonym  of  Xiimenopsis  iniirieifonnis) 
(synonyiii  ol  Xipm-nopsis  hiiriineiis) 
(.synonym  ol  Xiinuiu>psis  rornio/itiis) 

(synonym  ot  Xt/menopsis  iniiririfonnis) 

(synonym  of  Xymenopsis  subnodosus) 

(belongs  in  the  buccinid  genus  Fareuthria  Strebel,  1905) 

(synonym  of  Xynwnopsis  muririfonnis) 

(synonym  of  Xymenopsis  muririfonnis) 

(belongs  to  different  niuricid  genus,  not  Xymenopsis) 

(synonym  ot  Xymenopsis  miiricifonnis) 

(synonym  of  Xymenojisis  muriciformis) 

(synonym  o(  Xymenopsis  corrugatiis) 

(synonym  of  Xymenopsis  muriciformis) 

(synonym  of  Xymenopsis  cornigatus) 

(new  name  for  F.  textiliosus  Honibron  and  Jacquinot.  1854,  not 

F.  textiliosus  Deshayes.  18.35) 
(synonym  of  Xymenopsis  miiricifonnis) 
(syiionyiii  of  Xymenopsis  miiricifonnis) 
(sviionym  of  Xymenopsis  muriciformis) 

(synonym  of  Xymenopsis  miiricifonnis) 

(synonym  of  Xymenopsis  cornigatus) 

(synonym  of  Xymenopsis  muriciformis) 

(synonym  of  Xymenopsis  muriciformis) 

(belongs  in  the  buccinid  genus  Fareuthria  Strebel,  1905) 

(synonym  of  Xymenopsis  nuiriciformis) 

(synonym  of  Xymenopsis  buccineus) 

(belongs  in  the  buccinid  genus  Pareiithria  Strebel,  1905) 

(synonym  of  Xymenopsis  cornigatus) 

(synonym  of  Xymenopsis  buccineus) 
(synonym  of  Xymenopsis  miiricifonnis) 


be  treated  as  a  subgenus  of  Xijmene.  Despite  reducing 
their  taxonomic  rank.  Ponder  (1972)  recognized  5  dis- 
tinctive groups  within  the  Xijmenc  complex,  correspond- 
ing to:  1)  Xymene  +  Xijinenclla;  2)  Zcatrophon;  3)  a 
group  consisting  of  2  fossil  species  (1  Eocene,  1  Mio- 
cene) with  features  expected  in  the  "ancestor  of  all  the 
species  of  Xymene";  4)  Axymciw;  and  5)  Xi/menopsis. 

Beu  and  Maxwell  (1990:39)  reported  Xymene  to  range 
from  the  Kaiatan  (=  Bartonian,  Late  Eocene)  to  the 
Recent  in  New  Zealand.  The  fossil  record  of  Xymenopsis 
is  thus  far  limited  to  4  species  recently  described  from 
the  Late  Miocene  Entrerriense  Formation  along  the  Val- 
des  Peninsula  in  northern  Patagonia,  Argentina  (Bninet, 
1997).  While  there  is  little  doubt  that  Xymene  and  Xy- 
menopsis are  closely  related,  these  lineages  are  treated 
here  as  separate  genera. 

In  the  present  study,  we  review  and  discuss  all  Recent 
nominal  species  that  have  been  included  in  Xymenopsis 
(Table  1)  based  on  an  examination  of  available  type  ma- 
terial. Type  specimens  of  all  taxa  referable  to  Xt/men- 
opsis that  could  be  located  are  illustratetl,  some  for  the 
first  time.  Data  on  shell  ultrastructure,  radular  mor- 
phology, and  gross  anatomy  are  provided  for  Xymenopsis 


miiricifonnis,  the  type  species  oi  Xymenopsis.  Compar- 
ative data  are  provided  for  Xymene  plcbeiiis,  the  type 
species  of  Xymene,  and  for  the  other  Recent  species  of 
Xymenopsis. 

MATERIALS  AND  METHODS 

The  majority  of  specimens  examined  in  this  study  are  in 
the  collections  of:  the  National  Museum  of  Natural  His- 
tory, Smithsonian  Institution,  Washington,  DC  (USNM); 
Museo  de  La  Plata,  La  Plata,  Argentina  (MLP);  Museo 
Argentino  de  Ciencias  Naturales  "Bernardino  Rivada- 
via",  Buenos  Aires  (MACN);  and  the  Los  Angeles  Coun- 
ty Museum  of  Natural  History  (LACM).  Type  material 
is  housed  in  the  following  museimis:  The  Natural  His- 
tory Museum,  London,  (BMNH);  Zoologisches  Institut 
und  Zoologisches  Museum  der  Universitat  Hamburg, 
(ZMH);  Swedish  Museum  of  Natural  History,  Stock- 
holm, (NHRM);  Museum  national  d'Histoire  naturelle, 
Paris  (MNHN);  Museo  Nacional  de  Historia  Natural, 
Santiago,  Chile  (MNHNS);  and,  Loebbecke- Museum 
und  Aquazoo,  Dusseldorf,  (LM). 

Dissections   were   performed   on   ethanol   preserved 


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THE  NAUTILUS,  Vol.  114.  No.  2 


G.  Pastorino  aiul  M.  G.  Harasewych,  2000 


Page  41 


specimens  to  studv  their  gross  anatomy,  with  emphasis 
on  the  nioiphologN'  of  the  anterior  alimentary  system, 
and  the  palli;il  portions  ol  the  male  ami  female  repro- 
ductiye  systems.  Radulae  were  prepared  according  to 
the  method  described  by  Solem  (1972)  and  observed 
using  a  LEO  440  scanning  electron  microscope  (SEM). 
Radular  terniinolog)  follows  Kool  (199.3a:  fig.  6B).  ,Shell 
ultrastnicture  data  were  procured  from  freshly  fractured 
collabral  sections  taken  from  the  central  portion  of  the 
lip  on  the  last  whorl  of  2  individuals  per  taxon,  whenever 
sufficient  material  was  available. 

Most  photographs  were  taken  using  a  Leaf  Lumina 
digital  scanning  camera.  Several  images  were  scanned 
from  black  and  white  .35  mm  negatives  using  a  Nikon 
Coolscan  III  slide  scanner.  All  images  were  processed 
with  the  software  Photoshop  4.01  and  5.02. 

SYSTEM  ATICS 

Class  Gastropoda  Cuvier,  1797 
Order  Neogastropoda  Wenz,  1938 
Family  Muricidae  Rafinesque,  1815 
Subfamily  Trophoninae  Cossniann,  1903 
Genus  Xymenopsis  Powell,  1951 

Xijinciiopsis  Powell,  1951:1.58;  Radwiii  and  D'Attilio,  1976:190. 

Xijnwne  iXtjmenopsis)  Ponder,  1972:474;  Yokes,  1991:10. 

Type  species:  Ftistts  Uratiis  Gould,  1.S49  (by  original  designa- 
tion), a  junior  synonym  of  Bticciiiuin  iniiricifonne  King 
and  Broderip,  18.32.  ' 

Diagnosis:  The  genus  Xi/incuopsis  contains  species 
characterized  by  a  small  to  medium-sized  (to  48  mm), 
fusiform,  high-spired  shell  with  a  tall,  conical,  multi- 
whorled  (==3  whorls)  protoconch  that  lacks  a  keel,  and 
a  teleoconch  with  a  rounded  shoulder,  sculpture  of 
rounded  axial  ribs  that  never  develop  lamellae,  and  nar- 
row to  broad  spiral  cords  that  may  or  may  not  produce 
beads  at  intersections  with  a.xial  ribs.  The  siphonal  canal 
is  short,  open,  and  broad.  Shell  ultrastructure  consists  of 
an  outer  amorphous  calcific  layer  and  an  inner,  crossed- 
lamellar  aragonitic  layer.  The  rachichan  tooth  of  the  rad- 
ula  has  a  basal  plate  that  is  simple  and  concave  anteri- 
orly, and  inner  lateral  denticles  that  are  completely  free 
from  the  central  and  lateral  cusps.  The  mantle  edge  is 
always  lobulated. 

Remarks:  In  his  treatment  of  the  Xi/mcne  complex. 
Ponder  (1972)  identified  the  oldest  known  member  of 
the  group  in  the  Late  Eocene  fauna  of  South  Island, 
New  Zealand.  He  recognized  that  Xi/inciw  apipagpdus 


Ponder,  1972,  characterized  by  a  small,  strongly  shoul- 
dered shell  with  ;L\ial  lamellae,  smooth  aperture,  and  a 
tall,  conical,  4y2-whorled  protoconch  with  a  sinuated  var- 
L\  indicative  of  a  pelagic  larval  stage,  had  features  from 
which  all  subsefjuent  Xi/mene  lineages  could  be  derived, 
and  suggested  that  several  lineages  evolved  from  this 
phenotype. 

One  hneage  compri.ses  the  Zcatrophoti  group,  which 
appeared  during  the  Late  Oligocene  {Xtfiiwrw  chatto- 
ncnsis  Ponder,  1972,  Duntroonian)  and  persists  in  the 
Recent  fauna  (e.g.,  Zcafrophou  amhitnius  Finlay,  1927, 
the  type  species  oi  Zcatwphon).  This  hneage  is  charac- 
terized l)y  a  tall,  strongly  shouldered  shell  with  axial 
nodes  or  weak  lamellae,  an  aperture  with  or  without  lir- 
ae,  and  a  protoconch  that  is  tall  and  conical,  yet  smaller 
and  with  fewer  (2%-3V4)  whorls  than  X  apipagodns. 
Within  this  hneage,  there  is  a  trend  for  a  spiral  keel  to 
develop  on  the  last  protoconch  whorl,  and  for  the  nu- 
cleus of  the  protoconch  to  increase  in  size.  A  second 
lineage  iXijmcnc  +  Xymcnclla  group)  also  appeared  in 
the  Late  OUgocene  (Trophon  lepidiis  Suter,  1917,  Dun- 
troonian) and  survives  in  the  Recent  fauna  (e.g.,  Ftistis 
pk'bciiis  Hntton,  1873,  the  tvpe  species  of  Xijmcnc,  and 
Trophon  pii.sillns  Suter,  1917,  the  type  species  of  X(/- 
iiicncUa).  Shells  are  small,  strongly  to  weakly  shouldered, 
with  strong  spiral  cords  and  axial  ribs,  but  without  la- 
mellae, and  with  apertures  that  are  usually  lirate.  Pro- 
toconchs  are  depressed,  with  a  trend  to  fewer  (2  whorls 
in  X.  plchcuis,  2  Vi  whorls  in  X.  gouldi  =  T.  piisiUus,  see 
Ponder,  1972),  more  inflated  whorls,  and  an  increase  in 
the  size  of  the  nucleus.  Another  hneage  (Axipnene 
group)  is  limited  to  Pliocene  and  Recent  species.  Shells 
tend  to  be  large,  strongly  shouldered,  with  axial  rilis  and 
spiral  cords,  but  without  lamellae,  and  generally  have 
lirate  apertiues.  Protoconchs  consist  of  1-2  roimded,  in- 
flated whorls,  with  a  large  nucleus. 

Ben  et  al.  (1997)  regarded  Xifmcnc  and  Xymcnclla  to 
be  distinct  genera,  and  reassigned  Triton  dauizcnhcrgi 
von  Ihering,  1897,  and  Urosalpinx  elegans  Ortmann, 
1900,  both  from  the  Monte  Leon  Formation  (Late  Oli- 
gocene-Early  Miocene)  of  Patagonia,  to  Xymcnc.  and 
Xymcnella  respectively.  Based  on  the  first  occurrences 
of  these  taxa  in  the  fossil  record,  they  concluded  that 
Xymcnclla  originated  in  New  Zealand  and  dispersed  to 
South  America  with  the  origin  of  the  Antarctic  Circum- 
polar  Current  at  the  time  the  Drake  Passage  opened 
(Late  Oligocene),  while  Xymene  evolved  in  South  Amer- 
ica and  dispersed  to  New  Zealand,  also  along  the  Ant- 


Figures  1-19.  Xi/incnopsis  luiiricifonnis  (King  and  Broderip,  18.32).  1-3.  Holotvpe  of  Fiisus  liratiis  Gould,  1S49,  USNM  5680, 
Orange  Harbor,  Tierra  del  Fuego.  4-6.  HoJotype  of  Trophon  loebheckei  Kobelt,  1878,  Loebbecke  Museum  unnumbered,  lacks 
locality  data.  7-8.  Lectot\pe  of  Trophon  violacetis  Mabille  and  Rochebmne,  1889,  MNHN  unnumbered,  Baie  Orange,  Tierra  del 
Fuego.  9-10.  Holotype  of  Fusiis  decolor  Philippi,  1845,  MNHNS  unnumbered.  Strait  of  Magellan.  11-12.  Lectotype  of  Trophon 
elegans  Strebel,  1904,  ZMH  unnumbered.  Port  Stanley,  Malvinas  (Falkland)  Is.  13.  Leetotype  of  Trophon  lebnini  Mabille  and 
Rochebrune,  1889.  MNHN  unnumbered,  Santa  Cruz,  Argentina.  14-16.  Leetotype  of  Biiccimim  niuricifonne  King  and  Broderip. 
1832,  BMNH  1837.12.15.2695,  Strait  of  Magellan.  17-19.  Leetotype  as  Bnrciniim  cnncrUnrioidcs  Reeve,  1847.  BMNH  19920.55, 
lacks  locality  data.  Scale  bar  =  1  cm  for  all  specimens. 


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THE  NAUTILUS,  Vol.  114,  No.  2 


G.  Pastorino  and  M.  G.  Harasewvch,  2000 


Page  43 


arctic  Circunipolar  (^iinciil,  diiriiit;;  (lie  Middle  Mio- 
cene. 

The  origin  of  the  Xt/incitc  complex  in  New  Zealand  is 
amplv  supported  In  tlic  fossil  record.  The  hvpothesized 
dispersal  ot  this  liiu'age  to  Southern  South  America  dur- 
ing the  Late  Oligocene  is  concordant  with  its  appearance 
in  the  fossil  record  of  the  region,  and  is  supported  bv 
the  prexalence  of  narrow,  polvgyrate  protoconchs,  inchc- 
ative  of  pelagic  lanal  stage  (see  Thorson,  19.50;  Shuto, 
1974;  Jablonsk)-  and  Lutz,  1980),  in  Paleogene  members 
of  this  complex.  Recent  and  fossil  species  of  Xijmenopsis 
are  most  similar  to  members  of  the  Xt/iiicuc  +  Xi/inc- 
nclla  group,  especially  to  early  members  of  this  lineage 
such  as  the  Oligocene  Xi/mcnc  Icpichis.  with  which  they 
share  a  sculptural  pattern  of  axial  ribs  and  spiral  cords, 
but  not  axial  lamellae,  and  a  conical,  .3  whorled  proto- 
conch  with  a  small,  apical  nucleus.  We  suggest  that  Xi/- 
meiwpsis  represents  a  radiation  endemic  to  the  Magel- 
lanic region  from  ancestors  of  the  Xijmcne  +  Xi/iucnclla 
group  that  dispersed  as  larvae  to  Patagonia  tluring  the 
Late  Oligocene. 

While  it  is  possible  that  multiple  lineages  were  intro- 
duced to  South  America,  or  that  1  lineage  subsequently 
dispersed  back  to  New  Zealand  while  becoming  extinct 
in  South  America,  evidence  for  such  hypothe.ses  is  more 
tentative  or  lacking.  Xi/mcnc  are  characterized  bv  a  trend 
to  protoconchs  with  fewer,  more  roimded  whorls  that 
are  indicative  of  lecithotrophic  rather  than  planktotroph- 
ic  development.  This  would  argue  against  their  abihty  to 
disperse  as  plank-tonic  laivae  from  South  America  east- 
ward on  the  Antarctic  Circunipolar  Current  to  New  Zea- 
land. Clearly,  a  rigorous  re-assessment  of  the  Xymene 
complex  in  a  phylogenetic  context  would  resolve  many 
of  the  systematic  and  biogeographic  questions  that  have 
plagued  this  group. 

Xijincnopsis  lack  the  strongly  shouldered  shell  of  the 
Xymene  +  Xymenella  group.  While  strong  apertural  lirae 
are  prevalent  in  Xi/mcnc,  they  are  rare  in  Xymcuopsis. 
occurring  only  in  the  X.  .siihnodosus  (Gray,  1839).  X(/- 
mcnop.sis  may  also  be  distinguished  from  Xtpticitc  on  the 
basis  of  shell  ultrastiiicture,  radular  moiphology  and  by 
features  of  their  mantle  edge  and  egg  capsules.  Xymen- 
opsis  (figures  45,  79,  80)  has  a  thick  (>  Vi  shell  thick- 
ness) outer  layer  of  calcite,  and  a  single  layer  of  crossed- 
lamellar  aragonite  in  which  the  crystal  faces  are  comar- 
ginal.  In  Xymene  (figure  50),  the  calcitic  layer  is  thin, 
the  co-marginal  layer  of  aragonite  thickest,  and  an  in- 
nermost layer  of  crossed-lamellar  aragonite,  with  civstal 
faces  perpendicular  to  those  of  the  co-marginal  layer,  is 


also  pre.sent.  The  radula  i)^  Xymene  pleheiiis  has  rachi- 
dian  teeth  (figure  51)  that  ari>  characteristically  chevron- 
shaped,  eac'h  with  its  apex  at  the  mid-point  of  the  an- 
terior (>dge  of  the  basal  plate,  which  has  a  concave  pos- 
terior edge.  Rachidian  teeth  oi'  Xymcuopsis  (figures  48- 
49,  75-76,  81-82,  102-103)  have  a  straight  to  sfightly 
concave  anterior  edge,  and  a  convex  posterior  edge  of 
the  basal  plate.  The  mantle  edge  of  Xymene  is  smooth, 
while  the  mantle  edge  oi'  Xymenopsis  is  lobulated.  While 
egg  capsules  ui' Xymene  (figure  43)  and  Xymenopsis  (fig- 
ure 42)  are  similar,  the  exit  aperture  is  apical  in  Xymene, 
Init  is  situated  at  the  base  of  a  shallow  depression  in 
Xymenopsis. 

There  are  no  direct  observations  on  development  in 
Xymenopsis.  Shuto  (1974)  reported  that  the  ratio  of  the 
protoconch  chameter  (D)  to  the  number  of  whorls  (V) 
is  a  reasonable  estimator  of  the  t\pe  of  development, 
and  that  a  ratio  below  0.3  is  characteristic  of  plankto- 
trophic  larvae,  especially  when  the  number  of  whorls  is 
s  3.  Both  Xymenopsis  muriciformis  (figure  41)  and  X. 
huecineus  (figures  77-78)  have  narrow,  conical  proto- 
conchs of  3  whorls.  The  DA'  ratios  for  these  species  are 
0.23  and  0.21  respectively,  predicting  that,  unlike  Xy- 
mene, Xymenopsis  retains  planktotrophic  larval  devel- 
opment. 

Powell  (1951)  provisionally  included  the  species  Fusus 
rosens  Hombron  and  Jac(juinot,  1854  and  Trophon  ean- 
didatiis  RochebiTine  and  Mabille,  1889  in  his  genus  Xy- 
menopsis. Ceniohorsky  (1977)  subsequently  transferred 
Fusus  ro.seus  to  the  buccinid  genus  Pareuthria  Strebel, 
1905.  He  proposed  the  new  name  Pareuthria  powelli  for 
this  species  because  Fusus  roseus  Hombron  and  Jac- 
quinot,  1854  was  preoccupied  by  Fusus  roseus  Anton, 
1838.  The  type  material  of  these  taxa  was  examined  by 
the  senior  author  at  the  Museum  national  d'Histoire  na- 
turelle,  and  both  are  clearly  referable  to  the  buccinid 
genus  Pareuthria.  The  type  material  of  Fusus  dispar 
Hombron  and  Jacquinot,  1854  was  also  examined 
(MNHN).  This  taxon  has  affinities  with  the  Antarctic 
Trophoninae,  but  does  not  belong  to  the  genus  Xymen- 
opsis. 


Xymenopsis  muriciformis  (King  and  Broderip,  1832) 
(Figures  1-42,  45-49,  83-95,  105-106) 

Biicciniiin  miiricifonne  King  and  Broderip,  18.32:.348. 
Fusus  muricifonuis  King  and  Broderip — Gray,  18.39:118. 
Fusus  huecineus  Sowerby  in  Gray.  18.39:15.5,  pi.  .36,  fig.  12. 
Fusus  decolor  Pfiilippi,  1845:68;  1846:118,  pi.  3,  fig.  3;  Hupe 


Figures  20-40.  Xymenopsis  muriciformis  (King  and  Broderip,  1832).  20-22.  Holot\pe  of  Trophon  ncuniinntus  Strebel.  1904, 
ZMH  unnumbered.  Punta  Arenas.  Strait  of  Magellan.  23-25.  Lectotype  of  Trophon  couthoui/i  Strebel,  1904,  ZMH  unnumbered. 
Port  Grappler,  Smyth  Gliannel.  26-28.  Lectotype  of  Trophon  paessleri  Strebel,  1904.  (specimen  coated  with  ammonium  cliloride), 
ZMH  unnumbered.  Port  Grappler,  Sm\th  Channel,  29-.31.  Holotvpe  oi'TropIion  pscudoehinaatus  Strebel.  1904,  ZMH  unnumbered, 
Ushuaia.  32-34.  Lectotype  olTroplion  ohciiis  Strebel,  1904,  ZMH  unnumbered,  Punta  Arenas,  Chile.  35-37.  Leetot^-pe  of  Troplwn 
elongatus  Strebel,  1904,  ZMH  unnumbered,  Puerto  Angosto.  Strait  of  Magellan.  38-40.  Holotyjie  of  Trophon  albus  Strebel,  1904. 
ZMH  unnumbered,  Ushuaia.  Scale  bar  =  1  cm  for  all  specimens. 


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THE  NAUTILUS,  Vol.  114,  No.  2 


G.  Pastorino  and  M.  G.  Harasewych,  2000 


Page  45 


in  Gav,  1854;162;  Homhroii  and  |a<.(|uin()t  in  Rousseau, 

1854:i08,  pi.  2.5,  figs.  6-8;  Kolx-lt',  1878:291,  pi.  74,  figs. 

5-6;  Rochebrune  aiid  Mabille,  1889;  H.54. 
Buccinum  cancellarioicles  T{ee\e,  lS47:pl.  14,  fig.  11.3. 
Fusiis  liratus  Gould,  1849:141. 
Trophon   liratus  Couthous- — Kobelt,    1878:.31();  Tryon,    1880: 

14.3,  pi.  31,  fig.  3.33;  Watson  1886:165;  Lamv.  1906:3. 
Trophon   Locbbcckci   Kobelt,    1878:294,   pi.   74, 'figs.    13,   14; 

TnoH,  1880:145,  pi.  31.  fig.  .3.35. 
Trophon  nuiricifonnis  King — Tnon.  1880:145. 
Trophon  Icbnini  Mabille  and  Rochebnnie  in  Rochebnine  and 

Mabille,  1889:  H..55. 
Trophon  violaceus  Mabille  and  Rochebnine  in   Rochebnine 

and  Mabille,  1889:  H.56,  pi.  2,  fig.  1. 
Trophon  paessleri  Strebel,  1904:213.  pi.  7,  figs.  56a-f. 
Trophon  paessleri  \'ar.  turrita  Strebel,  1904:215,  pi.  7,  fig.  57. 
Trophon  elon^atus  Strebel,  1904:217,  pi.  7,  figs.  .58a-f. 
Trophon  pseudoelong,atiis  Strebel,  1904:220,  pi.  7,  figs.  60a,b. 
Trophon  albus  Strebel,  1904:221,  pi.  7,  fig.  61. 
Trophon  acuminatus  Strebel,  1904:222,  pi.  7,  fig.  63. 
Trophon  obesus  Strebel,  1904:223,  pi.  7,  fig.  62. 
Trophon  fenestrattis  Strebel,  1904:225,  pi.  7,  fig.  59a-d. 
Trophon  couthoinji  Strebel,  1904:236,  pi.  7,  fig.  65a-e;  Pi.  8, 

fig.  76. 
Trophon  elegans  Strebel,  1904:241,  pi.  8,  fig.  71. 
Xt/menopsis  hratiis  (Gould) — Powell,   1951:158;  Radwin  and 

D'Attilio,  1976:190,  fig.  1.36. 
Xijmenopsis  decolor  (Philippi) — Dell.  1971:211. 
Xtjmenopsis  muricifonnis  (King  and  Brodenp) — Dell,  1972:8, 

figs.  31,  .32;  Yokes,  1991:  7,  unnumbered  fig.;  Yokes,  1992: 

3,  figs.  8a-c;  Castellanos  and  Landoni,  1993:16,  pi.  3,  figs. 

.39,  40C,  D. 

Diagnosis:  Axial  sculpture  of  irregular,  rounded  ribs, 
16-20  on  body  whorl.  Spiral  sculpture  of  flat  cords  (4— 
6  on  early  whorls,  14-20  on  body  whorl),  much  broader 
than  intervening  spaces,  which  appear  as  incised  fur- 
rows. 

Description:  Shell  large  tor  genus  (reaching  48  mm), 
fusiform,  slender,  chalky.  Protoconch  of  3  tall,  conical, 
whorls.  Teleoconch  of  up  to  6  weakly  shouldered  or 
rounded  whorls.  Spire  high,  =  V2  shell  length,  spire  an- 
gle ~  45°.  Suture  deeply  impressed;  subsutural  ramp 
narrow  in  early  whorls,  then  absent.  Aperture  subovoi- 
dal;  outer  lip  rounded,  anterior  half-reflected;  inner  Up 
curved,  adpressed;  interior  glossy  pinkish  to  brownish. 
Siphonal  canal  short  (<  V2  aperture  length),  obhcjue, 
open.  Umbilicus  absent.  Axial  sculpture  of  irregular, 
rounded  ribs,  16-20  on  body  whorl.  Spiral  sculpture  of 
flat  cords  (4-6  on  early  whorls,  14-20  on  body  whorl), 


much  broader  than  intei-v'ening  spaces,  which  appear  as 
incised  furrows.  (Growth  lines  irregular,  faint  to  obsolete 
throughout  shell.  Shell  composed  of  2  layers:  outer  layer 
thick  (0.6.5  slicll  thickness),  of  amoqihoiis  calcitc;  inner 
layer  (0..35  shell  thickness),  of  crossed-lamellar  aragouite 
with  crystal  planes  oriented  perpendicular  to  growing 
edge.  Operculum  (figure  47)  oval,  subpolvgonal,  v\ath 
terminal  nucleus.  External  surface  covered  bv  irregular 
growth  hues.  Inner  surface  with  .3—4  horseshoe-sliapt'd 
lines;  posterior  rim  with  slight  callus,  glazed  in  adult  and 
young  specimens. 

Aniiual  large.  Gephalic  tentacles  medium  in  size, 
blunt,  thick,  with  small  black  eyes.  Mantle  edge  with  flat, 
rounded  lobules  (figure  S3,  1).  Siphon  (figure  83,  s)  of 
medium  length.  Accessory  boring  organ  (figure  95,  abo) 
situated  along  ventral  midline  of  foot,  shares  opening 
with  ventral  pedal  gland  in  females.  Osphradium  (figure 
84,  os)  <  V2  ctenidium  length,  thin,  veiy  asymmetrical, 
with  45-50  leaflets  per  side.  Ctenidium  (figure  84,  ct) 
3-4  times  as  wide  as  osphradium,  with  120-135  trian- 
gular leaflets.  Pleuroembolic  proboscis  very  long,  broad. 
Radular  ribbon  (figures  48^9)  small,  short  (0.31  X  ap- 
erture length  vs.  0.78  X  aperture  length  in  Trophon  gev- 
ersianus),  extends  just  beyond  rear  of  buccal  mass  (fig- 
ure 94,  ra).  Rachidian  tooth  wide  (to  230|xm),  central 
cusp  large,  needle-shaped,  lateral  cusps  outwardly  point- 
ed, shorter  than  central  cusp,  inner  lateral  denticle 
smaU,  thin,  separate  from  lateral  cusps.  Basal  plate  of 
rachidian  tooth  rectilinear  or  gently  curved,  with  low, 
broad,  marginal  areas.  Lateral  teeth  large,  each  with  sin- 
gle, long,  scythe-shaped  cusp  and  broad  basal  plate.  Sal- 
ivary glands  (figure  94,  sg),  ascinous,  white  (yeUow  in 
fixed  animals)  veiy  large,  flanking  retracted  proboscis. 
Salivary  ducts  join  esophagus  just  anterior  to  valve  of 
Leiblein  (figure  94,  vl).  Accessory  sahvars'  glands  (figure 
94,  asg)  small,  pyriform,  white  to  brownish,  situated  be- 
low salivary  glands.  Glande  framboisee  absent.  Gland  of 
Leiblein  (figure  94,  gl)  large,  broad  anteriorlv,  tapering 
posteriorly  to  form  long,  sinuous,  blind  duct  with  small, 
terminal  ampulla  (figure  94,  dgl).  Anterior  aorta  (figure 
94,  aa)  passes  through  fold  in  gland  of  Leiblein  en  route 
to  buccal  mass.  Stomach  (figure  94,  sto,  89)  U-shaped, 
small,  almost  completely  enveloped  bv  digestive  gland. 
Both  digestive  gland  ducts  (figure  89,  dd)  near  esopha- 
geal opening,  situated  in  deep  pouch  (figure  89,  dp), 
separated  from  posterior  mbdng  area  (figure  89,  pma)  by 
small  horizontal  fold  (figure  89,  hf).  Two  tyj^hlosoles  (fig- 


Figures  41—42,  45-49.  Xijmenopsis  muricifonnis  (King  and  Broderlp,  1832).  41.  Protoconch,  USNM  901629,  Strait  of  Magellan, 
53°.39.4'S-70°55.5'W,  in  24  m.  Scale  bar  =  100  (xm.  42.  Apical  and  lateral  views  of  the  egg  capsule.  Scale  bar  =  800  [im.  45. 
Shell  ultrastructure,  fracture  surface  comarginal.  Scale  bar  =  30  \x.m.  46.  Penis,  critical-point  dried.  Scale  bar  =  40  |xni.  47. 
Operculum,  external  (left)  and  internal  (right)  \iews.  Scale  bar  =  1  cm.  48-49.  Radula.  USNM  9016.35,  Strait  of  Magellan, 
5.3°39.8'S-70°54.9'W,  in  27-73  m.  48.  Dorsal  \new  of  radular  ribbon.  Scale  bar  =  40  fxm.  49.  Lateral  \iew  of  rachidian  teeth.  Scale 
bar  =  20  (jim.  Figures  43-44,  50-51.  Xi/ineiw  picbeius  (Hutton,  1873).  43.  Lateral  and  apical  \iews  of  egg  capsule,  USNM 
6815.38,  Spit  Beach,  Dunedin  Harbor,  New  Zealand.  Scale  bar  =  1  mm.  44.  Operculum,  external  (left)  and  internal  (right)  \iews, 
same  specimen  as  43.  Scale  bar  =  1  cm.  50.  Shell  iiltrastnicture,  fracture  surface  comarginal.  AK  80295,  Northland,  Bay  of  Islands, 
Parekuta  Bay,  New  Zealand.  Scale  bar  =  100  |xm.  51.  Dorsal  view  of  radular  ribbon,  same  specimen  as  43.  Scale  bar  =  20  (jim. 


Page  46 


THE  NAUTILUS.  Vol.  114,  No.  2 


Figures  52—36.  Xi/menopsis  bticcinetis  (Lamarck,  1816).  52-54.  Neotvpe  of  Fiistis  biiccincus  Lamarck,  L816.  USNM  870410, 
5.3°.32'S,  64°.57'\\',  in  119-124  m,  55-56.  Holohpe  of  Ftistis  texiiliostis  Hombroii  and  |ac(juinot,  1854,  MNHN  unnumbered.  Strait 
of  Magelkui,  Figures  57-71.  Xi/rnenopsis  comioatus  (Reeve,  1848).  57-59.  Holot\pe  of  Fusus  cornigatiis  Reeve,  BMNH 
1874. 12. 11. 15.3,  lacks  localits  data.  60-62.  Lectohpe  of  Trophon  hoijlei  Strebel,  1904,  ZMH  mmumbered.  Port  Stanlev,  .VlaKinas 
(Falkland)  Is.  63-65.  Holotvpe  of  Trophon  falklandicus  Strebel,  1908,  NHRM  1049,  5r35'S-57°56'\\',  Islas  MaMnas '(Falkland). 
66-68.  USNM  368350,  Port' Stanley,  MaKinas  (Falkland)  Is.  69-71.  Holotvpe  of  Trophon  ornatns  Strebel,  1904,  ZMH  unnumbered, 
Port  Stanley,  Malvinas  (Falkland)  Is.  Figures  72-74.  Xymrne  plebeius  (Hiitton,  1873).  USNM  681538.  Spit  Beach,  Dunedin  Harbor, 
New  Zealand.  Scale  bar  =  1  cm  for  all  specimens. 


G.  Pastorino  and  M.  G.  Harasew\'ch,  2000 


Page  47 


Figures  75-79.  Xi/iiii-nopsis  biiccinciis  (Lamarck,  1S16).  75.  Dorsal  view  ol  radnlar  nbhoii  from  specimen  in  figures  52-54.  Scale 
bar  =  40  p-ni.  76.  Oblique  (45°)  lateral  Wew,  of  radular  ribbon,  MACN  12553,  Tierra  del  Fuego.  Scale  bar  =  40  |jim.  77-78. 
Protoconch,  MACN  25012,  54°26'S-64°53'W,  in  112  m.  77.  Apical,  and  78.  lateral  views.  Scale  bars  =  ISO  jxm.  79.  Shell  ultra- 
structure,  fracture  surface  comarginal.  Scale  bar  =  40  |j.m.  Figures  80-82.  Xi/menopsis  comigatus  (Reeve,  1848).  80.  Shell 
ultrastnicture,  fracture  surface  comarginal.  USNM  368350,  Port  Stanlev,  MaKinas  (Falkland)  Is.  Scale  bar  =  40  |j,m.  81-82.  Radula. 
USNM  421884,  Port  W'iUiams,  York  Bay,  MaKinas  (Falkland)  Is.  81.  Dorsal,  and  82.  obliijue  lateral  view.  Scale  bars  =  20  (xm. 


Page  48 


THE  NAUTILUS,  Vol.  114,  No.  2 


G.  Pastorino  and  M.  G.  Harasewvcli,  2{)0() 


Page  49 


nre  S9,  tl,  t2)  flank'  intestinal  i^rooN-e  (figure  89,  ig)  sep- 
arating it  ironi  expantled  eonipacting  area  (figure  89,  ea). 
Intestine  (figures  89,  94,  i)  runs  anteriorly  alongside  pal- 
lial  gonoduct,  forming  rectum  with  cylindrical  papilla 
(figin-es  86,  90  ap)  o\'er  anus  (figiu'es  86,  90  a).  Rectal 
gland  (figure  93,  rg)  small,  inconspicuous,  confined  to 
anterior  region  ot  rectum.  Prostate  gland  (figures  86-88, 
pg)  rvms  alongside  rectum  in  mantle  cavity,  lacking  open- 
ings to  mantle  cavitv"  (unlike  Trophoii  <yi:crsianus,  see 
Kool,  199.3h:  fig.  57).  Vas  deferens  (figure  86,  vd)  leads 
from  anterior  end  of  prostate  to  base  of  penis  (figures 
46,  83,  p).  Penis  large  (>  3  X  tentacle  length),  wide, 
elongate,  flattened  to  semitriangular  in  cross-section, 
with  centrallv  situated  sperm  duct  (figure  85,  pvd),  dor- 
sal iilood  sinus  (figure  85,  ds).  Penis  tapers  ahruptly  to 
form  small,  cylindrical  papilla  (figures  46,  83).  Pallial  oyi- 
duct  broad  posteriorly,  tapered  anteriorly.  Albumen 
gland  (figure  90,  ag)  situated  at  rear  of  mantle  cavity, 
joins  posterior  of  capsule  gland  (figure  90,  eg).  Left  and 
right  lobes  of  capsule  gland  similar  to  T.  ^eversianus 
(Harasewych,  1984:  fig.  22;  Kool,  1993b),  Nucella  lapil- 
lits  (Fretter  and  Graham,  1994:  fig.  171),  but  ventral  and 
dorsal  glandular  areas  proportionally  much  smaller  (fig- 
ure 92).  Bursa  copulatri.x  (figures  91,  93,  be)  joins  an- 
terior of  capsule  gland,  leads  to  vaginal  opening  (figures 
90,  91,  vg),  situated  below,  posterior  to  anal  opening, 
papilla  (figure  90). 

Egg  capsules  of  Xi/mcnopsis  Duihciftinnis  were  dis- 
cussed but  not  illustrated  by  D'Asaro  (1991:54).  Cap- 
sules are  attached  to  the  dorsal  surface  of  empty  shells 
of  X.  miiricifonnis,  densely  grouped,  overlapping  each 
other.  Egg  capsule  (figure  42)  bulliform,  approximately 
6-7  mm  in  diameter,  2  mm  high,  outline  subcircular, 
may  vary  according  to  substrate.  Preformed  exit  aperture 
oval,  situated  at  center  of  dorsal  surface  in  shallow,  sub- 
ovate  depression,  sealed  by  a  mucus  plug  until  hatching. 
Suture  between  capsule,  plug  weakly  defined,  nmning 
parallel  to  large  axis  of  depression.  Pattern  of  non-con- 
centrically  oriented  fibers  visible  on  upper  surface  of 
capsules.  External  morphology  of  egg  capsules  veiT  sim- 
ilar to  that  observed  in  dry  specimens  of  Xi/mcnc  plc- 
bciits  (figure  43),  which  differ  in  having  aperture  situated 
apically  rather  than  in  depression. 


Type  material:  iHiirriniiin  iiiuricifoniw]  Lectotvpe 
(figures  14-16),  BMNII  1837.12.15.2695,  6  paralecto- 
t>pes,  BMNII  1992056,  all  from  Strait  of  Magellan:  [Fii- 
sii.s  decolor]  Ilokitype  (figures  9-10),  MNHNS  unnum- 
bered. Strait  of  Magellan;  [Buccinum  caiiccllarioklcs] 
Lectot\pe  (figures  17-19),  BMNII  1992055-1,  here  des- 
ignated, I  paralectotype  BMNH  1992055-2,  lack  locality 
data;  [Fusus  liratus]  Holotype  (figures  1-3),  USNM 
5680,  Orange  Harbor,  Tierra  del  Fuego:  [Trophon  loeh- 
bcckci]  II()Iot\pe  (figures  4—6),  Lobbecke-Museum,  no 
catalog  ninnber,  lacks  locafity  data;  [Tropliou  Ichnini] 
Lectotype  (figure  13),  MNHN  unnumbered,  here  des- 
ignated, 2  paralectotypes,  all  from  Santa  Cruz,  Argenti- 
na; [Trophon  violaceiis]  Lectotype  (figures  7-8),  MNHN 
unnumbereil,  figured  .syntyjae  here  designated  as  lecto- 
type, from  Bale  Orange,  Tierra  del  Fuego;  [Trophon 
paessleri[  Lectotype  (figures  26-28),  ZMH  unnumbered, 
here  designated,  4  paralectotypes,  all  from  Port  Grap- 
pler,  Smvih  Channel,  Strait  of  Magellan;  [Troi)hon  paes- 
sleri  var.  turiita]  Original  tspe  material,  from  Cape 
Horn,  Tierra  del  Fuego,  was  destroyed  (Hausdorf  hi 
Utt.).  The  lectotvpe  of  Bucchmm  nuiiiciformc  (BMNH 
1837.12.15.2695)  is  here  designated  as  neotype  of  Tro- 
plion  pac.'islcri  var.  titrrita.;  [Trophon  elon^iatus]  Lecto- 
tvpe (figures  35-37),  ZMH  unnumbered,  here  designat- 
ed, from  Puerto  Angosto,  Strait  of  Magellan,  4  paralec- 
totypes, 1  from  Navarino  Is.,  Chile,  1  from  Puerto 
Bueno,  1  from  Eden  Harbor,  1  from  Puerto  Angosto, 
Strait  of  Magellan:  [Trophon  pseudoclongatns]  Holotype 
(figures  29-31),  ZMH  unnumbered,  Ushuaia;  [Trophon 
alhus]  Holotvpe  (figures  38—40),  ZMH  unnumbered,  Us- 
hutiia;  [Troplion  acuinimitiis]  Holot\pe  (figures  20-22), 
ZMH  unnumbered,  Punta  Arenas,  Strait  of  Magellan; 
[Trophon  obcsus]  Lectotype  (figures  32-34),  ZMH  un- 
ninubered,  here  designated,  from,  Punta  Arenas,  Chile, 
1  p;u;ilectot\pe  from  Punta  Arenas,  Chile;  [Trophon  fe- 
nc.stratu.s]  Original  t\pe  material,  from  GregoiT  Bank, 
Strait  of  Magellan,  was  destroyed  (Hausdorf //i  htt.).  The 
lectotype  of  Buccinum  muriciforme  (BMNH 
1837.12.15.2695)  is  here  designated  as  neot\pe  of  Tro- 
phon fcncstratus;  [Trophon  confhouyi]  Lectotvpe  (fig- 
ures 23-25),  ZMH  unnumbered,  here  designated,  4 
paralectotypes,  all  from  Port  Grappler,  Smvth  Channel; 


Figures  83-95.  Xymcnopsi.s  iiiioicifoniiis  (King  and  Broderip,  18.32).  83.  Anterior  piirtion  of  mule  specimen,  mantle  reHected. 
84.  Ctenidium  and  osphradium.  85.  Transverse  section  of  the  penis,  (figure  83  at  1-1').  86.  Male  pallial  gonoduct.  87.  Transverse 
section  (figure  86  at  2-2').  88.  Transverse  section  (figure  86  at  .3-3').  89.  Stomach,  opened  dorsally.  90.  Female  pallial  gonoduct. 
91.  Transverse  section  through  capsule  gland  (figure  90  at  4—4').  92.  Transverse  section  (figure  90  at  6-6').  93.  Transverse  section 
through  bursa  eopulatrix  (figure  84  at  5-.5').  94.  Diagrammatic  representation  of  the  alimentary  system.  95.  Sagittal  section  throua;h 
the  anterior  portion  of  the  foot. 

a  =  anus;  aa  =  anterior  aorta;  abo  =  accessory  boring  organ;  ag  =  albumen  gland;  ap  =  anal  papilla;  asg  =  accessory  salivarv^ 
gland;  be  =  bursa  eopulatrix;  ca  =  compacting  area;  eg  =  capsule  gland;  ct  =  ctenidium;  dd  =  ducts  to  digestive  diverticula;  dgl 
=  ampulla  of  gland  of  Leiblein;  dp  =  duct  pouch;  ds  =  dorsal  blood  sinus;  e  =  esophagus;  gl  =  gland  of  Leiblein;  hf  =  horizontal 
fold;  hg  =  hypobranchial  gland;  i  =  intestine;  ig  =  intestinal  groove;  1  =  lobes  along  meuitle  edge;  leg  =  left  lobe  of  capsule  gland; 
mo  =  mouth;  nr  =  nerve  ring;  os  =  osphradium;  p  =  penis;  pg  =  prostate  glanil;  pma  =  posterior  niLxing  area;  pro  =  propodial 
groove;  pvd  =  penial  vas  deferens;  r  =  rectum;  ra  =  radula;  rcg  =  right  lobe  of  capsule  gland;  rg  =  rectal  gland;  s  =  siphon;  sg 
=  salivary  gland;  sto  =  stomach;  tl  and  t2  =  tyjihlosoles;  vd  =  vas  deferens;  vg  =  vagina;  vl  =  valve  of  Leiblein. 


Page  50 


THE  NAUTILUS,  Vol.  114,  No.  2 


Figures  96-103.  Xijincnopsis  subnodosus  (Gray,  1839)  96-97.  Holotype  of  Fusns  cancelVmus  PhiLippi,  1845,  and  11601)^6  of 
Buccinum  suhnodosn  Gray,  18.39.  MNHNS  unnumbered.  Strait  of'  Magellan.  98-100.  BMNH  19990.3.30,  4.3°.3S.9'17"S. 
7.3°37.2'.38"\\'.  101.  Operculum,  external  (left)  and  internal  (right)  Ndews,  same  specimen  of  98-100.  Scale  bar  =  1  cm.  102.  Dorsal 
view  of  radular  ribbon  from  specimen  in  figures  98-100.  Scale  bar  =  .50  fj.m.  103.  Oblique  (45°)  lateral  view,  of  radular  ribbon. 
Scale  bar  =  25  |j,m. 


[Trophon  elegans]  Lectotype  (figures  11-12),  ZMH  un- 
numbered, here  designated,  from  Port  Stanley,  Mahinas 
(Falkland)  Is.,  2  paralectotypes  from  Port  Stanley,  Mal- 
vinas  (Falkland)  Is. 

Additional  material  examined:  AMNH  50211,  Strait 
of  Magellan;  AMNH  8.5428,  Canadon  de  las  Vacas; 
AMNH  99610,  West  Malvinas  (Falkland)  Is.;  AMNH 
137057,  Beagle  Channel;  AMNH  17929.3,  Puerto  De- 
seado;  AMNH  271757,  53°37.7'S,  69°54.6'W,  in  42  ni; 
AMNH  271758,  54°10.2'S,  65°57.5'W,  in  101  m;  MACN 


10043-1,  Ushuaia;  MACN  12374,  Punta  Arenas;  MACN 
12376,  53°10'S,  70°55'W;  MACN  12448,  Puerto  Harris. 
Dawson  Is.;  MACN  12552,  Ri'o  Grande;  MACN  12555, 
Rio  del  Fuego,  Tierra  del  Fuego;  MACN  13120,  Punta 
Arenas;  MACN  13568,  Ushuaia^i  MACN  1.3569,  Ushuaia; 
MACN  17754,  Puerto  Deseado;  MACN  22567,  Punta 
Colnet,  Isla  de  los  Estados;  MACN  22717,  54°48'S, 
64°55'W,  in  103  ni;  MACN  23943,  20°07'S,  66°33'W,  in 
82  m;  MACN  30876,  Lapataia,  Tierra  del  Fuego;  USNM 
96228,  Strait  of  Magellan;  USNM  96842,  Puerto  Mayiie, 


G.  Pastorino  and  M.  G.  Harasewvch,  2000 


Page  51 


104 


105 


Figure  104.  Original  illustration  of  Fusiis  biiccineiis  La- 
marck, 1816  (pi.  427,  figs.  3a,  b). 

Figure  105.  Sowerby  in  Gray's  (1839,  pi.  36.  fig.  12)  illus- 
tration of  Fusus  buccineus.  The  specimen  depicted  is  Xipnen- 
opsis  mtiricifonnis. 

Chile;  USNM  126899,  Tierra  del  Fuego,  Intertidal; 
USNM  348720,  Puerto  Harberton,  Tierra  del  Fuego; 
USNM  .368273,  Port  William.s,  Malviiias  (Falkland)  Is., 
in  18  m;  USNM  368401,  Port  Stanley,  MaKdnas  (Falk- 
land) Is.;  USNM  368639,  Punta  Arenas,  Chile;  USNM 
368641,  Punta  Arenas,  Chile,  intertidal;  USNM  368654, 
Punta  Arenas,  Chile;  USNM  368761,  Punta  Arenas,  In- 
tertidal, Chile;  USNM  381693.  Bahia  San  Sebastian,  Tie- 
rra del  Fuego;  USNM  710029,  Puerto  Deseado,  Santa 
Cruz,  Intertidal;  USNM  870002,  53°50'54"S, 
70°30'42"W,  in  15-17  m;  USNM  870165.  54°05'S. 
58°52"W,  in  119  m;  USNM  886188,  53°51'32"S. 
70°25'52"W,  in  2-3  ni;  USNM  886190,  53°51'32"S. 
70°25'52"W,  in  2-3  ni;  USNM  886737,  53°39'S, 
70°55'30"W,  in  20  ni;  USNM  886745,  53°39'S, 
70°55'30"\V,    in    15-18    ni;    USNM    901628,    53°39'S, 


Figure  106.  Localities  at  which  Xi/menopsis  species  (0 
A',  muricifonnis;  O  =  X.  cornigatiis:  #  =  A',  buccineus;  -k 
X.  subnodosus )  were  collected  off  Chile  and  Argentina. 


70°55'30"W,  in  20  m;  USNM  901629,  Strait  of  Magellan, 
53°39.4'S,  70°55.5'W,  in  24  m;  USNM  901630, 
53°39.4'S,  70°55.0'W,  in  82  m;  USNM  901631, 
54°47.25'S,  64°18.3'W.  in  35  m;  USNM  901632. 
53°33.9'S,  69°58.8'W,  in  82-91  m;  USNM  901633, 
53°39.3'S,  70°55.4'W,  38-42  m;  USNM  901634, 
53°17'S,  6S°13'W,  in  0-1  m;  USNM  901635,  53°39.8'S, 
70°54.9'W,  in  27-73  m;  LACM  69170,  Islas  Guarello, 
Chile,  52°40'S,  73°41'W;  LACM  10503.  MoKiiieux 
Sound.  Strait  of  Magellan.  Chile;  LACM  118051,  Lively 
Island,  East  Malvinas  (Falkland)  Is.,  52°00'S,  58°2S'W; 
LACM  71-262,  KN  Hkko  Cruise  712.  Sta.  672.  W  of 
Puerto  Aiio  Nuevo.  Isla  de  los  Estados.  Tierra  del  Fue- 
go, Argentina,  54°45.1'S,  64°07.3'W,  in  50  m;  LACM  71- 
305,  IW  Hkro  Cruise  715,  Sta.  690,  8  miles  E  Ensena- 
da  Patagones,  SE  Tierra  del  Fuego,  Argentina.  54°52'S, 
65°05'W,  in  144  m;  LACM  71-337,  R>V  Hkho  Cruise 
715,  Sta.  888,  W  of  Bahia  York,  Isla  de  los  Estados,  Tie- 
rra del  Fuego,  Argentina,  54°49.5'S,  64°19.6'W,  in  56- 
63  m;  LACM  73-66,  IW  Hkro,  Cabo  Colnett,  Isla  de 
los  Estados,  Tierra  del  Fuego,  Argentina,  54°43'27"S, 
64°14'18"W,  in  15  ni;  LACM  73-68,  IW  Hkho.  Beagle 
Canal.  C;ileta  Aw;iiakirrh,  Chile,  55°0'S,  69°02.2'W,  in 
17  m;  LACM  73-69,  IW  Hero,  Punta  Valparaiso.  Canal 
Cockburn,  Chile,  54°22.2'S,  7r21.7'W.  in  15  ni;  LACM 
73-70,  Isla  Carlos  III,  Strait  of  Magellan.  Chile, 
53°39.4'S.  72°14.8"W.  in  11-12  ni;  LACM  73-71.  Punta 
Dashwood.  Canal  Smyth,  southern  Chile,  52°24'S, 
73°39.7'W,  in  12  ni;  LACM  73-72,  Bahia  Tom,  Chile, 


Page  52 


THE  NAUTILUS,  Vol.  114,  No.  2 


5()°11.3'S,  74°47.9'W,  in  14  m;  LACM  73-107,  RA^ 
Hero,  Colling%vood,  Chile,  5r52'S,  73°43.6'W,  intertid- 
al  to  shallow  subtidal;  LACM  75-49,  Puerto  el  Hambre, 
Brunswick  Peninsula,  Strait  of  Magellan,  ChLle,53°37'S, 
70°56'W,  intertidal;  LACM  78-84,  IW  Hero  Sta,  B  4- 
6,  N  of  Cabo  Buen  Tiempo,  Rio  Gallegos,  Santa  Cruz 
Prov.,  Argentina,  51°16-20'S,  68°50-54"W,  in  30  m,  mud 
bottom:  LACM  86-268.5,  Canal  Oeste,  S  side  Isla  madre 
de  Dios,  Ultima  Esperanza,  Magallanes  Prov.  Chile, 
50°27.7'S,  75°11.5'W,  in  30  m. 

Literature  records:  RA'  Discoxerv:  Sta. 52.  7.4  ca- 
bles N,  17°E  of  Naw  Point,  Port  William,  East  Mahinas 
(Falkland)  Islands,  in  17  ni;  Sta.  1230,  6.7  miles  N,  62°W 
of  Dungenes  Light,  Strait  of  Magellan,  in  27  m;  Sta. 
WS784,  N  of  Malvdnas  (Falklandy  Islands,  49°47.75'S, 
61°05'W,  in  170  m  (Powell,  1951);  Sta.  24,25,  Puerto 
Eden  in  10-12,  6-7  m  respectively;  30,33  Puerto  Eden, 
in  8-10  and  11-12  m;  Sta.  Caleta  Lackawana  in  4-6  m; 
Sta.  43  Paso  Indio;  Sta.  50,  63,  65  Puerto  Williams,  Nav- 
arino  Is.  (Dell,  1971). 

Distribution  (figure  106):  Knov\ni  from  45°  S  (Chonos 
Archipelago),  Southern  Chile  {fide  Dell,  1971  as  X.  de- 
color); Tierra  del  Fuego,  Strait  of  Magellan,  Beagle 
Channel  (Argentina)  and  the  Malvinas  (Falkland)  Is- 
lands. The  bathvmetric  range  for  X.  imihciformis  is  0- 
170  m. 

Remarks:  Xiimcuop.sis  miuiciformis  has  a  long  and 
comphcated  taxonomic  histoiy.  The  oldest  name  to  have 
been  applied  to  the  Xymenopsis  characterized  by  spiral 
sculpture  that  appears  as  incised  grooves  is  Buccinum 
luuriciformc  King  and  Broderip,  1832.  There  were  2  lots 
of  svntvpes,  BMNH  1837.12.15.2695  (2  specimens)  and 
BMNH  1992056  (5  specimens),  both  from  Str;iit  of  Ma- 
gellan. One  specimen  BMNH  1837.12.15.2695  (figures 
14-16),  was  illustrated  by  Dell  (1972:8,  figs.  31,32)  as 
the  holotvpe  (an  inads'ertent  lectotype  designation).  The 
holotvpe  of  Fh.si/.s  decolor  Philippi,  1845  (figures  9-10), 
iilso  from  the  Strait  of  Magellan,  clearly  represents  a 
worn  specimen  of  X  mttiicifonnis.  The  next  name  to  be 
apphed  to  this  species  is  Buccinum  cancellarioidcs 
Reeve,  1847,  based  on  2  syntyjjes  lacking  locafity  data. 
One  specimen  is  here  designated  as  lectotype  (figures 
17-19).  Powell  (1951)  recognized  B.  cancellarioidcs  to 
be  an  earher  name  for  Ftisus  lirafus  Gould,  1849  (Ho- 
lot\-pe,  figures  1-3).  Because  he  was  unsure  if  Reeve's 
(February,  1847)  name  was  a  junior  homonvan  of  Buc- 
cinum cancellaroides  Basterot  in  Grateloup,  given  as 
1847  by  Sherborn  (1924:1030).  he  ret;iined  the  use  of 
X.  liratus.  The  holotvpe  of  Trophon  loebbeckei  Kobelt, 
1878  (figures  4-6),  from  an  unkniown  locality,  also 
matches  closely  the  holotype  of  B.  muricifimnc,  as  do 
the  lectotypes  of  Trophon  lebnini  and  Trophon  viola- 
ceus,  both  Mabille  and  Rochelinme  in  Rochebrune  and 
MabiUe,  1889.  Trophon  Ichruni  was  based  on  3  .svntypes, 
MNHN  unnumbered,  all  from  Santa  Cruz,  Argentina. 
One  specimen  is  here  designated  as  the  lectoty|3e  (figure 
13).  The  figured  s)'ntype  of  Trophon  violaceus  MNHN 


unnumbered,  from  Baie  Orange,  Tierra  del  Fuego  is 
here  designated  lectotvpe  (figures  7-8). 

In  his  publication  on  the  molluscan  fauna  of  the  Ma- 
gellanic Province,  Strebel  (1904)  proposed  numerous 
taxa  based  on  minor  phenotypic  variants.  Among  these 
were  16  species  and  varieties  of  Trophon  that  were  sub- 
sequentlv  referred  to  Xipncnopsis  bv  Powell  (1951).  Ac- 
cording to  Dance  (1986:227),  Strebel's  collections  were 
housed  in  the  Hamburg  Museum  and  "totally  destroyed " 
during  World  War  II.  However,  this  appears  to  be  true 
only  for  the  "dry"  (shell  only)  specimens.  The  alcohol 
preserved  material  on  which  much  of  Strebel's  (1904) 
Magellanic  publication  was  based,  survived  the  war  and 
is  in  the  collections  of  the  Zoologisches  Institut  und 
Zoologisches  Museum  der  Universitat  Hamburg.  Mate- 
rial collected  b\-  the  Swedish  Sudpolar-Expedition,  in- 
cluding types  of  taxa  described  by  Strebel  (1908),  is  in 
the  Swedish  Museum  of  Natural  History  in  Stockholm. 

The  senior  author  visited  both  these  museums  in  the 
course  of  this  study,  and  was  able  to  locate  much  of 
Strebel's  tspe  material.  Examination  of  the  t\pes  ot;  Tro- 
phon paessleri,  T  clongatus,  T.  pscudoelongatus,  T  al- 
bus,  T  acuminahis,  T  couthouiji,  T  elegans  and  T.  obesus 
leaves  no  doubt  that  these  are,  at  most,  minor  moq:)ho- 
logical  variants  oi  Xipnenopsis  nuiriciformis.  The  taxon 
Trophon  paessleri  is  based  on  6  syiitypes.  ZMH  unnum- 
bered, 5  from  Port  Grappler,  Smyth  Channel,  Strait  of 
Magellan  and  1  from  Punta  Arenas.  One  specimen  (fig- 
ures 26-28)  from  Port  Grappler  is  here  designated  as 
lectotype.  The  type  material  of  Trophon  clongatus  con- 
sists of  7  syntypes,  ZMH  unnumbered,  2  from  Puerto 
Angosto,  Strait  of  Magellan,  3  from  Puerto  Bueno,  1 
from  Navarino  Is.  and  I  from  Eden  Harbor  One  spec- 
imen (figures  35-37)  from  Puerto  Angosto  is  here  des- 
ignated as  lectotype.  Trophon  couthouiji  is  based  on  3 
syntypes,  ZMH  unnumbered,  from  Port  Grappler, 
Smvth  Channel.  One  specimen  (figures  23-25)  is  here 
designated  as  lectotvpe.  Holotvpes  are  illustrated  for 
Trophon  pscudoelongatus  (figures  29-31),  ZMH  unnum- 
bered, and  T.  albus  (figures  38—40),  ZMH  unnumbered, 
both  from  Ushuaia;  and  T.  acuminatus  (figures  20-22), 
ZMH  unnumbered,  from  Punta  Arenas,  Strait  of  Ma- 
gellan. Trophon  elegans  is  based  on  3  specimens  from 
Port  Stanley,  MiJvinas  (Falkland)  Is.  One  specimen  (fig- 
ures 11-12)  ZMH  unnumbered,  is  here  designated  as 
lectotype.  Trophon  obesus  is  represented  by  2  syntypes 
from  Punta  Arenas,  Chile;  one  (figures  32-34),  ZMH 
unnumbered,  is  here  designated  as  lectotype. 

The  type  material  of  Trophon  paessleri  var.  turrita  and 
of  T  fenestratus  could  not  be  found,  and  was  among  the 
material  destroyed  during  World  War  II  (Hausdorf  in 
litt.).  Based  on  Strebel's  (1904)  descriptions  and  figures, 
there  is  no  doubt  that  both  these  taxa  are  conspecific 
with  Xymenopsis  muricifonnis.  We  therefore  designate 
the  lectotype  of  Buccinum  muriciforme  (BMNH 
1837.12.15.2695)  as  the  neotype  for  both  Trophon  paes- 
sleri var  turrita  and  T  fenestratus,  making  both  these 
taxa  objective  junior  synomins  of  Buccinum  muricifiir- 


G.  Pastorino  and  M.  G.  Harasewych,  2000 


Page  53 


Xijinenopsis  comi^atiis  (Reeve,  1848) 
(Figures  57-71,  80-82,  106) 

Fusus  cornif^atiis   Kee\e,   1848:  pi.  20  figs.  84  a.  h.   Kolielt, 

1878:  293,  pi.  74.  fi<is.  1.5-16. 
Trophon  corni'^^dliis  Ki'tne — Tnon,  1880:  145.  pi.  3.3.  fig.  .352; 

Rocliehniiii'  ami  Maliille,  1889:  II  ,54. 
Trophon  miiricifonnis  Sowerby,  1880:  66,  pi.  3,  fig.  41.  [not 

Bucriiiiim  muricifonne  King  and  Broderip,  1832] 
Trophon  hot/lei  Strebel,  1904:  227.  pi.  8,  figs.  68  a-f,  69  a-c; 

MelviU'and  Standen,  1907:  1.37. 
Trophon  hnicci  Streliel,  1904:  2.30.  pi.  8.  fig.  72;  MeKill  and 

.Stiuiden.  1907:  1.35. 
Trophon  ornntus  Strehel.  1904:  231.  pi.  8.  fig.  73. 
Trophon  standeni  Strehel.  1904:  2.32,  pi.  7,  fig.  67. 
Trophon  A— Strehel,  1904:  2.34,  PI.  8,  fig.  78. 
Trophon  B— Strebel,  1904:  2.35,  Pi.  8,  fig.  79. 
Trophon  falklanclicus  Strebel,  1908:  .39,  PI.  1,  figs.  8a-c. 
Xymenopsis  falklandicus  Strebel — Powell,  1951:159,  PI.  9,  figs. 

46,  47;  Castellanos  and  Landoni,  1993:18,  pi.  3.  figs.  37.  A. 
Xijmenopsis  cornigattis  (Reeve) — Ceniohorskv.  1977:118,  fig. 

19. 

Diagnosi.s:  Axial  .sculpture  of  regular,  thin,  .slender, 
rounded  rih.s,  19-24  on  body  whorl.  Spiral  sculpture  of 
rounded  cords  (4-6  on  penultimate  whorl,  19-26  on 
body  whorl),  slightly  broader  to  slightly  narrower  than 
intervening  spaces,  producing  weakly  cancellate  surface 
at  intersection  with  axial  ribs.  Spiral  threads  present  be- 
tween adjacent  cords.  Outer  lip  without  apertural  teeth. 

Description:  Shell  of  medium  size  (2.5-.32  mm),  thin, 
fusiform.  Protoconch  worn  on  all  specimens,  but  clearly 
multispiral,  conical.  Teleoconch  of  up  to  6  rounded 
whorls.  Spire  high,  <V2  shell  length.  Spire  angle  =  45°. 
Suture  deeply  impressed;  subsutural  ramp  small,  but 
present.  Aperture  semicircular;  outer  hp  rounded,  dis- 
tinctly crenulated,  reflecting  spiral  cords  at  lip  edge;  in- 
ner hp  cur^'ed.  adpressed;  interior  glossv  brownish.  Si- 
phonal  canal  short,  oblique,  open.  Umbilicus  absent.  A.x- 
ial  sculpture  of  regular,  thin,  slender,  rounded  ribs,  19- 
24  on  body  whorl.  Growth  lines  present  throughout 
shell,  clearly  defined  over  the  nodes.  Spiral  sculpture  of 
rounded  cords  (4-6  on  penultimate  whorl,  19-26  on 
body  whorl),  slightly  broader  to  shghtly  narrower  than 
intervening  spaces,  producing  weakly  cancellate  surface 
at  intersection  with  axial  ribs.  Finer  spiral  threads  pre- 
sent between  adjacent  cords.  Shell  ultrastructure  as  in 
X.  miiricifonnis,  composed  of  2  layers:  outer  layer  (0..55 
shell  thickness)  ot  amorphous  calcite,  with  ribs,  cords 
confined  to  this  layer;  inner  layer  (0.45  shell  thickness), 
crossed-lamellar  aragonite  with  ciTstal  planes  oriented 
perpendicular  to  growing  edge,  of  relatively  constant 
thick-ness.  Operculum,  gross  anatomy,  as  in  X.  murici- 
fonnis.  Radula  (figures  75-76)  very  similar  to  that  of  X. 
miiricifonnis.  Rachidian  teeth  narrower  (<  100|xm),  in 
proportion  to  smaller  size  of  this  species.  Central,  lateral 
cusps  more  conical,  less  cyhndrical  that  in  X.  nutricifor- 
mis,  inner  lateral  denticles  of  X.  cornigattis  proportion- 
ally larger. 

Type  material:     [Fiisiis  comifiafiis]  Hoiotvpe  (figures 


57-59),  BMNH  74.12.11.1,5.3,  lacks  locality  data;  \Tro- 
plnni  hoijh'i]  Lectotvpe  (figure  60-62).  ZMH  unnum- 
bered, here-  designated,  with  4  paralectotvpes,  all  from 
Port  Stanley,  Isias  Malvinas  (Falkland);  {Trophon  brucei] 
Tyi^e  material  could  not  be  located  at  ZMH.  No  locaUty 
exphcitly  associated  with  this  species  l)v  Strebel.  The  ho- 
iotvpe of  Fusus  cormgatus  (BMNH  74.12.11.1.53)  is 
here  designated  as  neotype  of  Trophon  brucei;  [Trophon 
onmtus]  Holotyj^e  (figures  69-71)  ZMH  tmnumbered, 
from  Port  Stanley,  Malvinas  (Falkland)  Is;  [Trophim 
standcni]  Original  tyj^e  material,  from  Malvinas  (Falk- 
land) Is.,  could  not  be  located  at  ZMH.  The  holotype  of 
Fusus  corrufiatiis  (BMNH  74.12.11.1.53)  is  here  desig- 
nated as  neotype  oi  Trophon  standcni;  [Trophon  falklan- 
dicus] Holot\pe  (figures  6.3-65),  NHRM  1049,  Berkeley 
Sound,  Malvinas  (Falkland)  Is.,  (51°.35'S,  .57°.56'W),  in 
2.5-30  m.  Station  49. 

Additional  material  examined:  MACN  10136,  Mal- 
Ndnas  (Falkland)  Is.,  m  0  ni;  USNM  173062,  Malvinas 
(Falkland)  Is.;  USNM  368288,  Port  Stanley;  USNM 
368292,  Port  Stanley,  shore;  USNM  368311,  between 
Pembrooke  Is  and  Tussock  Is.  Malvinas  (Falkland)  Is., 
in  27  m;  USNM  .368342,  Port  Stanley,  shore;  USNM 
368347,  Port  Stanley,  shore;  USNM  368.3.50,  Port  Stan- 
ley, Malvinas  (Falkland)  Is.;  USNM  368360,  Port  Stan- 
ley, shore;  USNM  3683,89,  Port  Stanley,  shore;  USNM 
368422,  Port  Stanley,  shore;  USNM  .368432,  Port  Stan- 
ley; USNM  421877,'  Port  Stanley;  USNM  421884,  Port 
Williams,  York  Bay,  Malvinas  '(Falkland)  Is.;  USNM 
421890,  Port  Stanley;  USNM  6.5.5342,  Port  Stanley  area; 
USNM  702124,  Port  William,  in  16  ni;  LACM  15394, 
LACM  15395,  LACM  55989,  LACM  61583,  all  from 
Fox  Bay,  East  Malvinas  (Falkland)  Is. 

Literature  records:  [as  X.  falklandicus]  WW  Disco\- 
ERV:  Sta.  .55,  2  cables  S,  24°E  of  Naw  Point,  Entrance 
to  Port  Stanley,  East  Falkland  Island,  in  10-16  m;  Sta. 
56,  1.5  cables  N,  50°E  of  Sparrow  Point.  Sparrow  Cove, 
Port  William,  East  Falkland  Island,  in  10.5-16  m  (Pow- 
ell, 1951). 

Distribution:  All  specimens  belonging  to  this  species 
that  we  were  able  to  examine  were  from  the  Malvinas 
(Falkland)  Islands.  The  bathymetric  range  of  X.  corru- 
gatiis  is  0-27  m. 

Remarks:  The  holoUpe  of  Fusus  cornigatus  BMNH 
1874.12.11.153,  was  illustrated  by  Cernohorsky  (1977, 
fig.  19).  Trophon  ornatus  is  based  on  1  specimen  (figures 
69-71)  ZMH  unnumbered,  from  Port  Stanley,  Malvinas 
(Falkland)  Is.  The  tvpe  material  oi  Trophon  bnicei,  and 
T.  standcni  could  not  be  found  at  ZMH.  In  the  original 
description,  Streliel  wrote  that  this  material  was  sent  to 
him  from  Manchester  Museum  by  Bruce.  According  to 
D.  Penney  {in  litt.),  these  specimens  never  reached 
Manchester  again.  We  therefore  consider  the  tvpe  ma- 
terial ol  both  species  to  be  lost.  These  taxa  are  consid- 
ered sviionyms  ol  Xijmenopsis  cornigatus  based  on  Stre- 
bel's  (1904)  descriptions  and  figures.  The  holotype  of 
Fusus  corrugatus  (BMNH  1874.12.11.1.53)  is  here  des- 


Page  54 


THE  NAUTILUS,  Vol.  114,  No.  2 


ignated  as  the  neotype  for  both  Trophon  bntcei,  and  T. 
standeni  making  both  these  taxa  objective  junior  syno- 
nyms of  Fiisus  cornigatus. 

Xijmenopsis  huccineus  (Lamarck,  1816) 
(Figures  52-56,  75-79,  104,  106) 

Fusus  bucciiH'us  Lamarck,  1816:  pL  427,  figs.  .3a,  b,  Liste:  6. 

Ftisus  albidus  Philippi,  1846:  119  pi  .3.  fig.  5. 

Fusus  textiliosus  Honibron  and  Jactjuinot  in  Rousseau,  1854: 
108,  Pi  25,  fig.  9-10,  non  Deshayes,  1835. 

Fusus  textilosus  Honibron  and  Jacquinot  [error  for  F.  textilio- 
sus]. Hupe  in  Gay,  1854:  162;  Philippi,  1855:  206. 

Fusus  Jacqtiinoti  Philippi,  18.55:  206,  new  name  for  F.  textilio- 
sus Honibron  and  Jacquinot.  1854,  not  F.  textiliosus  De- 
shayes. 1835. 

Trophon  albidus  Phifippi — Kobelt,  1878:  293,  pi  74,  figs.  11- 
12;  Strebel  1904:  222.  pi  7.  figs.  64  a-cl 

Trophon  nuiricifonnis  Sowerby,  1880:  66,  pf  3,  fig.  40.  [not 
Buccinuin  muriciforme  King  and  Broderip,  1832] 

Trophon  buccineus  Gray— Tryon,  1880:146,  p!.  39,  fig.  490. 

Trophon  textiliosus  (Honibron  and  Jacquinot) — Rochebrune 
and  Mabille,  1889:  H..55. 

Trophon  rinff'i  Strebel,  1904:  242,  pi.  8,  fig.  77. 

Xijmenopsis  albidus  (Philippi)— Powell,  1951:159.  fig.  M.  90; 
Yokes,  1991:  7.  fig.  ininunibered;  Castellanos  and  Lan- 
doni,  1993:  17,  pl."3,  figs.  .388,  41-42. 

Diagnosis:  Axial  sculpture  of  irregular,  \'erv  weak  ribs 
(25-28  on  body  whorl).  Irregular,  obsolete  growth  lines 
present  throughout  shell.  Spiral  sculpture  of  flattish 
cords  (9  on  penultimate  whorl,  17-20  on  body  whorl)  as 
broad  as  intei"vening  spaces,  forming  weakly  cancellate 
sculpture  at  intersections  with  axial  ribs.  No  spiral 
threads  between  adjacent  cords. 

Description:  Shell  of  medium  size  (s  30  mm),  thin, 
fusiform,  cUrty  white  to  brownish  tan.  Protoconch  taU, 
conic,  of  .3  whorls.  Teleoconch  of  up  to  6  convex  whorls, 
spire  high,  =  Va  shell  length.  Suture  deeply  impressed. 
Aperture  subovate;  outer  lip  evenly  rounded,  weakly 
crenulated;  inner  Up  gently  concave;  interior  glossy, 
whitish.  Siphonal  canal  long,  narrow  for  genus,  oblique, 
open.  Umbilicus  absent,  pseudoumbilical  chink  in  some 
specimens.  A,xial  sculpture  of  irregular,  veiy  weak  ribs 
(25-28  on  body  whorl).  Irregular,  obsolete  growth  lines 
present  throughout  shell.  Spiral  sculpture  of  flattish 
cords  (9  on  penultimate  whorl,  17-20  on  bodv  whorl)  as 
broad  as  inten'ening  spaces,  forming  weakly  cancellate 
sculpture  at  intersections  with  a.xial  ribs.  Shell  ultrastnrc- 
ture  of  two  layers,  as  in  other  Xymenopsis,  but  outer, 
calcific  laver  thinner  (0.35  shell  thickness),  than  inner 
aragonitic  laver  (0.65  shell  thickness).  Operculum,  gross 
anatomy,  as  in  X.  mtiriciformis.  Radula  (figures  75-76) 
more  similar  to  that  of  X.  cornigatus  than  to  X.  miiri- 
ciformis  in  that  lateral  cusps  broader,  more  triangular  in 
outline,  inner  lateral  denticles  more  pronounced.  Weak 
outer  lateral  denticles  present  in  some  individuals  of  X. 
buccineus  (figure  75,  arrow). 

Type  material:  \Fusus  buccineus]  Lost,  not  in  Paris 
or  Geneva  (Bouchet  in  litt.  and  Finet  in  lift.),  type  lo- 


cality unspecified.  Neotype,  (figures  52-54),  USNM 
870410,  here  designated,  5.3°32'S,  64°57'W,  in  119-124 
m;  [Fusus  albidus]  Probablv  lost,  not  in  MNHNS,  type 
localits'  unspecified;  [Fusus  textiliosus  Hombron  and  Jac- 
quinot, 1854,  not  F.  tcxtdiosus  Deshayes,  1835]  Holotype 
(figures  55-56),  MNHN  no  catalog  number,  Strait  of 
Magellan;  [Trophon  lingei]  Original  tvpe  material,  fnnn 
Strait  of  IVIagellan,  destroyed,  (B.  Hausdorf,  in  litt.).  The 
Neotype  of  Fusus  buccineus,  USNM  870410,  is  here 
designated  as  the  neotype  of  Trophon  ringei. 

Additional  material  examined:  MACN  12553,  Ti- 
erra  del  Fuego;  MACN  2.3862,  5r46'S,  68°45'W,  in  22 
m;  MACN  25012  +  MACN  25013,  54°26'S,  64°53'W, 
in  112  m;  USNM  110731,  Strait  of  Magellan,  East  En- 
trance, in  10  m;  USNM  110782,  Strait  of  Magellan,  East 
Entrance,  in  18  m;  USNM  152S92,  Santa  Cruz,  Pata- 
gonia; USNM  870481,  53°40'S,  66°20'W,  in  81  m; 
USNM  870491,  53°42'S,  66°19'W,  in  81  m;  USNM 
881098,  54°34'S.  63°50'VV.  in  118  m;  USNM  881924, 
54°27'S,  66°12'W,  in  0  m;  USNM  881961,  5.3°32'S, 
64°57'W,  in  119-124  m;  USNM  881966,  52°30'S, 
67°14'W,  in  82  m;  USNM  901625,  53°39.8'S,  70°54.9'W, 
in  27-73  m;  USNM  901626,  5.3°()6'S,  67°04'W,  in  86  m; 
USNM  901627,  54°59.9'S,  64°50'W,  in  438  m  (dead 
specimens);  LACM  71-302,  IW  Hero  Cruise  715,  sta. 
687,  5  miles  E  of  San  Mauricio,  SE  end  Tierra  del  Fue- 
go, Argentina,  54°45'S,  64°04.6'W,  in  75  m;  LACM  71- 
313,  IW  Hero  Cruise  715,  sta.  853,  9  miles  N  of  Cabo 
Colnett,  Isla  de  los  Estados,  Tierra  del  Fuego,  Argentina, 
54°34'S,  64°20'W,  in  91  m.  LACM  71-320,  RA^  Hero 
Cruise  715,  sta.  865,  4  miles  E  of  Isla  Observatorio,  Isla 
de  los  Estados,  Tierra  del  Fuego,  Argentina,  54°39'S, 
64°00'W,  in  75  m;  LACM  71-324",  IW  Hero  Cruise  715, 
sta.  870,  6  miles  NE  of  Isla  Observatorio,  Isla  de  los 
Estados,  Tierra  del  Fuego,  Argentina,  54°34'S, 
64°00.3'W,  in  84  m;  LACM  71-327,'  IW  Hero  Cruise 
715,  sta.  873,  9  miles  N  of  Cabo  San  Juan  E  end  of  Isla 
de  los  Estados,  Tierra  del  Fuego,  Argentina,  54°.34'S, 
63°50"W,  in  118  m;  LACM  71-329,  IW  Hero  Cruise 
715,  sta.  875,  5  miles  S  of  Punta  Ventana,  S  side  of  Isla 
de  los  Estados,  Tierra  del  Fuego,  Argentina,  54°54.5'S, 
63°56'W,  in  771-903  m  (dead' specimens);  LACM  71- 
330,  IW  Hero  Cmise  715,  sta.  876,  14  miles  N  of  Cabo 
San  Juan,  Isla  de  los  Estados,  Tierra  del  Fuego,  Argen- 
tina,'54°29'S,  63°50'W,  in  112  m;  LACM  71-333,  IW 
Hero  Cmise  715,  sta.  881,  Puerto  Vancouver,  Isla  de  los 
Estados,  Tierra  del  Fuego,  Argentina,  54°39'S,  64°00'W, 
grab;  LACM  71-341,  RA'  Hero  Cmise  715,  sta.  894,  5 
miles  SE  of  Cabo  Kempe,  Isla  de  los  Estados,  Tierra  del 
Fuego,  Argentina,  54°54.9'S,  64°19.5"W,  in  263-285  m 
(dead  specimens);  LACM  71-348,  RA^  Hero  Cmise  715, 
sta.  903,  13  miles  N  of  Baliia  Crosslev,  Isla  de  los  Es- 
tados, Tierra  del  Fuego,  Argentina,  54°34.3'S,  64°40'W, 
in  84-85  m;  LACM  71-349,  RA^  Hero  Cmise  715,  sta. 
904,  8  miles  N  of  Baliia  Crosslev,  Isla  de  los  Estados, 
Tierra  del  Fuego,  Argentina,  54°39.0'S,  64°40'W,  in  84- 
85  m;  LACM  71-351,  RA'  Hero  Cmise  715,  sta.  906, 
14  miles  N  of  Cabo  San  Antonio,  Isla  de  los  Estados, 


G.  Pastorino  and  M.  G.  Harasewvch,  2()()() 


Page  55 


Tierra  del  Fuego,  Argentina,  54°29'S,  64°29.2'W,  in 
122-124  m;  LACM  71-352,  RA'  Hemo  Cniise  715,  sta. 
907,  9  miles  N  of  Cabo  San  Antonio,  Isla  de  lo.s  Estado.s, 
Tierra  del  Fuego.  Argentina,  54°34'S,  64°3()'W,  in  73- 
76  m;  LACM  71-353,  RA'  Hi;ho  Cruise  715,  Sta.  908, 
5  miles  NNE  of  Cabo  San  Antonio,  Isla  de  los  Estados, 
Tierra  del  Fuego,  ArgenHna,  54°38.9'S,  64°30'VV,  in  60 
ni. 

Literature  records:  RA'  Disccn  krv:  Sta.  WS71,  6 
miles  N  60°  E  of  Cape  Pembroke  Liglit,  East  Falkland 
Island,  in  82  m;  Sta.  WS88,  Off  Staten  Island  (Isla  de 
los  Estados),  Tierra  del  Fuego,  54°S,  65°W  to  54°S. 
64°55'W,  in  118  m;  Sta.  VVS97,' Between  Malvinas  (Falk- 
land) Islands  and  Patagonia,  49°S,62°W  to  49°01'S, 
erse'W,  in  145-145  m;  sta.  WS222,  SE  of  Puerto  De- 
seado,  Patagonia,  48°23'S,  65°W,  in  100  m;  Sta.  WS243, 
Off  Santa  Cruz,  Patagonia,  51°06'S,  64°30'W,  in  144- 
141  m;  Sta.  WS750,  NE  of  MaKdnas  (Falkland)  Islands 
51°50'S,  57°15.2'W,  in  135-144  ni;  Sta.  WS805,  Be- 
tween Malvinas  (Falkland)  Islands  and  Patagonia, 
50°11'S,  63°27'W  to  50°09.5'S,  63°31'W,  in  150-148  m: 
Sta.  SW808,  Off  Santa  Cruz,  Patagonia,  49°40.25'S, 
65°42'W.  in  109-107  m;  Sta.  WS829,  Between  MaKinas 
(Falkland)  Islands  and  Patagonia,  50°51'S,  63n3.5'W,  in 
155  m;  Sta.  WS834,  Off  Baliia  Grande,  Patagonia, 
52°57.75'S,  68°08.25'W,  in  27-38  m;  Sta.  WS838,  Be- 
tween Malvinas  (Falkland)  Islands  and  Patagonia, 
53°11,75'S,  65°W,  in  148  m;  Sta.  WS861,  SE  of  Puerto 
Deseado,  Patagonia,  47°40'S,  64°12'W,  in  117-124  m; 
Sta.  WS863,  Between  Malvinas  (Falkland)  Islands  and 
Patagonia,  49°05'S,  64°09'W,  in  127-117  ni;  Sta.  WSS65, 
Between  Malxdnas  (Falkland)  Islands  and  Patagonia, 
50°03'S,  64°14'W,  in  126-128  m;  Sta.  WS867,  Between 
Malvinas  (Falkland)  Islands  and  Patagonia,  51°10'S, 
64°15'W,  in  137-144  ni;  Sta.  WS869,  Between  MaKinas 
(Falkland)  Islands  and  Patagonia,  52°15.5'S,  64°13.75'W, 
in  187-0  [.sic]  m. 

Distribution:  KnowTi  from  Southern  Chile  and  Ar- 
gentina; Tierra  del  Fuego,  Strait  of  Magellan,  Beagle 
Channel,  MaKinas  (Falkland)  Is.  The  vertical  distribu- 
tion oi  X.  biiccinettf;  ranges  between  0  and  155  m.  Rec- 
ords from  greater  depths  are  dead  shells.  The  batliy- 
metric  and  geographic  ranges  of  this  species  are  nearly 
identical  with  those  of  of  X  muriciformis,  and  the  two 
species  have  been  collected  together  in  some  trawls. 
(e.g.  USNM  901625  (X.  biux-iiicu.s)  and  USNM  901635 
(X  muricifonnis)  both  from  53°39.8'S,  70°54.9'W,  in 
27-73  m.) 

Remarks:  Ftistis  buccincus  (Lamarck,  1816)  was  pub- 
lished without  localitv  or  detailed  description.  The  t\pe 
specimen  is  lost  (Finet,  in  ht.,  Bo\ichet  ni  Ut.).  Although 
Sowerby  [in  Gray,  1839)  labelled  a  figure  F.  buccincus, 
the  specimen  illustrated  (figure  105)  is  clearly  X  muri- 
cifonnis, and  bears  little  resemblance  to  Lamarck's  fig- 
ure of  K  buccincus  (fig.  104).  Strebel  (1904:237)  hkewise 
noted  that  F.  buccincus  of  Gray  [actually  Sowerby  in 
Gray]  was  a  different  species  than  F  buccincus  Lamarck. 


The  specimen  illustrateil  bv  Sowerby  is  also  lost  (K.  Way, 
(■/(  lift.). 

As  Fusus  bucciiwus  is  the  oldest  name  to  be  applied 
to  anv  species  of  Xi/mcnopsis,  we  designate  as  neotyi^e 
(figures  52-54)  a  specimen  (USNM  870410)  that  rea- 
sonably conforms  to  Lamarck's  (fig.  104)  illustration  of 
this  species.  This  serves  to  remove  anv  ambiguit\'  re- 
garding the  identity  of  Fusus  buccincus  and  thus  stabi- 
lize the  nomenclature  of  the  Recent  Xyinenopsis.  The 
t\pe  locLllit\-  for  Fusus  buccincus  becomes  the  continen- 
tal shelf  east  of  Tierra  del  Fuego  (53°32'S,  64°57'W),  in 
119-124  m. 

The  next  available  name  for  this  species  is  F(/.s7/.s  al- 
bidus  Philippi,  1846,  for  which  no  type  locality  was  des- 
ignated. As  the  type  specimen  could  not  be  located  at 
MNIINS,  it  is  included  in  the  synonymy  based  on  Phi- 
hppis  description  and  figure.  We  have  not  designated  a 
neotyj^e  for  Fusus  albidus  because  there  remiiins  a  pos- 
sibility- that  the  type  will  yet  be  located  at  MNHNS.  The 
ty|3e  of  Fusus  tcxtiliosus  Hombron  and  Jacquinot,  1854 
from  the  Strait  of  Magellan,  housed  in  MNHN,  is  illus- 
trated in  figures  55-56,  and  undoubted!\-  belongs  to  this 
species.  The  type  materiid  of  Trophon  lingci  could  not 
be  found,  and  was  among  the  material  destroyed  during 
World  War  II  (Hausdorf  (>i  lift. ).  This  taxon  is  considered 
to  be  a  svnonvm  of  X.  buccincus  based  on  Strebel's 
(1904)  descriptions  and  figures.  We  therefore  designate 
the  neotype  of  Fusus  buccincus  (USNM  870410)  to 
serve  as  the  neotype  of  Trophon  ringei.  Trophon  ringei 
thus  becomes  an  objective  synonym  of  Fusus  buccincus. 

Xyincnopsis  subnodosus  (Gray,  1839) 
(Figures  96-103,  106) 

B[uccimnn]  suhtiodosa  Gray,  1S39:  118. 

Fu.ms  cancellinus  Philippi,  1845:  67;  1846:  117,  pi.  3,  fit;.  2; 

Hupe  in  Gay,  1854:  163. 
Trophon  cnncellinii.i  Philippi — Kobelt,  1878:  291,  pi.  74,  figs. 

7,  8;  Rochehnme  and  Mabille,  1889:  H.59 

Diagnosis:  Axial  sculpture  of  strong,  irregular  ribs 
(12-16  on  body  whorl),  nearly  as  broad  and  intenening 
spaces.  Spiral  sculpture  of  dense,  evenly  rounded  cords, 
(6  on  penultimate  whorl,  22-24  on  body  whorl),  broader 
than  intervening  spaces.  Spiral  threads  present  between 
adjacent  cords.  Outer  lip  with  apertural  teeth. 

Description:  Shell  large  (to  37  mm),  solid,  slender, 
fusiform,  dirty  white,  chalkT.  Protoconch  miknown.  Te- 
leoconch  of  up  to  6  gently  convex  whorls.  Spire  high, 
=  Vi  shell  length.  Spire  angle  ==  45°.  Suture  strongly 
impressed.  Aperture  small,  ovate;  outer  lip  thickened, 
with  8-10  weak  teeth;  inner  fip  concave,  thickly  glazed; 
interior  whitish.  Siphonal  canal  long,  obfique,  open. 
PseudoumbiUcal  chink,  small  but  present.  Axial  sculp- 
ture of  strong,  irregular  ribs  (12-16  on  body  whorl), 
nearly  as  broad  and  intenening  spaces.  Spiral  sculpture 
of  dense,  evenK'  roundeil  cords,  (6  on  penultimate 
whorl,  22-24  on  body  whorl),  broader  than  intei-vening 
spaces.  General  morphology-  of  animal  and  operculum 


Page  56 


THE  NAUTILUS,  Vol.  114,  No.  2 


(fi2;ure  101)  as  in  X.  miiriciformi.s.  Osphradium  <  V2  cte- 
nidium  length,  asymmetrical,  with  48  and  32  leaflets  on 
the  external  and  internal  side  respectively.  Ctenidium  2- 
.3  times  as  wide  as  osph/adium,  witli  85-90  triangular 
leaflets.  Radular  ribbon  small,  short  (0.23X  aperture 
length).  Radula  (figures  102-103)  similar  to  those  of  X. 
corntgattis  and  X.  huccincus,  but  with  single  weak  den- 
ticle along  outer  edge  of  each  lateral  cusp. 

Type  material:  [Buccinum  stibnodosa]  Original  type 
material  from  the  Pacific  Ocean  lost  (K.  Way,  in  lift.). 
The  holotype  of  Fusits  cancellintis,  MNHNS,  is  here 
designated  as  neot\pe  of  Bticciniim  subiuxlo.m.  [Fiisiis 
cancellintis]  Holot)'pe,  MNHNS  unnumbered.  Strait  of 
Magellan. 

Additional  material  examined:  BMNH  19990330, 
43°38.9'17"S,  73°37.2'38"W,  East  side  Traiguen  Is.,  Es- 
tero  Elefantes,  Region  11,  Chile.  Dredged  in  5-15  m  on 
bottom  of  colibles  and  dead  \enerids;  sheltered  bay. 

Distribution:  Known  from  the  Str;ut  of  Magellan,  and 
Traiguen  Island,  Chile. 

Remarks:  Gray  (1839)  introduced  Buccinum  suhno- 
closa  as  a  variety  of  Buccinum  muriciformc  (=Xi/mcn- 
opsis  muiiciformis  King  and  Broderip)  in  a  two  Une  de- 
scription without  illustration.  As  Gray's  type  material  is 
lost  (K.  Wav,  in  Jitt.)  this  leaves  some  doubt  as  to  the 
identitv'  of  this  species.  Grav's  description  refers  to  "al- 
ternate narrow  low  and  wider  convex  spiral  ridges"  that 
produce  tubercles  as  they  rise  over  the  axial  ribs,  fea- 
tures characteristic  of  both  X.  corrugatus  and  a  species 
subsequently  named  Fu.su.s  canccUinus.  Grav  does  not 
mention  the  prominent  apertural  teeth  that  are  diagnos- 
tic of  X.  canccUinus,  but  it  is  possible  that  he  had  a  sub- 
adult  specimen  lacking  apertural  teeth.  Gray  states  that 
B.  subnodoso  inhabits  the  Pacific  Ocean.  Fusus  cancel- 
linus  occurs  along  the  Pacific  Coast  of  Chile,  while  X. 
corrugatus  is  known  only  from  the  Malvinas  (Falkland) 
Islands  and  Isla  de  los  Estados.  In  order  to  stabihze  the 
nomenclature  of  Recent  Xi/mcnopsis.  we  designate  the 
holot^pe  oi  Fusus  canccUinus  Phihppi,  1845  (MNHNS) 
to  serve  as  neotvpe  of  Buccinum  subuodosa  Grav,  1839. 
Fusus  canccUinus  thus  becomes  a  junior  objective  syn- 
onym of  Buccinum  subnodosa. 

We  know  of  only  two  specimens  of  Xi/mcnopsis  sub- 
nodosus.  incluchng  the  neotspe  (shell  onK )  housed  at  the 
Museum  of  Santiago,  Chile.  Xtjmenopsis  subnodosus  re- 
sembles both  X.  muiiciformis  and  X.  corntgatus  in  size, 
general  shape,  and  in  the  coarseness  of  its  axial  ribs.  It 
differs  from  X.  muricifonnis  in  ha\ing  a  thicker  shell, 
narrower  more  numerous,  and  more  rounded  spiral 
cords,  and  shares  with  X.  corrugatus  the  presence  of  fine 
spiral  threads  interspersed  between  adjacent  cords.  Xy- 
menopsis  subnodosus  is  larger  has  a  higher  spire,  and 
coarser  sculpture  than  X.  buccincus^  The  most  distinctive 
features  of  X.  subnodosus  are  the  thickening  oi  the  outer 
hp,  and  the  presence  of  weU  defined  apertural  teeth, 
which  occur  in  New  Zealand  members  of  the  Xi/mcnc 
complex,  but  not  in  any  other  Xymcnopsis.  Xymcnopsis 


subnodosus  is  most  similar  to  X.  corntgatus.  which  is 
restricted  to  the  Mahinas  (Falkland)  Islands.  We  provi- 
sionally retain  this  taxon,  recognizing  that  it  may  prove 
to  be  a  subspecies  or  rare  variant  of  X.  corrugatus  char- 
acterized by  an  atavistic  expression  of  a  rare  allele  or 
alleles  for  apertural  dentition. 

SYSTEMATIC  KEY  TO  THE  GENUS  XYMENOPSIS 

1.   Shell  large,  with  strong,  well  defined  axial  ribs  .  .     2 
Shell  small,  with  weak,  poorly  defined  axial  ribs         3 

2.  Shell  thin,  with  spiral  sculpture  of  flat  cords 
much  broader  than  intervening  spaces,  which 
appear  as  incised  furrows X.  muiicifoiinis 

Shell  thick,  spiral  sculptine  ot  dense,  evenly 
rounded  cords,  (6  on  penultimate  whorl,  22- 
24  on  body  whorl),  broader  than  intervening 
spaces.  Outer  lip  thickened,  with  8-10  weak 
teeth X.  subnodosus 

3.  Spiral  sculpture  of  rounded  cords  slightly 
broader  to  sUghtlv  narrower  than  intervening 
spaces,  producing  weakly  canceUate  tubercles 
at  intersection  with  axial  ribs.  Weak  spiral 
threads  present  between  adjacent  spiral  cords 
X.  corrugatus 

Spiral  sculpture  of  flattish  cords  as  broad  as 
interx'ening  spaces,  forming  weakly  cancellate 

sculpture  at  intersections  with  axial  ribs 

X.  buccinciis 

ACKNOWLEDGMENTS 

We  thank  the  following  people  for  access  to  material  in 
their  collections:  K.  Wav  (BMNH);  A.  Waren  (NHRM); 
P  Bouchet  and  V.  Heros  (MNHN):  J.  H.  McLean 
(LAGM):  P  Mikkelsen  (AMNH);  B.  Hausdorf  (ZMH); 
O.  Galvez  Herrera  (MNHNS);  J.  Boscheinem  (Loeb- 
becke-Museum);  H.  Irigoyen  and  A.  Tablado  (MAGN); 
C.  Ituarte  (MLP).  R.  Prasad  (Auckland  Museum)  pro- 
vided specimens  oi  Xt/mcnc  plcbcius.  P.  Louzet  took  the 
photographs  of  specimens  at  MNHN.  D.  Reid  (BMNH) 
provided  the  presened  specimen  of  X.  subnodosus.  Yu. 
Kantor  provided  useful  discussions  during  the  course  of 
this  work.  V.  Ivanov  prepared  the  anatomical  drawings 
of  X.  muricifonnis.  R.  Petit  provided  information  on 
nineteenth  century  hterature.  E.  H.  Yokes  provided 
helpful  discussions  of  muricid  systematics.  We  are  grate- 
ful to  A.  Beu  and  J.  H.  McLean  for  thorough  reviews 
and  helpful  suggestions  that  improved  this  paper 

This  study  was  conducted  during  a  Postdoctoral  Fel- 
lowship granted  to  the  senior  author  by  the  Consejo  Na- 
cional  de  Investigaciones  Cientificas  v  Tecnicas  (CON- 
ICET),  Argentina,  to  work  at  the  National  Museum  of 
Natural  History,  Smithsonian  Institution,  Washington. 
DC.  It  was  supported  in  part  by  a  Research  Award  from 
the  NSF-USAP  United  States  Antarctic  Program  [Con- 
tract No.  OPP-9509761]  and  a  grant  in  aid  from  the 
Conchologists  of  America  and  the  Walter  E.  Sage  Me- 
moriid  Award. 


G.  Pastorino  and  M.  G.  Harasewvch,  2000 


Page  5" 


LITERATURE  CITED 

Ben,  A.  G.  and  P.  A.  Mawvell.  1990.  Cenozoic  Molluscaof  New 
Zealand.  New  Zealand  Geoloi;ical  Suncx  Palcontological 
Bulletin  58:1-518. 

Bnniet,  R.  1997.  New  species  of  Mollusca  tiDin  the  Entrer- 
riense  Formation  (upper  Miocene)  of  Chuhnt  Province, 
Argentina  and  species  not  pre\iously  reported  from  this 
foniiation;  Part  II,  Gastropoda.  Tulane  Studies  in  Geolog) 
and  Paleontologv-  30:61-98. 

Castellanos,  Z.  J.  A.  de  and  N.  A.  Landoni.  1993.  Muricidae  y 
Thiiisidae.  25  pp.,  4  pis.  In:  Castellanos,  Z.  J.  A.  de,  N.  A. 
Landoni  and  ].  R.  Dadon.  1988-1993.  Catalogo  Desciip- 
tivo  de  la  Malacofamia  Marina  Magallanica.  Comision  de 
Investigaciones  Cientffi'cas  de  la  Provincia  de  Buenos  Ai- 
res. 

Cemohorsky,  W.  O.  1977.  The  taxonomy  of  some  Southeni 
Ocean  Mollusca  (Gastropoda)  mainly  Antarctic  and  Suh- 
antarctic.  Records  of  the  Aukland  Institute  and  Museum 
14:105-119. 

Cossmann,  M.  1903.  Essais  de  Paleoconchologie  comparee. 
Volume  5.  F.  R.  de  Rude\al,  Paris,  151  pp. 

Cuvier,  G.  1797.  Tableau  elementaire  de  I'liistoire  naturelledes 
animau.x.  Paris,  .\-\i  +  710  pp.,  14  pi. 

Dance,  S.  R  1986.  A  histon'  of  shell  collecting.  E.  J.  Bnll, 
Leiden,  265  pp. 

D'Asaro,  C.  N.  1991.  Gunnar  Thorson's  world  wide  collection 
of  prosobranch  egg  capsules:  Muricidae.  Ophelia  351:1- 
101. 

Dell,  R.  K.  1971.  The  marine  Mollusca  of  the  Roval  Society 
Expedition  to  southeni  Chile,  1958-1959.  Records  of  the 
Dominion  Museum  7:155-233. 

Dell.  R.  K.  1972.  Notes  on  the  nomenclature  of  some  Mollusca 
from  Antarctica  and  southeni  South  America.  Records  of 
the  Dominion  Museum  8:21—42. 

Dell,  R.  K.  1990.  Antarctic  Mollusca  with  special  reference  to 
the  fauna  of  the  Ross  Sea.  Bulletin  of  tlie  Royal  Societv 
of  New  Zealand  27:1-311. 

FinlaN',  H.  J.  1926.  New  shells  from  New  Zealand  Tertiary  beds, 
part  2:  Transactions  of  the  New  Zealand  Institute  56:227- 
258. 

Finlay,  H.  J.  1926  [1927].  A  further  commentary  on  New  Zea- 
land molluscan  svstematics.  Transactions  of  the  New  Zea- 
land Institute  57':.320-4S5. 

Fretter,  \'.  and  A.  Graham.  1994.  British  Prosobranch  Molluscs. 
Their  functional  anatonn  and  ecologN.  Re\ised  and  up- 
dated edition.  The  Ray  Society,  London,  820  pp. 

Gmelin,  J.  F.  1791.  Venues.  In:  Caroli  a  Linnei,  Systenia  Na- 
turae per  Regiia  Tria  Naturae.  .  .  Editio  decinia  tertia,  auc- 
ta,  reformata.  Tomo  1  (6).  Beer,  Leipzig,  pp.  3021-3910. 

Gould.  A.  A.  1849.  Shells  collected  by  the  U.  S.  E.x-ploring 
Exjiedition  under  the  command  of  Charles  Wilkes.  Pro- 
ceedings of  the  Boston  Societv  of  Natural  Historv  3:140- 
144. 

Gould,  A.  A.  1852-1861.  Mollusca  and  shells.  In:  United  States 
Exploring  Exjjedition  during  the  years  1839-1842  under 
the  command  of  Charles  Wilkes.  Boston,  510  pp.  [1852]; 
addenda  and  corrigenda,  Philadelphia  [1856];  Atlas.  Phil- 
adelphia, 16  pp.  +  52  hand-colored  pis.,  1856  [1861]. 

Gray,  ].  E.  1839.  Molluscous  Animals.  In:  The  Zoologx' of  Cap- 
tain Beechey's  Voyage:  compiled  from  tlie  cf)llections  and 
notes  made  by  Captain  Beechey,  the  officers  and  naturalist 
of  the  expedition.  During  a  voyage  to  the  Pacific  and  Beh- 
ring's  Straits  performed  in  his  Majesty's  ship  Blossom,  un- 


der the  command  of  Captain  F  W.  Beechev,  in  the  years 
1825.  26,  27  and  28,  H.  G.  Bohn,  London.pp.  10.3^142. 

Hara.sewych.  M.  G.  1984.  Comparative  anatomy  of  four  prim- 
iti\e  muricacean  gastropods.  Implications  for Trophoninae 
phylogeny.  American  Malacological  Bulletin  3:11-26. 

Hupe.  L.  H.  i854.  Fauna  Chilena.  Mollusca.  In:  C.  Gay  (ed.), 
Historia  Fi'sica  v  Politica  de  Chile.  Tomo  8.  Zoologia.  Pub- 
lished pri\'ately  and  b\  Vluseo  de  Historia  Natural  de  San- 
tiago, Paris  and  Santiago,  499  pp. 

Ihering.  H.  \on  1905-1907.  Les  Mollusques  fossiles  du  Ter- 
tiaire  et  du  Gretace  superieur  de  I'Argentine.  Anales  del 
Museo  Nacional  de  Buenos  Aires,  Serie  3,  7:1-611.  [pub- 
lished in  38  parts,  Nov.  1905-sep.  1907] 

Jablonski.  D.  and  R.  A.  Lutz.  1980.  Molluscan  lanal  shell  mor- 
phology': ecological  and  paleontological  applications.  In: 
D.  C.  Rhoads  and  R.  A.  Lutz  (eds.)  Skeletal  growth  of 
aquatic  organisms.  Plenum  Press,  New  York,  pp.  323-377. 

King.  P.  P.  and  W.  J.  Broderip.  1832.  Description  of  the  Cir- 
rhipedia,  Conchifera  and  Mollusca  in  a  collection  formed 
by  the  officers  of  H.M.S.  Adventure  and  Beagle  employed 
between  the  years  1826  and  1830  in  surveying  the  south- 
ern coasts  of  South  America,  including  the  straits  of  Ma- 
galhaens  and  the  coast  of  Tierra  del  Fuego.  The  Zoological 
Joumal  5:.332-.349. 

Kobelt,  W.  1878.  Die  geschiiwiizten  und  bewehrten  Piiqiur- 
schnecken  (Murex.  Ranelln,  Tritonhim,  Trophon.  Hindsia). 
In  Abbildungen  nach  der  Natur  mit  Beschreibungen.  Be- 
gonnen  von  H.  C.  Kiister,  fortgesetzt  und  beendet  \on  W. 
Kobelt.  In:  Systematisches  Conchylien-Cabinet  von  Mar- 
tini und  Chemnitz.  Neu  herausgegeben  und  vervollstan- 
digt  \on  H.  C.  Kiister  (nach  dessen  Tode  fortgesetzt  von 
W.  Kobelt  und  H.  C.  Winkauff).  Verlag  von  Bauer  and 
Raspe,  Niiniberg,  336  pp. 

Kool,  S.  P.  1993a.  Phylogenetic  analysis  of  the  Rapaninae  (Neo- 
gastropoda:  Muricidae).  Malacologia  35:155-259. 

Kool,  S.  P.  199.3b.  The  systematic  position  of  the  genus  Nucella 
(Prosobranchia:  Muricidae:  Ocenebrinae).  The  Nautilus 
107:43-57. 

Lamarck.  J.  B.  P.  A.  de  M.  de.  1816.  Tableau  encvclopedique 
et  niethodique  des  trois  regnes  de  la  nature,  vingt-troisie- 
me  partie.  Mollusques  et  polypes  divers.  Paris.  Chez 
Mnie.  Veuve  Agasse. 

Lamy  E.  1906.  Gastropodes  Prosobranches  et  Pelec)podes: 
Expedition  Antarcticjue  Fran^aise  (190.3-1905)  comnian- 
dee  par  le  Dr  Jean  Charcot.  Sciences  Naturelles:  Docu- 
ments Scientifiques.  Masson  et  Cie.  Paris,  pp.  1-20. 

Mabille  J.  and  A.  T  Rochebnme.  1889.  [see  Rochebnnie,  A. 
T  and  J.  Mabille,  1889] 

MeMIl,  ].  C.  and  R.  Standen.  1907.  The  marine  Mollusca  of 
the  Scottish  National  Antarctic  Expedition.  Transactions 
of  the  Royal  Society  of  Edinburgh  46:119-157. 

MeMll,  J.  C.  and  R.  Standen.  1912.  The  marine  Mollusca  of 
the  Scottish  National  Antarctic  Expedition.  Part  11.  Trans- 
actions of  the  Royal  Societx'  of  Edinburgh  48:333-366. 

Molina,  G.  1.  1810.  Saggio  sulla  storia  naturale  del  Chile  (2nd 
ed.).  Bologna,  .306  pp. 

Montfort.  P.  D.  de.  1810.  Conchyliologie  systematique.  et  clas- 
sification niethodique  des  coquilles;  offrant  leurs  figures, 
leur  arrangement  generique.  leurs  descriptions  caracter- 
istiques,  leurs  noms;  ainsi  que  leur  s\non\inie  en  plusieurs 
langues.  Coquilles  Uni\aKes,  non  cloisonnees.  Tome  2.  F 
Schoell,  Paris,  676  pp. 

Philippi.  R.  A.  1845.  Diagnosen  einiger  neuen  Conclnlieu.  Ar- 
chiv  fiir  Naturgeschichte  11:50-71. 


Page  58 


THE  NAUTILUS,  Vol.  114,  No.  2 


Philippi,  R.  A.  1846.  Abbildungen  unci  Beschreibungen  neuer 
oder  wenig  gekannter  Conclnlien  herausgegeben.  Tomo 
2,  Cassel. 

Philippi,  R.  A.  1855.  Observaciones  sobre  las  conchas  de  Ma- 
gallanes.  Anales  de  la  Universidad  de  Chile  5:203-213. 

Philippi,  R.  A.  1868.  Conchylia  nova  potissimum  niageljanica. 
Malakozoologische  Blatter  15:22.3-226. 

Ponder,  W.  F.  1972.  A  re\iew  of  the  genus  Xijmcnc  Iredale  of 
New  Zealand  (Mollusca:  Muricidae).  Journal  of  the  Royal 
Society  of  New  Zealand  2:471-499. 

Powell,  A.  W.  B.  1951.  Antarctic  and  subAntarctic  Mollusca: 
Pelecypoda  and  Gastropoda.  Discovery  Reports  26:47- 
196. 

Powell,  A.  VV.  B.  1971.  New  Zealand  nioUuscan  svsteniatics 
with  descriptions  of  new  species:  part  7.  Record  of  the 
Auckland  Institute  and  Museum  8:209-228. 

Radwin,  G.  E.  and  A.  D'Attilio.  1976.  Murex  Shells  of  the 
world:  an  illustrated  guide  to  the  Muricidae.  Stanford  Uni- 
versity Press,  284  pp. 

Rafinesque,  C.  S.  1815.  Analyse  de  la  Nature,  on  Tableau  de 
rUnivers  et  des  corps  organises.  Barravecchia,  Palermo, 
224  pp. 

Reeve,  L.  1847-1848.  Conchologia  Iconica,  or  Illustrations  of 
the  shells  of  molluscous  animals.  Volume  4,  containing 
monographs  of  the  genera  Chama.  Chiton,  Chitonelliis, 
Ficula,  Pi/nda,  Tiirbinella.  Fasciolaria.  Fiisiis.  Pahtdoimm. 
Turbo.  Ree\e,  Benhan,  and  Ree\e,  London,  238  pp. 

Rochebrune,  A.  T.  de  and  J.  Mabille.  1889.  Mollnsques.  In: 
Mission  Scientifique  du  Cap  Honi.  1882-1883.  Volume  6 
Zoologie,  Part  2.  Gauthier-\'illars  et  fils,  Paris,  126  pp. 

Rousseau,  L.  1854.  Description  des  Mollusques  coquilles  et 
zoophytes. /«.■  Voyage  an  Pole  Sud,  et  dans  I'Oceanie  sur 
les  cor\'ettes  I'Astrolabe  et  la  Zelee:  execute  par  ordere  du 
roi  pendant  les  annees  1837-1838-18.39-1840,  sous  le  com- 
niandement  de  M.  J.  Dumont-D'Ur\ille,  Capitaine  de 
vaisseau.  T.  5,  Zoologie.  E.  Thunot  et  Cie.,  Paris,  132  pp. 

Sherbom,  C.  D.  1924.  Index  Animalium,  sive  index  nominum 
quae  ab  A.  D.  MDCLVIII  generibus  et  speciebus  ani- 
malium imposita  sunt.  Section  2,  1801-1850  part  5.  Index 
C-Ceyl.  Oxford  Uni\-ersity  Press,  London,  pp.  945-1196. 

Shuto,  T.  1974.  Lanal  ecology  of  prosobranch  gastropods  and 
its  bearing  on  biogeography  and  paleontology.  Lethaia  7: 
239-256. " 

Smith,  E.  A.  1881.  Account  of  the  Zoological  Collections  made 
during  the  Sur\'ey  of  H.M.S.  'Alert'  in  the  Straits  of  Ma- 
gellan and  on  the  Coast  of  Patagonia.  Mollusca  and  Mo- 
luscoida.  Proceedings  of  Zoological  Societv  of  London 
1881:22-44. 
Smith,  E.  A.  1907.  Mollusca  2,  Gastropoda.  In:  National  Ant- 
arctic Expedition  1901-1904.  Natural  History,  2  Zoology 
(\'ertebrata:  Mollusca:  Cnistacea).  'W.  Clowes  and  Sons, 
Ltd.,  London,  pp.  1-12  pp. 


Smith,  E.  A.  1915.  Mollusca  Part  1.  Gastropoda  Prosobranchia, 
Scaphopoda  and  Pelecvpoda.  In:  British  Antarctic  { "Terra 
Nova")  Expedition,  1910.  Natural  Histoiv  Reports.  Zool- 
ogy, volume  2,  part  4.  British  Museum,  London,  pp.  61- 
112. 

Solem,  G.  A.  1972.  Malacological  apphcations  of  scanning  elec- 
tron microscopy.  II.  Radular  stnicture  and  functioning. 
The  Veliger  14:327-336. 

Sowerby,  G.  B.  1839.  In:  J.  E.  Gray,  Molluscous  Animals.  In: 
The  Zoology  of  Captain  Beechev's  Voyage;  compiled  from 
the  collections  and  notes  made  by  Captain  Beechey,  the 
officers  and  naturalist  of  the  expedition,  during  a  voyage 
to  the  Pacific  and  Behring's  Straits  performed  in  his  Maj- 
esty's ship  Blossom,  under  the  command  of  Captain  F.  W. 
Beechey,  in  the  years  1825,  26,  27  and  28,  H.  G.  Bohn, 
London,  pp.  14:3^155,  pis.  33-44. 

Sowerby,  G.  B.  II.  1880.  Monograph  of  the  Subgenus  Trop/ion. 
Thesaurus  Conchyliorum,  or  Monographs  of  Genera  of 
Shells.  4(:35):59-67,  pis.  1-3. 

Strebel,  H.  1904.  Beitriige  zur  Kenntnis  der  MoUuskentauna 
der  Magalhaen-Pro\dnz.  Zoologischen  Jahrbiichem  21: 
171-248" 

Strebel,  H.  1908.  Die  Gastropoden.  In:  W'issenschaftliche  Er- 
gebnisse  der  Schwedischen  Siidpolar-Ex-pedition  1901- 
1903  Unter  Mitwirkung  Zahlreicher  Fachgenossen  Her- 
ausgegeben von  Otto  Nordenskjold  Leiter  der  Expedition. 
Tome  6,  Zoologie  2(1).  Stockliolm,  pp.  1-112. 

Thorson,  G.  1950.  Reproductive  and  lar\'al  ecology  of  marine 
bottom  imertebrates:  Biological  ReMie  25:1—45. 

Tryon,  G.  \\'.  Jr  1880.  Manual  of  Conchology;  Stnictural  and 
Systematic,  with  illustrations  ot  the  species.  Vol.  2.  Mur- 
icinae,  Puqjurinae.  Philadelphia,  289  pp,  70  pis. 

Vokes,  E.  H.  1971.  The  geologic  histoiy  of  the  Muricinae  and 
the  Ocenebrinae.  The  Echo  4:.37-.54. 

Vokes,  E.  H.  1991a,  Collecting  trophons  in  Argentina,  Tierra 
del  Fuego.  Part  1.  American  Conchologist  19(1):7-10. 

Vokes,  E.  H.  1991b.  Collecting  trophons  in  Argentina.  The 
N'aldes  Peninsula.  Part  2.  American  Conchologist  19(2):8- 
11. 

Vokes,  E.  H.  1992.  Argentine  trophons  revisited  or  Dr  Powell, 
I  owe  you  an  apology.  American  Conchologist  20(1):.3— 4. 

■Watson,  R.  B.  1886.  Report  on  the  Scaphopoda  and  Gastero- 
poda collected  by  H.M.S.  Challenger  during  the  years 
1873-1876.  In:  Reports  on  the  Scientific  Results  of  the 
Voyage  of  the  H.M.S.  Challenger  during  the  years  1873- 
1876.  Zoolog)'  \olume  15,  part  41,  pp.  1-608;  part  42,  pp. 
609-756,  pis.'  1-.50  -I-  1-3  [Caecidae]. 

Wenz,  W.,  1938-1944.  Gastropoda,  Teil  I:  Allgemeiner  Teil 
und  Prosobranchia,  Handbuch  der  Palaozoologie.  Verlag 
\'on  Gebriider  Bomtraeger,  Berlin,  1639  pp. 

Zinsmeister,  ^^'.  ].  1982.  Late  Cretaceous-Early  Tertian-  mol- 
luscan  biogeograph\  of  the  southern  circum-Pacific.  Jour- 
nal of  Paleontology'  56:84-102. 


THE  NAUTILUS  114(2):59-68.  2000 


Pa^e  59 


Copulahijssia  riosi,  a  new  deep-sea  limpet  (Gastropoda: 
Pseudococculinidae)  from  the  continental  slope  off  Brazil  with 
comments  on  the  systematics  of  the  genus 


Jose  H.  Leal 

The  BaileN-Mattliews  Shell  Muscinii 
P.O.  Bo.x  1580 
Sanibel,  FL  33957  USA 
jleal@gate.net 


Luiz  Ricardo  L.  Sinione 

Museu  lie  Zoologia  da  Uiii\ersidade 

de  Sao  Paulo 
Cai.xa  Postal  42694 
04299-970  Siio  Paulo 
BRAZIL 
lrsimoiie@usp.br 


ABSTRACT 

A  new  bathval  species  of  the  family  Pseudococculinidae  is  de- 
scribed from  samples  dredged  in  1987  along  the  continental 
slope  of  southeastern-eastern  Brazil  b>  the  French  research 
vessel  Marion-Diifresne.  The  new  species,  allocated  in  the  ge- 
nus Copulahijssia.  differs  from  the  4  other  known  congeners 
by  characters  of  shell,  external  morphology,  mantle  cavity,  and 
radula.  Descriptions  of  internal  moqihology  based  on  micro- 
dissections are  provided,  including  the  digestive  system  and  the 
complex  muscular  apparatus  ot  the  odontopliore.  The  distri- 
bution of  characters  ot  taxonomic  significance  at  the  species 
level  in  Copulabyssia  is  given  in  tabular  format  and  discussed. 

Key  words:  MoUusca,  Cocculiniformia,  Vetigastropoda,  Lepe- 
telioidea,  bathval,  anatomy,  taxonomy,  odontopliore. 


INTRODUCTION 

Bathyal  samples  dredged  off  E-SE  Brazil  yielded  spec- 
imens of  an  unnamed  species  of  the  genus  Copulabi/ssia, 
family  Pseudococculinidae.  Two  live  specimens  and  one 
empty  shell  were  collected  at  three  stations  performed 
between  960-L320  m.  The  Pseudococcuhnidae  and  oth- 
er families  comprising  the  superlamily  Lepetelloidea 
Dall,  1882,  are  included  amongst  the  mollusks  living  at 
greatest  depths  in  the  ocean  (Leal  and  Harasewych, 
1999;  Lesicki,  1998;  Marshall,  1986;  Wolff,  1979).  The 
new  taxon  exhibits  the  diagnostic  characters  of  the  genus 
Copulabyssia  Haszpmnar,  1988.  Copulabyssia  riosi  new 
species  is  herein  compared  with  the  4  other  species  of 
the  genus  named  to  date  (Lesicki,  1998):  C.  corni^ata 
(Jeffreys.  1883)  (NE  Atlantic);  C.  ii,mda1a  (Marshall, 
1986)  (SW  Pacific);  C.  Icptah-a  (Verrill,  1884)  (N  Atlan- 
tic); C.  similaris  Hasegawa,  1997  (Japan). 

MATERIALS  AND  METHODS 

Two  live-collected  specimens  and  one  empty  shell  were 
available  for  studv.  A  single  specimen  was  dissected.  Mi- 


crodissection was  performed  with  the  specimen  im- 
mersed in  70%  ethanol.  The  apical  (distal)  region  of  vis- 
ceral mass  was  damaged;  adjacent  structures  were  de- 
scribed and  are  shown  herein  based  on  reconstruction. 
After  dehydration  in  a  standard  alcoholic  series,  the  sec- 
ond specimen  was  chemically  dried  tor  10  min  in  hex- 
amethykUsilizane  (HMDS).  Shells,  radula,  and  chemi- 
cally dried  soft  parts  were  coated  with  carbon  and  gold 
and  e.xamined  and  photographed  under  Hitachi  S-57() 
(at  National  Museum  of  Natural  History,  Smithsonian 
Institution),  Leo  440  (at  MZSP),  or  Philips  XL30 
ESEM-FEG  (Center  for  Advanced  Microscopy,  Univer- 
sity of  Miami)  electron  microscopes.  Institutional  abbre- 
viations used  are;  MNHN,  Museum  national  dTIistoire 
naturelle,  Paris,  France;  MZSP,  Museu  de  Zoologia  da 
Universidade  de  Siio  Paulo,  Brazil;  NMNH,  Museum  of 
New  Zealand  Te  Papa  Tongarewa,  Wellington;  NSMT, 
National  Science  Museum,  Tokyo,  Japan;  SMNH,  Natur 
Historika  Riksmuseet,  Stockliolm,  Sweden;  USNM,  Na- 
tional Museum  of  Natural  History,  Smithsonian  Institu- 
tion, Washington.  Other  alilirexiations  and  text  conven- 
tions are:  1,  5,  lateral  teeth  (lateral  5  =  pluricuspid 
tooth);  1,  2,  marginal  teeth;  ac,  anterior  cartilages  of 
odontophore;  an,  anus;  bm,  buccal  mass;  br,  subradular 
membrane;  bs,  Iniccal  sphincter;  df,  dorsiil  folds  oi  buc- 
cal mass;  dg,  digestive  gland;  ep,  epipodium;  es,  esoph- 
agus; et,  epipodial  tentacles;  gi,  gill;  go,  gonad;  gp,  gen- 
ital pore;  if,  inner  mantle  fold;  in,  intestine;  ir,  insertion 
of  m4  in  radular  sac;  is,  insertion  ot  m5  in  radular  sac; 
ki,  kidney;  It,  left  cephalic  tentacle;  mb,  mantle  edge; 
mj,  jaw  and  peribuccal  muscles;  mo,  mouth;  mp,  me- 
sopochum;  m2,  pair  of  buccal  mass  retractors;  m3,  inner 
buccal  mass  protractors;  m4,  pair  of  antero-dorsal  ten- 
sors; mS,  pair  of  postero-dorsal  tensors;  ni6,  horizontal 
muscle;  ni7,  pair  of  odontophore  approximators;  mS, 
pair  ot  cartilage  approximators;  no,  nuchal  ca\its:  of, 
outer  mantle  fold;  pa,  mantle  papillae;  pc,  posterior  car- 
tilages of  odontophore;  pr,  protuberance  on  right  ten- 


Page  60 


THE  NAUTILUS,  Vol.  114,  No.  2 


tacle;  re,  rectum;  ra,  radula;  m,  radiilar  caecum;  rs, 
radular  sac;  rt,  right  cephalic  tentacle;  sc,  subradular 
cartilage;  sf,  fold  along  gill;  sg,  seminal  groove;  sm,  shell 
muscle;  sn,  snout;  st,  stomach;  vg,  vestigial  left  gill  leaf- 
let. The  odontophore  muscles  examined  in  this  studv 
were  also  described  bv  Haszpnuiar  (1988:  165-167,  fig. 
.3)  for  Kluilahijssia  vcnczuclcnsis  McLean,  1988.  Ter- 
minology used  herein  (boldface)  corresponds  to  the  one 
used  bv  HaszpiTuiar  as  follows:  mj  =  br  -I-  otr;  bs  =  bs; 
in2  =  dr2;  in4  =  rd2;  m5  =  rr;  m6  =  ho;  m7  =  sr3 
(?);  mS  =  ci.  The  remaining  small  muscles  described  by 
Haszpnmar  (1988)  could  not  be  observed  in  the  single 
specimen  a\';ulable  for  microdissection. 

SYSTEMATICS 

Class  Gastropoda  Cuvier,  1797 

Subclass  Orthogastropoda  Ponder  and  Lindberg,  1996 

Order  Vetigastropoda  Salvini-Plawen,  1980 

Remarks:  The  superfamilies  Cocculinoidea  Dall, 
1S82,  and  Lepetelloidea  Dall,  1882  have  been  united 
under  the  order  Coccuhformia  Haszprunar,  1987.  Re- 
cent studies  of  gastropod  phylogeny  based  on  moi"pho- 
logical  characters  (Ponder  and  Lindberg,  1996;  1997) 
suggest  that  CoccuMnoidea  is  sister  taxon  to  Neritopsina, 
and  that  Lepetelloidea  is  part  of  the  order  Vetigastro- 
poda (see  also  Haszpioinar,  1998:  664).  This  arrangement 
is  followed  herein.  Harasewvch  ct  al.  (1997)  indicated, 
based  on  partial  18S  rDNA  sequences,  that  CoccuHni- 
formia  may  not  constitute  a  clade.  These  latter  authors, 
however,  suggested  that  Cocculinoidea  and  Lepetello- 
idea are  more  closely  related  to  each  other  and  to  Pa- 
teUogastropoda  than  to  Neritopsina  or  Vetigastropoda. 

Superfamily  Lepetelloidea  Dall,  1882 
Family  Pseudococcuhnidae  Hickman,  1983 
Subfamily  Caymanabyssinae  Marshall,  1986 
Genus  Copiilabijssia  Haszprunar,  1988 

T\pe  species:  CoccuUna  oirn/gflffl  Jeffreys,  1883,  p. 
394,  pi.  44,  figs.  2-2a.  Type  locaUty-;  59°40'N,  7°21'W, 
930  m,  Triton  Ex]:)ecUtion  station  10. 

Diagnosis:  Protoconch  sculpture  of  microscopic  pris- 
matic crvstals;  right  cephalic  tentacle  exceptionallv  large, 
"swollen";  right  cephalic  tentacle  with  deep,  open  glan- 
dular seminal  groove  situated  postero-dorsallv;  large 
pedal  gland;  gills  palhal,  several  well-developed  gill  leaf- 
lets on  right  side,  a  single  one  on  left  side. 

Copidabi/ssia  riosi  new  species 
(Figures  1-24,  Table  1) 

Diagnosis:  Teleoconch  at  first  with  radial  sculpture 
then  with  closely  set  concentric  ribs  (interspaces  about 
25  (Jim  in  width);  internally  with  narrow  but  well-defined 
anterior  apical  septum;  radial  sculpture  only  on  early  te- 
leoconch; cutting  area  of  rachidian  tooth  with  blunt 
cusp;  left  side  of  subpalhal  cavity  with  small  structure 


that  could  represent  a  vestigial  gill  leaflet;  right  cephalic 
tentacle  with  prominent  subterminal  protuberance;  left 
cephahc  tentacle  small  for  genus;  inner  branches  of  epi- 
podial  tentacles  filamentous. 

Description:  Shell  (Figures  1-6):  Small  (length  to 
3.50  nun),  thin,  arched,  elevated  (height/length  about 
0.5),  with  apex  at  posterior  quarter  pointing  in  posterior 
direction.  Anterior  slope  convex,  about  88%  of  shell 
length.  Posterior  slope  concave,  about  30%  of  shell 
length.  Shell  surface  usually  lightlv  eroded,  milky  white. 
Protoconch  (figure  4)  length  about  200  |jLm,  surface  bad- 
ly eroded  in  type  material.  Teleoconch  sculpture  of  con- 
centric (commarginal)  ribs  separated  by  intervals  of 
about  25  |jLm.  Concentric  ribs  narrow  (about  10  jim)  and 
crossed  bv  fine  radial  lines  on  earlv  part  of  teleoconch 
(figure  5).  Concentric  ribs  on  latter  part  of  teleoconch 
proportionally  thicker,  radial  lines  absent  (figure  6).  In- 
ternally with  narrow,  but  well-defined,  anterior  apical 
septum.  Aperture  elhptical  (figure  3),  not  conforming  to 
a  planar  surface,  convex.  Shell  muscle  scar  continuous, 
horseshoe-shaped. 

Head-foot  (Figures  7,  8,  13,  14,  17):  Head  prominent 
and  large  (about  %  foot  length)  (figures  7,  13,  14,  17). 
Cephahc  tentacles  (figures  7,  13,  14,  17,  rt.  It)  on  dorso- 
lateral surface  of  head.  Right  tentacle  about  4  times  larg- 
er  than  left  tentacle,  modified  as  copulatory  organ  (de- 
scribed below).  Eyes  apparently  lacking  (unpigmented?). 
Snout  (figures  7,  13,  14,  16,  17,  sn)  very  large,  cyhndrical 
on  base,  gradualK'  becoming  slightK'  flattened  dorso-ven- 
trally.  Anterior  extremity  of  snout  almost  flat,  broader 
than  region  immediately  behind  it,  with  pair  of  small 
lateral  projections.  Foot  large  and  flat  (figures  7,  13,  14), 
smaller  than  shell  aperture  in  presened  animal.  Meso- 
pochum  (figures  7,  13,  14,  mp)  flattened.  Epipodium 
(figures  7,  13,  14,  ep)  flange-Uke,  surrounthng  entire  me- 
sopodium,  larger  on  anterior  edge,  which  covers  baso- 
ventral  region  of  snout.  Pair  of  epipodial  tentacles  (figure 
7,  9,  13,  14,  et)  well-developed,  projecting  from  poste- 
rior region  at  both  sides  of  longitudinal  axis  of  animal, 
slightly  dorsal  to  epipodium,  on  intersection  between 
mantle  and  shell  muscle.  Each  epipothal  tentacle  bifid; 
outer  branch  shorter  and  with  broad  tip;  inner  branch 
longer  and  pointed.  Shell  muscle  (figures  15-17,  sm) 
horseshoe-shaped  (concaNdty  towards  anterior  region); 
anterior  extremities  broader,  pointing  away  from  shell 
apex.  Almost  all  head-foot  muscles  converge  toward  in- 
sertion of  shell  muscle. 

Mantle  cavity  (Figures  12-15,  17):  Mande  edge  (fig- 
ures 12-14,  17,  mb)  thick,  with  2  folds;  outer  fold  (fig- 
ures 12,  14,  17,  ofO  covered  by  slender  and  very  small, 
translucent  papillae  (figure  12);  inner  fold  (figures  14, 
17,  if)  about  twice  as  broad  as  outer  fold,  richlv  glan- 
dular, white.  A  small,  transparent,  yellowish  protuber- 
ance (figure  13,  vg)  on  inner  margin  of  inner  told,  in 
region  adjacent  to  left  cephafic  tentacle,  may  represent 
vestigial  left  gill  leaflet.  Mantle  cavity  surrounding  entire 
head-foot,  somewhat  deep,  i.e.,  with  ample  space  be- 


J.  H.  Leal  and  L.  R.  L.  Simone,  2000 


Page  61 


Figures  1-6.  Coptilabt/ssia  riosi  new  species:  shell  of  holotrvpe,  MZSP  32150.  1.  Dorsal  view.  2.  Lateral  \iew.  3.  X'entral  view. 
Scale  line  =  1.0  mm.  4.  Protoconch.  Scale  line  =  100  p-m.  5-6.  Shell  sculpture.  5.  On  early  teleoconch.  Scale  hne  =  50  jxm.  6. 
Close  to  shell  margin.  Scale  line  =  100  [xm. 


tween  mantle  edge  and  shell  muscle.  Nuchal  cavitv  (fig- 
ures 14,  15,  17,  nc)  deeper  than  remainder  of  mantle 
cavity.  Gill  (figures  13,  14,  17,  gi)  small,  situated  on  right 
side  of  mantle  cavity,  somewhat  parallel  to  mantle  edge 
and  extending  for  about  half  of  foot  lenijth,  consisting  of 
aliout  14  successively  smaller  leaflets.  Anterior  extremity 
oi  gill  (figure  5,  gi)  just  posterior  to  head-foot  Umit.  Each 
leaflet  low  and  triangular,  relatively  thick.  Most  posterior 
leaflets  situated  peqjendicularlv  to  mantle  edge,  gradu- 
ally becoming  obfique  to  it  on  anteri(jr  region.  A  small, 
narrow,  and  low  fold  (figure  14,  sf)  nms  between  gill 
and  shell  muscle  along  posterior  half  of  gill.  Anus  (fig- 
ures 15-17,  an)  and  genital  pore  (figure  l7,  gp)  on  right 
side,  at  head  and  foot  hniit,  just  anterior  to  right  extrem- 
ity of  shell  muscle. 

Circulatonj    and   excretory    sijstem.'i    (Figures    15-17): 


Heart  not  observed.  Large  blood  vessels  ;ilong  thick- 
mantle  border  Kidney  (figures  15,  17,  la)  verv  small, 
white,  sohd,  slightly  triangular;  situated  in  right-posterior 
region  ot  head,  just  posterior  to  rectum. 

Visceral  mass  (Fifiures  15,  16):  Round,  surrounded  by 
following  stmctures:  laterally  and  in  posterior  direction 
by  shell  muscle,  in  anterior  direction  by  buccal  mass, 
ventraliv  by  inner  surface  of  mesopodium,  and  dorsallv 
by  central  region  of  mantle  and  shell.  Gonad  (figure  15, 
go)  and  digestive  gland  (figure  15,  dg)  cream  in  color, 
intersected  by  several  intestinal  loops  (figure  16).  Gonad 
situated  mainly  on  left  region  and  digestive  gland  mainlv 
on  central  region. 

Digestive  si/sfem  (Figures  7.  13-24):  Mouth  (figures  7, 
13,  14,  18,  mo)  on  middle-ventral  region  oi  anterior  sur- 


Page  62 


THE  NAUTILUS,  Vol.  114,  No.  2 


Figures  7-12.  Copulabi/ssit]  riosi  new  species;  paraUpe,  MNHN  unnumbered,  gross  external  moipholoi^  and  radula.  7.  N'entral 
\ie\v  ot  animal.  Scale  line  =  .500  p,m.  8.  Detail  of  right  cephalic  tentacle  (copulatorN-  organ).  9.  Inner  branches  ot  epipodial  tentacles 
(damaged  during  preparation).  Scale  Unes  =  100  |j.m.  10-11.  Radula.  10.  View  of  5  rows.  Scale  hue  =  50  (jim.  11.  Detail  of  lateral 
and  first  marginal  teeth.  Scale  line  =  10  [j.m.  12.  Papillae  on  outer  mantle  fold.  Scale  line  =  .50  |xm. 


face  oi  snout,  oriented  transversally.  Buccal  mass  (figure 
16,  bm)  somewhat  large,  occupying  about  entire  head 
cavit\'.  Buccal  sphincter  (figure  18,  19,  bs)  well  devel- 
oped. Superficial  protractor  muscle  of  buccal  mass  rel- 
atively narrow,  connected  to  lateral  and  dorsal  regions  of 
buccal  sphincter  and  adjacent  region  of  snout,  iiinning 
along  lateral  and  dorsal  sinface  of  buccal  mass,  con- 
nected to  lateral  and  ventral  region  of  buccal  mass.  Dor- 
sal wall  of  buccal  mass  ver\'  thin,  with  pair  of  thick  lon- 
gitudinal folds  (figure  19,  df)  that  join  together  in  an- 
terior direction  at  median  fine,  becoming  gradiuillv 
weaker  toward  esophagus  (figures  16,  19,  es).  Odonto- 
phore  (figures  18-24)  large,  comprising  most  of  buccal 
mass,  oval  in  outline.  Odontophore  muscles  (figures  18- 
24):  (ml)  (not  illustrated)  several  small  and  short  fibers 


connecting  buccal  mass  to  adjacent  inner  lining  of  snout, 
more  concentrated  on  dorsal  surface;  (m2)  (figures  18, 
19)  pair  of  narrow  retractor  muscles  of  buccal  mass  con- 
nected to  inner  ventral  lining  of  snout  just  posterior  to 
buccal  mass,  running  parallel  to  radular  sac  (figures  19- 
23,  rs),  connected  to  postero-dorsal  surface  of  buccal 
mass  on  lateral  surface  of  beginning  of  esophagus;  (m3) 
(figure  21,  22)  pair  of  thin  inner  protractor  muscles  of 
buccid  mass  connected  to  inner  lateral  lining  ot  mouth, 
running  on  (and  covering)  lateral  surface  of  odonto- 
phore, connected  to  antero-ventral  margin  of  posterior 
cartilages;  (m4)  (figures  20-24)  pair  of  antero-dorsal  ten- 
sor muscles  connected  in  part  to  outer  ventral  surface 
of  anterior  cartilages  and  in  part  to  lateral  surface  of 
posterior  cartilages,  running  along  (and  covering)  pos- 


J.  H.  Leal  and  L.  R.  L.  Simone,  2000 


Page  63 


Figures  13-17.  Copiilabi/ssin  nasi  new  species;  paratope,  MZSP  32149,  gross  inorphologv.  13.  \'entral  \ne\v.  14.  Lateral  \iew, 
right  side,  margin  of  mantle  deflected  to  show  its  inner  surface.  15.  Dorsal  \iew.  16.  Digestixe  system,  dorso-lateral  \iew,  left  side, 
shown  by  transparency,  only  dorsal  contour  and  part  of  shell  muscle  shown.  17.  Detail  of  head,  lateral  view,  right  side,  adjacent 
mantle  removed  by  means  of  an  incision  around  shell  muscle.  Scale  lines  =  500  jjim. 


Page  64 


THE  NAUTILUS,  Vol.  114,  No.  2 


mo 


J.  H.  Leal  and  L.  R.  L.  Siiiioiu',  2()()() 


Page  65 


Table  1.  Diagnoses  of  species  oi  Copulnl}yssia.  "Shell  length"  is  miLximinn  size  for  species,  "Rih  interspaces"  is  the  distance  between 
concentric  ribs  on  posterior  shell  slope.  "Lett  gill  leaflet"  is  size  of  single-leaflet  gill  in  relation  to  right  gill  leaflets;  medinni  wonld 
be  about  the  same  size  as  right  gill  leaflets.  "Right  cephalic  tentacle"  =  copulator>-  organ  ("RT").  Unknown  character  states  indicali'd 
b\'  "NA",  Degree  of  de\elopment  of  cephalic  tentacles  is  relatixe  to  other  congeneric  species. 


C.  riosi 

C.  Icptalea 

C.  t^iriddta 

C.  cornii^ata 

C.  similah.s 

Di.stribution 

Off  SE,  E  Brazil 

NW  Atlantic 

Off  New  Zealaml 

NE  Atlantic.  Med 

japan 

Character 

Shell  leiiijth  (nun) 

3.50 

2.45 

1.95 

1.35 

3.36 

Rib  interspaces  (jxni) 

±25 

±40 

±40 

±130 

±50 

Radial  sculpture 

early  Icleo  only 

strong 

lacking 

present 

very  weak 

Postition  of  apex 

anterior  2/10 

anterior  .3/10 

anterior  .'VIO 

anterior  1/10 

variable 

Apical  septum 

present 

absent 

absent 

absent 

absent 

Rachidian 

broad 

broad,  rounti  sides 

broad 

elongate 

broad 

Racliidian  cutting  area 

blunt  cusp 

long,  pointed  cusp 

cuspless,  rounded 

cuspless,  s(juarish 

cuspless,  round 

Cusjis  lateral  tooth  5 

2-3 

4 

NA 

2 

2 

Epipodial  tentacles 

filamentous 

NA 

NA 

stubby 

present 

Right  ce  ihalic  tentacle 

well-developed 

NA 

swollen  base 

swollen  base 

"bilobed" 

Left  cep  lalic  tentacle 

vei-y  reduced 

NA 

normal 

normal 

normal 

RT  protuberance 

well-deyeloped 

NA 

absent 

absent 

absent 

Left  gill  leaflet 

?vestigial 

NA 

medium 

small 

Pabsent 

References 

this  work 

Waren,  1991 

Marshall.  19.S6 

Haszpnuiar  1988 

Hasegawa,  1997 

McLean  and  Hara- 

Ilaszprunar,  1988 

Dantart  and 

sewych,  1995 

Luque,  1994 

tenor  surface  of  odontophore  (figure  24),  connected  to 
ventral  surface  of  radular  sac  on  region  posterior  to 
emergence  of  radula;  (m5)  (figures  20-24)  pair  of  pos- 
tero-dorsal  tensor  muscles  connected  to  postero-ventral 
surface  of  posterior  cartilages,  ninning  dorsallv  and  me- 
dially along  (and  covering)  posterior  surface  ot  odonto- 
phore, connected  to  radular  sac  just  posterior  to  m4  in- 
sertion; m6)  (figures  20,  24)  horizontal  muscle  relatively 
narrow,  connected  to  antero-ventral  margin  (jf  anterior 
cartilages  along  about  %  oi  their  length;  m7)  (figiu'es  20, 
22,  24)  pair  of  narrow  ventral  tensor  muscles  connected 
to  meso-ventral  margin  of  posterior  region  of  anterior 
cartilages,  running  along  meso-ventral  surface  of  odon- 
tophore, connected  to  antero-ventral  e.xtremih  of  sub- 
radular  cartilage  (figures  21-23,  sc);  m8)  (figures  10,  12) 
pair  of  broad  approximator  muscles  of  cartilages  con- 
nected to  antero-lateral  surface  of  posterior  cartilages 
(figures  22,  24,  pc),  nmning  toward  anterior  cartilages, 
connected  along  lateral  surface  of  anterior  cartilages  (fig- 
ure 24,  ac).  Anterior  cartilages  ot  odontophore  flattened, 
long,  narrower  in  anterior  direction,  broader  in  posterior 
direction.  Posterior  cartilages  short  (about  1/5  of  anterior 
cartilage  length),  semi-spherical.  Anterior  and  posterior 
cartilages  joined  together  on  small  area  close  to  median 
fine  (figure  24).  Esophagus  (figures,  16,  19,  es)  narrow 
and  simple,  running  toward  postero-ventral  region  of  vis- 


ceral mass,  where  it  suddenly  bends  in  anterior  direction 
to  its  insertion  in  stomach  (figure  16,  st).  Stomach  large, 
cylindrical,  directed  toward  shell  apex;  inner  surface  uni- 
form, iridescent,  greenish.  Single  duct  to  digestiye  gland 
situated  close  to  shell  apex.  Stomach  suddenly  narrows 
giving  origin  to  intestine  (figures  16,  in),  which  runs  to- 
ward the  right  and  then  in  posterior  direction,  surround- 
ing dorsal  margin  of  shell  muscle,  gradually  turning  ven- 
trallv  and  in  anterior  tlirection  to  buccal  mass,  where  it 
suddenly  curves  in  posterior  cbrection,  running  along 
dorsal  surface  of  mesopodium  dorsalK  to  its  posterior 
edge,  looping  dorsally  to  the  right,  nmning  ventrally 
back  to  preceding  loop,  near  posterior  region  of  head, 
where  it  crosses  transversally  from  right  to  left  dorsallv 
to  left  extremit)-  f)f  shell  muscle,  suddenly  turning  suc- 
cessively in  posterior,  dorsal,  and  anterior  directions;  last 
loop  represented  by  rectum  (figures  15-17,  re)  crossing 
transversally  from  left  to  right  through  kidney  along  pos- 
terior hmit  of  mantle  ca\iU'.  Anus  (figure  15-17,  an) 
small  and  simple,  located  on  posterior  right  side  of  man- 
tle cax-ity. 

Radula  (Fifiiirvs  10-11):  A.s)anmetrical,  narrow,  rela- 
tively short  (about  tsvice  odontophore  length).  Rachidian 
tooth  rhomboid,  broad  for  genus,  broader  at  mid-sec- 
tion, and  blunt  (worn?)  cusp.  First  lateral  tooth  trian- 


Figures  18-24.  Coptilabi/ssin  riosi  new  species:  paratvpe,  MZSP  32149,  buccal  mass  and  odontophore.  18-19.  Buccal  mass 
extracted  from  snout.  18.  Neutral  \iew,  19.  Dorsal  \ie\v.  20-24.  Odontophore.  20.  Neutral  \iew,  supei^ficial  layer  ot  muscles  and 
membranes  partially  excised.  21.  Dorsal  \iew  22.  Neutral  \iew,  superficial  la\er  of  nniscles  anil  membranes  fully  excised,  23.  Dorsal 
view.  24.  Dorsal  view,  radular  ribbon  and  radula  sac  extracted,  most  of  muscles  and  cartilages  deflected  to  expose  inner  surfaces, 
right-side  structures  more  deflected  than  those  on  left  side.  Scale  hnes  =  500  (jlui. 


Page  66 


THE  NAUTILUS,  Vol.  114,  No.  2 


Figures  25-28.  Dorsal  and  lateral  views  of  shells  of  other  species  o{  Copulabijssia.  25.  C.  leptalea  (Verrill,  1884).  USNM  757345. 
NE  of  Norfolk,  \'irginia,  3080-,3090  m,  RA'  Gillis  cniise  7508,  station  36.  26.  C.  cornioata  (Jeffrevs,  1883).  SMNH  unnumbered, 
SE  of  Reykjanes  Ridge,  off  southern  Iceland,  2.50-350  m,  on  sunken  wood.  27.  C.  gracfnto'( Marshall,  1986).  NMNZ  M.  75007, 
holohpe,  off  \\'hite  Island,  New  Zealand,  107.5-1100  m,  on  sunken  wood.  28.  C.  similaris  Hazegawa,  1997.  NSMT-Mo  70822.  off 
Toi,  Suniga  Bay,  Japan,  4.30-710  m,  on  sunken  wood.  Scale  lines  =  1.0  mm. 


gular  with  outer  basal  projection  that  fits  into  a  depres- 
sion on  second  lateral  tooth,  and  pointed  single  cusp. 
First  lateral  tooth  (figure  11,  1),  broad,  triangular.  Sec- 
ond, third,  and  fourth  lateral  teeth  strongly  ciu^ed.  with 
two  short  cusps.  Fifth  lateral  (pluricuspid)  tooth  (figure 
11,  5)  massive,  club-like,  with  3  subterniinal  denticle- 
hke  cusps.  Latero-marginal  plate  present.  Inner  marginal 
teeth  pointed,  with  two  denticle-like  lateral  cusps  and 
thick  shafts  (e.g.,  figure  11,  1.  2),  decreasing  in  size  out- 
ward.  Second   marginal  tooth  largest.   Outer  marginal 

o  o  o 

teeth  with  serrations. 

Reproductive  system  (Figures  7,  8,  13,  14,  15,  17):  Go- 
nad (figure  15,  go)  relatively  small,  on  posterior  left  re- 
gion of  visceral  mass,  surrounded  bv  left  branch  of  shell 
muscle,  right  surface  of  digestive  gland,  and  some  intes- 
tinal loops.  A  ver)'  narrow  gonoduct  on  right  side  (not 


fully  examined),  running  in  anterior  direction  and  api- 
calK'  toward  right  side  of  mantle  cavity,  where  it  opens. 
Genital  pore  (figure  17,  gp)  turned  ventrally,  just  pos- 
terior to  anus.  Seminal  groove  (figure  17.  sg)  beginning 
on  genital  pore,  running  on  integument  surface  for  short 
distance,  up  to  posterior  region  of  base  of  right  tentacle. 
Right  tentacle  broad,  sfightlv  flattened  dorso-ventnilly, 
graduallv  tapering  to  flat  tip  (figures  7,  S,  13,  14,  17,  rt). 
Edges  of  seminal  groove  on  posterior  surface  of  tentacle, 
ending  on  posterior  region  of  tip.  A  well-developed  sub- 
terminal  protuberance  present  on  anterior  surface  of 
tentacle  (figure  14). 

Type  material:  Holotype  MZSP  32150,  3.25  mm 
length  X  2.50  mm  width  X  1.60  mm  height,  from  t\pe 
localitv,  P.  Bouchet,  J.  Leal,  and  B.  Metivier,  27  May 
1987,  dead  shell.  Paratypes:  MNHN  unnumbered,  3.3.5 


J.  H.  Leal  and  L.  R.  L.  Simone,  2000 


Page  67 


mm  length  X  2.35  mm  width  X  1.75  mm  height,  MZSP 
32149  (same  specimen,  soft  parts  onlv),  E  of  Caho  Siio 
Tome,  off  Rio  de  Janeno  .State,  Brazil,'2r24'S,  39°56'W, 
1320°1360  m  depth,  RA'  Mahoit-Dufrcsnc  cruise 
MD55,  station  CP-04,  R  Bouchet,  J.  Leal,  B.  Metivier, 
9  Mav  1987,  bottom  of  basaltic  gravel  and  o.>ddized  iron 
pelibles;  MZSP  32151,  3.50  mm  length  X  2.35  mm 
width  X  1.90  mm  height,  Doce  Ri\er  Canvon,  off  Es- 
pinto  Santo  State,  Brazil,  19°38'S,  38°43'W,  960  m 
depth,  RA'  Marion-Diifrcsnc  cruise  MD55,  station  CB- 
95,  R  Bouchet,  J.  Leal,  B.  Metivier  30  May  1987,  bottom 
of  basaltic  gra\el  and  o>dcbzed  iron  pebbles  (shell  broken 
during  SEM  session  in  Miami). 

Tjpe  locality:  Continental  slope  SE  off  Abrolhos  coral 
reef  system,  off  Bahia  State,  Brazil,  19°00.4'S, 
37°48.8'W,  950-1050  m  depth,  compact  dark  mud  and 
shell  hash  bottom,  (RA'  Marion-Dufrcsne  cruise  MD55, 
station  DC-72). 

Geographic  distribution:  Continental  slope  off 
E-SE  Brazil. 

Bathymetry:     960-1320  m. 

Et\Tnology:  Named  after  Prof.  EUezer  de  C.  Rios,  en- 
thusiastic Brazilian  author  and  mentor  to  an  entire  gen- 
eration oi  malacologists. 

DISCUSSION 

Specimens  of  Copniabi/.ssia  rio.si  were  found  apparently 
removed  from  the  actual  organic  substrate  ol  the  species 
(see  Lesicld,  1998,  for  a  Usting  of  species-specific  sub- 
strates and  food  preferences  in  the  family).  The  new 
species  differs  morphologically  from  the  other  four  spe- 
cies allocated  in  the  genus  Copiilabyssia  (see  figures  25- 
28,  table  1)  by  the  presence  of  an  apical  septum  on  the 
inner  shell  surface,  narrow  sculptural  interspaces,  fila- 
mentous, longer  epipodial  tentacles,  presence  of  a  stnic- 
ture  on  the  left  side  of  subpallial  cavity  that  could  proxe 
to  be  a  vestigial  left  gill  leaflet,  and  by  the  presence  of 
an  anterior,  subterminal  protuberance  on  the  right  ce- 
phalic tentacle.  The  double  insertion  of  the  m4  and  the 
expanded  insertion  of  the  m5  are  character  states  unique 
to  Copiilabyssia  nasi  when  compared  to  other  specie.s 
of  Coccuhnoidea  and  Lepetelloidea  for  which  these 
characters  are  known  (see  Haszpnmar,  1987;  Simone, 
1996);  however,  the  states  of  these  characters  are  so  far 
unknown  for  the  other  four  species  ol  the  genus  Co- 
piilabyssia. 

The  mantle  organs  of  Copiilabyssia  riosi  are  typical  of 
the  genus,  with  nuchal  cavity  and  gill  leaflets  situated  in 
the  right  side  of  the  mantle  cavity.  Copulabyssia  riosi 
shows  a  satellite  fold  in  the  inner  margin  ol  the  gill,  and 
a  small  protuberance  that  could  represent  a  vestigial  gill 
leaflet  in  the  left  region  of  the  mantle  border.  In  the 
superfamiK'  Lepetelloidea,  similar  structures  are  also 
present  in  the  Addisonia  cnodis  Simone,  1996  (Lepetel- 
loidea, Addisoniidae).  The  arrangement  of  loops  of  the 
digestive  system  looks  similar  to  that  descrilied  by  Has- 


zpmnar  (1988:  167-168,  fig.  4)  kr  Amphiplica  ktiiidseni 
McLean,  1988  (Lepetelloidea:  Pseudococculinidae),  but 
differs  by  the  reduction  of  salivarv  glands,  jaw  plates,  and 
e.sophageai  pouthes. 

The  complex  odontopliore  oi  Copiilabyssia  riosi  pre- 
sents a  combination  oi  apparently  plesiomorphic  and  au- 
tapomori^hic  characti'rs  The  presence  of  4  odontophore 
cartilages  and  their  approximator  muscle  (m8)  repre- 
sents apparently  plesiomoq^hic  states  present  in  several 
species  oi  Patellogastropoda,  Vetigastropoda  (including 
Lepetelloidea),  Cocculinoidea,  and  Neritopsina  (L.  R.  L. 
Simone,  personal  observation).  The  wefl-developed  sub- 
radular  cartilage,  the  partial  connection  between  the  an- 
terior and  posterior  odontophore  cartilages  and  the  hor- 
izontal muscle  (m6)  connected  only  in  a  side  of  the  an- 
terior cartilages  seem  to  represent  apomorphic  charac- 
ters (at  least  in  relation  to  Patellogastropoda). 

ACKNOWLEDGMENTS 

Gerhard  Haszprunar  (Zoologische  Staatssammlung 
Miinchen,  Germany),  Anders  Waren  (SMNH),  and 
Bruce  Marshall  (NMNZ)  critically  reviewed  the  manu- 
script. We  are  grateful  to  Philippe  Bouchet  and  Virginie 
Heros  at  the  original  repositoiy  institution  (National  mu- 
seum d'Histoire  naturelle,  Paris)  for  making  availalile  the 
type  material.  We  are  also  indebted  to  Suzanne  R.  Bra- 
den  (National  Museum  of  Natural  History,  Smithsonian 
Institution),  for  help  with  some  of  the  SEM  illustrations. 
M.  G.  Harasewych  (National  Museum  of  Natural  His- 
tory, Smithsonian  Institution),  Anders  Waren  (SMNH), 
Bruce  Marshall  (NMNZ),  and  Kazunori  Hasegawa 
(NSMT)  kindlv  sent  photographs  or  negatives  of  relevant 
species  of  Copiilabi/ssia  for  illustration.  This  study  was 
supported  in  part  Iiy  FAPESP  (Funda^ao  de  Amparo  a 
Pesquisa  dcj  Estado  de  Sao  Paulo,  Brazil)  Grant  #  96- 
6756-2  to  L.  R.  L.  Simone. 

LITERATURE  CITED 

Dantart,  L.  and  A.  Luque.  1994.  Cocculiniforniia  and  Lepeti- 
dae  (Gastropoda:  Archaeogastropoda)  from  Iberian  wa- 
ters. Journal  of  Molluscan  Studies  60:277-.31.3. 

Harasewych,  M.  G.,  S.  L.  Adamkewicz,  J.  A.  Blake,  D.  Saudek, 
T  Spiiggs  and  C.  ].  Bult.  1997.  Phylogeny  and  relation- 
sliips  of  pleurotoniariid  gastropods  (Mollusca:  Gastro- 
poda): an  assessment  based  on  partial  ISR  rDNA  and  ci- 
tochrome  c  oxidase  I  sequences.  Marine  Molecular  Biol- 
ogy and  Biotechnology  6: 1-20. 

Hasegawa,  K.  1997.  Sunken  wood-associated  gastropods  col- 
lected from  Suniga  Bay,  Pacific  side  of  Central  Honshu, 
Japan,  with  descriptions  of  12  new  species.  Natural  Sci- 
ence Museum  Monographs  12:.59-123. 

Haszpnmar,  G.  1987,  Anatomy  and  affinities  of  cocculinid  lim- 
pets (Mollusca,  Archaeogastropoda).  Zoologica  Scripta  16: 
30.5-324. 

Haszpnmar,  G.  1988.  Anatomy  and  affinities  of  pseudococcu- 
linid  limpets  (Mollusca,  Archaeogastropoda).  Zoologica 
Scripta  17:161-179. 

Hasz|5nmar,  G.  1998.  Superorder  Cocculiniforniia.  In:  Beesley, 
P.  L.,  G.  J.  B    Ro.ss  and  A.  Wells  (eds.)  Mollusca:  The 


Page  68 


THE  NAUTILUS,  Vol.  114,  No.  2 


Southern  Synthesis.  Fauna  of  Australia,  volume  5,  part  B. 
CSIRO  Puijlishing,  Melbourne,  pp.  65.3-664. 

Leal,  J.  H.  and  M.  G.  Harasewych.  1999.  Deepest  Atlantic 
mollnsks:  hadal  limpets  (MoJlusca,  Gastropoda,  Cocculin- 
ifornua)  from  the  northern  boundaiT  of  the  Caribbean 
plate.  Invertebrate  Biologv'  118:116-1.36. 

Lesicki,  A.  1998.  Checklist  of  gastropod  species  referred  to  the 
order  Cocculiniformia  Haszprunar,  1987  (Gastropoda: 
Cocculinoidea  et  Lepetelloidea)  with  some  remarks  on 
their  food  preferences.  Folia  Malacologica  6:47-62. 

Marshall,  B.  A.  1986.  Recent  and  Tertiary  Cocculinidae  and 
Pseudococculinidae  (Mollusca:  Gastropoda)  from  New 
Zealand  and  New  South  Wales.  New  Zealand  Journal  of 
Zoology-  12:50.5-546. 

McLean,  J.  H.  and  M.  G.  Harasewych.  1995.  Reydew  of  west- 
ern Atlantic  species  of  cocculinid  and  pseudococculinid 
limpets,  with  descriptions  of  new  species  (Gastropoda: 


Cocculiniformia).  Contributions  in  Science  of  the  Natural 
History'  Museum  of  Los  Angeles  County  453:I-.33. 

Ponder,  W.  F  and  D.  R.  Lindberg.  1996.  Gastropod  phyloge- 
ny — challenges  for  the  90s.  In:  Taylor,  ].  D.  (ed.)  Origin 
and  evolutionarv  radiation  of  the  Mollusca.  Oxford  Uni- 
versity Press,  O.xford,  pp.  1.35-154. 

Ponder,  W.  F.  and  D.  R,  Lindberg.  1997.  Toyvards  a  phylogeny 
of  gastropod  molluscs:  an  analysis  using  morphological 
characters.  Zoological  Journal  of  the  Linnean  Society  of 
London  119:8.3-265. 

Simone,  L.  R.  L.  1996.  Addisonia  enodis,  a  new  species  of 
Addisoniidae  (Mollusca,  Archaeogastropoda)  from  the 
southern  Brazilian  coast.  Bulletin  of  Marine  Science  58: 
775-785. 

Waren,  A.  1991.  New  and  little  kiioyvn  Mollusca  from  Iceland 
and  Scandinayna.  Sarsia  76:53-124. 

Wolff,  T.  1979.  Macrofaunal  utiUzation  of  plant  remains  in  the 
deep  sea.  Sarsia  64:117-1.36. 


THE  NAUTILUS  114(2):69-73,  2()()() 


Page  69 


Invasion  of  the  exotic  freshwater  mussel  Limnoperna  fortiinei 
(Dunker,  1857)  (Bivalvia:  MytiHdae)  in  South  America 


G.  Darrigran 

Departamento  Cientifico  de  Zoologi'a 

de  linertebrados 
P'atultad  Ciencias  Naturales  v  Museo 
1900  La  Plata 
ARGENTINA 
in\asion@wav.coni.ar 


L  Ezcurra  de  Drago 

Instituto  National  de  Liiimologfa 

(INALI/CONICET) 
Maciii  1933,  3016  Santo  Tome 
Santa  Fe 
ARGENTINA 


ABSTRACT 

We  traced  the  im'asion  and  iinestigated  the  (.hstiibntion  of  the 
Asian  bivaKe  Limnopenui  foriiinci  (Dnnker.  1857)  in  South 
America.  In  addition,  we  comment  on  the  problems  caused  bv 
this  in\'asive  species.  In  1991,  L.  fortiinei  was  introduced  into 
la  Plata  Basin  (currently  the  onlv  American  continent  drainage 
system  invaded  bv  this  species)  through  Ri'o  de  la  Plata  River 
B\'  the  end  of  1994  and  during  1995  this  species  dispersed 
throughout  the  Argentine  coast  of  the  Rfo  de  la  Plata  River 
and  was  reported  from  the  Uruguayan  coast.  In  199.5  and  1996, 
it  was  first  discovered  in  the  Parana  Ri\'er,  in  Santo  Tome  ( Sal- 
ado  del  Norte  River),  and  later  up  in  Goya.  In  November, 
1996,  L.  fortiinei  was  reported  in  Cerrito  Island,  where  the 
Paraguay  River  joins  the  Parana  River  In  April  1997,  it  was 
collected  in  Paraguay  River  on  Asuncion  Harbor,  Paraguay.  In 
1999,  if  was  detected  in  Brazil  (Itapuii  Beach,  MunicipaUty  of 
Viamao,  Rio  Grande  do  Sul  State).  These  recent  records  doc- 
ument tlie  fast  and  ongoing  e.xjiansion  ot  the  species  geograph- 
ic range,  as  well  as  its  prompt  adaptation  to  the  ditterent  en- 
\ironments  of  the  la  Plata  Basin 

Ki'ij  words:  Soutli  America;  Neotropical  Region;  la  Plata  Basin; 
invasive  species;  distribution;  biofouling. 


INTRODUCTION 

The  la  Plata  Basin  is  one  of  the  most  important  hydro- 
graphic  systems  in  South  America.  It  has  a  drainage  area 
of  approximately  3X10''  km-,  and  comprises  4  main  sub- 
hasins  (Bonetto,  1994):  (1)  Uniguav  River,  (2)  Parana- 
Paraguay  system,  (3)  Rio  de  la  Plata  River,  and  (4)  the 
Andean  tributaries  (figure  1). 

A  diverse,  native  molluscan  fauna  occurs  along  the  Ar- 
gentine shore  of  the  Rio  de  la  Plata  estuan'  (Darrigran, 
1994),  including  27  gastropod  and  21  bivalve  species. 
Only  4  of  the  bivalves  are  strictly  freshwater  and  intaunal 
species:  Anodontites  tenebricosus  (Lea,  1834),  Diplodon 
parancnsis  (Lea,  1834),  MtisciiUum  orfictitimnn 
(d'Orhigny,  1835)  and  Pisiclimn  stcrkiaiiinn  Pilsbi"v, 
1897.  The  only  epifaunal  species  on  the  Argentine  coast 
of  Rio  de  la  Plata  River  is  the  estuarine  Mi/tclla  clior- 


niana  (d'Orbigny,  1842).  Until  1993,  A/,  charniana  was 
the  only  Mytilidae  found  in  Rio  de  la  Plata  River,  in 
Punta  Piedras,  Argentina  and  Montevideo,  Uruguay 
(Darrigran  and  Pastorino,  1995a). 

In  the  1970s,  2  asiatic  freshwater  bi\aKes,  Corhicida 
largiUiciii  (Philippi,  1844)  and  C.  fliiininca  (Miiller, 
1774)  (Corbicuhdae),  entered  South  America  along  the 
Argentine  shores  of  Ri'o  de  la  Plata  River  (Ituarte,  1981). 

In  1991  we  detected,  for  the  first  time  in  South  Amer- 
ica, the  presence  of  a  mytilid,  Limnoperna  foiinnci 
(Dunker,  1857),  in  the  Rio  de  la  Plata  River  shore  at 
Baghardi  Beach  (Pastorino  et  al.,  1993).  This  is  an  eu- 
ryhaline  freshwater  species,  native  to  China  and  to 
southeastern  Asia  (Morton,  1977),  which  resembles  ma- 
rine mussels  (Mytifidae)  in  its  moiphology  and  biology. 

Commerce  beUveen  Argentina  and  the  countries 
where  L.  fortiinei  is  supposedlv  native  takes  place  m;unly 
by  sea.  Following  Carlton  (1992)  and  Carlton  and  Geller 
(1993),  Darrigran  and  Pastorino  (1995a)  hypothesized 
that  L.  fortiinei  and  other  invading  organisms  may  have 
arrived  in  ballast  water  of  ships  coming  from  southeast- 
em  Asia. 

Other  authors  have  inthcated  that  L.  foi-fiinei  was  in- 
troduced in  and  recorded  from  Hong  Kong  in  1966 
(Morton,  1975,  1996)  and  Japan  in  1991  (Kimura,  1994). 
In  the  Americas,  L.  foiiiinei  is  known  to  have  invaded 
only  Rio  de  la  Plata  Basin.  VIorton  (1973)  suggested  that 
the  morpho-functional  features  of  this  species  allow  for 
rapid  expansion  of  its  distribution  range,  in  similar  fash- 
ion to  the  invasive  abilities  of  the  zebra  mussel,  Drei.s- 
sena  polijmorpha  (Pallas,  1771),  in  the  Northern  Hemi- 
sphere. The  species  exhibits  2  features  that  mav  contrib- 
ute to  its  abilities  as  an  invader:  (1)  it  is  the  onlv  relevant 
mollusk  in  the  freshwater  littoral  ha\ang  a  bvssus,  and 
(2)  has  no  known  native  competitors  for  physiciil  space. 
As  a  result  of  the  high  rates  of  invasion,  as  we  further 
chscuss  below,  L  foiiiinei  causes  serious  biofouling  of 
municipal  drinking  water  and  industrial  water  svstems. 
As  we  demonstrate  below,  populations  of  Limnopenui 
fortiinei  are  not  onlv  rapidly  increasing  in  densit\-  in  sev- 


Page  70 


THE  NAUTILUS,  Vol.  114,  No.  2 


tial  localities  but  are  also  quickly  expanding  geographi- 
cidly,  mostly  in  northward  direction. 

Since  1991,  when  it  was  first  found  at  Bagliardi  Beach, 
until  the  end  of  1993,  the  species  was  found  only  in 
mLxohaline  (e.g.,  Punta  Piedras  and  Punta  Indio)  and 
eur\'haline  environments  (e.g.,  Magadalena  Beach,  Ata- 
laya  Beach,  Punta  Blanca,  La  Balandra  Beach).  At  that 
time,  the  species  had  not  been  collected  farther  north 
than  the  freshwater  environment  of  Punta  Lara  Beach, 
and  was  apparently  absent  from  all  other  countries  and 
rivers  comprising  the  la  Plata  Basin  (Darrigran  and  Pas- 
torino,  1993). 

The  objectives  of  this  paper  are  to  investigate  the  in- 
vasion of  L.  foiiunci  in  South  America,  to  provide  an 
update  of  its  cbstribution  in  the  Neotropical  Region,  and 
to  emphasize  the  importance  of  this  invasive  bivalve  in 
biofouling  processes  in  the  la  Plata  Basin. 

MATERIALS  AND  METHODS 

Collections  were  made  along  the  Rio  de  la  Plata  River 
shore  at  low  tides.  In  the  Parana  Ri\'er,  samples  were 
collected  with  a  319  cm-  "Tamura"  dredge  (Marchese 
and  Ezcurra  de  Drago,  1992).  Periph\tic  populations 
were  sampled  by  hand.  In  the  mouth  of  Paraguay  River, 
"macrofouling"  samplers  (Stupak  et  al,  1996)  were  used. 
Environmental  parameters  taken  into  account  were: 
temperature,  pH,  and  conductivity.  The  study  localities 
are  showni  in  figure  1.  Specimens  from  the  Malacological 
Collection  of  La  Plata  Museum,  Argentina,  (MLP)  were 
examined.  Samphng  for  larvae  was  made  using  a  mesh 
of  25  (xm  in  Umguav  River  (towns  of  Colon  and  Con- 
cepcion  del  Uiiiguay),  at  3  stations  in  the  Carcaraiia  Riv- 
er, and  Correntoso  River 


RESULTS  AND  DISCUSSION 

Liinnopenw  foifunei  was  first  found  in  South  America 
in  1991  at  Bagliardi  Beach  (Argentina)  with  densities  of 
5  individualsm-  (Pastorino  et  al,  1993).  In  1992,  the 
maximum  densitv'  recorded  was  about  3X10^  indixidu- 
als-m  -.  In  1993,  in  the  same  locality,  the  maximum  den- 
sity recorded  was  SXIO'  individualsm  -.  In  1998,  den- 
sities of  about  15X10'  individuals-m"-  were  commonly 
recorded,  values  equivalent  to  those  for  mytilid  densities 
at  the  seashore  in  Mar  del  Plata,  Argentina  (Penchasz- 
adeh,  1973). 

In  1994  and  1995,  L.  fotiiinei  was  found  in  the  water 
intakes  of  the  town  of  Bemal  (MLP  5204)  and  Retiro, 
and  on  Buenos  Aires  Harbor  (MLP  5205).  Scarabino 
and  Verde  (1994)  reported  the  species  from  the  town  of 
Colonia  del  Sacramento,  Uruguay  (MLP  5202).  All  these 
localities  are  on  the  Rio  de  la  Plata  estuaiy.  During  1996, 
the  species  expanded  to  the  town  of  Goya  (29°10'S, 
59°16'W),  in  Corrientes  Province  (Di  Persia  and  Bonet- 
to,  1997)  and  nearby  lotic  bodies,  and  was  collected  in 
the  middle  Parana  River,  near  the  city  of  Santa  Fe, 


where  the  river  has  a  \'erv 
width;  Drago,  1990). 


wide  alluvial  plain  (25  km  in 


Figure  1.  Studied  localities  on  the  la  Plata  Basin.  A,  Punta 
Piedras;  B,  Punta  Indio;  C,  Magdalena  beach;  D,  Atalaya 
Beach;  E.  Pmita  Blanca;  F,  La  Balandra  Beach;  G,  Bagliardi 
Beach;  H,  Pnnta  Lara  Beach;  L  Benial  Beach;  ].  Buenos  Aires 
Harbor;  K,  Atncha  I  Nuclear  Power  Plant;  L,  City  of  Rosario; 
M,  Town  of  Santo  Tome;  N,  City  of  Santa  Fe;  O,  Esquina;  P, 
Goya;  Q,  Cerrito  Island;  R,  Town  of  Formosa;  S,  Asuncion 
Harbor;  T,  Posadas  Harbor;  U,  Praia  de  Itapna  (Rio  Grande 
do  Sul  State,  Brazil);  V,  Town  of  Colonia  del  Sacramento;  1, 
Town  of  Colon;  2,  Town  of  Concepcion  del  Umguav;  3,  Car- 
caraiia River. 


Litnnopcrna  foiiunci  was  first  collected  in  Parana  Riv- 
er at  Vuelta  del  Este,  Zarate,  on  the  Parana  de  Las  Pal- 
mas  River  (MLP  5206),  and  at  Paso  Burghi,  city  of  Ro- 
sario, on  the  lower  Parana  River  (MLP  5207)  toward  the 
end  of  1995  (figure  1).  The  species  settles  on  all  kinds 
of  substrates  across  the  entire  floodplain.  It  was  found 
in  the  main  channel,  on  the  left  bank  at  town  of  Parana 
and  associated  alluvial  plain,  in  San  Javier  and  Corren- 
toso (MLP  5287)  rivers,  near  the  city  of  Santa  Fe.  It  was 
also  found  on  the  right  bank  of  the  lower  Salado  del 
Norte  River  (MLP  5285),  in  the  town  of  Santo  Tome 
(31°40'S,  60°45"W). 

Limnopcnia  foiiinwi  was  part  of  the  epifauna  present 
on  Solanaceas  and  Paspahun  .sp.  (MLP  5286)  in  all  hab- 
itats observed,  but  was  not  found  on  the  sandy,  moving 
bars  (Drago,  1997)  in  the  main  channel.  Perhaps  due  to 
the  unfavorable  sandy  substrate,  the  central  part  of  the 


G.  Darrigran  and  I.  Ezcurra  de  Drago,  2000 


Page  71 


main  channel  pro\ides  haliitat  to  onK-  a  few  henthic  spe- 
cies (Marchese  and  Ezcurra  de  Drago,  1992).  In  the 
Correntoso  River  (a  secondary  channel  of  the  middle 
Parana  Ri\er\  L.  foiiiiiici  i-olonized  the  silt\  cla\  lieds 
at  all  locations  sanipleil 

The  eun,haline  attributes  of  L.  foiinnci  probably  fa- 
cilitates its  rapid  spread.  The  mean  salinitv  at  the  main 
channel  of  the  Parana  River  is  0.05  %r,  and  in  Corren- 
toso River  it  ranges  bet\veen  ().()fi4-().345  %c.  In  Salado 
del  Norte  Rixer,  which  mi.\es  with  the  waters  of  the  Pa- 
rana River,  sahnitv  varies  from  0.5  to  4  %c  (Ezcurra  de 
Drago,  personal  obser\'ation).  Mean  pH  is  7.4  in  the  Pa- 
rana m;un  channel  and  7.2  in  secondaiy  courses  (Mar- 
chese and  Ezcurra  de  Drago,  1992),  \s'hile  in  Salado  tlel 
Norte  River  pH  reaches  8.7. 

In  1996,  this  species  was  collected  on  "fouling"  sam- 
plers in  Cerrito  Island  (27°20'S,  5S°43'W),  near  the  con- 
fluence of  the  Paraguay  and  Parana  rivers  (MLP  5340). 
In  April  1997,  it  was  collected  on  Asuncion  Harbor 
(25°17'21"S,  57°38'0S"W),  in  the  Paraguay  River,  in  Par- 
aguay (figure  1).  It  was  also  collected  during  the  last 
months  of  1998,  at  Posadas  Harbor,  on  the  upper  Parana 
River.  In  November  1999,  L  foiiimci  was  detected  in 
Brazil  on  Itapua  Beach,  Municipality  of  Viamao,  Rio 
Grande  do  Sul  State  (MLP  5550).  No  larvae,  juveniles 
or  adults  have  been  found  at  anv  of  the  locahties  ex- 
amined in  UiTiguav-  River  (Colon,  Concepcion  del  Uru- 
guay) or  Carcarana  River  (Carcaraiia  and  la  Ribera). 

The  biofouling  problems  caused  by  L.  fortunei  in 
South  America  are  similar  to  those  described  for  Drcis- 
scna  pohjinoiyha,  the  zebra  mussel,  in  the  Northern 
Hemisphere  (Darrigran,  1995). 

The  principal  problems  caused  by  zebra  mussel  in- 
vasion, settlement,  and  maturity,  into  water  distribution 
systems  (Nalepa  and  Schloesser,  1993)  are  listed  below 
(the  problems  which  have  already  been  detected  in  Ar- 
gentina resulting  from  invasion  bv  L  foiiuiici  are 
marked  with  (X)). 

•  ( X )  Reduction  of  pipe  diameter 

•  ( X )  Blockage  of  the  pipeUne 

•  Decreased  water  velocitv  caused  bv  friction  (turbulent 
flows). 

•  (X)  Accumulation  of  enipt\'  shells 

•  Contamination  of  water  pipelines  by  mass  mortality 

•  ( X )  Filter  occlusion 

These  problems  have  occurred  in  the  intakes  of  water 
treatment  plants  in  the  city  of  La  Plata;  water  treatment 
plants  along  the  Corrientes  River  (MLP  5365),  and  also 
in  industrial  and  power-generating  plants.  They  cause  an 
increase  in  the  operational  costs  of  these  facilities  due 
to  reduced  pump  efficiency,  increased  tube  corrosion 
(caused  by  proUferation  of  bacteria  and  fungi),  and  in- 
creased frequenc)'  of  shutdownis  for  cleaning  and  filter 
changes. 

Absence  of  relevant  records  in  the  numerous  benthic 
studies  conducted  prior  to  1991  (Darrigran,  1991,  1994; 
Marchese  and  Ezcurra  de  Drago,  1992)  indicate  that 
colonization  by  L.  foiiiinci  is  a  recent  event.  By  late  1996 


and  early  1997,  the  first  cases  of  fouling  bv  L.  fniiunei 
in  nearby  industries  were  reported  in  the  Parana  River 
(e.g.,  Atucha  I  Nuclear  Power  Plant  and  San  Nicolas  de 
Los  Arroyos  Electric  Plant  (MLP  5300),  Buenos  Aires 
Province). 

Based  on  the  facts  above  described,  it  appears  that 
the  geographic  range  of  L.  foi-tunci  is  in  continuous  ex- 
pansion in  South  America,  and  that  the  species  is  ap- 
parently adapting  to  different  environments  along  its  in- 
vasion routes.  Between  1991-1999  this  species  invaded 
4  c(juntries  (Argentina,  Uruguay,  Paraguay,  and  Brazil); 
3  of  the  Ri'o  de  la  Plata  Basin  main  rivers  (la  Plata,  Pa- 
rana, and  Paraguay).  It  traveled  upstream  at  a  speed  of 
about  240  kmvear  '.  Throughout  its  distribution  range, 
L.  fdiiiinci  inhabits  a  wide  variety  of  environments:  from 
euryhaline  waters  in  the  La  Plata  estuary  to  strictly 
freshwater  habitats  along  the  Paraguay  and  Parana  riv- 
ers. 

This  expansion  is  proiiably  favored  by  the  moqiho- 
functional  features  of  the  species,  and  by  the  apparent 
absence  of  competitive  interactions  along  the  httoral  of 
la  Plata  Basin.  Lininopcnio  foiitinci  is  the  onlv  relevant 
species  using  bvssal  epifaunal  attachment  among  local 
freshwater  species. 

Although  the  distribution  of  L.  foiiunci  is  currently 
still  limited  to  la  Plata  Basin,  there  are  no  apparent  con- 
straints that  could  prevent  the  invasion  of  other  drainage 
systems.  Moreover,  this  expansion  may  be  favored  by  the 
increase  in  trade  between  Argentina,  Uruguay,  Paraguay 
and  Brazil,  as  a  consequence  of  the  recently  establi,shed 
MERCOSUR  international  economic  consortium  (Dar- 
rigran, 1995;  Darrigran  and  Pastorino,  1995b).  Addition- 
al dispersal  of  the  species  could  also  be  favored  by  the 
international  project  HIDROVIA,  which  consists  of  a 
wateiAvay  connecting  the  mtiin  hvdrographic  systems  of 
South  America.  The  project  comprises  a  complex  fluvial 
network  system  that  involves  and  interconnects  Rio  de 
la  Plata,  Parana,  and  Paraguay  rivers,  and  includes  plans 
for  extensive  basin  dredging,  harbor  construction,  and 
improvement  of  the  harbors  and  highways.  Connections 
among  the  main  South  American  basins  (e.g.,  Amazonas, 
Orinoco)  are  likely  to  be  artificially  established  in  the 
future.  Up  to  now  no  effective  control  mechanisms  have 
been  proposed  to  prevent  further  spread  oi  L.  fortunei. 

The  impact  of  L.  foiiintci  will  not  be  restricted  onlv 
to  the  economy,  but  will  also  affect  the  diversit>-  of  the 
native  molluscan  communities.  Darrigran  et  al.  (1998) 
showed  that,  since  the  introduction  o{L.  fortunei  at  Bag- 
liarili  Beach,  populations  of  2  common  gastropods  have 
been  displaced:  Ciiilina  jiuminea  (Maton,  1809)  is  no 
longer  found,  while  Gundhiehia  conccnfiica  (d'Orbigny, 
1835)  is  becoming  rare  (figure  2). 

In  contrast,  several  benthic  species,  uncommon  or  ab- 
sent before  the  occurrence  of  L.  fortunei.  are  now  pre- 
sent. These  include  Annelida  Ofigochaeta  (8  species), 
Aphanoneura  (1  species)  and  Himdinea  (8  species);  plus 
assorted  Crustacea  and  Insecta  (Darrigran  et  ai,  1998). 
Additionallv,  we  observed  the  epizoic  colonization  and 
snujthering  of  native  bivalves  (e.g.,  Anodontite.s  trapesi- 


Page 


THE  NAUTILUS,  Vol.  114,  No.  2 


DENSITY 


Helobia  piscium 


10' ■• 


Chilina  fluminea 


I    . 


]«|«jB|*|«|«|«|*|B|«|*|B| 


I     ■     I     ■     I     ■     I     ■    I    ■ 


Gundlachia  concentnca 


I'i'i'i'i'i — I  *  I ■ I ■  I  ■  I 


Limnoperna  fortune! 


H 1 1 1 1 


-•-h ( 1 1 1 (-1 — I 1 h 

■3       7       8      9      10     12      1       2       3       5      8      10     12    |l       2       3      4       6      9      10     11  I  3 
88  I    92  |93  194  I  95 

MONTH/YEAR 

Figure  2.  Temporal  \ariation  of  density  (individuals-m"-, 
represented  bv  mean  and  standard  deviation)  of  some  native 
mollusks  associated  with  the  byssus  of  L,  fortiinei  in  Bae;liardi 
Beach  (34°55'S,  57°49'W),  Wentina. 


alis  (Lamarck,  1819)  and  A.  tcncbricosus  (Lea,  1834))  by 
L.  foiiiinei,  in  a  behavior  similar  to  that  shown  by  Dreis- 
seno  poli/inoi-f)ha  on  unionids  (Parker  ct  al,  1998). 

Last  but  not  least,  further  expansion  of  the  natural 
range  ot  Limnoperna  foi-tunci  and  the  possibilitv  of  in- 
vasion of  parts  of  Noith  America  by  this  species  should 
not  be  rejected.  As  Ricciardi  (1998)  obserx'ed,  "Given 
that  shipping  traffic  from  both  Asia  and  South  America 
has  alreadv  resulted  in  recent  introduction  of  exotic  bi- 
valves to  the  USA,  a  future  North  American  invasion  bv 
L.  fortunci  is  highly  probable." 

ACKNOWLEDGMENTS 

The  work  was  supported  by  the  grants  of  the  Consejo 
Nacional  de  Investigaciones  Cienti'ficas  y  Tecnicas  (PEI 
N°  548/97),  Facultad  Ciencias  Naturales  v  Museo 
(UNLP),  and  Agencia  Nacional  Promocion  Cientifica  y 
Tecnologica  (PICT98  No.  01-03453).  We  thank  Dr.  Ri- 
chard Sparks  and  Ruth  Sparks  for  the  revision  of  the 
manuscript.  Dr.  Edmundo  Drago  and  M.  C.  Dambore- 
nea  for  helping  in  assorted  tasks,  and  Lionel  Mehaudy 


for  the  technical  assistance.  Maria  CrisHna  Dreher  Man- 
sur  collected  and  provided  the  specimens  from  Itapua 
Beach,  Rio  Grande  do  Sul,  Brazil. 


REFERENCES 

Bonetto,  A.  A.  1994.  Austral  rivers  of  South  America.  In:  Mar- 
galef,  R.  (ed.)  Limnology  now:  a  paradigm  of  planetary 
problems.  Elsevier  Science,  Amsterdam,  pp.  425-472. 

Carlton,  ],  T.  1992.  Introduced  marine  and  estuarine  mollusks 
of  North  America:  an  end-of-the-20'''  centurv  perspective. 
Journal  of  Shellfish  Research  11:489-505. 

Carlton,  ].  T  and  J,  B.  Geller  199.3.  Ecological  roulette:  the 
global  transport  of  nonindigenous  marine  organisms.  Sci- 
ence 261(51 17):7S-S2. 

Danigran,  G.  1991.  Aspectos  ecologicos  de  la  malacofauna  h- 
toral  del  Ri'o  de  la  Plata,  Republica  Argentina.  Facultad 
Ciencias  Naturales  y  Museo.  Tesis  no.  568.  Universidad 
Nacional  de  La  Plata,  Argentina,  238  pp.  [unpublished 
dissertation] 

Darrigran.  G.  1994.  Composicion  de  la  malacofauna  litoral  del 
estuario  del  Rio  de  la  Plata.  Republica  Argentina.  Tankav 
1:147-149. 

Darrigran,  G.  1995.  Limnopenin  fortiinei.  jun  problema  para 
Ids  sistemas  naturales  de  agua  dulce  del  Mercosur?.  Mu- 
seo l(5):85-87. 

Darrigran,  G.  and  G.  Pastorino.  1993.  Bi\al\x)S  Invasores  en  el 
Rio  de  la  Plata,  Argentina.  Comunicaciones  de  la  Sociedad 
Malacologica  del  Uniguav  7(64-65 ):309-313. 

Darrigran,  G.  and  G.  Pastorino.  1995a.  The  recent  introduction 
of  a  freshwater  asiatic  bivalve,  Limnoperna  foiinnei  (My- 
tilidae)  into  South  America.  The  Veliger  38:171-175. 

Darrigran,  G.  and  G.  Pastorino.  1995b.  Linmopema  fortiinei 
(Dunker,  1857)  (BivaKia:  Mytilidae),  potencial  problema 
en  las  cuencas  dulceacuicolas  de  America  del  Sur.  II  Con- 
greso  Latinoamericano  de  Malacologi'a.  Porto  Alegre. 
Brasil.  Resumenes,  p.  55. 

Darrigran,  G.,  S.  M.  Martin,  B.  Gullo  and  L.  Annendariz. 
1998.  Macroinvertebrates  associated  with  Limnoperna 
fortiinei  (Dunker,  1857)  (BivaKia,  M\tilidae)  in  Rio  de  la 
Plata,  Argentina.  Hvdrobiologia  367:22.3-230. 

Di  Persia,  D.  and  A.  A.  Bonetto.  1997.  Nuevas  citas  de  Lim- 
noperna fortiinei  para  la  cuenca  del  no  Parana,  Argentina. 
Neotropica  43(109-1 10):  119-120. 

Drago,  E.  1990,  Geomoi"pholo2;\  ot  large  alluvial  rivers:  Lower 
Paragna)'  and  Middle  Parana.  Interciencia  15(6):378— .387. 

Drago,  E.  1997.  Campana  "Keratella  I"  a  lo  largo  del  rio  Pa- 
rana Medio.  II:  Fonnas  del  lecho  en  su  cauce  principal. 
Revista  Asociacion  Cientifica  Litoral  8:57-62. 

Ituarte,  C.  1981.  Primera  noticia  acerca  de  la  introduccion  de 
pelecfpodos  asiaticos  en  el  area  rioplatense  (Mollusca: 
Corbicuhdae).  Neotropica  27(77):79-S3, 

Kimura,  T  1994.  Moqjhological  identification  of  Limnoperna 
fortunei  (Dunker)  and  Limnoperna  joiiunci  kiknchii 
Habe.  Chiribotan  25:.36-40. 

Marchese,  M.  and  I.  Ezcurra  de  Drago.  1992.  Benthos  of  the 
lotic  environments  in  the  middle  Parana  River  system: 
transverse  zonation.  Hydrobiologia  237:1-13. 

Morton,  B.  1973.  Some  aspects  of  the  biology  and  funcional 
morphology'  of  the  organs  of  feeding  and  digestion  of  Lim- 
noperna fortiinei  (Dunker)  (BivaKia:  Mvtilacea).  .Malacol- 
ogia  12:26.5-281. 

Morton,  B.  1975.  The  colonization  of  Hong  Kong's  raw  water 


G.  Darrigran  and  I.  Ezcurra  de  Drago,  2000 


Page  73 


supply  system  by  Lininopcrnd  foiiunri  (Dunker)  (Bivalvia: 
M)tilacea)  from  China.  Malacolopcal  He\iew  8:91-105. 

Morton,  B.  1977.  Freshwater  tonling  bivalves.  Proceedings  of 
the  First  International  C'orhicula  SNinjiosium.  Texas 
Christian  Uni\ersit\',  pp.  1-14. 

Morton,  B.  1996.  The  a(|uatic  nuisance  species  problem;  A 
global  perspecti\e  and  re\iew.  In:  F.  M.  D'ltri  (ed.).  Zebra 
Mussels  and  Aquatic  Nuisance  Species.  Ann  Arbor  Press, 
Inc.  Chelsea,  Michigan. 

Nalepa,  T,  and  W  Schloesser.  1993.  Zebra  mussels:  biolog)', 
impacts,  and  control.  Lewis  Pviblishers,  Boca  Raton.  508  pp. 

Parker,  B.  C  M.  A.  Patterson  and  R.  J.  Neves  1998  Feeding 
interactions  between  native  freshwater  mussels  (BivaKia: 
Unionidae)  and  zebra  mussels  {Drcisscna  i>oli/morplia)  in 
the  Ohio  Ri\er  American  Malacological  Bulletin  14:17.3- 
179. 

Pastorino,  G.,  G.  Darrigran,  S.  Martin  and  L,  Lunaschi.  1993. 


Liinnopentri  foriiiitci  (Dunker,  1857)  (Mytilidae),  umcm) 
bixalvo  invasor  en  aguas  del  Rio  de  la  Plata.  Neotropica 
39(101-102):34. 

Pencha.szadeh,  P.  1973.  Ecologfa  de  la  comunidad  del  mejillin 
(Brachyclontes  rodriguezi  d'Orb.)  en  el  medioiitoral  ro- 
coso  de  Mar  del  Plata  (Argentina):  el  proceso  de  recolon- 
izacion.  Physis  32A(84):5I-64. 

Ricciardi,  A.  1998.  Global  range  expansion  of  the  Asian  nnissel 
Limnoprnin foiiumi  (Mvtilidae):  another  fouling  threat  to 
freshwater  .systems.  Biofouling  13:97-106. 

Scarabino.  F.  and  M.  N'erde.  1994.  Limnoperna foiii/iwi  (Dtm- 
ker  1857)  en  la  costa  unigua\a  del  Rio  de  la  Plata  (Biv- 
alvia, Mytilidae).  Connmicaciones  Sociedad  Malacologica 
del  Uruguay  7 (66-67): 374-375. 

Stupak,  M.,  M,  Perez,  M.  Garcia  Sola,  A.  Leiva  Azuaga,  A. 
Mendivil  and  G.  Niveyro.  1996.  Preliminary  study  of  the 
biofouling  of  the  Parana  Rix'er  (Argentina).  Corrosion  Re- 
news 14:145-155. 


THE  NAUTILUS  114(2):74-79,  2000 


Page  74 


Location  of  allospermatozoa  in  the  freshwater  gastropod 
Biomphalaria  fenagophila  (d'Orbigny,  1835)  (Puhiionata: 
Planorbidae) 


Warton  Monteiro 

Depaitiimento  de  Ecologia 
Uiu\ersidade  de  Brasilia 
Brasilia,  DF 
BRAZIL 
warton@uol.c-oni.br 


Toshie  Kawano' 

Laboratorio  de  Parasitologia 
Instituto  Butantan 
A\'.  Vital  Brasil,  1500 
0550.3-900  Sao  Paulo 
BRAZIL 


The  freshwater  puhnonate  Biomphalaria  fenagophila  is 
a  simultaneous  hermaphrodite.  The  route  ot  allosper- 
matozoa in  the  receiving  partner  was  used  in  the  analysis 
of  resource  allocation  in  the  male  function  in  this  spe- 
cies. Allospermatozoa  enter  the  vagina,  mav  or  may  not 
bvpass  the  spermatheca,  continuing  through  nidamental 
gland,  o\'iduct-muciparous  gland,  albumen  gland,  oNds- 
permioduct,  seminal  vesicle,  ovotestis  collecting  canal, 
and  folhcles  of  ovotestis.  A  new  technique  is  proposed 
to  label  spermatozoa  xaa  injection  oi  tritiated  th\iiiitline 
through  the  pneumostome  (under  anesthesia).  Labeled 
allospermatozoa  were  located  in  the  spermatheca,  ni- 
damental gland,  oviduct-muciparous  gland,  albumen 
gland,  seminal  vesicle,  and  follicles  of  ovotestis.  No  re- 
lation was  found  between  the  duration  of  pairing  and 
movement  of  the  allospermatozoa  through  the  repro- 
ductive system.  It  is  suggested  that  the  final  destination 
of  allospermatozoa  are  the  folhcles  of  ovotestis.  Their 
presence  in  the  spermatheca,  where  thev  ma\'  be  dis- 
solved, could  be  fatal.  Presence  of  allospermatozoa  in 
the  nidamental  gland,  oviduct-muciparous  gland,  or  ;il- 
bumen  gland  may  be  a  consequence  of  unilateral  cop- 
ulation (snail  plavs  only  a  recei\ing  role),  which  packs 
the  seminal  vesicle  with  autospermatozoa.  Allosperma- 
tozoa may  wait  in  the  interior  of  the  female  tract  until 
the  snail  becomes  a  donor  in  a  subsequent  pairing.  From 
then  on,  allospermatozoa  may  pass  through  an  empty 
seminal  vesicle  toward  their  final  destination,  the  folh- 
cles of  ovotestis. 

Key  words:  Reproduction,  spermatozoa,  hermaphrocUt- 
ic,  pulmonates. 


INTRODUCTION 

The  location  of  spermatozoa  within  the  reproductive  sys- 
tem of  inseminated  puhnonate  gastropods  plavs  an  im- 

'  Author  for  correspondence;  tkawano@usp.br 


portant  role  in  the  processes  of  self-fertilization,  cross- 
fertilization  and,  in  particular,  preferential  cross-fertiliza- 
tion in  these  mollusks  (Larembergue,  1939;  Paraense, 
1959). 

In  the  genus  Phijsa,  2  or  3  possible  chambers  for  fer- 
tilization were  described.  Although  these  chambers  open 
in  the  oviduct,  the  actual  site  of  fertifization  may  be  lo- 
cated in  the  hermaphrodite  region,  close  to  the  carrefour 
(Duncan,  1969).  In  planorbids,  the  seminal  vesicle  has 
been  cited  as  the  main  location  of  spermatozoa  (Lar- 
ambergue,  1939).  The  fertilization  chamber,  in  the  car- 
refour, where  the  spermatozoa  received  from  the  part- 
ner are  stocked,  is  also  a  possible  ferhhzation  site  (Ab- 
del-Malek,  1954;  Duncan,  1975).  The  oviduct  (Fretter 
and  Graham,  1964)  and  the  upper  part  of  the  o\isper- 
mioduct  mav  be  sites  where  the  oocytes  are  fertilized  by 
the  allospermatozoa  (Abdel-Malek,  1954).  Some  authors 
consider  the  spermatheca  as  a  location  where  sperma- 
tozoa remain  immecfiatelv  after  copulation,  proceeding 
subsequently  through  the  female  ducts  (Duncan,  1975). 
Notwithstanding,  the  spermatheca  in  B.  glahrata  also 
behaves  as  a  site  of  lysis  of  spermatozoa  (Horstmman, 
1955;  Jong-Brink,  1969;  Kitajima  and  Paraense,  1983). 
In  planorbids,  the  spermatheca  is  a  blind,  pvriform  or- 
gan, opening  in  the  vagina  through  a  short  duct  (Par- 
aense, 1975;  Kitajima  and  Paraense,  1983).  This  is  also 
called  bursa  copulatrix  (Duncan,  1975),  or  seminal  re- 
ceptacle. In  euth\aieuran  gastropods,  the  evolutionary 
acquisition  of  reproductive  s\stems  such  as  functional 
simultaneous  hermaphrocUtism  has  been  associated  with, 
among  other  traits,  fundamental  changes  in  the  structure 
of  the  spermatozoa  (Thompson,  1973). 

A  past  experiment  iuNolving  incorporation  ot  '"Fe  by 
BioinpJialaria  tAahrata  took  place  in  water  containing 
this  rachoisotope  (Gazzinelli  ct  al.  1970).  Incoiporated 
isotope  was  transferred,  during  pairing,  from  labeled  B. 
glabrata  to  non-labeled  partners.  The  presence  of  the 
rachoisotope  in  snails  receiving  spermatozoa  was  record- 
ed chiefly  in  the  spermatheca  and  collecting  canal  of  the 
ovotestis  (Paraense,  1976). 


W.  Monteiro  and  T.  Kawano,  2()()() 


Page  75 


Bioinphdldhd  <iliil>i'iitti  is  known  tu  pivfcTfntiallv  con- 
centrate iron  in  the  (notestis,  as  well  as  in  non-repro- 
tluetive  organs  such  as  chgestive  gland  and  mantle  (He- 
neine  et  al,  1969).  Therefore,  it  is  plausible  that  ^'^Fe 
transferred  from  labeled  to  non-labeled  snails  could  be 
incoiporated  in  the  ovotestis  of  receptor  snails,  indepen- 
dentK  of  spermatozoa  being  the  carrier  of  ''Fe  within 
the  receptor  suiiil. 

On  the  other  hand,  the  use  of  DNA-specific  labels  can 
be  an  efficient  wav  of  tracking  the  location  of  allosper- 
matozoa  in  the  inseminated  snail  partner.  Tlie  route  of 
spermatozoa  heads  in  the  inseminated  snail  can  be  de- 
termined by  the  use  of  tritiated  thymidine  ('H  Td).  This 
method  also  allows  for  the  detection  of  possible  areas  of 
storage  and  sites  of  cross-fertilization. 

MATERIALS  AND  METHODS 

One  hundred  and  six  specimens  of  Biomphalaria  tcna- 
gophila  were  used  both  of  the  wild  txpe  and  albino  mu- 
tants. Specimens  were  collected  in  Bom  Retiro,  Joinville, 
Santa  Catarina  State,  Brazil.  They  were  kept  for  over  4 
vears  under  laboratorv'  conditions.  Each  aquarium  con- 
tained a  tablespoon  of  red  earth  sterilized  in  <\t\  incu- 
bator at  90°C  for  12  h.  One  hundred  and  eightv  ml  fil- 
tered water,  saturated  with  calcium  carbonate,  was  add- 
ed to  each  aquarium.  Snails  were  fed  daily  with  fresh 
lettuce  and  twice  a  week  with  a  meal  consisting  of  equal 
amounts  of  red  earth,  powdered  whole  milk,  wheat  germ 
flakes,  powdered  calcium  carbonate,  and  dry  powdered 
alfalfa.  To  a  5  1  amount  of  diis  mixture  350  mg  of  vitamin 
E  was  added.  Except  in  some  cases,  snails  were  kept  in 
isolation  from  sexual  immaturity  onward. 

Two  suppfies  of  aqueous  solutions  of  'H  Td  were  used 
in  the  ex-periments.  Ten  pi  of  one  (USP,  kindlv  offered 
by  Dr.  Mariano  Amabis,  University  of  Sao  Paulo)  yielded 
218687  counts  per  min.  (cpm);  the  other  commercial 
solution  (\VM)  was  purchased  as  1  millicurie,  or  .37  me- 
gabequerel/ml;  5|xl  from  this  solution  \ielded  314850 
cpm  in  a  Beckmann  Liquid  Scintillator  7200,  using  "pro- 
gram 2  for  tritium".  Each  of  the  106  specimens  was 
anesthetized  by  immersion  in  a  0.05%  nembutal  solution 
for  3^  h,  depending  on  snail  size.  Aftei-ward,  they  were 
inoculated  with  5-10|jl1  of  'H  Td  injected  using  micro- 
syringe  into  the  lung  cavity,  through  the  pneumostome. 
Recovery  from  anesthesia  took  place  in  a  humid  cham- 
ber, for  3—4  h,  with  each  snail  placed  on  its  right  side  in 
a  plate  with  water  covering  half  shell.  After  this  period, 
each  inoculated  snail  was  returned  to  its  aquarium  for 
completion  of  recovery  from  anesthesia  and  radioisotope 
incoiporation. 

Between  4—29  days  after  treatment,  each  inoculated 
snail  (IC)  was  paired  with  a  non-inoculated  snail  (NIC) 
for  a  period  of  6  —  984  h.  NIC  snails  were  dissected  un- 
der stereoscopic  microscope  within  a  period  of  0—24  h 
after  separation.  The  following  organs  were  extracted: 
spermatheca,  oothecal  gland  +  muciparous  gland,  o\i- 
duct  or  oothecal  gland  by  itself,  but  muciparous  gland 
with  oviduct,  part  of  digestive  gland,  carrefour,  albinnen 


Figure  L  Diagram  of  tlie  reproductive  system  of  Biomphal- 
aria tcnaf^ophiln.  1.  Hind-part  of  ovotestis;  2.  Follicles  of  the 
ovotestis;  3.  Ovotestis  collecting  canal;  4.  Seminal  vesicle;  5. 
Albumen  gland;  6.  0\'idiict;  7.  Muciparous  gland;  8.  Oothecal 
gland;  9.  Nidamental  gland;  10.  Spermatheca;  11.  Vagina;  12. 
Spermioviduct;  13.  Prostate;  14.  Deferens  canal;  15.  Penis 
sheath;  16.  Prepntiiim 


gland,  seminal  vesicle  and  ovotestis  (Figure  1).  Counting 
in  a  section  of  the  digestive  gland  determined  the 
amount  of  incorporation  of  radioisotope  outside  the  re- 
productixe  system,  the  background  radioisotope  incor- 
poration. The  ovotestis  was  macerated  releasing  material 
from  the  collecting  canal  and  follicles.  These  contents 
were  collected  separately  from  the  rest  of  the  macerate. 
Each  sample  was  transferred  to  a  vial  containing  0.3  or 
0.5  ml  of  a  solution  of  3  %  sodiiun  hvpochloride  (com- 
mercial bleach).  The  material  was  repeatedly  refluxed 
with  a  Pasteur  pipette  until  completely  dissolved.  Prep- 
aration of  filter  paper  strips  imbibed  with  samples  for 
measuring  in  the  liijuid  scintillator  was  based  on  the 
methodology  of  Byfiekl  and  Scherbaum  (1966). 

Though  spermatozoa  may  be  found  and  counted  in 
chfferent  organs  of  the  reproductive  system  during  and 
after  the  pairing  process  (Monteiro  and  Kawano,  1998), 
in  this  work  the  occurrence  of  allosperTnatozoa  in  the 
examined  organs  was  accounted  tor  only  by  counting  in 
the  hquid  scintillator 

Some  controls  were  used  to  test  the  efficiencv  of  the 
method.  Processing  of  each  series  of  paper  strips  with 
samples  included  a  paper  strip  without  sample,  a  "blank" 
that  allowed  us  to  investigate  the  amount  of  free  racho- 
active  material  that  could  be  absorbed  from  solutions 
where  the  papers  with  samples  were  being  treated  (Ta- 
ble 1). 

The  radiation  in  the  water  where  the  snail  recovered 
from  anesthesia  was  measured  by  scintillation  to  evaluate 
the  efficiency  of  the  inoculation  in  the  lung  cavitv  and 
the  draining  of  H  Td  into  the  mechum  during  the  re- 
coveiy  period.  The  amount  of  radiation  injected  into 
each  snail  was  also  measured  (Table  1). 

RESULTS  AND  DISCUSSION 

The  technique  of  inoculation  in  the  lung  ca\it\  through 
the  pneumostome  of  the  anesthetized  snail  was  here 
used  for  the  first  time.  The  evaluation  of  this  technique 


Page  76 


THE  NAUTILUS,  Vol.  114,  No.  2 


Table  1.   Experiments  with  different  supplies  of  tritiated  th^inidine  (USP  and  WM  supplies) 
minute.  Blanks  =  data  not  available 


Cpni  =  scintillation  counts  per 


Exp.  1 

Exp.  2 

Exp.  3 

Exp.  4 

No.  snails 

5 

5 

5 

5 

Microliters  inoculated/snail 

lOUSP 

lOUSP 

lOUSP 

5WM 

Inoculation-pairing  (days) 

7 

7 

11 

6 

Duration  of  pairing  (h) 

24 

24 

24 

24 

Pairing-dissection  (h) 

52 

48 

216 

40 

Separation-dissection  (h) 

28 

24 

192 

16 

Inoculation  to-dissection  (approx.  days) 

9 

9 

20 

8 

Diameter  of  snails  (mm) 

12-14 

12-13 

12-13 

13-14 

Reference  (cpm) 

27507 

26911 

23534 

Background  (cpm) 

71 

65 

84 

75 

Blank  (cpm) 

105 

119 

0\nduct  (cpm) 

109 

89 

78 

0\'otestis  collecting  canal  (cpm) 

91 

84 

76 

105 

Follicles  of  ovotestis  (cpm) 

174 

84 

107 

121 

Spermatheca  (cpm) 

81 

114 

114 

110 

Nidamental  gland  (cpm) 

96 

890 

884 

Oothecal  gland  (cpm) 

135 

Seminal  vesicle  (cpm) 

1184 

63 

139 

121 

Digesti\'e  gland  (cpm) 

87 

104 

Carrefour  (cpm) 

76 

107 

Albumen  gland  (cpm) 

427 

0\iduct  -muciparous  gland  (cpm) 

102 

°  USP,  supplv  kindh'  offered  by  Mariano  Amabis,  Universitv  of  Sao  Paulo:  WM,  supplv  purchased  by  one  of  the  authors. 


was  carried  out  in  18  snails  inoculated  with  a  total  oi  90 
jjlI  of  ^H  Td.  Retention  of  inoculate  in  the  lung  cavity 
was  tested  through  measurements  of  the  relative  amount 
of  radioisotope  that  drained  during  the  4  h  recovery 
from  anesthesia.  The  snails  recovered  in  27.5  ml  of  water 
that  yielded  81152.5  cpm.  As  each  5  |xl  of  the  applied 
radioisotope  solution  counted  314850  cpm,  there  was, 
on  average,  a  1.4  %  loss  of  radioactive  material  from 
snails  to  the  water.  This  result  encourages  the  use  of  this 
inoculation  method  in  tests  of  prospective  toxicology,  se- 
lection of  mutagens,  and  of  the  mechanism  of  action  of 
water  pollutants. 

The  results  of  1 1  out  of  a  total  of  30  e.xperiments  are 
shown  in  Table  1.  The  remaining  experiments  did  not 
record  high  counts  for  any  of  the  examined  organs.  The 
low  counts  in  all  examined  organs  may  have  occurred  in 
sniiils  that  did  not  act  as  females  in  relation  to  the  in- 
oculated partner.  An  arbitraiy  criterion  was  established; 
to  consider  onlv  the  experiments  in  which  the  counting 
in  the  most  labeled  organ  was  at  least  twice  as  those  in 
the  least  labeled  one. 

As  a  natural  consequence  of  metabolism,  the  long  du- 
ration of  an  e.xperiment  might  cause  the  incoqjoration 
of  the  radioisotope  in  an  organ  absolutely  lacking  sper- 
matozoa. This  does  not  seem  to  be  the  case  here,  even 
in  experiments  lasting  for  216  and  948  h  (e.xperiments 
3,  10,  and  11),  from  pairing  to  dissection  time.  In  these 
conditions,  the  most  labeled  organs  were  the  nidamental 
gland  (884  cpm),  the  albumen  gland  (210  cpm)  and  fol- 
licles of  ovotestis  (365  cpm),  respectively  (Table  1).  We 
looked  for  possible  transfer  of  the  radioisotope  label 
from  the  reproductive  system  to  a  highly  metabolically 


active  organ  such  as  the  chgestive  gland.  A  tragment  of 
digestive  gland,  similar  in  volume  to  the  ovotestis,  was 
simultaneously  tested  in  experiments  2  and  3  (respec- 
tively 48  and  216  h  after  pmring).  The  relatively  low 
counts,  87  and  104  cpm  respectively,  discarded  that  pos- 
sibilit)'  (Table  1). 

After  a  specimen  of  B.  glabrata  behaves  as  female 
during  pairing,  its  spermatheca  is  packed  with  sperma- 
tozoa. That  organ  is  site  ot  lysis  of  spermatozoa  (Kitajima 
and  Paraense,  1983).  Suiprisingly,  among  all  the  e.xper- 
iments, only  one  presented  relatix'ely  elevated  countings 
for  spermatheca  (206  cpm):  experiment  8  (Table  1).  Be- 
sides, in  experiment  9,  where  the  time  lapse  from  pair- 
ing to  dissection  was  only  7  h,  the  spermatheca  counting, 
87  cpm,  was  equivalent  to  the  blank  (Table  I).  Given  the 
countings  in  the  digestive  gland  (458  cpm),  it  is  possible 
that  the  spermatozoa  have  either  bypassed  or  spent  a 
verv  brief  time  in  the  spermatheca  on  their  way  to  the 
digestive  gland.  Also,  clue  to  the  relatively  short  time 
elapsed  from  pairing  to  dissection,  7  h,  the  hypothesis 
of  radioisotope  transfer  from  spermatheca  to  digestive 
gland  was  not  taken  into  account  in  the  analysis  of  the 
data. 

No  relation  was  found  between  the  duration  of  pairing 
and  progression  of  the  allospermatozoa  along  the  repro- 
ductive ducts.  After  only  54  h  from  the  beginning  of 
pairing,  in  experiment  6,  the  allospermatozoa  were  in 
the  folhcles  of  the  most  distant  organ  from  the  vagina, 
the  ovotestis  (585  cpm).  But  216  h  after  the  beginning 
of  p;iiring,  the  iillospermatozoa  were  in  the  nidamental 
gland  (216  cpm,  experiment  3),  the  organ  closest  to  the 
vagina  (Table  1). 


W.  Monteiro  and  T. 

Kawano,  2000 

Page  77 

Tabic  1.   Exteiuled 

Exp.  5 

Exp.  6 

E.\p.  T 

Exp.  S 

Exp.  9 

Exp.  10 

Exp.  11 

4 

5 

3 

.5 

3 

2 

lOUSP 

5\\M 

lOVVM 

lOW'M 

lOWM 

lOWM 

lOWM 

19 

7 

/ 

11 

19 

29 

29 

96 

24 

8 

9 

6 

984 

984 

120 

54 

8 

31 

6 

984 

24 

30 

0 

22 

0 

? 

0 

24 

9 

-' 

12 

19 

70 

12-14 

12-14 

9-13 

9-11 

11-12 

12-13 

11-12 

24765 

24383 

24357 

24357 

24027 

24585 

75 

103 

72 

99 

99 

55 

65 

119 

103 

110 

87 

87 

92 

77 

103 

117 

89 

76 

68 

68 

787 

585 

106 

89 

73 

59 

365 

168 

93 

172 

206 

87 

76 

147 

133 

120 

74 

77 

78 

71 

84 

111 

105 

275 

74 

57 

75 

68 

115 

166 

119 

92 

458 

210 

63 

314 

131 

9.3 

84 

81 

S7 

65 

The  simultaneous  occurrence  of  high  counting  in  the 
toUicles  of  o\otestis  and  in  the  oviduct-muciparous  gland 
in  a  single  experiment  (experiment  5),  after  96  h  of  pair- 
ing, suggests  the  occurrence  of  double  copulation  (Dias, 
1995).  Spermatozoa  transferred  in  the  initial  copulation 
could  be  located  in  the  organ  most  distant  from  the  va- 
gina, the  folHcles  of  ovotestis.  The  spermatozoa  trans- 
ferred in  a  subsequent  copulation  may  be  located  in  an 
organ  closer  to  the  vagina,  such  as  the  oviduct-mucipa- 
rous gland  (Table  1).  This  explanation,  however,  could 
be  inconsistent.  Even  if  the  last  copulation  happened  at 
the  end  of  pairing,  the  24  hours  that  passed  after  the 
separation  of  the  partners  would  be  enough  tor  these 
spermatozoa  to  advance  along  the  genital  tract,  bxpass- 
ing  the  oviduct-muciparous  gland.  In  a  shorter  period  of 
time,  from  pairing  to  dissection,  allospermatozoa  pro- 
ceeded to  the  seminal  vesicle  and  albumen  gland,  re- 
spectively, in  experiments  7  and  9.  The  results  of  exper- 
iment 3,  however,  show  that  it  is  possible  that  the  sper- 
matozoa remain  in  the  nidamental  gland  for  up  to  192 
h  after  the  separation  of  the  partners.  Therefore,  under 
the  experimental  conditions,  the  scintillation  counting 
may  indicate  that,  in  a  single  or  in  diiferent  snails,  from 
a  group  of  4  (experiment  5,  Table  1),  the  spermatozoa 
may  be  located  in  sites  as  separated  as  the  ovotestis  and 
organs  closer  to  the  vagina  such  as  the  oviduct-mucipa- 
rous gland.  It  is  possible  that,  rather  than  suggesting  ran- 
dom differences  in  locomotion  abilit),  these  results  could 
indicate  strategic,  differential  placement  of  allosperma- 
tozoa. 

All  the  occurrences  of  allospermatozoa  along  the  re- 
productive system,  registered  in  the  nidamental  gland, 
seminal  vesicle,  albumen  gland,  and  iolhcles  of  the  ovo- 


testis are  difficult  to  explain  At  first  sight,  the  nidamen- 
tal gland  seems  to  be  a  transient  site  ot  allospermatozoa 
on  their  way  to  their  final  destination,  where  the\-  will 
fertilize  the  oocytes  (Paraense,  1959).  However,  their 
stay  in  the  nidamental  gland  for  216  h  from  pairing  to 
dissection  time  (experiment  3,  Table  1),  a  period  of  time 
much  longer  than  that  necessary  for  the  spermatozoa  to 
reach  the  follicles  of  ovotestis  (experiments  5  and  6:  120 
h  and  54  h,  respectivelv;  Table  1),  contradicts  this  pre- 
mise. Considering  that  the  seminal  vesicle  of  the  donor 
may  be  empty  after  pairing  (Monteiro  and  Kawano, 
1998),  the  presence  of  labeled  allospennatozoa  in  this 
organ  (experiments  1  and  7)  suggests  that  autosperma- 
tozoa  were  replaced  b\'  allospermatozoa.  This  inteq^re- 
tation  apparently  conflicts  with  the  occurrence  ot  labeled 
allospermatozoa  both  in  the  folficles  of  ovotestis  and  in 
the  albumen  gland,  both  within  short  and  long  intervals 
from  pairing  to  dissection  (e.xperiments  4,  5,  6,  9,  10, 
and  11). 

No  relation  was  found  between  the  duration  of  pairing 
and  progression  of  the  allosperm  through  the  reproduc- 
tive system.  The  labeled  allospermatozoa  were  located 
in  the  spermatheca,  nidamental  gland,  oviduct-mucipa- 
rous gland,  albumen  gland,  seminal  vesicle  and  follicles 
of  ovotestis.  It  seems  that  the  destination  of  allosper- 
matozoa are  the  follicles  of  the  ovotestis.  Their  occur- 
rence in  the  spermatheca,  where  Ksis  takes  place,  may 
be  highly  detrimental  or  fatal.  Copulation  in  B.  tcna^o- 
phila  may  be  unilateral  or  reciprocal  (Dias,  1995).  Con- 
sequently, the  length  of  stav  of  labeled  allospermatozoa 
in  the  nidamental  gland,  oxiduct-muciparous  gland,  or 
albumen  gland  may  be  the  result  of  imilateral  copula- 
tion, when  the  snail  played  only'  a  receiving  role,  which 


Page  78 


THE  NAUTILUS.  Vol.  114.  No.  2 


resulted  in  a  seminal  \esitle  full  of  autospermatozoa.  Al- 
lospermatozoa  may  wait  within  the  female  tract  until  the 
snail  plays  a  donor  role  in  a  subsequent  pairing,  which 
can  be  unilateral  or  reciprocal.  Then,  allospermatozoa 
ma\'  pass  through  an  empt)^  seminal  \esicle  toward  the 
follicles  of  the  ovotestis.  Within  an  o\otestis  follicle,  in 
the  oocyte  formation  site  and  co-existing  with  immature 
autospermatozoa  (Homan,  1972),  allospermatozoa  may 
fertihze  the  oocytes  (Laramberg,  1939;  Paraense,  1955). 
The  many  possible  locations  for  strategic  storage  of  al- 
lospermatozoa suggests  displacement  of  autospermato- 
zoa, which  makes  possible  the  appropriate  deplo\nnent 
of  allospermatozoa. 

Allospennatozoa  were  present  in  ovotestis  follicles  in 
experiments  5,  6,  and  11,  but  were  not  found  in  the 
collecting  canal  of  ovotestis  in  any  of  the  11  experiments. 
NoUvithstanding.  Paraense  (1976),  labeUng  snails  with 
'"Fe,  found  e\idence  of  allospermatozoa  in  the  collecting 
canal.  A  possible  explanation  for  this  result  is  the  re- 
incorporation of  iron.  In  fact,  the  ovotestis  has  a  noted 
avidity  for  iron  (Heneine  ei  a/.,  1969;  1970a;  1970b).  Of 
all  these  locations,  the  one  more  consistent  with  data 
from  the  hterature  is  the  alliumen  gland,  although  all 
searches  for  spermatozoa  in  the  canals  of  the  gland  and 
nearby  areas  during  the  preparation  of  the  present  work 
have  yielded  negative  results  (not  even  spermatozoa 
heads  were  found).  Nevertheless,  a  chamber  of  fertiliza- 
tion, in  the  carrefour,  next  to  the  albinnen  gland,  is  often 
referred  as  the  final  site  for  storage  of  spermatozoa  in 
studies  carried  out  on  several  other  species  of  moUusks 
(Duncan,  1958;  1960;  Lind,  1973;  Geraerts  and  Joosse, 
1984). 


CONCLUSIONS 

Inoculation  of  a  solution  of  tritiated  thymidine  through 
the  lung  cavity  of  individuals  of  Biomphalaria  tcnago- 
phila  is  an  adequate  method  to  label  spermatozoa  for 
determining  their  location  in  the  partner  after  pairing. 
Labeled  spermatozoa  were  detected  in  the  recei\ing 
snail,  through  significant  levels  of  scintillation  counting, 
in  the  spennatheca,  seminal  vesicle,  folhcles  of  ovotestis, 
nidamental  gland,  albumen  gland,  and  oviduct-mucipa- 
rous  gland.  Material  extracted  from  the  collecting  canal 
of  the  ovotestis  did  not  show  countings  for  labeled  sper- 
matozoa. The  presence  of  labeled  spermatozoa  simul- 
taneouslv  in  the  follicles  of  the  o\'otestis  and  in  the  o\i- 
duct-muciparous  gland  suggests  options  for  tlifferential 
storage  locations,  in  a  process  that  includes  displacement 
of  autospermatozoa  to  allow  for  adequate  positioning  of 
allospermatozoa. 

ACKNOWLEDGMENTS 

We  thank  Dr.  Mara  Liicia  Ferreira  Dias  for  valuable  sug- 
gestions and  criticism  to  this  work,  and  to  fellowships 
from  CAPES  (Coordena9ao  e  Aperfei^oamento  de  Pes- 
soal  de  Nivel  Superior,  Brazil). 


LITERATURE  CITED 

Abdel-Malek.  E.  T.  1954.  Moqshological  studies  on  the  family 
Planorbidae  (Mollusca:  Pulmonata).  II.  Genital  organs  of 
Biomphalaria  boissiji  (Subfamily  Planorbmae,  H.A.  Pils- 
bn,  1934).  Transactions  of  American  Microscopical  Soci- 
et>'  7.3:285-296. 
Byfield,  J.  E.  and  O.  H.  Scherbaum.  1966.  A  rapid  radioassay 
technique  for  cellular  suspensions.  Analytical  Biochemis- 
try 17:433-443. 
Dias,  M.  L.  F.  1995.  Padroes  comportamentais  da  reprodufao 
de  Biomphalaria  tcnngophila  (Orbigny)  (Mollusca,  Pla- 
norbidae)  ( Reproducti\e  behavioural  patterns  of  Biom- 
phalaria tenagophila  (Orbigny)  (Mollusca.  Planorbidae).) 
University  of  Sao  Paulo,  Doctor  of  Science  Thesis. 
Duncan,  C.  J.  1958.  The  anatomy  and  phvsioIog\-  of  the  repro- 
ductive system  of  the  freshwater  snail  Phtjsa  fontinahs 
(L.).  Proceedings  of  the  Zoological  Societ)' of  London  131: 
.55-66. 
Duncan,  C.  J.  1960.  The  genital  systems  of  the  freshwater  Ba- 
sonimatophora.  Proceedings  of  the  Zoological  Socieh'  of 
London  138:339-356. 
Duncan,  C.  ].  1975.  Reproduction.  In:  Fretter,  \'.  and  ].  Peake, 
(eds.)  Pulmonates.  Academic  Press,  New  York,  pp.  309- 
365. 
Fretter,  V.  and  A.  Graliam.  1964.  Reproduction.  In:  Wilbur,  K. 
M.  and  C.  M.  Yonge,  (eds.)  Physiology  of  Mollusca.  Aca- 
demic Press,  New  York,  pp.  127-164. 
Gazzinelli,  G.,  F  J.  Ramalho  Pinto  and  B.  Gilbert.  1970.  Up- 
take of  '"'Fe  as  a  tool  for  study  of  the  crowding  effect  in 
Biomphalaria  glabrata.  The  American  Journal  of  Tropical 
Medicine  and  Hygiene  19:10.34-1037.  ' 
Geraerts,  VV,  P.  M.  and  J.  Joosse.  1984.  Freshwater  snails  (Ba- 
sommatophora).  In:  Tonipa,  A.  S.,  N.  H   \'erdonk  and  J. 
A.  M.  van  den  Biggelaar  (eds).  The  Mollusca,  vol.  7,  Re- 
production. Academic  Press,  New  York,  pp.  142-208. 
Heneine,  I.  F,  G.  Z.  GazzinelH  and  W.  L.  Tafuri.  1969.  Iron 
metabolism  in  the  snail  Biomphalaria  glahrata:  uptake, 
storage  and  transfer  Comparative  Biochemistry  and  Phys- 
iology 28:391-399. 
Heneine,  L.  E.,  M.  C.  dos  Santos  and  G.  Z.  Giizzinelli.  1970a. 
Transferencia  de  ferro  em  Biomphalaria  glahrata.  Anais 
da  Academia  brasileira  de  Ciencias  42  (suplemento):171- 
175. 
Heneine,  L  E.,  F  G.  Ramallio-Pinto,  J.  Pellegrino  and  B.  Gil- 
bert.  1970b.  Uptake  of  "Fe  as  a  tool  for  study  of  the 
crowding  effect  in  Biomphalaria  glahrata.  The  American 
Journal  of  Tropical  Medicine  and  Hygiene  19:10.34-1037. 
Homan.  J.  A.  1972.  The  ultrastructure  and  activity  of  spenu  in 
Acanthodoris  pilosa.  a  nudibranchiate  Mollusk.   Marine 
Biolo,2\-  12:179-183. 
Horstmann,  H.  J.  1955.  Untersuchungen  zur  Physiologic  der 
Begattung  und  Befruchtung  der  Schlammschnecke  Li/»i- 
naea  stagnalis  L.  Zeitschrift  fiir  Morphologic  und  Okolo- 
gie  der  Tiere  44:  222-268. 
Jong-Brink,  M,   1969.  Histochemical  and  elecron  microscope 
observations  on  the  reproductive  tract  of  Biomphalaria 
glahrata   {Aiistrahvbi.s  glabratus).   Intermediate  host  of 
Schistosoma    mansoni.    Zeitschrift   Zellforschungen    102: 
507-542. 
Kitajima,  E.  W.  and  VV.  L.  Paraense.  1976.  The  nltrastnicture 
of  mature  spemis  of  the  freshwater  snail  Biomphalaria 
glahrata   (Mollusca,  Gastropoda).  Transactions  of  the 
American  Microscopical  Society  95:1-10. 
Kitajima,  E.  W.  and  W.  L.  Paraense.  1983.  The  ultrastructure 


W.  Monteiro  and  T.  Kavvano,  2()()() 


Page  79 


of  the  spermatheca  of  Bioiiij>h(il/iria  <J/ihnitti  (Gastropoda, 
Puliiionata).  Jonnial  of  Moqiliologv  176:211-220. 

Laianibergiie,  M.  1939.  Etude  de  I'autot'econdatioii  cliez  les 
Gasteropodes  Pulmoiies.  Recherches  sur  I'apliallie  et  la 
fecondation  chez  Bulimis  (Isidora)  contortus  Michaud. 
Bulletin  Biologique  de  France  et  Belgique  73:19-1.31. 

Lind,  H,  1973  The  functional  sipiificance  of  the  spermato- 
pliore  and  the  fate  of  spermatozoa  in  the  tjenital  tract  of 
hh'lix  pomatia  (Gastropoda:  Stsloniniatophora).  )ouniai  of 
Zooiogv-  169:39-64. 

Monteiro.  W.  and  T.  Kawano.  1998.  Investimento  na  fvui^ao 
niasculina  em  Biomplmlaria  tena^ophila  (ORBIGNY) 
(Mollusca,  Planorhidae)  a\aliado  pelo  numero  de  esper- 
niatozoides.  Revista  brasileira  de  Biologia  58(3):4(Sl-490. 


Paraense,  W.  L.  1955.  Self  and  cross-fertilization  in  Australor- 
his  glabrattis.  Memorias  do  Instituto  Oswaldo  Gruz  53: 
277-284. 

Paraense,  W.  L.  1959.  One-sided  reproductive  isolation  be- 
tween geograpliicallv  remf)te  populations  of  a  planorbid 
snail.  The  American  Naturalist  93:93-101. 

Paraense,  W.  L.  1975.  Estado  atual  da  sistematica  dos  planor- 
bideos  brasileiros.  Arquivos  do  Museu  Nacional  55:105- 
128. 

Paraense,  W.  L.  1976.  The  sites  of  cross  and  self-fertilization 
in  planorbid  snails.  Rev-ista  Brasileira  de  Biologia  .36:5.35- 
5.39. 

Thompson,  T  E.  197.3.  Euthyneuran  and  other  molluscan  sper- 
matozoa. Malacologia  14:167-206  (plus  addendum,  44.3- 
444). 


THE  NAUTILUS  114{2):S0-83,  2000 


Paee  80 


Status  of  the  tidewater  mucket,  Leptodea  ochracea  (Say,  1817) 
(Bivalvia:  Unionidae),  in  Halfway  Pond,  Massachusetts,  USA 


Jay  Cordeiro 

Department  of  Invertebrates 
American  Museum  of  Natural  History 
Central  Park  West  at  79th  Street 
New  York,  NY  10024  USA 
corcleiro@amnh.orCT 


ABSTRACT 

The  tidewater  mucket,  Leptodea  ochracea  (Say,  1817),  is  dis- 
tributed along  the  Atlantic  coast  of  North  America  and  is  often 
found  on  sandy  substrates  with  little  or  no  flow.  Halfwav  Pond, 
in  PKinouth,  Massachusetts,  is  one  of  the  few  remaining  hab- 
itats in  Massachusetts  for  tliis  species,  but  li\e  specimens  have 
not  been  collected  there  since  1981.  In  summer  1995,  the 
pond  was  sur\'eyed  in  an  attempt  to  determine  if  the  tidewater 
mucket  had  been  extiipated.  Of  213  li\'e  unionids  collected, 
comprising  6  different  species,  only  1  specimen  was  identified 
as  L.  ochracea.  The  single  specimen  was  found  in  the  Agawam 
River  outflow  area  in  the  southwest  comer  of  the  pond  along 
with  4  of  the  5  other  species  of  unionids  occurring  in  the  pond. 
This  high  flow  area  had  a  high  proportion  of  benthic  macro- 
phvtes  in  a  sanch'  substrate — hpical  habitat  for  L.  ochracea. 
Other  species  found  in  the  pond  include  EUiptio  coiuplanata 
(Lightfoot,  1786),  LampsiUs  radiata  radiata  (Gmelin,  1791), 
Alasmidonta  undulata  (Say,  1817).  Strophitiis  undidatus  (Say, 
1817),  and  Pijganodon  cataracta  (Say,  1817).  This  study  con- 
firms the  decline  of  L.  ochracea  in  Halfway  Pond,  Massachu- 
setts, over  the  last  15  years.  If  not  already  gone,  this  species 
may  disappear  from  the  pond  once  the  remaining  indi\iduals 
die. 

Keij  words:  Unionoidea,  freshwater  clams,  freshwater  mussels, 
extirpation 


INTRODUCTION 

The  tidewater  mucket,  Leptodea  ochracea  (Say,  1817), 
is  a  freshwater  mussel  (family  Unionidae)  distributed 
along  the  Atlantic  coast  of  North  America  from  Nova 
Scotia  to  Georgia  (Johnson,  1947;  1970;  Strayer  and  Jir- 
ka,  1997).  Populations  are  often  found  on  sandy  sub- 
strates with  httle  or  no  flow  (Smith,  1981),  usually  in 
water  bodies  near  the  ocean  but  not  necessarily  con- 
nected to  it. 

In  Massachusetts,  Leptodea  ochracea  is  found  in 
coastal  plain  ponds  larger  dian  four  hectares,  along  the 
southeastern  portion  of  the  state,  from  the  South  Shore 
to  Buzzard's  Bay  to  Cape  Cod  (Smith,  1991).  Because 
of  its  highly  restricted  geographic  distribution  and  de- 


creasing abundance  in  ponds  throughout  its  range,  the 
tidewater  mucket  is  listed  as  a  species  of  special  concern 
(Smith,  1981;  Williams  et  al,  1993;  MDFW,  1997). 

Over  the  past  65  years,  Leptodea  ochracea  was  found 
in  Halfvvay  Pond  in  PlvTnouth  County  on  several  occa- 
sions (Table  1).  Since  1981,  however,  it  has  not  been 
recorded  there  (Nature  Conservancv,  unpublished  data). 
In  the  summer  of  1995,  the  author  surveved  Half\yay 
Pond  in  an  attempt  to  determine  the  distribution  status 
of  L.  ochracea  and  other  unionids. 

Halfwav  Pond  is  a  94  ha  freshwater  pond  with  a  4.9 
ha  island  preserve  in  its  center  located  just  south  of 
Plymouth,  Massachusetts,  next  to  Miles  Standish  State 
Forest.  The  island  preserve  is  a  candidate  for  designa- 
tion as  a  National  Natural  Landmark  bv  the  Department 
of  the  Interior  mainly  because  it  contains  one  of  the 
oldest  forests  rem;uning  in  Massachusetts  (Nature  Con- 
servancy, unpublished  data)  and  is  located  in  the  area 
with  the  fastest  development  rates  in  the  state  (Living- 
ston, 1987). 


MATERIALS  AND  METHODS 

Live  mussels  were  collected  by  hand  with  a  meter-long 
dip  net  with  2.5  cm  mesh.  Surveys  were  supplemented 
by  snorkeUng  to  a  depth  of  1.5  meters.  The  net  handle 
was  used  to  delineate  a  circular  quadrat  for  each  sample 
area  (1  m-  area).  Surveys  were  conducted  at  2  to  4  m 
intervals;  30  samples  were  taken  at  the  first  2  sites  and 
15  samples  at  the  remaining  4  sites  around  the  lake.  For 
each  specimen,  species  name,  total  length  in  miUimeters, 
degree  of  dissolution  of  the  shell  (shell  wear),  collection 
localit\-,  and  sechment  t\pe  (estimated  visually  as  sand, 
mud,  or  cobble)  were  recorded.  Specimens  were  ex- 
amined for  identification  with  a  6  X  hand  lens  and  dis- 
secting microscope  to  30  X ,  when  necessary.  Specimens 
were  only  dissected  when  internal  characters  were  re- 
quired for  identification.  Species  were  identified  using 
Smith  (1991)  as  a  primarv'  .source  and  McM;Jion  (1991) 
and  Peckarsky  et  al.   (1990)  as  supplemental  sources. 


J.  Cordeiro,  2000 


Page  81 


Tabic  1.  Li'ptodcn  ochmcca  in  H;ilf\\':i\'  Pond.  Massacliusetts:  specimens  in  major  American  mnsenni  collections 


No.  specimens 

Localih 

Collector!  s) 

Date 

Collection  No. 

1 

Haltwas-  Pond 

ex-MCZ 

USNM  656539 

3 

Halt\va\  Pond 

R.  I.  Johnson 

USNM  592091 

1 

alonCT  Mast  Roail.  Hal 

K\a\   Pond 

ex-MCZ 

[pre-1975] 

OSUM  38073 

4.5 

HalR\a\-  Pond 

R.  E.  M..  W.  F. 

C. 

(William  F. 

Clapp?)  ex-MCZ 

Aug.  19,  1933 

OSUM  26550 

9 

HalKvaN-  Pond 

R    I    Johnson 

Jul.  1,  1941 

AMNH  72936, 
DMNH  48551, 
USNM  600325 

3 

Halfway  Pom!  outlet 

R.  I.  Johnson 

June  1943 

MCZ  134.841. 
MCZ  159150 

4 

Halfway  Pond 

M.  K.  Jacohson 

Aug.  22,  1952 

AMNH  121013 

4 

South  shore,  Half\va\'  Pond 

M.  K.  Jacohson 

Aug.  23,  1952 

AMNH  127888 

12 

Agawani  River  outlet. 

Halfway  Pond 

D.  G.  Smith.  A. 

E. 

Pratt 

July  1981 

UMAMZ  5.39 

Voucher  .specimens  ior  Lcptodca  ochracca  were  not  .sub- 
mitted due  to  the  scarcity  of  specimens. 

Over  the  past  300  years  there  has  been  some  contu- 
sion as  to  the  proper  nomenclature  o( Lcptoclca  ochracca 
(Johnson,  1947;  1970;  Morrison,  1975;  Bereza  and  Ful- 
ler, 1975),  since  its  original  description  as  Pectunculus 
fluviatilis  Lister,  1685.  A  forthcoming  manuscript  (Cor- 
deiro, in  preparation)  should  clarifv  manv  of  the  complex 
nomenclatural  issues  inxoKing  this  species.  In  hght  of 
this  uncertainty  in  nomenclature,  the  author  herein  will 
refer  to  the  species  as  Leptodea  ochracca  (Say,  1817)  as 
per  its  listing  in  Turgeon  ct  al.  (1998). 

Institutional  abbreviations  used  are:  AMNH,  Ameri- 
can Museum  of  Natural  History,  New  York;  DMNH, 
Delaware  Museum  of  Natural  History,  Greenville,  Del- 
aware; MCZ,  Museum  of  Comparative  Zoology,  Harvard 
University,  Cambridge,  Massachusetts;  OSUM,  Museum 
of  Biological  Diversity,  Ohio  State  Universitv,  Colimibus, 
Ohio;  UMAMZ,  Universit\-  of  Massachusetts  Museum  of 
Comparative  Zoology,  Amherst,  Massachu.setts;  USNM, 
National  Museum  of  Natural  History,  Smithsonian  In- 
stitution, Washington,  DC. 


RESULTS 

The  freshwater  mussels  EUiptio  complanata  (Lightfoot, 
1786),  Laiupsilis  radiata  radiata  (Gmelin,  1791),  A]as- 
mkhmta  iiudulata  (Sav,  1817),  Strophitiis  iindulatus  (Say, 
1817),  Pijganodon  cataracta  (Say,  1817),  and  Leptodea 
ochracca  were  collected  along  the  shores  of  Halfway 
Pond  and  the  Halfwav  Pond  Island  Preserve.  Table  2 
depicts  results  of  sur\eys  at  the  6  sites  around  the  lake. 
Survey  site  1  is  sandy  with  no  benthic,  macrophvtic 
plants  and  frequently  used  for  sport  fishing.  All  mussels 
collected  there  were  identified  as  E.  complanata.  Site  2 
was  also  sandy  and  clear  Only  2  live  specimens  of  E. 
complanata  were  found.  Site  3  had  a  sandy  substrate  that 
was  hghtly  (15%)  plant-covered.  Foi+y-seven  live  speci- 
mens of  E.  complanata  and  6  L.  radiata  radiata  were 
found.  Site  4  was  sandy  and  moderatelv  (50%)  plant- 
covered  with  80  live  .specimens  of  E.  complanata,  5  L. 
radiata  radiata,  and  1  A.  undulata.  Site  5  at  the  Agawani 
River  outflow  was  choked  with  macrophytes  amidst  oc- 
casional spots  of  bare  sand.  Although  not  measured,  cur- 
rent flow  was  estimated  to  be  higher  than  in  the  other 


Table  2.  Sampling  results:  samples  taken,  species,  and  specimen  number  obtai 


ned  m  sur\e\s  o 


f  Half\vav  Pond,  Massachusetts. 


Site 


Locality 


Samples 


Species 


No. 
specimens 


1  0.3  km  S  of  Mast  Rd.,  west  bank 

2  N  end  Mast  Rd.,  northernmost  shore 

3  NW  comer  Island 
Preserve 

4  SW  shore  Island 
Preser\'e 

5  Agawam  River  outflow,  SW  corner 


6  SW  comer,  at  pumping  station,  NW  of  site  5 


30 

£,  complanata 

14 

30 

E.  complanata 

2 

15 

E.  complanata 

47 

L.  radiata  radiata 

6 

15 

E.  complanata 

80 

L   radiata  radiata 

5 

A   iindnlata 

1 

15 

E  ci>mplanata 

22 

L.  radiata  radiata 

2 

S.  undulatus 

1 

P.  cataracta 

1 

L.  ochracca 

1 

15 

E.  complanata 

31 

Page  82 


THE  NAUTILUS,  Vol.  114,  No.  2 


sun^eyed  areas  in  the  pond.  Maximum  depth  was  less 
than  1  m.  Twenty-two  specimens  of  E.  coinplaiuita.  2  L. 
radiata  radiata,  1  S.  iiiuhdatus,  1  P.  cataracta.  1  un- 
known juvenile  mussel,  and  the  only  specimen  of  the 
target  species,  L.  ochracca.  from  the  entire  survey,  were 
found.  The  water  on  survey  site  6  was  cloudv;  the  bot- 
tom at  this  site  was  lightly  covered  with  plants  and  sandy. 
Thirty-one  specimens  plus  1  unknov^ai  juvenile  were 
identified  as  E.  complanata. 

A  total  of  213  live  specimens  comprising  6  different 
species  were  collected  in  6  sites  around  the  lake.  AH 
specimens  showed  evidence  of  shell  dissolution,  mostly 
light  wear  (nacre  exposed  on  25%  or  less  of  the  shell 
surface).  Mean  specimen  length  for  Elliptio  coinphmata, 
the  only  species  collected  in  significant  numbers  for 
analysis,  was  76.2  mm  (n  =  191),  although  this  figure  is 
most  hkelv  inflated  due  to  bias  in  sampling  methodology. 
At  sample  site  5,  the  only  area  where  the  target  species 
was  found,  specimens  of  E.  complanata  were  11.6%  larg- 
er than  in  the  other  sites.  In  addition,  the  greatest  di- 
versity (5  species)  was  found  at  this  site. 

DISCUSSION 

The  density  oi  Lcptodea  ochracca,  is  decreasing  in  Half- 
way Pond.  This  study  confirms  general  observations  of 
dechne  over  the  past  15  years.  The  single  specimen 
found  in  this  survey  is  the  only  documentation  of  the 
existence  of  L.  ochracca  in  Halfway  Pond  since  1981 
when  Doug  G.  Smith  and  A.  E.  Pratt,  University  of  Mas- 
sachusetts Museum  of  Comparative  Zoology,  collected 
12  specimens  (UMAMZ  5.39).  Agricultural  nmoff  from 
nearby  cranberr}'  bogs  recently  contributed  to  decreased 
water  clarity  accompanied  by  fish  kills  and  potential  de- 
crease in  mussel  densitv  (Douglas  G.  Smith,  personal 
communication).  Lcptodea  ochracca  is  typically  found  at 
low  densities  across  its  range  in  northeastern  North 
America  (Smidi,  1991;  Strayer  et  al,  1994;  Strayer  and 
Smith,  1996;  Strayer  and  Jirka,  1997).  The  single  speci- 
men represents  0.47%  of  all  specimens  found  (n  =  213). 
Strayer  ct  al.  (1994)  found  onlv  33  in  their  study  of  the 
Hudson  River  estuary  in  eastern  New  York  and  report 
the  species  constituting  appro.ximately  5%  of  the  unionid 
community  of  over  one  biUion  animals.  Repeated  sam- 
phng  of  the  same  area  in  1993-1995,  subsequent  to  ze- 
bra mussel  (Drcisscna  polipiu>if)ha  (Pallas,  1771))  inva- 
sion, resulted  in  only  16  specimens  found.  At  the  time 
of  this  studv  (1995),  zebra  mussels  had  not  reached  Half- 
way Pond. 

The  Lcptodea  ochracca  found  in  this  stucK'  was  in  the 
pond  outflow  in  an  area  of  high  benthic  macrophyte  cov- 
er and  sandy  substrate  that  is  typical  habitat  for  this  spe- 
cies (Johnson,  1947;  1970).  Increased  species  diversih- 
in  this  area  (5  of  6  total)  and  increased  mean  length  of 
the  most  common  species,  Elliptio  complanata,  supports 
the  prevision  that  the  outflow  area  is  a  vital  microhabitat 
for  freshwater  mussels  in  Halfway  Pond.  Increased  den- 
sities of  other  freshwater  mussel  species  (Nalepa  and 
Gauvin,   1988)  and  filter-feeding  insects  (/Vllan,   1995; 


142)  have  also  been  observed  at  other  lake  outflows.  Fa- 
vorable conditions  may  include  increased  nutrient  flow 
across  mussel  ctenidia  induced  by  the  increased  stream 
flow,  or  protection  from  fish  and  mammal  predators 
amidst  the  macrophvtic  plants. 

Whatever  the  cause,  the  Agawam  Ri\er  outflow  area 
is  an  important  refugial  microhabitat  for  the  continued 
survival  oi  Lcptodea  ochracca  in  Halfway  Pond.  The  out- 
look for  this  species  in  the  pond,  however,  is  not  good. 
With  such  a  low  density,  reproductive  potential  may  not 
be  high  enough  to  maintain  a  population.  Surveys  of  sim- 
ilar areas  in  other  coastal  ponds  should  be  conducted 
and  appropriate  protective  measures  taken  if  L.  ochracca 
is  to  remain  as  a  species  of  special  concern  or  upgraded 
to  endangered  status  in  Massachusetts. 


LITERATURE  CITED 

Allan,  ].  D.  1995.  Stream  Ecolog\'.  Stnicture  and  Function  of 
Running  Waters.  Cliapman  and  Half  London,  388  pp. 

Bereza,  D.  J.  and  S.  L.  H.  Fuller.  1975.  Notes  on  "LampsiUs" 
ochracca  (Say)  (Mollusca;  BivaMa).  Association  of  South- 
eastern Biologists  Bulletin  22:42. 

Johnson,  R.  I.  1947.  Lampsilis  cariosn  Say  and  Lampsilis 
ochracca  Say.  Occasional  Papers  on  Mollusks  1:14.5-1.56. 

Johnson,  R.  I.  1970.  The  systematics  and  zoogeography  of  the 
Unionidae  of  the  southern  Atlantic  slope  region.  Bulletin 
of  the  Museum  of  Comparative  Zoologv  140:26.3—449. 

Livingston,  T.  (ed.)  1987.  Massachusetts  and  Rhode  Island  Pre- 
serve Guide.  Nature  Conservancy  Press,  Boston,  56  pp. 

MDFW  (Massachusetts  Division  of  Fisheries  and  Wildlife). 
1997.  Massachusetts  list  of  endangered,  threatened,  and 
special  concern  species.  321  CMR  10.60.  Natural  Heri- 
tage and  Endangered  Species  Program,  Massachusetts  Di- 
vision of  Fisheries  and  Wildlife,  Westborough, 

Morrison,  J.  P.  E,  1975.  Maryland  and  Virginia  Mussels  of  List- 
er. Bulletin  of  the  American  Malacological  Union  for 
1974:.36-.39. 

Nalepa,  T.  F  and  J.  M.  Gauvin.  1988.  Distriljution,  abundance, 
and  biomass  of  freshwater  mussels  (Bivalvia:  Unionidae) 
in  Lake  St.  Clair.  Journal  of  Great  Lakes  Research  14(4): 
411-419. 

Peckarsky,  B.  L.,  P.  R.  Fraissinet,  M.  A.  Penton  and  D.  J.  Conk- 
lin.  Jr  1990.  Freshwater  Macroinvertebrates  of  North- 
eastern North  America.  Cornell  University  Press,  Ithaca, 
442  pp. 

Smith,  D.  G.  1991.  Keys  to  the  Freshwater  Macroinvertebrates 
of  Massachusetts,  v.  1.2.  Universih'  of  Massachusetts 
Press,  Amherst,  Massachusetts,  236  pp. 

Smith,  D.  G.  1981.  Selected  freshwater  invertebrates  proposed 
for  special  concern  status  in  Massachusetts.  Massachusetts 
Department  of  Environmental  Quality  Engineering,  Di- 
vision of  Water  Pollution  Control,  Westborough,  26  pp. 

Strayer,  D.  L.,  D.  C.  Hunter,  L.  C.  Smith  and  C.  K.  Borg, 
1994.  Distribution,  abundance,  and  roles  of  freshwater 
clams  (Bivalvia:  Unionidae)  in  tlie  freshwater  tidal  Hudson 
River  Freshwater  Biology  31:239-248. 

Strayer,  D.  L.  and  K.  J.  Jirka.  1997.  The  Pearl  Mussels  of  New 
York  State.  The  New  York  State  Education  Department, 
New  York,  113  pp. 

Straver,  D.  L.  and  L.  C.  Smith  1996.  Relationship  between 
zebra  mussels  (Dreissena  polipnorpha )  and  unionid  clams 


J.  Cordeiro,  2000 


Page  83 


(liiriiii;  the  earlv  stages  of  tlie  zebra  mussel  imasiiiii  dlllic 
Hudson  Rh'er.  Fresliwater  Biolog)'  36:771-779. 
Turgeon,  D.  D.,  J.  F.  Quinii  |r.,  A.  E.  Bogan,  E.  \'.  Coan,  F. 
G.  Hochberg,  VV.  G.  Lyons,  P.  M.  Mikkelsen,  R.  J.  Neves, 
C.  F  E.  Roper,  G.  Rosenberg,  B.  Roth,  A.  Sche'ltenia,  F. 
G.  Tlionipson,  M.  Vecchione  and  W'ilhanis,  ].  D.  199S. 
Common  and  Scientific  Names  of  Aquatic  In\ertel)rates 


troni  the  United  States  and  Canada:  Molhi.sks.  2nd.  Edi- 
tion. American  Fisheries  Society  Special  P\iblication,  Be- 
thesda,  509  pp. 
Williams,  J.  D.,  M.  L.  Warren  Jr,  K.  S.  Cummings,  J.  L.  Harris 
and  R.  J.  Neves.  1993.  Consen'ation  status  of  freshwater 
nnissels  of  the  United  States  and  Canada.  Fisheries  18(9): 
6-22. 


In  Memoriam 
Ruth  D.  Turner     Rudolf  Stohler 
1914-2()()()  1901-2()()() 


THE  NAUTILUS  114(2):84,  2000 


Page  84 


Notices 


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Thank  vou  for  your  support  and  understancfing  as  we  institute  these  changes  in  the  subscription  rate.  We  look  forward 
to  continuing  to  provide  you  with  cutting-edge  information  on  cfiverse  aspects  of  the  systematics,  biology,  and  ecology 
of  mollusks. 

Sincerely, 

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Editor 


V 


Florida  Department  of  State 

Katherine  Harris 

Secretary  of  State 

Florida  Arts  Council 

Division  of  Cultural  Affairs 

This  program  is  sponsored 

in  part  by  the  State  of  Florida, 

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Division  of  Cultural  Affairs,  and 

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THEt7NAUTILUS 


CONTENTS 


.^    '^^^  Vb/u/JK?  114,  Number  3 

°°'''''°"'-September5,2000 

ISSN  0028-1344 

Phylogeny  of  some  gastropod  mollusks  derived  from  18S 

rDNA  swquences  with  emphasis  on  the  Euthyneura 85 

Size-depth  patterns  in  two  bathyal  turrid  gastropods: 

Benthoinangelia  antonia  (Dall)  and  Oenopota  ovalis 

(Friele) 93 

Four  new  genera  for  northeastern  Pacific  gastropods 99 

ObscuraneUa  papyrodcs,  a  new  genus  and  species  ot 

abyssal  tonnoidean  gastropod  from  Antarctica 103 

Host-tree  selection  by  Florida  tree  snails,  Liguus  fasciatus 
(Miiller,  1774),  in  Big  Cypress  National  Preserve,  Florida, 
USA 112 

The  development  of  three  heterobranch  mollusks  from 

California,  USA 117 

Diminishing  species  richness  of  mollusks  in  Oneida  Lake, 

New  York  State,  USA 120 


Sook  Hee  Yoon 
Won  Kim 


Alisabet  J.  Clain 
Michael  A.  Rex 


James  H.  McLean 


Yuri  L  Kantor 
M.G.  Harasewych 


Robert  E.  Bennetts 
Steven  A.  Sparks 
Deborah  Jansen 


Rachel  Collin 


Willard  N.  Harmann 


THE  NAUTILUS  114(3):84-92,  2000 


Page  85 


Phylogeny  of  some  gastropod  mollusks  derived  from  18S  rDNA 
sequences  wdth  emphasis  on  the  Euthyneura 


Sook  Hee  Yoon 

Department  ot  Biological  Sciences 
College  of  Natural  Sciences 
Sung  Kmui  Kwan  University 
Suwon  440-746,  KOREA 


Won  Kim' 

Department  of  Molecular  Biology 
College  ot  Natural  Sciences 
Seoul  National  Universitv' 
Seoul  151-742.  KOREA' 
W()nkim@plaza.snu.ac.kr 


ABSTRACT 

The  phvlogenetic  relationships  among  gastropod  subgroups, 
with  emphasis  on  the  Euthviieura,  were  investigated  through 
the  analyses  of  nearly  complete  ISS  rDNA  sequences  of  29 
representati\e  gastropods.  Neighbor-joining,  maximum-likeli- 
hood, and  maximum-parsimonN'  methods  were  used  ni  the  con- 
stniction  of  phvlogenetic  trees.  The  18S  rDNA  data  support 
the  monophyK'  of  Vetigastropoda,  the  \etigastropod  clade  Tro- 
choidea,  and  Caenogastropoda.  However,  the  monophylies  of 
two  caenogastropod  subgroups,  Neotaenioglossa  and  Neogas- 
tropoda,  are  not  supported.  The  basal  position  of  Neritopsina 
is  confirmed.  Within  the  Euthyneura,  the  Stvlommatophora 
and  the  Systellommatophora  are  monophyletic,  but  the  Opis- 
thobranchia,  the  Pulnionata,  and  the  Basonnnatophora  are  not. 
The  present  study  supports  the  inclusion  of  Succineidae  within 
Stylommatophora.  However  the  phylogenetic  position  of  Sys- 
tellommatophora within  Gastropoda  remains  unresolved. 

Additional  key  words:  Mollusca,  molecular  phylogeny,  Apo- 
gastropoda,  Caenogastropoda,  Opisthobranchia,  Pulnionata, 
Vetigastropoda,  Stylommatophora,  Basonnnatophora,  Systel- 
lommatophora, Archaeopulmonata. 


INTRODUCTION 

Of  the  molluscan  classes.  Gastropoda  is  the  most  diverse 
and  the  most  ubiquitous  group.  It  has  successfully  adapt- 
ed to  most  habitats,  including  marine,  freshwater,  and 
terrestrial  emdronments. 

Many  comparative  studies  were  published  based  on 
morjDho-anatomical  characters,  including  those  of  shell, 
pallial  complex,  and  nervous,  reproductive,  and  digestive 
systems  (for  recent  reviews,  see  Haszpnmar,  198Sa;  Bie- 
ler,  1992;  Ponder  and  Lindberg,  1997).  However,  the 
status  of  knowledge  of  the  phylogenetic  relationships 
among  and  within  the  gastropod  subgroups  is  still  con- 
troversial (e.g.,  Golikov  and  Starobogatov,  1975;  Gra- 
ham, 1985;  Haszprunar,  1988a;  Bieler,  1992;  Ponder  and 
Lindberg,  1996;  1997).  This  uncertainty  is  largely  due  to 
the  lack  of  informative  morphological  characters  com- 

'  Author  for  correspondence. 


men  to  the  different  taxa  and  the  presence  of  the  high 
level  of  phenotvpic  diversity  observed  in  the  Gastropoda. 

In  atkUtion  to  moiphological  characters,  molecular  se- 
quences have  proven  to  be  very  useful  for  in  phyloge- 
netic reconstructions.  18S  rDNA  sequences  are  amongst 
the  most  informative  molecular  characters  along  a  broad 
range  of  taxa  within  the  Mollusca  (e.g.,  Steiner  and 
Miiller,  1996;  Winnepenninckx  ct  al.,  1996;  Winnepen- 
ninckx  ct  a].,  1998a;  Winnepenninckx  et  al.,  1998b;  Ada- 
mkewicz  ct  al.,  1997;  Bargues  and  Coma,  1997;  Canapa 
ct  al.,  1999)  and  other  animal  phyla.  Several  studies  were 
pubhshed  on  the  molecular  phylogeny  of  Gastropoda 
based  on  the  sequence  data  of  28S  rDNA  (Rosenberg 
ct  al,  1994;  1997;  Tillier  ct  al..  1994;  Tillier  et  al,  1996) 
and  18S  rDNA  (Winnepenninckx  ct  al,  1996;  Winne- 
penninckx ct  al.  1998a;  Harasewych  et  al,  1997a;  b; 
1998).  Winnepenninckx  ct  al  recently  (1998a)  investi- 
gated the  phylogeny  of  gastropod  groups  below  the  class 
rank  using  the  complete  18S  rDNA  sequences  from  18 
gastropod  species. 

To  further  address  gastropod  phylogeny  with  empha- 
sis on  Euthyneura  (Opisthobranchia  +  Pulnionata),  a 
group  that  has  not  been  examined  or  discussed  in  detail 
from  18S  rDNA  data,  we  determined  the  complete  18S 
rDNA  sequences  for  five  representative  gastropods. 
They  include  the  first  complete  sequence  data  from  Ce- 
phalaspidea  (within  Opisthobranchia)  and  Archaeopul- 
monata (within  Pulnionata),  and  additional  sequence 
data  from  Vetigastropoda  and  Stylommatophora  (this  lat- 
ter within  Pulmonata).  These  sequences  were  analyzed 
in  conjunction  with  previously  published  sequences  of 
24  other  gastropotls.  We  focus  on  testing  the  monophyly 
of  each  of  the  euthvneuran  subgroups,  the  Opisthobran- 
chia (e.g.,  Boettger,'  1955;  Ghisehn,  1965;  Gosfiner,  1981; 
1985;  1991;  Goshner  and  Ghisehn,  1984;  Pouhcek  et  al, 
1991;  Ponder  and  Lindberg,  1997),  the  Basommatopho- 
ra  (e.g.,  Tilher,  1984;  Haszpnmar  and  Huber,  1990; 
Nordsieck,  1992),  the  Stylommatophora  (e.g.,  Nord- 
sieck,  1992),  and  the  Systellommatophora  (Salvini-Plaw- 
en,  1980;  Chnio,  1980;  Tilher,  1984;  Haszpnmar  and 
Huber,  1990;  Nordsieck,  1992).  We  also  discuss  the  phy- 


Page  86 


THE  NAUTILUS,  Vol.  114,  No.  3 


logenetic  position  of  the  Succineidae  in  the  Stylomma- 
tophora  (Rigby,  196.5;  Solem,  1978;  Tillier,  1989;  Nord- 
sieck,  1992).  In  addition,  we  examine  the  monophylies 
of  the  Vetigastropoda  (Salvini-Plawen,  1980;  Salvini- 
Plawen  and  Haszprunar,  1987;  Haszprunar,  1988a;  b; 
Ponder  and  Lindberg,  1996)  and  the  vetigastropod  clade 
Trochoidea  (Haszprunar,  1988a). 

MATERIALS  AND  METHODS 

Specimens  An,\lyzed 

The  188  rDNA  sequences  of  two  vetigastropods  (Nor- 
dotis  disciis,  1858  base  pairs,  from  Cheju  Island  and  Ba- 
tilhis  comutus,  1859  base  pairs,  from  Mara  Island),  one 
opisthobranch  {Bidlacta  cxarata,  1849  base  pairs,  from 
Inchon),  and  two  pulmonates  {EUobium  chinensis,  1845 
base  pairs,  from  Tamjin  River,  and  Acusta  dcspccta  sie- 
boldiana,  1847  base  pairs,  from  the  Campus  of  Seoul 
National  University).  The  material  above  was  collected 
in  Korea,  and  their  sequences  are  described  for  the  first 
time  in  the  present  study.  The  sequences  of  two  neo- 
gastropods  (Rapana  venosa  and  Rcishia  bronni),  one 
pulmonate  (Antliosiphonaria  .siriti.s),  and  one  chiton 
(Lcpidozona  (Lcpidozona)  corcanica)  were  reported  in 
our  previous  study  (Yoon  et  al,  1996)  and  the  sequences 
of  the  remaining  21  other  gastropods  and  2  bivalves 
were  obtained  from  GenBank. 

The  nearly  complete  18S  rDNA  sequences  were  an- 
alyzed for  the  29  representative  gastropods  (one  neri- 
toid,  three  vetigastropods,  nine  caenogastropods,  two 
opisthobranchs,  and  14  pulmonates).  two  bivalves,  and 
one  chiton  species.  Of  these,  the  poK-placophoran  Lcp- 
idozona {Lcpidozona)  corcanica  was  used  as  an  out- 
group,  as  the  class  Polyplacophora  (included  in  the  Acu- 
Ufera)  is  currently  accepted  as  the  stem  group  of  the 
classes  Gastropoda  and  Bivalvia  (included  in  Conchi- 
fera),  from  studies  based  on  morphological  characters 
(see  Salvini-Plawen,  1980;  1990;  Haas,  1981;  Runnegar 
and  Pojeta,  1985;  Brusca  and  Brusca,  1990;  Ponder  and 
Lindberg,  1996)  as  well  as  molecular  results  (Adamke- 
wicz  et  al,  1997;  Bargues  and  Mas-Coma,  1997;  Hara- 
sewych  et  al,  1997b).  Table  1  Usts  the  studied  taxa  and 
GenBank  accession  numbers  for  the  respective  sequenc- 
es. The  baseUne  classification  used  in  this  work  follows 
Haszprunar  (1988a)  for  Streptoneura,  Vaught  (1989)  for 
Opisthobranchia,  and  Boss  (1982)  for  Pulmonata. 

DNA  Extraction,  PCR  Amplification,  and 
Sequencing 

Total  nucleic  acids  were  extracted  from  foot  muscle  of 
hve-collected  and  ethanol-preserved  snails  bv  modifica- 
tions of  standard  procedure  of  Sambrook  et  al  (1989). 
The  18S  rDNAs  were  amphfied  using  the  polymerase 
chain  reaction  (PCR)  with  two  oUgonucleotide  primers 
corresponding  to  conserved  sequences  proximal  to  5' 
and  3'  termini  of  metazoans  (Nelles  ct  al,  1984:  1-19, 
5'-CCTGGTTGATCCTGCCAG-3';  1848-1868,  5'- 
TAATGATCCTTCCGCAGGTTA-3':  the  numbers  cor- 


Table  1.   Gastropod  species  used  in  the  present  study,  with 
GenBank  accession  nvuiihers  for  sequences. 


NERITOPSINA 
NERITOIDEA 
Neritidae 

Nerita  albicilla  X91971 
VETIGASTROPODA 
HALIOTOIDEA 
Haliotidae 
'Nordotis  disais 

AF082177 
TROCHOIDEA 
Trochidae 

Monodoiita  labia  X94271 
Turhinidae 
'Btitillus  comutus 
AF  16.5.3 11 
CAENOGASTROPODA 
NEOTAENIOGLOSSA 
LITTORINOIDEA 
Littorinidae 
Littorina  littorea 

X91970 
Nodilittorina  punctata 

Y11755 
CALYPTRAEOIDEA 
Calvptraeidae 
Crepidula  adunca 

X94277 
TONNOIDEA 
Bursidae 

Bursa  rana  X94269 
NEOGASTROPODA 
MURICOIDEA 
Muricidae 

Reishia  bronni  X98827 
Rapana  venosa  X98826 
Buccinidae 

Pisania  striata  X94272 
Nassariidae 
Nassarius  singuijorensis 

X94273 
Fasciolariidae 
Fasciolaria  lignaria 
X94275 
EUTHYNEURA 
OPISTHOBRANCHIA 
CEPHALASPIDEA 
PHILINOIDEA 
Haniineidae 
'BuUacta  exarta 
AF  18867.5 


APLYSIOMORPHA 
APLYSIOIDEA 

Aplysiidae 
Aphisia  sp.  X94268 
PULMONATA 

ARCHAEOPULMONATA 
ELLOBIOIDEA 
Ellobiidae 
'EUobium  chinensis 

AF190452 
BASOMMATOPHORA 
SIPHONARIOIDEA 
Siphonariidae 
Anthosiphonaria  sirius 

X98S28 
Siplionaria  algesirae 

X91973 
LYMNAEOIDEA 
Lviunaeidae 

Ltjmnaea  glabra  Z739S2 
Bakerihpnnaea  ctibensis 

Z83831 
STYLOMMATOPHORA 
MEASURETHRA 
CLAUSILIOIDEA 
Clausihidae 

Balea  biplicata  X94278 
HETERURETHRA 
Succineidae 
Oxijloma  sp.  X94276 
Omalomjx  matheroni 

AF047199 
Athoracophoridae 
Athoracophonis  bitenta- 

culatus  AF047198 
SIGMURETHRA 
ACHATINOIDEA 
Achatinidae 
Limicolaria  kambcxd 

X66374 
HELICOIDEA 
Bradvbaenidae 
'Acusta  despecta  sieboldi- 

ana  AF190453 
Hehcidae 

Helix  aspcrsa  X91976 
SYSTELLOMMATOPHO- 

RA 
ONCHIDIIOIDEA 
Onchidiidae 
Oncliidella  celtica 

X70211 
VERONICELLOIDEA 

X'eroniceUidae 

Laevicaulis  alte  X94270 


Note:  Classification  follows  Haszimniar  (1988a)  for  Strepto- 
neura, Vaught  (1989)  for  Opisthobranchia,  and  Boss  (1982)  for 
Pulmonata.  "  New  sequences  marked  with  asterisk. 


S.  H.  Yoon  and  W.  Kim,  2()()() 


Page  87 


respond  to  positions  of  liunian  ISS  rDNA).  PCR  ampli- 
fications were  performed  with  Taij  DNA  pol\merase  for 
30  cycles  (94°  C  for  1  min,  52°  C  for  2  min,  and  72°  C 
for  3  min).  The  ends  of  the  amplified  DNA  fragments 
were  modified  for  blunt-ended  ligation  using  T4  kinase 
and  T4  polymerase.  The  hlunt-endeil  18S  rDNAs  were 
hgated  into  pGEM-3zf(-)  plasmid  vector  and  trans- 
formed into  DH5-a  cell  fines.  Sequencing  primers  used 
in  this  study  were  reported  in  a  previous  paper  (Moon 
ct  «/.,  1996).  ISS  rRNA-coding  regions  were  completely 
sequenced  in  both  directions  with  complete  overlap.  The 
DNA  sequencing  was  performed  by  the  dideoxynucleo- 
tide  ch;iin-termination  method  (Sanger  ct  al.,  1977)  us- 
ing a  Taq-Track  kit  (Promega  Co.),  accorcfing  to  the 
manufacturer's  instructions.  Eli'ctrophoresis  of  sequenc- 
ing reaction  mixtures  was  periormed  on  butler-gradient 
6%  polyacrylamide  gels  and  examined  by  autoradiogra- 
phy. 

Piivi,ck;enetic  Analysis  of  18S  rDNA  Sequences 

The  sequences  were  initially  aligned  with  the  CLUSTAL 
W  multiple-aUgnment  program  (Thompson  ct  al.,  1994) 
and  the  alignment  refined  manualK'.  A  data-set  of  align- 
ment-stable positions  was  produced  by  excluding  those 
positions  that  differed  between  alignments  (Gatesy  cf  aZ., 
1993).  Analyses  were  limited  to  refiablv  aligned  regions, 
which  included  a  total  of  1754  nucleotide  positions.  Phv- 
logenetic  reconstructions  were  performed  using  the 
neighbor-joining  (NJ),  maximum-likelihood  (ML),  and 
maximum-parsimony  (MP)  methods.  PHYLIP  version 
3.572c  (Felsenstein,  1995)  was  used  for  the  neighbor- 
joining  (Saitou  and  Nei,  1987)  analyses.  The  distance 
analyses  were  done  using  Kimura  (1980)  and  Jukes  and 
Cantor  (1969)  matrices  as  input  for  the  neighbor-joining 
analyses.  Maximum-likelihood  analvses  were  performed 
using  the  HKY  (Hasegawa  ct  al.  1985)  model  in  PAUP 
4.0b2  (Swofford,  1999).  For  the  quartet  puzzling  meth- 
od (the  number  of  puzzling  steps  is  1000),  empirical  nu- 
cleotide frequencies,  and  transition/transversion  ratio  of 
1.5  were  estimated.  Parsimony  analvses  were  also  per- 
formed using  the  computer  program  PAUP  version 
4.0b2  with  closest  stepwise  addition  options.  The  anal- 
yses employed  a  heuristic  search  using  TBR  branch 
swapping  with  random  taxon  addition.  Branch  length 
was  optimized  according  to  the  ACCTRAN  option. 
Bootstrap  analyses  (Felsenstein,  1985)  of  one  hundred 
rephcates  were  performed  to  examine  the  confidence  of 
nodes  in  NJ,  ML,  and  MP  analyses. 

RESULTS 

Figure  lA  shows  the  phylogenetic  tree  resulting  from 
the  neighbor-joining  (NJ)  analysis  using  the  Kimura 
(1980)  distances  of  an  alignment  of  complete  18S  rDNA 
sequences  of  29  gastropod  species.  The  polyplacophoran 
Lepidozona  (Lcpidozoua)  corcanica  was  used  as  out- 
group.  The  same  tree  topology  was  also  obtained  using 
Jukes    and    Cantor    (1969)    distances.    The    Neritoidea 


branches  off  first  and  the  Vetigastropoda  (Trochoidea  -I- 
Haliotoidea)  diverges  next  as  an  independent  clade  be- 
fore the  clade  Apogastropoda  (Caenogastropoda  +  Eu- 
thvTieura).  The  monophyly  of  the  Vetigastropoda  and  its 
subclade  the  Trochoidea  (represented  by  Monodonta 
and  BatiUus)  is  clearly  shovvni  in  the  tree,  with  ven'  high 
boostrap  support  (100%:  100%).  The  Caenogastropoda 
shows  a  sister  group  relationship  with  the  Euthyneura 
with  very  high  bootstrap  support  (94%).  Monophyly  of 
the  Caenogastropoda  is  supported  (boostrap  value  = 
100%),  though  neither  the  Neotaenioglossa  (=  Meso- 
gastropoda)  nor  the  Neogastropoda  emerged  as  mono- 
phyletic  clades. 

There  is  strong  bootstrap  support  (100%)  for  the 
monophvlv  of  Euthviieura  (Opisthobranchia  +  Pulmon- 
ata),  though  the  monophyhes  of  Opisthobranchia  (Ce- 
phalaspidea  +  Anaspioidea)  and  Pulmonata  are  not  sup- 
ported. The  Basommatophora,  consisting  of  the  Siphon- 
arioidea  and  Lymnaeoidea,  did  not  emerge  as  a  mono- 
phyletic  group.  On  the  other  hand,  there  is  good  support 
for  the  monoph)'fies  of  hvo  additional  groups  in  the  Eu- 
thyneura, the  Stylommatophora  (boostrap  value  =  84%) 
and  the  Systellommatophora  (boostrap  value  =  87%). 
Bootstrap  values  strongly  support  the  position  of  the 
Succineidae  (Oxi/loma  and  Omalonijx)  within  the  sty- 
lommatophoran  clade.  However,  the  position  of  the  Sys- 
tellommatophora within  the  Gastropoda  was  not  posi- 
tively determined  in  the  present  study. 

The  resulting  tree  from  maximum-fikefihood  (ML) 
analyses  of  the  same  data  set  is  shown  in  figure  IB.  The 
ML  tree  confirms  all  the  major  results  of  the  NJ  tree 
(figure  lA),  with  the  exception  that  clades  within  Eu- 
thyneura lack  significant  bootstrap-support.  The  new 
branching  order  of  Systellommatophora  {Onchidclla  and 
Laevicaulis) ,  Aplysiomoipha  (Aplijsia),  Archaeopulmon- 
ata  (E//oi>((/)7()-Siphonarioidea  {Siphonaria  and  Antho- 
siphonaiia),  Cephalaspidea  (BuUacta).  and  Stvlomma- 
tophora  in  the  clade  chffers  from  the  order  shouni  in  the 
NJ  tree  (figure  lA),  Aplysiomoipha-Svstellommatopho- 
ra-Siphonarioidea  and  Archaeopulmonata-Cephalaspi- 
dea-Stylommatophora. 

The  maximum  parsimony  (MP)  analyses  produced  a 
single  tree  with  minimum  length  of  950  steps  (figure 
IC).  Generally  speaking,  MP  analyses  also  yielded  sim- 
ilar results  except  for  minor  differences  in  topologies 
among  groups  within  the  caenogastropod  and  the  euthv- 
neuran  clades.  Maximum  parsimony  shows  topological 
shifts  within  the  caenogastropod  clade,  e.g.,  the  new 
branching  order  of  Bursa,  Na.ssariiis,  and  Pisania-Fas- 
ciolaiia-Crcpidiila  instead  of  the  order  shown  in  the  NJ 
tree  (figure  lA),  Crcpidida,  Bursa,  Pisania,  Nassarius, 
and  Fasciolaria.  Within  the  Euthviieura,  the  MP  tree 
differs  from  the  NJ  tree  only  in  those  branching  points 
\vith  low  bootstrap  values.  There  is  no  sister-group  re- 
lationship among  the  Cephala.spidea  [BuUacta).  Aply- 
siomorpha  {Aphjsia),  Archaeopulmonata  (Ellohium),  Si- 
phonarioidea,  Stvlommatophora,  and  Systellommatopho- 
ra. 

Ne.xt,   we   focused  on   the   Euthvneura  in   separate. 


Page  88 


THE  NAUTILUS,  Vol.  114,  No.  3 


KHlTl 


Omalonyx 

Athoracophorus 


Oxyloma 

r^Limicolaria 
^  Helix 
Acusta 

Bullacta 

— Ellobium 
ZT  Anthosiphonaria 
Siphonaria 
'gj~  Onchidelta 
•evicaulis 
Aplysia 
100  t—tymnea 

Bakerilymnea 
Fasciolaria 
Nassarius 
Pisania 
Bursa 
Crepidula 
Rapana 
■j^Reishia 
{r^Litlorina 
— Nodilittorina 
I 
100 


Siylommatophora 


Pulmonata 


Cephalaspidea  Opisthobranchia 

Archaeopulmonata 

J  Basommatophora 


J  Systellommalophora 

Aplysiomorpha 
J  Basommatophora 

Neogastropoda 
J  Neotaenioglossa 
J  Neogastropoda 
J  Neotaenioglossa 


Nerita 
[r~  Placopeclen 
Chlamys 
Lepidozona 


^Balii 

' fl 


Monodonta 
Batillus 

Nordotis 


Pulmonata 

Opisthobranchia 
Pulmonata 


Caenogastropoda 


Euth  /neura 


Vetigasiropoda 

Nentopsina 
J  Bivalvia 
Polyplacophora 


Apogastropoda 


B 


11  I An 


^'  I —  LimUolaria 

Helix 

Acusta 

Balea 

Omalonyx 

Athoracophorus 

Oxyloma 

Bullacta 

Antbosiphonaria 

Siphonaria 

Ellobium 

Aplysia 

Hj —  Onchidella 

—  iMevicaulis 
J       Lymnaea 

—  Bakerilymnea 

Fasciolaria 

Nassarius 

Pisania 

Bursa 

Crepidula 

U       Rapana 

—  Reishia 
Lj       Littorina 

—  Nodilittorina 
Monodonta 
Batillus 
Nordotis 
Nerita 


91 


Omalonyx 

Athoracophorus 

Oxyloma 

Limicolaria 

Helix 

Balea 

Acusta 

A  nrhosiphonaria 

Siphonaria 

Onchidella 

iMevicaulis 

Bullae  Ui 

Aplysia 

Ellobium 

Lymnaea 

Bakerilymnea 

Crepidula 

Fasciolaria 

Pisania 

Nassarius 

Bursa 

Rapana 

Reishia 

Littorina 

Nodilittorina 

Monodonta 

Batillus 

Nordotis 

Nerita 

Placopecten 

Chlamys 

Lepidozona 

Figure  1.  A.  Neighbor-joining  tree  detemiined  by  an  alignment  ol  29  nearly  complete  18S  rDNA  sequence  data  for  gastropods 
witli  Lepidozona  coreanica  (Polvplacophora)  as  outgroup.  Bootstrap  percentages  are  shown  abo\'e  branches  supported  in  at  least 
50%  of  100  replicates.  B.  Strict  consensus  tree  resulting  from  maximum-likelihood  analyses  of  :29  nearly  complete  gastropod  18S 
rDNA  sequences.  Quartet  puzzling  method  and  HKY  (Hasegawa  et  al,  1985)  setting  model  were  used.  Lepidozona  coreanica 
(Polyplacophora)  was  the  outgroup.  Bootstrap  analysis  was  performed  with  100  rephcates;  values  above  50%  are  indicated  above 
the  nodes.  C.  Strict  consensus  tree  of  maximum  parsimony  analyses  based  on  the  333  informative  sites  of  an  alignment  of  29  nearly 
complete  gastropod  18S  rDNA  sequences  (length  =  950;  CI  =  0.6611;  RI  =  0.8418).  Lepidozona  coreanica  (PoKplacophora)  was 
die  outgroup.  Bootstrap  values  higher  than  50%  are  indicated  above  the  nodes. 


IIJU| 


LI       Placopecten 

^~  Chlamys 
Lepidozona 


— 1 

c 

(.} 

551 

1 

_   85  1 

6i 

1 
"1 

1 

100  1 

85 

1 
54 

74 

57 

97 

<» 

"l 

1 
81, 

100 

100 

1 

100  1 

1 

mainly  because  the  use  of  outgroups  that  are  too  far 
removed  from  the  elade  in  studv  mav  give  origin  to  ad- 
ditional homoplasies  between  ingroup  and  outgroup  spe- 
cies. Figure  2A  shows  the  results  of  NJ  analyses  of  16 
nearly  complete  euthyneuran  18S  rDNA  sequences, 
with  the  caenogastropod  Littorina  littorca  as  outgroup. 


The  resulting  tree  exhibits  the  same  topology  as  the 
complete  NJ  tree  (figure  lA),  with  the  exception  that 
Ellohiitm  (Archaeopulmonata)  becomes  the  sister  group 
to  the  clade  Aplysiomoq^ha  (Aphfsia)  +  Systellomma- 
tophora-Siphonarioidea,  instead  of  clade  Cephalaspidea 
(Bullacta)  +  Stvlommatophora,  as  present  in  the  com- 


S.  H.  Yoon  and  W.  Kim,  2000 


Page  89 


0.01 


~NodUittorina 
~Littorina 


3ol 


84 


88  pOma/on^ijr 

'"*' ' Alhoraco^horus 

Oxytoma 
'    Balea 

_  Limicolaria 

so" 


Helix 
■Icuito 
Bullacta 
%  [~Antbosiphonaria 
Siphonaria 
Oncbideila 
icaulis 
—Aplysia 
'Ellobium 
^PQ[     Lymnaea 
Bakerilymnea 


hrA 

I Ont/i 

taeii 


Styloninialophora 


Cephalaspidea  Opisthobranchia 

Basomniatophora 

Systellommalophora 

Aplysiomorpha  Opisthobranchia 

Archaeopulmonala  - 


Basomniatophora 


Pulmonata 


Ncotacnioglossa 


Limicolaria 

Helix 

Acusta 

Balea 

Omalonyx 

Arhoracophorus 

Oxyloma 

Bullacta 

A  nthosiphonaria 

Siphonaria 

Aplysia 

Onchidella 

Laevicaulis 

Ellobium 

Lymnea 

Bakerilymnea 

Nodilittorina 

Littorina 


"  Omalonyx 

'  Athoracophorus 

'  Oxyloma 

"  Limicolaria 

'  Helix 

'  Balea 

~  Acusta 

'  Anthosiphonaria 

~  Siphonaria 

~  Lymnaea 

~  Bakerilymnea 

~  Onchidella 

~  Laevicaulis 

~  Aplysia 

~  Ellobium 

-  Bullacta 

~  Nodilittorina 

-  Littorina 


Figure  2.  A.  Euthviieuran  neighbor-joining  tree  detennined  by  an  alignment  of  16  nearly  complete  euth>Tieuran  18S  rDNA 
sequences,  using  Littorina  littorea  (Caenogastropoda:  Littorinidae)  as  outgroup.  Numbers  at  a  node  indicate  bootstrap  \alues  higher 
than  50%.  B.  Euthyneuran  maximum-likelihood  tree  detennined  by  an  ahgnment  of  16  nearly  complete  euthyneuran  18S  rDNA 
sequences,  using  Littorina  littorea  (Caenogastropoda:  Littorinidae)  as  outgroup.  Quartet  puzzling  method  and  HKY  (Hasegawapf 
ai.  1985)  setting  model  were  used.  Bootstrap  values  above  50%  are  indicated  above  the  nodes.  C.  Euthyneuran  maximum  parsimony 
tree  calculated  from  the  149  infomiative  sites  of  an  alignment  of  16  nearly  complete  euthyneuran  18S  rDNA  sequences,  using 
Littorina  littorea  (Caenogastropoda:  Littorinidae)  as  outgroup  (length  =  415;  CI  =  0.7494;  Rl  =  0.6750).  Only  bootstrap  values 
higher  than  50%  are  indicated. 


plete  NJ  tree.  These  two  clades  are  not  supported  by 
bootstrap  analysis  of  the  euthyneuran  clade.  When  the 
euthyneuran  ML  tree  (figure  2B)  is  compared  to  the 
entire  ML  tree  (figure  IB),  small  topological  shifts  are 
discernible.  The  Siphonarioidea  shows  a  sister  group  re- 
lationship with  Aplysiomorpha  (Aplysia)  instead  of  with 
Archaeopulmonata  {Ellobium),  and  this  Siphonarioidea- 
Aplysiomorpha  cluster  appears  as  sister  group  of  the  Sys- 
tellommatophora  instead  of  Cephalaspidea-Stylomma- 
tophora.  However,  bootstrap  values  that  support  these 


nodes  are  very  low.  Nevertheless,  the  euthyneuran  ML 
tree  strongly  supported  most  of  the  major  nodes  found 
in  the  entire  ML  tree,  with  generally  higher  bootstrap 
values.  Maximum  parsimony  analyses  based  on  the  149 
phylogenetically  informative  characters  of  the  alignment 
of  16  euthyneuran  species  produced  a  single  tree  with 
minimum  length  of  415  steps  (figure  2C).  In  the  euthy- 
neuran MP  tree,  the  first  branching  member  is  Cepha- 
laspidea (Bullacta)  rather  than  Lymnaeoidea  (Basom- 
matophora)  found  in  the  entire  MP  tree.  The  topologies 


Page  90 


THE  NAUTILUS,  Vol.  114,  No.  3 


for  the  remaining  taxa  are  generally  identical  to  the  ones 
in  the  tree  containing  all  taxa.  The  euthyneuran  trees 
resulting  from  NJ,  ML,  and  MP  analyses  confirmed  the 
topology  of  trees  generated  from  the  same  types  of  anal- 
yses but  based  on  all  ta.xa. 

DISCUSSION 

For  the  phylogenetic  relationships  among  and  within  the 
gastropod  major  subgroups,  our  study  supports  several 
aspects  of  the  studv  bv  Winnepennickx  ct  al.  (1998a). 
The  Neritoidea  diverged  first,  followed  by  the  Vetigas- 
tropoda.  Next,  the  clade  Apogastropoda,  comprising  the 
Caenogastropoda  and  Euthyneura  (each  well  supported 
as  monophyletic  groups),  appears  as  a  monophyletic 
group.  However,  the  Neotaenioglossa  (=  Mesogastro- 
poda)  and  the  Neogastropoda  (included  in  Caenogastro- 
poda) and  the  Pulmonata  (included  in  Euthyneura)  do 
not  appear  as  monophyletic  groups.  In  addition,  the  po- 
sition of  Systellommatophora  within  the  Gastropoda 
could  not  be  defined  in  the  present  study. 

In  contrast  to  previous  reports,  our  study  provides 
more  details  on  gastropod  phylogeny,  especially  with  re- 
gard to  the  phylogenetic  status  of  the  subgroups  within 
the  Euthyneura.  All  trees  resulting  from  NJ,  ML,  and 
MP  analyses  refute  the  monophyly  of  the  Opisthobran- 
chia.  This  result  supports  the  claims  of  many  opistho- 
branch  workers,  whom  observed  that  parallelism  and 
convergence  ha\'e  occured  in  most  major  organ  systems 
within  the  opisthobranchs  (Ghiselin,  1965;  Gosliner, 
19S1;  1985;  1991;  Gosliner  and  Ghiselin,  1984;  Pouhcek 
et  al,  1991)  and  that  the  high  degree  of  homoplasy  in 
many  characters  in  opisthobranchs  contribute  to  diffi- 
culties in  obtaining  robust  results  from  cladistic  analyses 
(Goshner  and  Ghiselin,  1984;  GosHner,  1985;  1991). 
Boettger  (1955)  maintained  that  the  Opisthobranchia  is 
paraphyletic.  Ponder  and  Lindberg  (1997)  also  suggest- 
ed that  the  Opisthobranchia  is  not  monophyletic,  a  \dew- 
point  in  agreement  with  the  results  of  TiUier  ct  al.  (1994) 
from  28S  rDNA  data.  It  is  not  surprising,  therefore,  that 
many  contradictory  phylogenies  and  classification 
schemes  have  been  suggested  for  the  Opisthobranchia 
(e.g.,  Boettger,  19.55;  Taylor  and  Sohl,  1962;  Ghisefin, 
1965). 

In  relation  to  the  phylogeny  of  the  Pulmonata,  we 
have  mentioned  that  the  Basommatophora  {scmtt  Ha- 
szprunar  and  Huber,  1990,  Siphonarioidea  -I-  Lymnaeo- 
idea),  is  not  monophyletic.  Tillier  (1984)  considered  that 
only  Lyinnaeoidea  belongs  to  Basommatophora  and  Si- 
phonarioidea to  Archaeopulmonata.  He  suggested  that 
pulmonates  radiated  into  freshwater  habitats  as  Basom- 
matophora and  into  marine  habitats  as  Archaeopulmon- 
ata. Tillier  et  al.  (1996),  based  on  28S  rDNA  data, 
showed  that  the  Basommatophora  is  not  monophyletic. 
However,  Haszprunar  and  Huber  (1990)  suggested  that 
both  Siphonarioidea  and  Lymnaeoidea  could  be  allocat- 
ed in  Basommatophora  due  to  the  presence  of  common 
morphological  characters  such  as  a  procerebrum  com- 
prising only  large  cells,  the  lack  of  a  contractile  pneu- 


mostome,  and  the  presence  of  an  osphradium  and  pallial 
ciliary  tracts.  Nordsieck  (1992)  based  on  the  presence  of 
an  anal  opening  shifted  to  the  posterior  mantle  lobe,  also 
considered  the  marine  Thalassophila  (Siphonarioidea)  to 
be  the  sister  group  of  the  limnic  Hygrophila  (Lymnaeo- 
idea). Therefore,  the  common  morphological  characters 
found  in  the  Basommatophoran,  rather  than  represent- 
ing synapomoiphies,  could  all  have  been  derived  by  con- 
vergence. Nordsieck  (1992),  based  on  morphological 
characters  found  in  tentacles,  kidney,  central  nervous 
system,  and  aspects  of  ontogeny,  considered  the  SKlom- 
matophora  to  be  a  monophyletic  group,  which  is  con- 
cordance with  the  results  derived  from  28S  rDNA  se- 
quence data  (Rosenberg  et  al,  1994;  1997;  TilUer  ct  al, 
1994;  Tillier  ct  al,  1996).  Although  there  is  instability  of 
branching  pattern,  the  results  for  the  euthvnieuran  clade 
in  the  present  study  indicate  that  the  Succineidae  (rep- 
resented by  Oxijloma  and  Omalomfx)  belongs  to  the  Sty- 
lommatophora  (e.g.,  Solem,  1978;  Tillier,  1989;  Nord- 
sieck, 1992).  These  results,  however,  contradict  the  view 
of  Rigby  (1965)  whom,  on  the  basis  ol  moq^hological 
similarities  of  the  digestive  and  reproductive  systems,  al- 
located the  Succineidae  in  the  Opisthobranchia.  Syna- 
pomorphic  characters  such  as  the  more  or  less  reduced 
shell  and  mantle,  and  a  visceral  ganglion  situated  cen- 
trally or  on  the  left  side  with  respect  to  the  axis  of  the 
central  nervous  system  (e.g.,  Salvini-Plawen,  1980; 
Nordsieck,  1992)  support  the  monophvlv  of  the  Systel- 
lommatophora (represented  by  Onchidudae  and  Veron- 
icellidae).  However,  this  monophyly  has  been  questioned 
by  several  authors  (e.g.,  Chmo,  1980;  Tilfier,  1984;  Ha- 
szprunar and  Huber,  1990).  Climo  (1980)  in  particular 
considered  the  systellommatophorans  as  a  polyphyletic 
assemblage  diverging  at  the  base  of  the  euthvmeurans. 
Tillier  (1984)  studied  the  morpho-anatomical  characters 
of  the  pallial  complex,  digestive  tract,  reproductive,  and 
central  nervous  systems,  and  divided  the  Pulmonata  into 
only  three  orders;  Archaeopulmonata,  Basommatophora, 
and  Stylommatophora.  He  included  the  Systellommato- 
phora in  the  Archaeopulmonata  and  suggested  that  the 
Onchidudae  (within  Systellommatophora)  is  more  close- 
ly related  to  the  Ellobiidae  (within  Archaeopulmonata) 
than  to  the  VeroniceUidae  (within  Systellonunatophora). 
However,  our  present  result  supports  the  monophyly  of 
Systellommatophora. 

Since  the  erection  of  the  Vetigastropoda  by  Salvini- 
Plawen  (1980),  the  presence  of  .synapomorphic  charac- 
ters such  as  ctenidial  sense  organs,  the  epipodial  sense 
organs,  and  the  special  structure  of  the  esophagus  have 
generally  supported  the  monophyly  of  this  clade  (Sal\'ini- 
Plawen  and  Haszprunar,  1987;  Haszpnmar,  1988a;  b; 
Ponder  and  Lindberg,  1996;  1997).  Other  vetigastropod 
features  include  the  dominant  presence  of  the  right  dor- 
soventral  retractor  muscle,  the  right  excretory  organ,  and 
bilamellate  ctenidia  with  skeletal  rods.  Previous  molec- 
ular data  using  partial  18S  rDNA  (Harasewvch  ct  al, 
1997a;  b)  and'  the  28S  rDNA  (Tillier  ct  a/.,' 1994)  se- 
quences also  supported  the  monophyly  of  the  Vetigas- 
tropoda, which  is  also  supported  in  the  present  study. 


S.  H.  Yoon  and  W.  Kim.  2000 


Page  91 


Moiioplnlv  of  the  Trochoidea  (represented  by  Mono- 
chmta  and  Batillus)  is  also  confirmed  herein.  The  Tro- 
choidea is  defined  hv  SNiiapomoqihies  such  as  loss  of  the 
right  ctenuhum  in  relation  to  the  loss  of  the  shell  slit 
(Haszpnniar,  IBSSa;  h)  and  the  nionophvK'  ol  the  gronp 
is  also  in  concordance  \\'ith  the  stud)-  based  on  28S 
rDNA  sequences  by  TilUer  et  al.  (1994). 

In  conchision,  the  ISS  rDNA  data  stronglv  support 
the  monoph\l\  of  the  following  higher  gastropod  clades: 
Vetigastropoda,  Trochoidea  (within  Vetigastropoda), 
Apogastropoda,  and  the  two  included  clades  Caenogas- 
tropotla  and  Euth\meura.  Within  the  euthyneuran  clade, 
both  the  Stslommatophora  and  the  Systellommatophora 
are  monophyletic.  However,  our  18S  rDNA  data  failed 
to  support  monophyly  of  the  Neotaenioglossa  and  the 
Neogastropoda  (within  Caenogastropoda),  Opisthobran- 
chia,  Pulmonata,  and  the  puhnoTiate  Basommatophora. 
These  non-monophvletic  subgroups,  therefore,  at  pre- 
sent can  be  considered  as  grades  rather  than  clades.  The 
basal  position  of  Neritopsina  is  confirmed  in  this  study. 
In  addition,  the  Succineidae  is  included  in  Stylomma- 
tophora.  Still,  the  Svstellommatophoran  position  within 
the  Gastropoda,  that  is,  its  immediate  relationship  to  ei- 
ther Opisthobranchia  or  Pulmonata,  or  to  any  other 
group  for  that  matter,  could  not  be  defined.  The  insta- 
bility of  topology  and  short  branch  lengths  within  the 
Caenogastropoda  and  the  Euthyneura  may  be  due  to  the 
fact  that  the  mollusks,  including  gastropods,  apparently 
rachated  in  an  "explosive"  fashion  during  a  relatively 
short  period  of  time.  Most  extant  major  groups  of  mol- 
lusks appeared  around  a  relatively  short  time  at  the  Pre- 
cambrianyCambrian  boundary  (e.g.,  Runnegar  and  Po- 
jeta,  19S5;  Winnepenninckx  et  al,  1996;  Adamkewicz  ct 
al.  1997;  Harasewych  ct  al,  1997a).  Futtire  studies  at- 
tempting to  define  phylogenetic  relationships  at  these 
levels  may  take  into  consideration  other  molecules  such 
as  cytochrome  c  oxidase  I  and/or  16S  rDNA.  Such  mol- 
ecules evolve  more  rapidly  than  18S  rDNA,  and  seem 
more  likelv  to  contain  information  needed  to  solve  phy- 
logenetic relationships  within  these  clades. 

ACKNOWLEDGMENTS 

This  work  was  supported  bv  a  grant  from  KOSEF  (95- 
0401-04-01-3)  for  years  1995-1998.  We  thank  Dr.  B.  L. 
Choe,  Mr.  J.  L.  Lee  (Sung  Kyun  Kwan  University,  Ko- 
rea), Dr.  J.  S.  Lee  (Kang  Won  University,  Korea),  and 
Mr  D.  G.  Min  (Shell  House,  Seoul,  Korea)  for  providing 
specimens.  We  are  also  indebted  to  Dr.  C.  B.  Kim  (Yale 
University,  USA)  and  Dr.  J.  K.  Park  (University  of  Mich- 
igan, USA)  for  providing  many  references.  We  thank  Ms. 
D.  W.  Jung  (Seoul  National  University)  for  proofreading 
the  manuscript.  Special  thanks  are  due  to  Dr.  S.  Y.  Moon 
(University  of  Koln,  Germany)  for  her  assistance  in  soft- 
ware analyses.  We  are  especially  grateful  to  Dr.  S.  Y 
Moon  and  to  Dr.  M.  G.  Harasewych  (Smithsonian  In- 
stitution, Washington,  USA)  for  their  comments  and 
suggestions. 


LITERATURE  CITED 

Adamkewicz,  S.  L,  M.  G.  Harasewvcli,  J.  A.  Bkike,  D.  Saudek, 
and  C.  ].  Bull.  1997.  A  molecular  plnlogeiiy  of  tlie  !)i\alve 
mollusks.  Molecular  Biology  and  Exolution  14:619-(i29. 

Bargues,  M.  D.  and  S.  Mas-Coma.  1997.  Phylogenetic  analysis 
of  limnaeid  snails  ba.sed  on  ISS  rDNA  sequences.  Molec- 
ular Biology  and  Evolution  14:.569-.577. 

Bieler,  R.  1992.  Gastropod  phylogeny  and  systematics.  Aiuuial 
Review  of  Ecologv'  and  Systematics  23:.311-.3.3S. 

Boettger,  C.  R.  19.5.5.  Die  Systematik  der  euthyneineii 
Schnecken.  Verliandlungen  der  Deutschen  Zoologisclien 
Gesellschaft  in  Tiihingen,  Supplement  18:253-280. 

Boss,  K.  |.  1982.  Mollusca.  In:  S.  R  Parker  (ed.)  Syniopsis  and 
classification  of  living  organisms.  McGraw-Hill  Book 
Company  New  York,  pp.  945-1166  (Volume  1);  1092-96 
(Volume  2). 

Bnisca,  R.  C.  and  G.  |,  Brusca.  1990.  Invertebrates.  Sinauer 
Associates,  Sunderland,  922  pp. 

Canapa,  A..  1.  Marota,  F.  Rollo,  and  E.  Olmo.  1999.  The  small- 
subunit  rRNA  gene  sequences  of  venends  and  the  phy- 
logeny of  Bixakia.  Journal  of  Molecular  Evolution  48:46.3- 
468. 

Climo,  F.  M.  1980.  Smeagolida,  a  new  order  of  gyninomorph 
mollusc  from  New  Zealand  based  on  a  new  genus  and 
species.  New  Zealand  Journal  of  Zoology  7:51.3-522. 

Felsenstein,  J.  1985.  Confidence  limits  on  the  phylogeny:  an 
approach  using  the  bootstrap.  Evolution  39:78.3-791. 

Felsenstein.  J.  1995,  PHYLIP;  phylogeny  inference  package, 
version  3.572c.  Deptartment  of  Genetics,  Universitv  of 
Washington,  Seattle. 

Gatesy,  J.,  R.  De  Salle,  and  W.  C.  Wheeler  1993.  Alignment- 
ambiguous  nucleotide  sites  and  the  exclusion  of  systematic 
data.  Molecular  Phylogenetics  and  Evolution  2:151-157. 

Ghiselin,  M.  T.  1965.  Reproducti\'e  fimction  and  the  phylogeny 
of  opisthobranch  gastropods.  Malacologia  3:327-378, 

Golikov.  A.  N.  and  Y.  I.  Starobogatov.  1975.  Systematics  of 
prosobranch  gastropods.  Malacologia  15:185-232. 

Goshner,  T.  M.  1981.  Origins  and  relationships  of  pnmitive 
members  of  the  Opisthobranchia  (Mollusca:  Gastropoda). 
Biological  Journal  of  Linnean  Society  16:197-225. 

Gosliner,  T.  M.  1985.  Parallelism,  parsimony  and  the  testing  of 
phylogenetic  hvpotheses:  the  case  of  opisthobranch  gas- 
tropods. In:  E.  S.  Vrba  (ed.)  Species  and  speciation.  Trans- 
vaal Museum  Monographs  number  4.  Tran.svaal  Musemn. 
Pretoria,  pp.  105-107. 

Gosliner,  T.  M.  1991.  Moipliological  parallelism  in  opistho- 
brancli  gastropods.  Malacologia  32:313-327. 

Gosliner,  T.  M.  and  M.  T.  Ghiselin.  1984.  Parallel  evolution  in 
opisthobranch  gastropods  and  its  implications  for  phylo- 
genetic methodology.  Systematic  Zoology  .33:255-274. 

Graham,  A.  1985.  Evolution  within  the  Gastropoda:  Proso- 
branchia.  In:  E.  R.  Tmeman  and  M.  R.  Clarke  (eds.)  The 
Mollu.sca.  Volume  10.  Academic  Press,  New  York,  pp. 
151-186. 

Harasewych,  M.  G.,  S.  L.  Adamkewicz,  J.  A.  Blake,  D.  Sandek. 
T.  Spriggs,  and  C.  J.  Bult.  1997a.  Neogastropod  phyloge- 
ny: A  molecular  perspective.  Journal  of  Molluscan  Studies 
63:327-351. 

Harasewych,  M.  G.,  S.  L.  Adamkewicz,  J.  A.  Blake,  D.  Saudek, 
T.  Spriggs,  and  C.  J.  Bult.  1997li.  Phylogeny  and  relation- 
ships of  pleurotomariid  gastropods  (Mollusca:  Gastro- 
poda): an  assessment  based  on  partial  18S  rDNA  and  cy- 
tochrome c  oxidase  I  sequences.  Molecular  Marine  Biol- 
ogy and  Biotechnology  6:1-20. 


Page  92 


THE  NAUTILUS,  Vol.  114,  No.  3 


Harasewych,  M.  G.,  S.  L.  Adamkewicz,  M.  Plassmeyer,  and  P. 
M.  Gille\et.  199S.  Phvlogenetic  relationships  of  the  lower 
Caenogastropoda  (Mollusca,  Gastropoda,  Architaenioglos- 
sa,  Campaniloidea,  Cerithioidea)  as  determined  bv  partial 
18S  rDNA  sequences.  Zoologica  Scripta  27:.'361-372. 

Hasegawa,  M.,  H.  Kishino,  and  T.  Yano.  1985.  Dating  of  hu- 
man-ape sphtting  by  a  molecular  clock  of  mitochondrial 
DNA.  Journal  of  Molecular  E\oIution  21:160-174. 

Hass,  W.  1981.  Evolution  of  calcareous  hardparts  in  primitive 
molluscs.  Malacologia  21:40.3-418. 

Haszpnmar,  G.  1988a.  On  the  origin  and  ex'olution  of  major 
gastropod  groups,  with  special  reference  to  the  Strepto- 
neura.  Journal  of  Molluscan  Studies  54:367^141. 

Haszprunar,  G.  1988b.  A  preliminary  phylogenetic  analysis  of 
the  streptoneurous  gastropods.  Malacological  Review. 
Supplement  4:7-16. 

Haszprunar,  G.  and  G.  Huber  1990.  On  the  central  nervous 
system  of  Smeagolidae  and  Rhodopidae,  two  families 
questionably  allied  with  the  Gvnnnomorpha  (Gastropoda: 
Euthviieura).  Journal  of  Zoolog)-  220:18.5-199. 

Jukes.  T.  H.  and  C.  R.  Cantor  1969.  Evolution  of  protein  mol- 
ecules. In:  H.  N.  Munro,  (ed.)  Mammalian  protein  me- 
tabolism. Academic  Press,  New  York.  pp.  21-132. 

Kiniura,  M.  1980.  A  simple  method  for  estimating  evolutionary 
rate  of  base  substitution  through  comparative  studies  of 
nucleotide  sequences.  Journal  of  Molecular  Evolution  16: 
111-120. 

Moon,  S.  Y.,  C.  B.  Kim,  S.  R.  Gelder,  and  W.  Kim.  1996.  Phy- 
logenetic positions  of  the  aberrant  branchiobdelUdans  and 
aphanoneurans  within  the  AnneHda  as  derived  from  18S 
ribosomal  RNA  gene  sequences.  Hydrobiologia  324:229- 
236. 

Nelles,  L.,  B.  L.  Fang,  G.  Volckaert,  A.  Vandenberghe.  and  R. 
De  Wachter  1984.  Nucleotide  sequence  of  a  crustacean 
18S  ribosomal  RNA  gene  and  secondary  stnicture  of  eu- 
karyotic  small  subunit  ribosomal  RNAs.  Nucleic  Acids  Re- 
search 12:8749-8768. 

Nordsieck,  H.  1992.  Phylogeny  and  system  of  the  Pulmonata. 
Archiv  fiir  MoUuskenknande  121:31-52. 

Ponder,  W.  F.  and  D.  R  Lindberg  1996.  Gastropod  phylogeny- 
challenges  for  the  90s.  In:  J.  D.  Taylor  (ed.)  Origin  and 
evolutionary  radiation  of  the  Mollusca.  O.xford  University 
Press,  Oxford,  pp.  135-154. 

Ponder,  W.  F.  and  D.  R.  Lindberg.  1997.  Toward  a  phylogeny 
of  gastropod  molluscs:  an  analysis  using  morphological 
characters.  Zoological  Journal  of  the  Linnean  Societ)- 119: 
8.3-265. 

Poulicek,  M.,  M.  F  Voss-Foucart,  and  C.  Jeuniau.x.  1991.  Re- 
gressive shell  evolution  among  opisthobranch  gastropods, 
Malacologia  32:223-232. 

Rigby,  J.  E.  1965.  Succinea  piitris:  A  terrestrial  opisthobranch 
mollusc.  Proceedings  of  the  Zoological  Society  of  London 
144:445-486. 

Rosenberg,  G.,  G.  S.  Kuncio,  G.  M.  Davis,  and  M.  G.  Hara- 
sewych. 1994.  Preliminary-  ribosomal  RNA  phylogeny  of 
gastropod  and  unionoidean  bivalve  moUusks.  The  Nauti- 
lus, Supplement  2:111-121. 

Rosenberg,  G.,  S.  Tillier,  A.  Tillier,  G.  S.  Kuncio,  R.  T  Hanlon. 
M.  Masselot,  and  C.  J.  WiUiams.  1997.  Ribosomal  RNA 
phylogeny  of  selected  major  clades  in  the  Mollusca.  Jour- 
nal of  Molluscan  Studies  63:.301-309. 

Runnegar,  B.  and  J.  Pojeta,  1985.  Origin  and  diversification  of  the 
Mollusca.  In:  E.  R.  Tnmian  and  M.  R.  Clarke  (eds.)  The 
Mollusca.  Volume  10.  Academic  Press,  New  York,  pp.  1-.57. 


Saitou,  N.  and  M.  Nei.  1987.  The  neighbor-joining  method:  a 
new  method  for  reconstructiong  phvlogenetic  trees.  Mo- 
lecular Biologv  and  Evolution  4:406^25. 

Salvini-Plawen,  L.  v.  1980.  A  reconsideration  of  systeniatics  in 
the  Mollusca  (phvlogeny  and  higher  classification).  Mala- 
cologia 19:249-278. " 

Salvini-Plawen,  L.  v,  1990.  Origin,  phylogeny  and  classification 
of  the  phylum  Mollusca.  Iberus  9:l-.33. 

Salvini-Plawen,  L.  v.  and  G.  Haszprunar  1987.  The  Vetigastro- 
poda  and  the  systeniatics  of  streptoneurous  Gastropoda 
(Mollusca).  Journal  of  Zoologv  211:747-770. 

Sambrook.  J.,  E.  F.  Fritsch,  and  T.  Maniatis.  1989.  Molecular 
cloning:  a  laboratory'  manual.  Cold  Spring  Harbor  Labo- 
rator)-  Press,  Cold  Spring  Harbor,  New  York. 

Sanger,  F,  S.  Nicklen,  and  A.  R.  Coulson.  1977.  DNA  sequenc- 
ing with  chain  termination  inhibitors.  Proceedings  of  the 
National  Academy  of  Sciences  74:546.'3-.5467. 

Solem,  A.  1978.  Classification  of  the  land  mollusca.  In:  Fretter, 
V.  and  J.  Peake  (eds.)  Pulmonates.  Volume  2A.  Academic 
Press,  New  York,  pp.  49-97. 

Steiner,  G.  and  M.  Miiller  1996.  What  can  18S  rDNA  do  for 
bivalve  phvlogenv?  Journal  of  Molecular  Evolution  43:58-70. 

Swofford,  D.  L.  1999.  PAUP  4.0b2  (a  commercial  test  version: 
completed  version  4.0  to  be  distributed  by  Sinauer,  Sun- 
derland, Mass). 

Taylor,  D.  W.  and  N.  F.  Sohl.  1962.  An  outline  of  gastropod 
classification.  Malacologia  l:7-.32. 

Tillier,  S.  1984.  Relationships  of  gvTiinomorph  gastropods  (Mol- 
lusca: Gastropoda).  Zoological  Journal  of  the  Linnean  So- 
ciety 82:345-362. 

Tillier,  S.  1989.  Comparative  morphologv,  phylogeny  and  clas- 
sification of  land  snails  and  slugs  (Gastropoda:  Pulmonata: 
Stvlommatophora),  Malacologia  30:1-303. 

TilUer'S.,  M.  Masselot,  J.  Guerdoux,  and  A.  Tillier  1994, 
Monophyly  of  major  gastropod  taxa  tested  from  partial 
28S  rRNA  sequences  with  emphasis  on  Euthyiieura  and 
hot-vent  limpets  Peltospiroidea.  The  Nautilus,  Supple- 
ment 2:122-140. 

Tillier,  S.,  M.  Masselot,  and  A.  TiUier.  1996.  Phylogenetic  re- 
lationships of  the  pulmonate  gastropods  from  rRNA  se- 
quences, and  tempo  and  age  of  the  stvlommatophoran  ra- 
diation. //!.  J  D.  Tavlor  (ed.)  Origin  and  evolutionary  ra- 
diation of  tile  Mollusca.  Oxford  University  Press,  Oxford, 
pp.  267-285. 

Thompson  J.  D.,  D  G.  Higgins,  and  T  J.  Gibson.  1994.  CLUS- 
TAL  W:  improving  the  sensitivitv  of  progressive  multiple 
sequence  alignment  through  sequence  weighting,  posi- 
tions-specific gap  penalties  and  weight  matrix  choice.  Nu- 
cleic Acids  Research  22:467.3-4680. 

\'aught,  K.  C.  1989  A  classification  of  living  Mollusca.  Amer- 
ican Malacologists,  Melbourne  (Florida),  189  pp. 

Winnepenninckx,  B.,  T  Backeljau.  and  R.  De  Wachter  1996. 
Investigation  of  molluscan  phvlogenv  on  tlie  basis  of  18S 
rRNA  Sequences.  Molecular  Biologv'  and  Evolution  13: 
1,306-1317. 

Winnepenninckx,  B.,  G.  Steiner,  T  Backeljau,  and  R.  De 
Wachter  1998a.  Details  of  gastropod  phvlogenv  inferred 
from  18S  rRNA  sequences.  Molecular  Phvlogenetics  and 
Evolution  9:.55-63, 

Winnepenninckx,  B.,  D.  G.  Reid,  and  T  Backeljau.  1998b.  Per- 
formance of  18S  rRNA  in  littorinid  phvlogenv  (Gastro- 
poda: Caenogastropoda).  Journal  of  Molecular  Evolution 
47:586-596.  " 

Yoon,  S.  H.,  S.  Y.  Moon,  B.  L.  Choe,  and  W.  Kim  1996.  Se- 
quence divergence  ot  18S  ribosomal  DNA  of  gastropods 
(molluscs).  Korean  Journal  of  Malacolgy  12:85-90. 


THE  NAUTILUS  114(3):93-98,  2000 


Page  93 


Size-depth  patterns  in  two  bathyal  turrid  gastropods: 
Benthomangelia  antonid  (Dall)  and  Oenopota  ovalis  (Friele) 


Alisabet  J.  Clain 

Department  of  Biologv' 
UiiiversiU'  of  Massachusetts,  Boston 
100  Morrissev  Boulexard 
Boston,  MA  02125  USA 


Michael  A.  Rex' 

Department  of  Biology 
Uni\ersit\'  of  Massachusetts,  Boston 
100  Morrissev  Boulexard 
Boston,  MA  02125  USA 
nnchael.rexCffumb.edu 


ABSTRACT 

During  the  past  decade,  there  has  been  a  resurgence  of  interest 
in  the  evolutionars'  and  ecological  significance  of  animal  body 
size.  Geographic  patterns  of  body  size  in  deep-sea  organisms 
remain  poorly  described.  In  this  paper,  we  analyze  depth-re- 
lated trends  of  larxiil  and  adult  shell  size  in  two  turrids  (Gas- 
tropoda) from  the  western  North  Atlantic.  The  upper  bathyal 
Oenopota  ovalis  (478-2022  m),  which  has  non-planktotrophic 
de\elopment,  shows  significant  positixe  size-depth  clines  for 
laryal.  but  not  adult  shells.  The  lower  bathyal  Benthomangelia 
antonia  (2359-3834  ni),  with  planktotrophic  development, 
shows  no  trend  for  lan'al  shells,  but  a  positive  relationship  of 
size  to  depth  for  adult  shells.  The  increase  in  size  with  depth 
confirms  earher  obser\'ations  of  size-depth  clines  in  deep-sea 
snails,  which  may  reflect  selection  for  greater  competitive  abil- 
it\'  and  foraging  efficiency  with  increased  depth.  Contrasts  in 
clina!  \ariation  between  lar\al  and  adult  shells  suggest  that  dif- 
ferent stages  of  life  history  in  deep-sea  snails  may  respond  in- 
dependently to  environmental  gradients. 

Additional  key  words:  Gastropoda,  Turridae,  deep  sea,  size- 
depth  patterns,  western  North  Atlantic. 


INTRODUCTION 

Animal  body  size  has  been  related  to  a  wide  range  of 
physiological  (Peters,  1983;  Chown  and  Gaston,  1999), 
evolutionary  (Stanley,  1979),  and  ecological  phenomena 
(LaBarbera,  1989).  There  is  also  considerable  new  in- 
terest in  the  imphcations  of  body  size  for  geographic 
range  (Brown,  1995),  species  diversity  (Finlay  et  al., 
1996;  Nee  and  Lawton,  1996),  abundance  (Siemann  et 
al,  1996)  and  conservation  of  biodiversity  (May,  1988; 
Marquet  ct  al,  1990;  Gaston  and  Blackbum,  1996).  One 
of  the  most  striking  characteristics  ot  the  deep-sea  fauna, 
including  moUusks,  is  the  small  size  of  most  organisms. 
Explanations  for  this  have  centered  primarily  on  mea- 
suring size-depth  trends  in  various  deep-sea  t;L\a  or  func- 
tional groups,  and  relating  these  to  food  availability, 
which  decreases  with  depth  (Thiel,  1975,  1979;  Gage 

'  Author  for  correspondence. 


and  Tyler,  1991).  Size-depth  relationships  appear  to  vary 
considerably  among  faunal  components  (Rex  and  Etter, 
1998).  It  is  unclear  the  degree  to  which  this  inconsisten- 
cy represents  biologically  meaningful  differences  in  how 
organisms  respond  to  environmental  gradients,  or  meth- 
odological differences  associated  with  how  size  is  mea- 
sured and  the  way  that  morphologically  and  taxonomi- 
cally  heterogeneous  groups  of  species  have  been  com- 
bined to  obtain  composite  estimates  of  size.  To  study 
size  as  an  adaptation,  it  is  important  to  standardize  mea- 
surements to  common  growth  stages  and  to  examine  pat- 
terns within  species  (Gould,  1969;  Rex  and  Etter,  1998). 
Rex  (1979)  and  Rex  and  Etter  (1990,  1998)  showed 
that  size,  standardized  to  specific  larval  and  adult  growth 
stages  within  individual  deep-sea  snail  species,  shows  a 
strong  tendency  to  increase  with  depth.  These  studies 
were  based  on  average  interpopulation  size  differences 
using  relatively  few  sampling  sites,  often  located  near  the 
extremes  of  the  species'  depth  ranges.  Here,  we  present 
detailed  analy,ses  of  size-depth  trends  in  populations  of 
two  species  of  deep-sea  snails,  using  larger  sample  sizes 
and  much  more  thorough  depth  co\'erage.  Results  gen- 
erally support  the  positive  size-depth  trends  reported 
earlier,  but  also  suggest  that  larxae  and  adults  may  re- 
spond differently  to  selective  differences  associated  with 
change  in  depth. 

MATERIALS  AND  METHODS 

This  study  focuses  on  two  common  and  taxonomically 
well-known  bathyal  turrids,  Benthomangelia  antonia 
(Dall,  1881)  and  Oenopota  ovalis  (Friele,  1877),  showni 
in  figures  1^.  They  were  collected  from  the  western 
North  Atlantic,  south  of  New  England  (table  1,  figure 
5),  using  epibenthic  sleds  (Hessler  and  Sanders,  1967), 
box  cores  (Hessler  and  Jumars,  1974),  Blake  deep-sea 
trawls  or  beam  trawls  (Tanner,  1897).  The  two  species 
were  chosen  primarily  because  they  were  abundant 
enough  in  existing  collections  to  allow  reasonable  statis- 
tical analyses  across  broad  depth  ranges. 

B   antonia  and  O.  ovalis  differ  in  their  mode  of  de- 


Paee  94 


THE  NAUTILUS,  Vol.  114,  No.  3 


Figure  1.  SEM  of  the  adult  shell  of  Oenopota  ovalis  (Station 
87,  1102  m,  4.01  mm  total  length).  Figure  2.  SEM  of  the 
adult  shell  of  Bcnthoinanoelia  antonia  (Station  76,  2862  m, 
6.51  mm  total  length).  Figure  3.  SEM  of  the  larval  shell  of 
Oenopota  ovalis  (Station  87,  1102m,  0.45mm  protoconch 
height).  The  indentation  before  the  first  readily  detectable 
adult  rib  (at  a  magnification  of  .30x)  was  used  as  the  lanal- 
adult  transition  (see  text  for  further  discussion).  Figure 
4.  SEM  of  the  larval  shell  oi  BenthomangeUa  antonia  (Station 
76,  2862  m,  1.18  nun  protoconch  height).  Horizontal  arrow 
indicates  the  lar\al-adult  transition  (see  text  for  further  discus- 
sion). 


velopnient  (Bouchet  and  Waren,  1980).  O.  ovalis  ap- 
pears to  haw  direct  or  leeithotrophic  development  in 
which  larvae  mature  in  a  protected  egg  capsule  and  ei- 
ther emerge  crawhng  or  undergo  a  brief  pelagic  or  de- 
mersal dispersal  phase  (e.g.,  Thorson  1935,  1944; 
Bouchet  and  Waren,  1980;  Shimek,  1983).  In  B.  antonia. 
larvae  hatch  at  a  small  size  and  develop  planktotrophi- 
cally.  Veligers  evidently  undergo  ontogenic  vertical  mi- 
gration in  the  water  column,  and  consequentlv  have 
more  dispersal  potential  than  do  the  laivae  of  O.  ovalis 
(Killinglev  and  Bex,  1985).  An  increa.se  in  the  proportion 
of  species  that  have  planktotrophic  development  with 
increased  depth  seems  to  be  a  general  pattern  in  deep- 
sea  turrids  of  the  western  and  eastern  North  Atlantic 
(Potter  and  Bex,  1992). 

Mollusks  are  useful  for  this  tvpe  of  study  because  their 
calcareous  shells  record  the  hfe  history  of  each  inchvid- 
ual,  and  are  not  subject  to  the  changes  in  form  that  are 
caused  by  preservation  in  soft-bodied  organisms.  We 


40°  - 


38° 


36 


207    73 


N13. 
^    W3 


.••62 


N4_ 
N10  _r-->    •^084 

2041*  2569  <^ 
»2037  ^/  "^ 


/ 
340 


'*\    -.^" 

L-1-^ 


2714 


,95  -  - 
'-'2038 


74° 


72° 


70° 


68° 


66° 


64° 


Figure  5.  Map  of  Northwest  Atlantic  showing  the  localities 
of  collection  stations.  See  table  1  for  station  data.  Depth  con- 
tours are  in  meters.  Diamonds  represent  stations  where  Oen- 
opota ovalis  was  collected;  circles  represent  sites  for  Benthom- 
anoelia  antonia. 


chose  only  specimens  that  were  collected  hve.  Measure- 
ments of  size  were  standarchzed  to  the  transition  be- 
tween laival  and  adult  shells  (figures  3  and  4).  If  this 
was  undetectable  due  to  erosion  or  breaking,  the  spec- 
imen was  not  used.  In  B.  antonia,  the  transition  is  clearly 
marked  by  changes  in  bodi  color  and  sculpture  (Bouchet 
and  Wart^n,  1980;  Bex  and  Etter,  1990).  In  O.  ovalis, 
both  larvae  and  adults  are  heaviK-  calcified  ami  are  the 
same  white  color  making  the  transition  less  chstinct.  The 
protoconch  is  smooth  and  the  adult  shell  has  fine  vertical 
ribs.  We  used  the  indentation  before  the  first  readily 
detectable  adult  rib  as  the  lar\'al-adult  transition  for  this 
species. 

Shells  were  measured  microscopically,  orienting  them 
by  the  method  described  in  Bex  and  Etter  (1990).  Each 
shell  was  mounted  on  a  disk  of  clav  and  then  placed  on 
a  4-diniensional  stage  of  an  Olvmpus  dissecting  micro- 
scope equipped  with  a  Sony  3  CCD  Color  Video  Camera 
mount.  The  camera  was  attached  to  a  NuVista  Imaging 
Board  in  a  Macintosh  Centris  650  computer.  Using 
NuVista  Capture+  Software,  the  images  of  the  mounted 
shells  were  digitized.  The  saved  images  were  then  load- 
ed into  the  NIH  Image  program.  Previously  digitized 
images  of  a  1.0  mm  stage  micrometer  with  0.01  mm 
gradations  were  used  to  calibrate  and  measure  the  shell 
images. 

Size-depth  trends  in  the  two  species  were  analv^zed 
using  three  variables:  standardized  lar\al,  and  adult  size, 
and  estimates  of  final  size  attained.  EarK'  protoconch 
whorls  for  one  species,  B.  antonia.  were  often  corroded, 
making  it  impossible  to  accurately  measure  protoconch 
height  in  all  specimens.  Consequentlv,  larval  size  was 
taken  as  protoconch  width.  Standardized  adult  size, 
hereafter  referred  to  as  Whorl- 1  Size,  was  measured  as 


A.  J.  Clain  and  M.  A.  Rex,  2000 


Pa^e  95 


Table  1.  Station  data  for  samples  of  O.  ovalis  and  B  antonia  measured  in  this  stud\.  Tlie  species  collected,  sampling  nietliodolog)-, 
and  number  of  specimens  used  in  this  study  are  also  given.  For  sampling  method:  ES  =  epibenthic  sled  (Hessler  and  Sanders, 
1967),  BC  =  box  core  (Hessler  and  Jumars,  1974),  BT  =  beam  trawl,  DSf  =  Blake  deep-sea  trawl  (Tanner,  1897).  ACSR  indicates 
Atlantic  Continental  Slope  and  Rise  Study  (Maciolek  et  ai,  1986). 


i:).-|)tli 

Saiiiplins: 

.Number  of 

Species 

Station 

Cruise 

1  m ' 

Latitude  °.\' 

Longitude  °  W 

method 

indi\iduals 

Oenopota  ovalis 

88 

Ch.mn 

478 

39°54.10' 

70°37.00' 

ES 

3 

96 

Cn.\iN 

498 

39°55.20' 

70°39.50' 

ES 

3 

N4 

ACSR 

550 

40°21.17' 

67°.32.18' 

BC 

1 

207 

Chain 

808 

.39°51.15' 

70°55.35' 

ES 

6 

87 

Chain 

1102 

.39°48.70' 

70°40.80' 

ES 

29 

NIO 

ACSR 

1220 

39°48.10' 

70°05.30' 

BC 

2 

N13 

ACSR 

1250 

39°48..35' 

70°54.94' 

BC 

1 

N13 

ACSR 

1250 

39°48.35' 

70°54.94' 

BC 

1 

7.3 

Atlantis 

1400 

39°46.50' 

70°43.30' 

ES 

15 

103 

Chain 

2022 

39°43.60' 

70°37.40' 

ES 

8 

BenthomangeUa 

2084 

Albatross 

•7.3,59 

40°16.S3' 

67°05.25' 

BT 

4 

antonia 

62 

Atlantis 

2496 

39°26.00' 

70°33.00' 

ES 

1 

2096 

Alb.atross 

2654 

39°22.33' 

70°52.33' 

BT 

3 

2221 

Alb.^tross 

2789 

39°05.50' 

70°44.50' 

BT 

9 

76 

Chain 

2862 

39°38..30' 

67°57.80' 

ES 

23 

72 

Atlantis 

2864 

38°16.00' 

71°47.00' 

ES 

12 

64 

Atlantis 

2886 

38°46.00' 

70°06.00' 

ES 

10 

2174 

.\lb.atross 

2915 

38°15.00' 

72°03.00' 

BT 

3 

2041 

Albatross 

2941 

39°22.83' 

6S°25.00' 

DST 

5 

2716 

Alb.atross 

2983 

38°29.50' 

70°57.00' 

BT 

14 

2037 

Alb.^tross 

3166 

38°53.00' 

69°23.50' 

BT 

3 

2569 

Albatross 

3259 

39°26.00' 

68°03.50' 

BT 

2 

.340 

Knorr 

3310 

38°16.00' 

70°21.55' 

ES 

13 

2570 

Alb.\tross 

3316 

39°54.00' 

67°05.50' 

BT 

4 

2714 

Alb.\tross 

3338 

38°22.00' 

70°17.50' 

BT 

4 

2038 

Albatross 

3718 

38°30.50' 

69°08.42' 

DST 

3 

95 

Atlantis 

3753 

38°33.00' 

68°.32.00' 

ES 

2 

126 

Atlantis 

3806 

39°.37.25' 

66°45.50' 

ES 

18 

"*"* 

Chain 

3806 

38°00.70' 

69°16.00' 

ES 

7 

78 

Chain 

3828 

.  38°00.80' 

69°18.70' 

ES 

8 

85 

Chain 

38.34 

37°59.20' 

69°26,20' 

ES 

8 

the  height  plus  width  of  the  first  post-lar\al  whorl 
(Gould,l969;  Re.\  and  Etter.  1990).  Final  size  attained 
by  adults,  hereafter  referred  to  as  Adult  Size,  was  mea- 
sured as  the  maximum  height  plus  width  of  the  post- 
larval  shell.  Adult  Size  can  not  be  standardized  to  a  com- 
mon de\elopment  stage,  because  the  species  appear  to 
have  indeterminate  growth.  The  distribution  of  Adult 
Size  in  a  population  reflects  the  recent  history  of  re- 
cruitment and  growth  more  than  size  as  an  adaptive 
property.  To  estimate  the  final  size  attained,  we  selected 
the  largest  two  indi\iduals  (from  anv  station)  found  in 
250  m  depth  increments.  These  three  measures  of  size 
(Protoconch  Width,  Whorl-l  Size,  and  Adult  Size  at- 
tained) were  regressed  against  depth  to  anal\-ze  bath\- 
metric  cUnal  patterns. 

RESULTS  AND  DISCUSSION 

Relationships  of  size  \'ariables  to  depth  are  showTi  in  fig- 
ure 6.  Regression  fines  are  fitted  onl\'  for  significant  re- 
lationships (see  table  2  for  regression  equations  and  their 


statistics).  All  significant  regressions  are  positi\e,  indi- 
cating an  increase  in  size  with  depth.  There  is  no  indi- 
cation that  size  decreases  significantly  with  depth  for  an\' 
variable  in  either  species.  In  O.  ovalis,  lanal  size  in- 
creases with  depth,  but  post-lar\'al  stages  show  no  pat- 
tern. In  contrast,  B.  antonia  shows  no  relationship  of 
lar\-al  size  to  depth,  but  both  Whorl- 1  Size  and  .\dult 
Size  do  increase  with  depth. 

These  results  largeK"  confinu  those  reported  bv  Rex 
and  Etter  (1998)  for  B.  antonia  and  O.  ovalis.  The  pre- 
sent study  is  more  statisticallv  accurate  in  that  it  includes 
larger  sample  sizes,  broader  depth  co\erage.  and  a  much 
more  continuous  distribution  of  samples  across  the 
depth  gradient.  For  Whorl-l  Size,  both  studies  detected 
a  strong  positi\e  increase  with  depth  in  B  antonia  and 
no  apparent  trend  in  O.  ovalis.  For  protoconch  size.  Rex 
and  Etter  (I99S)  reported  a  weaklv  significant  (p<0.051 
increase  with  depth  in  B.  antonia.  and  no  relationship 
in  O.  ovalis.  \\ith  more  extensi\e  sampling,  the  proto- 
conch size-depth  relationship  becomes  just  insignificant 
(p  =  0.05S9)  in  B    antonia.  and  strongly  significant  in 


Page  96 


THE  NAUTILUS,  Vol.  114.  No.  3 


1.50 


1500 


2000 


2500 


3000 


3500 


4000 


Depth  (m) 


A.  J.  Clain  and  M.  A.  Rex.  2(K)() 


Page  97 


Tabic  2.    Regression  siuiiiiuin   tor  rclatioiisliips  ol  sr/.i-  to  dt-ptli  in  fiilure  6.   For  eacli  regression,  tlie  niiiMt)er  of  iiuli\i(luals, 
regression  einiations,  F-\'alues.  r-  \alues,  ant!  tlie  sigiiitieanee  are  gi\en. 


Number  of 

Significance 

Measurement 

Species 

specimens 

Regression  equation 

r* 

F-\alue 

(p) 

Protoconcli  widtli 

B.  antonia 

156 

Y  =  0.982  +  2.69E  -  5x 

0.230 

3.622 

00589 

Protoconch  width 

O.  ovalis 

69 

Y  =  0.6.39  +  4..54E  -  5x 

0.147 

11.507 

0.0012 

Whorl  1  size 

B.  antonia 

135 

Y  =  2.219  +  1.78E  -  4x 

0.106 

15.850 

0.0001 

Whorl  1  size 

0.  ovalis 

62 

Y  =  2.016  +  7.44E  -  5x 

0.046 

2.863 

0.0958 

Adult  size 

B.  antonia 

156 

Y  =  7.644  +  l.OOE  -  .3x 

0.007 

1.116 

0.2925 

Adult  size 

0.  ovalis 

69 

Y  =  5.171  -  2.S8E  -  .5x 

0.000 

0.010 

0.9221 

Largest  adult 

B.  antonia 

14 

Y  =  1.694  +  4.00E  -  3x 

0.332 

5.963 

0.0310 

Largest  ailult 

O.  ovalis 

11 

Y  =  5.915  +  1.2.3E  -  5x 

0.000 

0.001 

0.9772 

O.  ovalis.  Rex  and  Etter  (1998)  did  not  e.stiniate  final 
adult  size.  As  with  studies  of  biodiversity  (Rex  et  al, 
1997)  and  life-history  characteristics  (Stuart  and  Rex, 
1994),  the  patterns  in  protoconch  size  that  are  revealed 
with  better  sampling  demonstrate  the  importance  of  us- 
ing large  databases,  and  broad  continuous  depth  gradi- 
ents to  establish  biogeographic  patterns  in  the  deep  sea. 
This  is  especiallv  relevant  for  species  like  B.  antonia  that 
show  high  variabihtv  in  shell  architecture  (figure  6). 

It  is  interesting  that  O.  ovalis,  with  non-planktotrophic 
development  shows  a  size-depth  cline  only  for  larvae; 
whereas  B.  antonia  with  plankiotrophic  development  ex- 
hibits no  trend  for  larvae,  but  a  significant  positive  cline 
for  Whorl- 1  Size  that  also  persists  at  final  size  attained. 
For  B.  antonia,  an  e.xplanation  might  be  that  larvae  show 
no  pattern  because  they  migrate  from  numerous  distant 
sites  that  may  exert  quite  different  selective  regimes,  and 
that  selection  for  size  at  settlement  along  the  depth  gra- 
dients is  not  strong  enough  to  overcome  the  mitigating 
effects  of  dispersal.  In  this  context,  it  is  interesting  that 
larvae  (and  adults)  of  B.  antonia  show  considerably  more 
variation  in  size  than  those  of  O.  ovalis,  possibly  reflect- 
ing multiple  geographic  origins  for  the  former  (figure  6). 
Selection  for  increased  size  with  depth  may  be  imposed 
largely  during  subsequent  growth  resulting  in  the  posi- 
tive size-depth  cfine  seen  in  adult  shells  of  B.  antonia. 
The  nature  of  the  relevant  depth-related  selective  gra- 
dient remains  unk-nown.  Rex  and  Etter  (1998)  suggested 
that  the  decrease  in  nutrient  input  with  depth  may  select 
for  larger  size  because  of  its  metabolic  and  competitive 
advantages. 

The  opposite  pattern  observed  in  O.  ovalis  is  difficult 
to  interpret  in  a  consistent  way.  Larger  size  at  hatching 
may  confer  advantages  of  lower  vulnerabifity  to  preda- 
tors, greater  locomotion  to  forage,  a  greater  range  of 
food,  and  more  abilitv  to  withstand  star\'ation  (Spight, 
1976).  However,  the  absence  of  a  size  cfine  at  adult  stag- 


es is  puzzUng.  In  general,  the  results  do  indicate  that 
different  life  stages  can  respond  differently  and  inde- 
pendently to  environmental  changes  associated  with 
depth.  Since  the  species  live  at  different  depths,  a  geo- 
graphically controlled  comparison  is  not  possible.  Also, 
because  we  are  analyzing  just  two  species,  the  results 
may  represent  idiosyncratic  effects  that  are  phenotypic 
plastic  responses  or  phvlogenetic  constraints,  rather  than 
general  patterns  that  are  related  to  selective  gradients 
and  developmental  modes. 

In  summary,  the  upper  bathyal  O.  ovalis  shows  a  sig- 
nificant positive  size-depth  cfine  for  larvae,  but  not 
adults.  The  lower  bathval  B.  antonia  reveals  no  depth- 
related  pattern  for  lai-val  shells,  but  a  significant  increase 
in  size  with  depth  for  adults.  The  exact  causes  of  these 
opposing  trends  are  necessarily  speculative;  but,  the  re- 
sults do  suggest  that  larvae  and  adults  may  respond  dif- 
ferently to  selective  gradients  in  species  with  contrasting 
modes  of  development  in  the  deep  sea  as  they  appear 
to  do  in  coastal  moUusks  (Pechenik,  1999;  Pechenik  ct 
al,  1996).  The  increase  in  size  with  depth  generallv  sup- 
ports earlier  studies  on  geographic  variation  in  deep-sea 
gastropods  (Rex  and  Etter,  1990,  1998;  Rexet  al.,  1999). 
However,  detailed  analyses  of  individual  species  high- 
light the  need  for  large  databases  and  good  sampUng 
coverage  throughout  species'  bathvmetric  ranges  to  ac- 
curately assess  biogeographic  patterns  in  deep-sea  or- 
ganisms. 

ACKNOWLEDGMENTS 

We  thank  Ron  Etter  Carol  Stuart,  John  Ebersole,  and 
Rob  Stevenson  for  reacfing  drafts  of  the  manuscript.  Bill 
Fowle  for  the  SEMs  and  Jack  Cook  for  the  map  of  the 
Northwestern  Atlantic  Ocean.  Howard  Sanders,  Fred 
Grassle,  Nancy  Maciolek,  James  Blake  and  Jerrv'  Hara- 
sewych  helped  provide  the  original  material  that  was 


Figure  6.  Relationships  of  Protoconch  Width  (top),  Whorl-1  Size  (middle),  and  Adult  Size  (bottom)  to  depth  in  populations  of 
Oenopota  ovalis  (diamonds)  and  Benthoinan^i'lia  antonia  (circles)  collected  in  the  deep  sea  of  the  western  North  Atlantic.  The 
solid  symbols  in  the  Adult  Size  graph  indicate  the  largest  two  individuals  found  in  each  250  m  interval.  Lines  indicate  significant 
regressions.  See  table  2  for  regression  statistics. 


Page  98 


THE  NAUTILUS,  Vol.  114,  No,  3 


measured  and  analv^zed  here.  This  research  was  sup- 
ported by  the  National  Science  Foundation  Grant  OCE- 
9301687  to  MAR,  and  by  the  University  of  Massachu- 
setts. 


LITERATURE  CITED 

Bouchet,  P.  and  A.  Waren.  1980.  Re\ision  of  the  north-east 
Atlantic  bathyal  and  abyssal  Turridae  (Mollusca,  Gastro- 
poda). Journal  of  Molluscan  Studies  Supplement  8:1-119. 

Brown,  J.  H.  1995.  Macroecology.  University  of  Chicago  Press, 
Chicago,  269  pp. 

Chown,  S.  L.  and  K.  J.  Gaston.  1999.  Exploring  links  between 
physiolog)'  and  ecolog\'  at  macro-scales;  the  role  of  respi- 
ratory metabolism  in  insects.  Biological  ReNiews  74;87- 
120. 

Finlay,  B.  J.,  G.  F.  Esteban  and  T  Fenchel.  1996.  Global  di- 
versity and  body  size.  Nature  383:132-13.3. 

Gage,  J.  D.  and  P.  A.  Tyler  1991.  Deep-sea  biology:  a  natural 
histors"  of  organisms  at  the  deep-sea  floor  Cambridge  Uni- 
versity Press,  Cambridge,  504  pp. 

Gaston,  K.  J.  and  T.  M.  Blackbum.  1996.  Rarity  and  body  size: 
importance  of  generahty.  Conservation  Biology  10:1295- 
1298. 

Gould,  S.  J.  1969.  An  evolutionarv  microcosm.  Pleistocene  and 
Recent  historv  of  the  land  snail  P  {Poecilozonites}  in  Ber- 
muda. Bulletin  of  the  Museum  of  Comparative  Zoologv 
138:407^532. 

Hessler  R.  R.  and  H.  L.  Sanders.  1967.  Faunal  diversity  in  the 
deep  sea.  Deep-Sea  Research  14:65-78. 

Hessler,  R.  R.  and  P.  A.  Juniars.  1974.  Abyssal  communitv  anal- 
ysis from  replicate  box  cores  in  the  central  North  Pacific. 
beep-Sea  Research  21:185-209. 

KiUinglev.  J.  S.  and  M.  A.  Rex.  1985.  Mode  of  lar\al  de\elop- 
ment  in  some  deep-sea  gastropods  indicated  bv  oxygen- 
18  values  of  their  carbonate  shells.  Deep-Sea  Research  .32: 
809-818. 

LaBarbera,  M.  1989.  Analyzing  body  size  as  a  factor  in  ecology 
and  evolution.  Annual  Rexoie  of  Ecology  and  Systematics 
20:97-117. 

Maciolek,  N.,  B.  Hecker,  C.  A.  Butman,  ].  F  Grassle,  W.  B. 
Dade,  P  D.  Boehm,  W.  G,  Steinhauer,  V  Starczk.  E  Bap- 
tiste,  R,  E.  Ruff,  and  B.  Brown.  1986.  Study  of  biological 
processes  on  the  United  States  North  Atlantic  Slope  and 
Rise.  Interim  Report  Prepared  for  the  United  States  De- 
partment of  the  Interior  Minerals  Management  Ser\dces. 
MMS86-0022,  201  pp. 

Marquet,  P  A„  S,  A,  Navarrete,  and  J,  C,  Castilla,  1990,  ScaHng 
population  density  to  body  size  in  rocky  intertidal  com- 
munities. Science  250:1125-1127. 

May,  R.  M.  1988.  How  many  species  are  there  on  earth?  Sci- 
ence 241:  1441-1449. 

Nee,  S.  and  ].  H.  Lawton.  1996.  Body  size  and  di\'ersity.  Na- 
ture 380:672-673. 

Pechenik,  J.  A.,  T.  J.  Hilbish,  L.  S.  Eyster,  and  D.  Marshall. 
1996.   Relationship  between  lar\'al  and  ju\enile  growth 


rates  in  hvo  marine  gastropods,  Crepidula  plana  and  C. 
fornicata.  Marine  Biology  125:119-127. 

Pechenik,  J.  A.  1999.  On  the  adx'antages  and  disad\'antages  of 
larval  stages  in  beuthic  marine  invertebrate  life  cycles. 
Marine  Ecologv'  Progress  Series  177:269-297. 

Peters.  R.  H.  1983.  The  ecological  implications  of  body  size. 
Cambridge  Uni\ersit)'  Press,  Cambridge,  329  pp. 

Potter,  E.  and  M.  A.  Rex.  1992.  Parallel  development-depth 
trends  in  deep-sea  turrid  snails  from  the  eastern  and  west- 
em  North  Atlantic.  The  Nautilus  106:72-75. 

Rex,  M,  A,  1979.  r-  and  K-selection  in  a  deep-sea  gastropod. 
Sarsia  64:29-32. 

Rex,  M.  A.  and  R.  J.  Etter  1990.  Geographic  variation  in  two 
deep-sea  gastropods,  Benthonella  tenella  (|effre\s)  and 
Bcnthonmng^elia  antonia  (Dalb.  Deep-Sea  Research  37: 
1229-1249. 

Rex,  M.  A.  and  R.  ].  Etter.  1998.  Bath\nnetric  patterns  of  body 
size:  impUcations  for  deep-sea  biodiversity.  Deep-Sea  Re- 
search II  45:10.3-127. 

Rex,  M.  A.,  R.  J.  Etter,  A.  J.  Clain.  and  M.  S.  Hill.  1999. 
Bathymetric  patterns  of  body  size  in  deep-sea  gastropods. 
Evolution  53:129.8-1.301. 

Rex,  M.  A.,  R.  J.  Etter  and  C.  T.  Stuart.  1997.  Large-scale 
patterns  of  biodiversity  in  the  deep-sea  benthos.  In:  R. 
Onnand,  ].  D.  Gage  and  M.  V.  Angel  (eds.)  Marine  bio- 
diversity: patterns  and  processes.  Cambridge  Uni\'ersity 
Press,  Cambridge,  pp.  94—121. 

Shimek,  R.  L.  1983.  The  biolog)'  of  northeastern  Pacific  Tm- 
ridae  II.  Oenopota.  Journal  of  Molluscan  Studies  49:146- 
163. 

Siemann,  E.,  D.  Tilman  and  J.  Haarstad.  1996.  Insect  species 
dix'ersity,  abundance  and  body  size  relationships.  Nature 
380:704-706. 

Spight,  T.  M.  1976.  Ecology  of  hatching  size  for  marine  snails. 
'  Oecologia  24: 283-294  ^ 

Stanley,  S,  M.  1979.  Macroexolution:  pattern  and  process.  \V. 
H.  Freeman  and  Co.,  San  Francisco,  332  pp. 

Stuart,  C.  T  and  M.  A.  Rex.  1994.  The  relationship  between 
development  pattern  and  species  diversity  in  deep-sea 
prosobranch  snails.  In:  C.  M.  Young  and  K.  J.  Eckelbarger 
(eds.)  Reproduction,  larval  biolog>',  and  recruitment  of  the 
deep-sea  benthos.  Columbia  Universitv'  Press,  New  York, 
pp.  1L8-136. 

Tanner,  Z.  L.  1897.  Deep-sea  exploration:  a  general  description 
of  the  steamer  Albatross,  her  appliances  and  methods. 
United  States  Fisheries  Commission  Bulletin  for  1896, 
Washington,  pp.  257—428, 

Thiel,  H.  1975,  The  size  structure  of  the  deep-sea  benthos, 
InteniaHonale  Re\aie  der  gesamten  Hydrobiologie  60: 
57.5-606. 

Thiel,  H.  1979.  Stnictural  aspects  of  the  deep-sea  benthos. 
Ambio  Special  Report  6:25-.31. 

Thorson,  G.  1935.  Studies  on  the  egg-capsules  and  develop- 
ment of  Arctic  marine  prosobranchs.  Meddelelser  om 
Gr0nland  100:1-71. 

Thorson,  G.  1944.  The  zoology  of  East  Greenland.  Marine 
Gastropoda  Prosobranchiata.  Meddelelser  om  Gronland 
121:1-181. 


THE  NAUTILUS  114(3):99-1()2,  2()()() 


Page  99 


Four  new  genera  for  northeastern  Pacific  gastropods 


James  H.  McLean 

Natural  Histors'  Museum  of  Los 

Angeles  Counh' 
900  Exposition  BKd. 
Los  Angeles.  CA  90007  USA 
jniclean@nhni.org 


ABSTRACT 

Four  new  genera  for  nortlieasteni  Pacific  Gastropoda  are  pro- 
posed; V'elutinidae:  Torcllivclutina.  type  species  "Torellia"  am- 
monia Dall,  1919;  Euliniidae;  Subniso.  type  species  "Chemnit- 
zia"  ranai  de  Folin.  1867;  Turridae.  Clathurellinae;  Retidrillia. 
tspe  species  "Suavodrillia"  willctti  Dall.  1919;  Mangeliinae; 
PcrimangeUa.  t\pe  species  "Mangelia"  intcrfossa  Carpenter, 
1864. 


Additional  kci/  words: 
lutinidae. 


Gastropoda,  Euliniidae,  Turridae,  V'e- 


INTRODUCTION 

This  is  the  third  in  a  series  of  papers  in  which  new  gen- 
era for  northeastern  Pacific  gastropods  are  introduced 
(see  McLean,  1995a,  b).  I  had  prexaously  indicated  (Mc- 
Lean, 1995a)  that  a  checklist  of  the  northeastern  Pacific 
gastropods  was  in  preparation;  however,  as  indicated  in 
McLean  (1996:  2),  I  am  preparing  a  more  complete 
work  rather  than  a  checklist.  Taxa  described  here  are  to 
be  used  in  an  illustrated  rexasion  of  all  shelled  gastropods 
of  the  northeastern  Pacific,  ranging  from  Arctic  Alaska 
and  the  Aleutian  Islands  to  central  I3aja  Cahfomia,  Mex- 
ico. Description  of  these  genera  in  advance  of  the  book 
allows  for  a  more  detailed  treatment  than  will  be  pos- 
sible in  the  larger  effort.  Further  papers  in  this  series 
will  describe  genera  in  which  the  tyjie  species  or  other 
included  species  are  new. 

To  distinguish  original  combinations  for  type  species 
of  the  new  genera  I  am  here  using  the  convention  of 
placing  all  citations  of  original  genera  within  quotation 
marks.  Illustrations  are  proxaded  here  for  type  species  of 
each  new  genus. 

Although  monotvpic  genera  are  not  encouraged  in 
phylogenetic  classifications,  I  do  not  refrain  from  intro- 
ducing a  few  such  genera  where  necessarv-,  allowing  that 
additional  living  or  fossil  species  may  yet  be  discovered 
that  would  render  those  genera  no  longer  monotypic. 

Museum  acronvms  are:  LACM,  Natural  History  Mu- 
seum of  Los  Angeles  Counb,';  USNM,  National  Museum 
of  Natural  History,  Washington. 


SYSTEM  ATICS 

Family  Velutinidae  Gray,  1840 
Genus  Torcllivclutina  new  genus 

Type  species:  "Torellia"  ammonia  Dall,  1919  (Figure 
1).  The  shriveled  liolotvpe  from  offshore  depths  at  the 
Aleutian  Islands,  Alaska,  was  illustrated  by  Waren  (1989: 
16,  fig.  llf).  A  more  recently  collected,  preserved  spec- 
imen from  the  Aleutian  Islands  is  illustrated  here. 

Included  species:     The  genus  is  monotypic. 

Diagnosis:  Shell  large  (to  30  mm  diameter),  apical 
whorl  depressed,  profile  nearly  planispiral,  of  two  rapidly 
inflated  whorls,  calcareous  layer  lacking  in  large  speci- 
mens except  for  that  which  borders  the  apertural  hp; 
shell  consisting  of  chitinous  periostracum  after  attaining 
3  mm  diameter;  protoconch  with  spiral  sculpture  (ac- 
cording to  original  description  of  Dall);  suture  deep,  um- 
bilicus broad;  mature  surface  composed  of  thick,  chitin- 
ous, colabral  a.xial  lamellae. 

Remarks:  Waren  (1989:  16)  examined  the  type  spe- 
cies for  purposes  of  comparison  with  his  then  described 
new  genus  and  species  Pscudotorcllia  fragilis  from  Ice- 
land. He  illustrated  the  radula  of  "Torellia"  ammonia 
(op.  cit.,  fig.  IIC),  which  he  found  to  agree  with  that  of 
other  velutinid  radulae,  and  reported  that  the  operculum 
and  other  trichotropine  features  are  lacking.  Although 
Waren  removed  "T."  ammonia  from  the  trichotropid  ge- 
nus Torellia,  and  transferred  it  to  the  Velutinidae,  he  did 
not  assign  it  to  a  genus.  Torellivehitina  is  therefore  here 
proposed  for  "T"  ammonia.  It  differs  from  Pseudotorcl- 
lia  in  not  having  a  sohd  calcified  shell  with  spiral  sculp- 
ture and  not  having  the  large,  projecting  protoconch  of 
Pseudotorellia. 

Because  the  calcareous  layer  is  lacking,  fresh  speci- 
mens of  Torellivehitina  ammonia  should  be  retiiined  in 
fluid  preservative  and  should  not  be  dried,  which  results 
in  the  shrinkage  and  buckling  of  the  thick  periostracal 
layer. 

Etymology:  The  name  is  a  compound  of  the  tricho- 
tropid genus  Torrellia  and  the  velutinid  genus  Vclutirm, 


Page  100 


THE  NAUTILUS,  Vol.  114,  No.  3 


A 


Figures  1^.  T)pe  species  of  new  genera.  1.  Torellivehttina  ammonia  (Dall,  1919),  apertural  and  spire  views.  LACM  152281, 
Kanaga  Pass,  between  Kanaga  Island  and  Tanaga  Island,  Andreanof  Islands,  Aleutian  Islands,  Alaska,  61  ni.  Maximum  diameter  .30 
mm.  2.  Subniso  rangi  (de  Folin,  1867).  LACM  1972-38.7,  Punta  Penca,  N  of  Bahia  Potrero,  Guanacaste  Pro\.,  Costa  Rica,  10  ni. 
Length  3.2  mm.  3.  Retidrillia  willetti  (Dall,  1919).  USNM  216409,  lectotype.  Forrester  Island,  southeasteam  Alaska,  dredged. 
Length  11.5  mm.  4.  Perimangelia  inferfossa  (Carpenter,  1864).  LACM  1959-13.42,  Granite  Creek,  Monterey  County,  California, 
10  m.  Length  8.5  mm. 


to  emphasize  that  it  represents  a  velutiiiid  with  the  su- 
perficial aspect  of  the  trichotropid  genus  Torrellia. 

Family  Euhmidae  Philippi,  1853 
Genus  Subniso  new  genus 

Type  species:  "Chemnitzia"  rangi  de  Fohn,  1867 
(Figure  2).  Type  locality:  Perlas  Islands,  Panama.  Waren 
(1992:  183)  could  not  locate  type  material,  but  the  spe- 
cies was  well  figured  bv  de  Folin  and  subsequently  by 
Waren  (1992,  figs.  17,  20-22,  25-27,  30,  31),  so  there  is 
little  doubt  as  to  its  identity.  The  species  occurs  from 
southern  Baja  Cahfomia,  Mexico,  to  Ecuador. 

Included  species:  Two  species,  the  type  species  and 
"Niso"  hipolitcnsis  Bartsch,  1917,  for  which  the  holotype 
from  Punta  San  Hipohto,  Baja  California,  Mexico,  was 
illustrated  by  Emerson  (1965,  fig.  9)  and  Waren  (1992: 
figs.  19,  23,  24).  Both  species  have  previously  been 
placed  in  Niso  Risso,  1826,  bv  Emerson  (1965)  and  War- 
en (1992). 

Diagnosis:  Shell  small  (length  to  4  mm),  slender 
(length  to  breadth  ratio  2.5-2.9),  non-umbilicate,  basal 
angulation  pronounced,  coloration  brown,  without  color 
pattern. 

Remarks:  The  two  species  of  Subniso  are  unUke  other 
species  of  Niso  in  size,  profile,  and  color.  The  two  spe- 
cies are  small,  non-umbilicate,  and  have  a  basal  angula- 
tion (strongly  projecting  in  Subniso  rangi)  and  a  uniform 
brown  coloration.  Waren  (1992:  185)  remarked:  "Both 
Niso  hipolitcnsis  and  N.  rangi  are  unusual  among  the 
species  of  Niso  in  their  small  size,  3^  mm  shell  height, 


whereas  most  species  of  the  genus  have  a  shell  that  is 
10-30  mm  high,  occasionally  even  higher."  Additionally, 
I  point  out  that  Niso  species  are  usually  broadly  umbil- 
icate,  except  for  the  large  N.  atiilloi  (Hertz  and  Hertz, 
1982),  usually  not  angulate  at  the  base,  and  have  color 
patterns  that  may  be  banded  or  variegated.  Taken  to- 
gether, these  differences  are  sufficient  to  justify'  generic 
recognition  of  Subniso. 

Waren  (1992:  1S3)  noted  that  a  starfish  host  is  known 
for  one  Indo-Pacific  species  of  Niso,  but  nothing  is 
known  of  the  host  echinoderm  for  the  two  species  of 
Subniso. 

Etymology:  The  generic  name  combines  the  generic 
name  Niso,  with  the  prefix  sub-,  to  emphasize  the  small 
size,  compared  to  Niso. 

Family  Turridae  Swainson,  1840 

Subfamily  Clathurelhnae  H.  and  A.  Adams,  1858 

Genus  Rctidrillia  new  genus 

Type  species:  ' Suavodrillia"  willetti  Dall,  1919  (Fig- 
ure 3).  Type  locality:  Forrester  Island,  Southeast  Alaska. 
Until  now  the  species  has  been  unfigured  (it  was  not 
illustrated  by  Kosuge,  1972).  The  here  selected  lectotype 
(Figure  7)  is  the  largest  (11.3  mm)  of  seven  original  syn- 
type  specimens  in  the  type  lot.  The  radular  tooth  of  the 
type  species  was  figured  by  McLean  (1971,  fig.  114) 
(from  LACM  66-66,  Graham  Island,  Queen  Charlotte 
Islands,  British  Columbia).  A  specimen  from  the  Rae 
Baxter  collection  in  the  LACM  extends  the  distribution 
to  Akutan,  Aleutian  Islands  (LACM  83-345.  263  m 
depth). 


J.  H.  McLean,  2000 


Page  101 


Included  species:  At  least  three  species,  the  t\pe  spe- 
cies and  two  species  from  the  northeastern  Atlantic  dis- 
cussed and  illustrated  bv  Bouchet  and  Waren  (1980:  32) 
in  the  genus  Drilliola  Locard,  1897:  "D. "  pntina  (Wat- 
son, 1881),  and  "D."  megalacmc  (Sykes,  1906). 

Diagnosis:  Shell  profile  with  concave  shoulder  and 
projecting  peripheral  carination  below  which  base  is 
rounded  and  has  strong  spiral  sculpture.  Anal  sinus  on 
concave  shoulder,  deep  and  broad.  Axial  sculpture  ex- 
pressed as  nodes,  strongest  at  peripheral  carination, 
more  f;iintlv  expressed  on  strong  spiral  cords  of  base. 
Peripheral  carination  of  early  whorls  at  midvvhorl.  Pro- 
toconch  paucispiral,  with  early  development  of  periph- 
eral carination.  Radular  tooth  long,  haqioon-like,  with 
broad  base. 

Remarks:  The  type  species  was  described  in  the  ge- 
nus SiiavodriUia  Dall,  1918  (tvpe  species:  "Drillia"  kcn- 
nicotti  Dall,  1871),  with  which  it  shares  the  long  har- 
poon-like radular  tooth.  Dall  (1921:  69)  indicated  doubt 
that  "S."  willetti  was  congeneric  by  placing  a  question 
mark  preceding  the  genus.  Differences  are  that  Reti- 
drillia  iLillctti  and  the  tvvo  additional  species  here  as- 
signed to  the  genus  are  half  the  size  of  S.  kcnnicotti,  and 
have  sculpture  that  is  axial  and  spiral,  rather  than  strictly 
spiral  as  in  SiiavodriUia.  The  newly  restricted  Siiavod- 
riUia is  monot\pic  for  S.  kcnnicotti,  a  species  broadly 
distributed  in  the  north  Pacific  from  Hokkaido,  Japan, 
the  Kurile  and  Aleutian  Islands,  and  the  Gulf  of  Alaska 
to  Southeastern  Alaska. 

Bouchet  and  Waren  (1980:  32)  retained  "a  wide  va- 
riety of  species"  in  Drilliola,  including  those  that  'look 
rather  different  but  have  a  similar  radula  and  opercu- 
lum." They  elected  "to  keep  them  in  Drilliola  rather 
than  placing  them  in  any  of  the  perhaps  more  similar, 
but  anatomically  unkiiown  genera  hsted  by  Powell 
(1966)  in  different  subfamihes. '  Species  tvpical  of  Dril- 
liola are  more  slender  than  those  of  Rctidrillia  and  do 
not  ha\e  the  broad,  excavated  shoulder.  The  proposal  of 
Rctidrillia  thus  provides  a  genus  with  a  type  species  hav- 
ing knowii  radular  characters  for  the  two  species  cited 
by  Bouchet  and  Waren  (1980). 

On  shell  characters,  Retidrillia  resembles  species  as- 
signed to  Plicisifrinx  Sysoev  and  Kantor,  1986,  in  which 
the  radular  tooth  is  of  the  modified  wishbone  t)pe,  in- 
dicative of  the  less  derived  turrid  subfamily  Cochlespi- 


30-4).  Synonym:  "Man^elia"  interlirata  Stearns,  1872. 
The  species  occurs  from  Clallnm  County,  Washington, 
to  Isla  San  Ceroninio,  Baja  Cahfomia,  Mexico,  based  on 
specimens  in  the  LACM  collection. 

Included  species:  Two  species,  the  type  species  and 
the  less  familiar  "Manficlia"  nitcns  Carpenter,  1864  (syn- 
type  figured  by  Palmer,  1958,  pi.  28,  fig.  1).  It  ranges 
from  Sonoma  County,  California,  to  Ensenada,  Baja  Cal- 
ifornia. 

Diagnosis:  Shell  relatively  small  and  slender  with 
dominant  axial  sculpture,  crossed  by  narrow  spiral  cords 
of  lesser  strength.  Protoconch  of  1.5  whorls,  strongly 
projecting,  smooth  at  first,  developing  fine  spiral  sculp- 
ture after  first  half  whorl,  followed  by  weaker  axials  (ax- 
ials  more  numerous  than  that  of  mature  sculpture), 
changing  imperceptibly  to  adult  sculpture. 

Remarks:  Earlier  (McLean,  1978:  74),  I  placed  the 
type  species  of  the  new  genus  ("Mangelia"  intcrfossa) 
along  with  "Daphnclla"  ftiscoligata  Dall,  1871  in  Clatli- 
romangclia  Monterosato,  1884.  I  now  consider  Clath- 
romangclia  (type  species  "Plcurotoma"  grantiin  Philippi, 
1844;  see  Powell,  1966:  106)  to  be  appropriate  for  two 
species  in  southern  Cahfomia:  "Daphnclla"  fuscoligata 
and  "Mangilia  iClafhmmangclia)"  rhi/ssa  Dall,  1919. 
These  two  species  have  coarse  clathrate  sculpture  and 
relatively  low,  paucispiral  protoconchs. 

Pcrimangclia  differs  from  the  two  species  of  Clath- 
romangclia  in  having  numerous,  narrow  spiral  cords 
overriding  the  dominant  axial  ribs,  rather  than  having 
coarsely  clathrate  sculpture  with  nodes  at  intersections. 
Protoconchs  differ:  the  paucispiral  protoconch  of  the 
two  species  of  Pcrimangclia  is  strongly  projecting, 
whereas  the  paucispiral  protoconch  of  the  two  species 
of  Clathromangclia  is  much  lower. 

Etymology:  The  name  combines  the  prefix  pcri- 
(near)  with  Mangelia  Risso,  1826,  one  of  die  oldest  gen- 
era in  the  subfamily  Mangeliinae. 

ACKNOWLEDGMENTS 

Photographic  prints  were  made  by  Michael  Eraser.  I  am 
grateful  to  Lindsey  Groves,  Daniel  Geiger,  Anders  War- 
en, and  a  further  anonymous  reviewer  for  offering  help- 
ful suggestions. 


Etymology:  The  name  is  a  compound  of  rcti-,  mean- 
ing net,  with  reference  to  the  axial  and  spiral  sculpture, 
and  Drillia  Gray,  1838,  one  of  the  earliest  named  of  tur- 
rid genera. 

Subfamily  Mangehinae  Fischer,  1883 
Genus  Pcrimangclia  new  genus 

Type  species:  "Mangelia"  intcrfossa  Caqienter,  1864 
(Figure  4).  Syiitypes  were  figured  by  Palmer  (1958,  pi. 
27,' figs.  5,  6)'  Also  figured  by  McLean  (1969;  1978,  fig. 


LITERATURE  CITED 

Bouchet,  P.  and  A.  Waren.  1980.  Revision  of  the  northeast 
Atlantic  bath\al  and  abyssal  Turridae  (Mollvisca,  Gastro- 
poda). Joiinial  of  Moiluscan  Studies,  Supplement  8:1-119. 

Dall,  W.  H.  1919.  Descriptions  of  new  species  of  mollusks  of 
tlie  family  Turritidae  from  the  west  coast  of  America  and 
adjacent  regions.  Proceedings  of  the  United  States  Na- 
tional Museum,  56(2288):  1-86,  pis.  1-24. 

Dall,  VV.  H.  1921.  Summary  of  the  marine  shellbearing  mol- 
lusks of  the  northwest  coast  of  America,  from  San  Diego, 
Califoniia,  to  the  Polar  Sea,  mostly  contained  in  the  col- 
lection of  the  United  States  National  Museum,  with  illus- 


Page  102 


THE  NAUTILUS,  Vol.  114,  No.  3 


tratdons  of  hitherto  unfigiired  species.  United  States  Na- 
tional Museum,  Bulletin  112,  217  pp.,  22  pis. 

Emerson,  W.  K.  1965.  The  eastern  Pacific  species  oiNiso  (Mol- 
lusca:  Gastropoda).  American  Museum  Novitates  2218:1- 
12. 

Kosuge,  S.  1972.  Illustrations  of  type  specimens  of  molluscs 
described  bv  William  Healev  Dall  (Northwestern  Pacific 
gastropods).  Special  publication  of  the  National  Science 
Museum,  Tokyo,  29  plates  and  unpaged  captions. 

McLean,  J.  H.  1969.  Marine  shells  of  Southern  California.  Los 
Angeles  County  Museum  of  Natural  History,  Science  Se- 
ries, no.  11,  104  pp. 

McLean,  J.  H.  1971.  A  revised  classification  of  the  family  Tur- 
ridae,  with  the  proposal  of  new  subfamilies,  genera,  and 
subgenera  from  the  Eastern  Pacific.  The  V'efiger  14:114- 
130. 

McLean,  J.  H.  1978.  Marine  shells  of  .southern  California,  Re- 
vised edition.  Natural  Histors'  Museum  of  Los  Angeles 
County,  Science  Series,  no.  24,  104  pp. 

McLean,  J.  H.  1995a.  Four  new  genera  for  northeastern  Pacific 
prosobranch  gastropods.  The  Nautilus  108:39-41. 

McLean,  J.  H.  1995b.  Three  additional  new  genera  and  two 


replacement  names  for  northeastern  Pacific  prosobranch 
gastropods.  The  Nautilus  108:80-83. 

McLean,  J.  H.  1996.  The  Prosobranchia.  In:  P  H.  Scott,  J.  A. 
Blake,  and  A.  L.  Lissner  (eds.)  Taxonomic  atlas  of  the 
benthic  fauna  of  the  Santa  Maria  Basin  and  western  Santa 
Barbara  Channel.  Volume  9.  The  MoUusca  Part  2 — The 
Gastropoda.  Santa  Barbara  Museum  of  Natural  History, 
Santa  Barbara,  v  -t-  160  pp. 

Palmer,  K.  V.  W.  1958.  Type  specimens  of  marine  Mollusca 
described  by  P.  P.  Carpenter  from  the  west  coast  (San 
Diego  to  British  Columbia).  The  Geological  Society  of 
America,  Memoir  76,  vi  -I-  .376  pp.,  35  pis. 

Powell,  A.  W.  B.  1966.  The  molluscan  families  Speightiidae  and 
Turridae.  Bulletin  of  the  Auckland  Institute  and  .Museum, 
no.  5,  184  pp.,  23  pis. 

Sysoev,  A.  V.  and  Y.  1.  Kantor.  1986.  [New  and  rare  abyssal 
species  of  the  tamilv  Turridae  (Gastropoda,  Toxoglossa)  in 
the  northern  part  of  the  Pacific  Ocean.]  Zoologicheskii 
Zhumal,  65(10):  1457-1469.  [In  Russian] 

Waren,  A.  1989.  New  and  little  known  Mollusca  from  Iceland. 
Sarsia  74:1-28. 

Waren,  A.  1992.  Comments  on  and  descriptions  of  eulimid 
gastropods  from  Tropical  West  America.  The  Veliger  35: 
177-194. 


THE  NAUTILUS  114(3):103-111,  2000 


Page  103 


Ohscuranella  papijrodes,  a  new  genus  and  species  of  abyssal 
tonnoidean  gastropod  from  Antarctica 


Yuri  I.  Kantor 

A.  N.  Se\ertzov  Institute  of  Problems 

of  E\oliition 
Russian  Academy  of  Sciences 
Leninski  Prospect  33 
Moscow  117071,  RUSSIA 
kantor@malaco-se\in.msk.ni 


M.G.  Harasevvych 

Department  of  In\ertel)rate  Zoology 
National  Museum  of  Natural  History 
Smitlisonian  Institution 
Washington,  DC.  20560-0118,  USA 
HanisevvTchS'nmnh. si.edu 


ABSTRACT 

The  new  genus  Ohscuranella  and  O.  papijrodes.  its  type  spe- 
cies, are  described  from  the  abyssal  plain  off  the  Ross  Sea, 
Antarctica.  Obsntranella  is  included  in  the  primarily  tropical, 
shallow  water  superfamiK  Tonnoidea  because  it  has  a  taenio- 
glossan  radula,  extensible  proboscis,  large  sali\'arv  glands  com- 
posed of  anterior  and  posterior  lobes,  salixary  ducts  that  pass 
through  die  nerve  ring,  and  an  undifferentiated  oesophageal 
gland.  This  taxon  is  precluded  from  the  families  Laubierinidae, 
Pisanianuridae,  and  Tounidae  because  it  lacks  a  monopectinate 
osphradium,  paired  proboscis  retractor  muscles  passing 
through  the  nerve  ring,  buccal  glands,  and  rachidian  teeth  with 
lateral  basal  denticles.  Ohscuranella  is  assigned  to  tlie  family 
Ranellidae  primarily  on  the  basis  of  shell,  radular,  and  oper- 
cular morphology.  This  is  the  first  report  of  the  Tonnoidea  in 
Antarctic  waters,  and  the  first  record  of  Ranellidae  from  abyssal 
depths. 

Additional  key  words:  Caenogastropoda,  Ranellidae,  anatomy, 
systemafics. 


INTRODUCTION 

In  the  course  of  examining  Antarctic  Buccinoidea  sam- 
pled under  the  auspices  of  the  United  States  Antarctic 
Program  (USAP)  and  housed  in  the  National  Museum 
of  Natural  Histoiy,  Smithsonian  In,stitution,  we  encoun- 
tered five  lots  of  gastropods  with  large  bucciniform  shells 
that  had  tentatively  been  attributed  to  the  buccinoidean 
genus  Bathijdomus  Thiele,  1912,  by  Dell  (1990:198- 
199).  Dell  identified  four  of  these  lots  as  Bathiidomus 
obicctus  Thiele,  1912,  but  considered  the  filth  lot,  con- 
sisting of  a  single  specimen,  to  represent  an  undescribed 
species  of  Bathijdomus.  Dissections  of  preserved  mate- 
rial clearly  demonstrate  that  these  gastropods  are  not 
referable  to  Buccinoidea,  nor  even  to  Neogastropoda, 
but  rather  represent  an  unnamed  genus  and  species  of 
the  superfamily  Tonnoidea.  This  is  the  first  report  of  this 
superfamily  in  the  Antarctic  nialacofaimu,  UTid  one  of 
very  few  records  of  the  Tonnoidea  from  the  abyssal  zone. 
The  family  Ranelfidae,  to  which  this  genus  is  assigned. 


has  not  prcNitjusly  been  reported  from  die  Antarctic  or 
from  abyssal  depths. 

In  this  paper,  we  provide  descriptions  of  this  new  ge- 
nus and  new  species,  and  infer  its  taxonomic  position 
within  the  Tonnoidea  by  comparing  its  anatomy,  radula 
and  operculum  with  published  anatomical  reports  (e.g. 
Weber,  1927;  Houbrick  and  Fretter,  1969;  Day,  1969; 
Beu,  1981;  Hughes  and  Hughes,  1981).  Waren  and 
Bouchet  (1990),  Riedel  (1995).  and  Beu  (1998)  each 
provide  useful,  if  not  entirely  congruent,  syntheses  of 
tonnoidean  families,  and  include  anatomical  descriptions 
as  well  as  numerous  illustrations  of  radulae,  opercula  and 
protoconchs.  Abbreviations  used  in  the  descriptions  of 
morphometric  characters  are  explained  in  Table  1. 

SYSTEMATICS 

Class  Gastropoda  CuNaer,  1797 
Superfamily  Tonnoidea  Suter,  1913 
Family  Ranelhdae  Gray,  1854 

Genus  OhsciirancUa  new  genus 

Type  species:  Ohscuranella  papijrodes  new  species 

Diagnosis:  Protoconch  unknown.  Teleoconch  large, 
pyriform,  thin,  with  large  aperture,  single,  rounded,  un- 
thickened  terminal  varix  adjacent  to  thin,  flared  outer  lip 
in  adult  specimens.  Shell  sculpture  limited  to  fine,  wide- 
ly spaced,  spiral  cords.  Operculum  very  small,  shaqily 
tapering  anteriorly,  with  anterior  terminal  nucleus.  Pro- 
boscis wall  extremely  thick,  paired  proboscis  retractor 
muscles  absent.  Rachidian  teeth  lacking  lateral  basal 
denticles.  Salivary  glands  large,  differentiated.  Osphra- 
dium bipectinate. 

Etymology:  Ohscunis  (L.) — obscure,  unclear  +  Ra- 
nclla — genus  of  Ranellidae. 

Ohscuranella  papijrodes  new  species 
(Figures  1-36,  Table  1) 

Bathijdomus  obtectus  Thiele,  1912— Dell,  1990:198-199,  figs. 
299-300. 


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THE  NAUTILUS,  Vol.  114,  No.  3 


Table  1.  Obscuranella  papifrodes.  new  species.  Measurements  of  shell  characters.  Linear  measurements  in  mm. 


USNM 

USNM 

USNM 

Character 

Holotvpe 

Paratype  1 

Paratype  3 

901317 

870610 

870610 

Shell  LensTth  (SL) 

63  + 

58  + 

33+ 

57.0 

43.6+ 

33.2+ 

Last  Whorl  Len.^th  (LWL) 

55.2 

51.8 

28.5 

47.3 

38.5 

30.4 

Aperture  Length  (AL) 

45.1 

43.3 

23.0 

38.4 

31.5 

25.5 

Shell  Width  (SW) 

40.8 

41.5 

19+ 

34.1 

24+ 

20.2+ 

Number  of  spiral  cords  on 

last  whorl 

14 

12 

14 

5 

14 

13 

Number  of  spiral  cords  on 

penultimate 

whorl 

6 

7 

7 

2 

5 

3 

Bathjdomns  sp.— Dell,  1990:199. 

Description:  Shell  large  (exceeding  63  mm),  very  thin, 
fragile,  ovate-pvrifonin.  Protoconch  and  upper  whorls 
missing  in  ;ill  t\pe  material.  Preserved  portions  of  teleo- 
conch  of  2V2  rapidly  expanding,  evenly  rounded  whorls. 
Shoulder  rounded,  indistinct.  Suture  adpressed,  shallow. 
Axial  sculpture  limited  to  fine,  straight,  weakly  prosocline 
growth  hues.  Adult  specimens  with  a  single,  weak,  hollow 
vailx  adjacent  to  thin,  flared  outer  Up  (figures  2,  7,  arrow). 
Spiral  sculpture  of  sharp,  narrow,  evenly  spaced  cords  (14 
on  last  whorl,  6  on  penultimate  whorl),  with  much  weaker 
sinuous  threads  (22-.30)  of  varying  width  between  adja- 
cent cords.  Aperture  large  [—0.7  shell  length  (SL)], 
broadly  ovate,  deflected  from  shell  axis  by  9-11°.  Outer 
lip  tliin,  evenly  rounded  in  upper  part  and  concave  at 
transition  to  siphonal  canal,  weakly  reflected.  Inner  hp 
consisting  of  long,  convex,  tnedially  indented  parietal  re- 
gion and  short,  smootli,  a.xial  columeOa  with  strong,  long 
siphonal  fold  that  crosses  cofling  axis  of  shell.  Siphonal 
canal  short,  broad,  weakly  recurved  dorsally.  Callus  of 
thin,  wliite,  porcellaneous  glaze  overlying  parietal  region, 
adapical  portion  of  broad,  nearly  axial  siphonal  fasciole. 
Shell  color  pale  ohve-tan,  confined  to  outermost  shell  lay- 
er. Aperture  white.  Periostracum  very  thin,  yeflowish 
brown,  with  densely  spaced  a.xial  lamellae,  occasional 
short  hairs  at  intersection  of  lamellae  with  spiral  cords. 
Operculum  (figures  4,  5)  very  small  (0.16  AL),  x'estigial, 
dark  yellow,  subtriangular,  with  straight  sides,  terminal 
nucleus.  Dorsal  surface  with  numerous,  closely  spaced 
growth  lines.  Ventral  surface  with  thin,  glazed  lateral  mar- 
gins. Operculum  attached  over  most  oi  its  surface. 

Shell  ultrastructure  (Figure  23):  Shell  thin  (101 
|xm),  composed  of  three  layers.  Outermost  laver  (figure 
23,  ca)  thinnest  (4  |xni),  composed  of  columnar  crystals. 
Middle  layer  (figure  23,  ccl)  thickest  (79  jjim),  composed 
of  collabrally  oriented  cross-lameUar  crystals.  Inner  layer 
(figure  23,  rcl)  thin  (IS  p-m),  composed  of  cross-lamellar 


crystals  oriented  perpendicular  to  growing  edge  of  the 
shell. 

Anatomy  (Paratype  1,  9):  Soft  tissues  comprising 
appro.ximately  3V2  whorls.  Mantle  cavity  spans  just  under 
V2  whorl,  nephridium  (figures  24,  25,  27,  n)  about  % 
whorl,  digestive  gland  (figures  24,  25,  dg)  2V^  whorls. 
Mantle  edge  (figures  24,  25,  27,  me)  thickened,  smooth, 
completely  covers  head.  Columellar  muscle  (figure  25, 
cm)  thick,  broad,  spanning  sUghtly  more  than  one  whorl, 
attached  to  shell  at  rear  of  nephridium.  Foot  short  in 
contracted  state  (LengthAVidth  ==  1.0),  with  conspicu- 
ous propodium.  Body  color  uniform  reddish-tan,  without 
pattern  in  alcohol  preserved  specimens.  Head  (figure  26) 
very  large,  as  wide  as  foot,  with  broad,  blunt,  tapering 
tentacles  (figure  26,  tn)  with  black  eyes  at  their  bases. 
Operculum  about  4  mm  long  (0.09  AL),  otherwise  sim- 
flar  to  that  of  holotvpe.  Paratype  1  (and  all  other  pre- 
served specimens)  with  proboscis  protruded  through 
very  wide  rhynchostome  (figure  26). 

Mantle  cavity  (Figure  27):     Mantle  cavity  as  deep  as 

broad  (—Vi  whorl).  Siphon  (figures  24,  25,  27,  s)  broad, 
muscular,  very  short,  extending  shghtly  beyond  mantle 
edge  (figures  24-27,  me).  Osphrachum  (figures  24—27, 
os)  situated  along  central  half  of  ctenidiuni,  bipectinate, 
nearly  symmetrical,  sUghtly  narrower  on  left  side  than 
right.  Ctenidium  (figures  24-27,  ct)  long,  spanning  near- 
ly entire  mantle  length,  formed  of  tall  triangular  lamel- 
lae, nearly  twice  as  high  as  broad.  Hvpobranchial  gland 
(figure  27,  hg)  poorly  developed,  lacking  distinct  folds. 
Rectum  (figure  27,  re)  runs  along  inner  surface  of  palfial 
oviduct,  narrow,  terminating  in  simple  anus  (figure  27, 
a)  behind  thickened  mantle  edge. 

Alimentary  system  (Figures  15,  19-22,  24,  32-35): 

Everted  proboscis  (figures  24-26,  pr)  —30  mm  long  (0.7 
AL),  unpigmented,  with  folded  walls  indicating  potential 
for  further  extension.  Proboscis  wall  very  thick,  —60% 


Figures  1-14.  Obscuranella  papyrodes  new  species.  1.  Apertural,  2.  right  lateral,  and  3.  dorsal  views  of  the  holotype,  USNM 
898683.  4.  External  and  5.  internal  views  of  the  operculum  of  the  holotype  (internal  view  coated  with  Ammonium  chloride  to 
emphasize  sculpture).  6.  Apertural  and  7.  dorsal  views  of  paratype  1,  USNM  896131.  8.  Apertural  and  9.  dorsal  views  of  the  larger 
shell,  10.  dorsal  view  of  smaller  shell,  USNM  870610.  11.  Apertural  and  12.  dorsal  views  of  paratype  3,  USNM  886105.  13. 
Apertural  view  and  14.  apical  view  of  early  whorls  of  USNM  901317.  Scale  bar  =  1  cm  for  all  shells;  Scale  bar  =  2  mm  for 
operculum;  Scale  bar  =  5  mm  for  protoconch. 


Y.  I.  Kantor  and  M.  G.  Hamsewvch,  2000 


Page  105 


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THE  NAUTILUS,  Vol.  114,  No.  3 


Y.  I.  Kantor  and  M.  G.  Harasew\'ch,  2000 


Page  107 


of  proboscis  radius,  composed  of  3  layers  of  muscles. 
Innermost  la\er  of  circular  nuiscles  (figure  32,  cm),  mid- 
dle layer  tliickest  (2/3  of  proboscis  wall),  of  longitudinal 
muscles  (figure  32,  Im),  outer  layer  of  circular  muscle. 
Buccal  mass  (figure  33,  bm)  small,  attached  to  proboscis 
walls  b\  numerous,  thin  tensor  muscles  (figures  15,  32, 

33,  tm),  as  is  the  anterior  oesophagus  (figures  15,  32, 
aoe).  Retractor  muscles  passing  through  ner\'e  ring  and 
joining  buccal  mass  and  columellar  muscle  absent. 
Mouth  (figure  32,  mo)  a  narrow,  vertical  slit.  Buccal  tube 
(figure  33,  bt)  short,  leading  to  cuticle-lined  buccal  cavity 
with  ventral  p;ur  of  semicircular  jaws  (figure  33,  j).  Jaws 
(figures  16-18)  dark  brown,  pappilate  along  outer  edge 
(figure  17).  Inner  surface  of  jaw  composed  of  small, 
closeK-  spaced  platelets  that  produce  "coiibled"  surface 
distally  (figure  18),  smooth  pro.ximally  (figure  16).  Odon- 
tophore  (figure  15,  od)  small,  oval,  lining  bottom  of  the 
buccal  cavity.  Walls  of  buccal  cavity  very  thick.  Proboscis 
nerves  (figure  32,  pn)  paired,  very  thick,  nuining  from 
cerebro-pIeur;il  gangfia  along  proboscis  length,  inner\'at- 
ing  buccal  mass  and  anterior  part  of  proboscis.  Anterior 
esophagus  divided  into  dorsal  and  ventral  channels  by 
prominent  longituchnal  folds  (figures  15,  34,  35,  If)  that 
extend  from  the  buccal  cavitv  to  the  posterior  edge  of 
esophageal  gland.  The  right  fold  overlaps  the  left  (figure 
34).  Radular  ribbon  (figure  19)  short  (5.8  mm,  0.13  AL), 
nearly  twice  as  long  as  cartilages,  narrow  (~  580  |xm, 
0.013  AL),  consisting  of  45  rows  of  teeth,  posteriormost 
4  rows  nascent.  Rachidian  tooth  (figure  21,  rt)  with  large, 
broad  median  cusp,  flanked  by  5-9  denticles  per  side. 
Base  broad,  strongly  concave  posteriorly,  lacking  cusps 
along  tooth  base.  Lateral  teeth  (figures  21,  22,  It)  narrow, 
with  long,  thin,  cusp  flanked  by  3—4  denticles  on  inner 
edge,  4—6  denticles  on  outer  edge.  Two  long,  recurved, 
distally  flattened  marginal  teeth  (figures  20;  22,  mt)  per 
side,  outer  tooth  longer  than  inner  Inner  distal  edges 
serrated  with  2-7  cusps.  Salivarv'  glands  large,  irregularly 
shaped,  completely  covering  the  esophageal  gland  (fig- 
ure 32,  oeg).  Right  salivary  gland  more  elongated,  slight- 
ly larger  than  left.  Each  gland  consists  of  two  lobes.  Pos- 
terior lobe  (figure  32,  plsg)  massive,  composed  of  curved 
radially  oriented  bfind  tubules.  Anterior  lobe  (figure  32, 
alsg)  smaller,  acinous,  ventral.  Salivary  ducts  (figures  32- 

34,  sd)  thick,  extending  from  posterior  lobes,  becoming 
attached  to  oesophagus  wafls  before  passing  through 
nerve  ring.  We  were  not  able  to  identify  connections 
between  the  salivary  ducts  and  the  anterior  lobes  of  the 
salivary  glands,  as  reported  for  Cymafium  intermedium 
(Pease,  1869)  by  Andrews  ct  al.  (1999).  Salivary  glands 


attached  to  oesophagus  by  thin  muscular  and  connective 
tissue  fibers  and  innervated  by  several  nei^ve  branches. 
Esophageal  gland  formed  of  deeply  glandular  dorso-ven- 
tral  folds,  small  relative  to  salivary  glands,  of  the  same 
color  as  surrounding  tissues  and  esophagus.  Stomach 
large,  U-shaped,  similar  to  that  of  Ci/mathiin  nicolxiri- 
cum  (Houbrick  and  Fretter,  1969)  in  external  moqihol- 
ogy,  too  poorly  preserved  to  reveal  details  of  internal 
morphology,  except  that  the  ducts  of  the  digestive  gland 
are  paired,  closely  spaced,  and  situated  in  the  middle 
region  of  the  stomach.  Posterior  duct  twice  the  diameter 
of  anterior  duct.  After  leaving  stomach  intestine  curves 
dorsally,  passing  under  then  along  posterior  edge  of  ne- 
phridium  towards  rear  of  mantle  cavity  (figures  24,  25, 
m). 

Female  reproductive  system  (Figures  25,  27-30): 

The  only  specimen  dissected  (Paratype  1)  was  a  mature 
female.  Pallial  oviduct  consists  of  small  albumen  gland 
(figure  25,  ag)  along  outer  wall  of  nephridium,  large  cap- 
sule gland  (figures  25,  27,  eg)  fining  right  wall  of  pallial 
cavity.  Lumen  of  capsule  gland  small  anteriorly  (figure 
29),  broad  posteriorly  (figure  28),  at  juncture  to  albumen 
gland.  No  seminal  receptacles  were  identified,  possibly 
due  to  poor  fixation.  Bursa  copulatrix  (figures  27,  29,  30, 
be)  large,  long,  spanning  more  than  half  length  of  cap- 
sule gland.  Pallial  oxiduct  opens  (figures  27,  29,  go)  by 
long,  narrow  slit  at  midlength. 

Male  reproductive  system  (Figure  31):  The  repro- 
ductive system  of  a  male  specimen  lacking  shell  (para- 
type 2)  was  examined.  Testes  occupy  upper  half  of 
whorls  of  visceral  mass  above  posterior  border  of  stom- 
ach, giving  rise  to  seminal  duct.  Seminal  duct  forms 
large,  convoluted  seminal  vesicle  before  entering  mantle 
cavity.  Duct  descends  to  floor  of  mantle  cavity  at  mid- 
length,  opening  to  form  groove  with  thickened,  obvious- 
ly glandular  walls  leading  to  penis  base.  Penis  (figure  31. 
p)  broad,  dorsoventrally  flattened,  with  seminal  groove 
(figure  31,  sgr)  nmning  along  inner  edge,  around  the 
bluntly  rounded  distal  edge,  halfvvay  down  the  outer 
edge,  terminating  in  a  verv  small,  inconspicuous  papilla 
(figure  31,  pap). 

Type  locality:  Victoria  Land,  Balleny  Islands,  Sturge 
Island,  RA'  Eltanin.  .st.  1949,  66252'S,  164a32'E,  \n 
2507-2525  m,  5  February  1967. 

Type  material:  Holotype,  USNM  898683  (figures  1- 
5,  shell  and  operculum  only,  soft  parts  not  present),  from 
the  type  locahty;  paratype  1,  USNM  896131  (figures  6- 


Figures  15-23.  Obsairanella  papijrodes  new  species.  Parat\pe  1,  USNM  8961.31.  15.  Distal  end  of  the  proboscis  opened  dorsally. 
16-18.  Left  jaw.  16.  Entire  inner  surface  of  jaw.  17.  Enlarged  section  of  the  upper  edge.  18.  Enlarged  section  of  the  surface  in 
the  middle  part  of  the  jaw.  19-22.  Radiila.  19.  View  spanning  width  of  radular  ribbon.  20.  Left  marginal  teeth.  21.  Rachidian  anil 
lateral  teeth.  22.  Right  lateral  and  marginal  teeth,  tilted  to  .30°.  23.  Ultrastnicture  of  the  shell  (Paratvpe  3.  USNM  886105). 

aoe,  anterior  oesophagus;  If,  lateral  folds;  od,  odontophore;  ca.  columnar  shell  layer;  ccl,  comarginal  crossed  lamellar  shell  layer;  j, 
jaw;  It,  lateral  teeth;  mt,  marginal  teeth;  r,  radula;  rcl,  radial  crossed  lamellar  shell  layer;  rt,  racliidian  tooth;  tm,  tensor  muscles. 


Page  108 


THE  NAUTILUS,  Vol.  114,  No.  3 


ov 


1  cm 


Figures  24-31.  Ohscuranella  papijrodes  new  species.  Paratope  1,  USNM  896131.  24.  Neutral  and  25.  dorsal  news  of  animal 
removed  from  the  shell.  26.  Antero-dorsal  \iew  of  the  head.  27.  Mantle  complex,  opened  from  left  side  and  reflected.  28-30. 
Schematic  sections  through  the  pallial  gonoduct.  31.  Penis  (Parat)'pe  2). 

a,  anus;  ag,  albumen  gland;  be,  bursa  copulatrix;  eg.  capsule  gland;  cm,  columellar  muscle;  cnie,  cut  mantle  edge;  ct.  ctenidium; 
dg,  digestive  gland;  go,  genital  opening;  hg,  hypobranchial  gland;  in,  intestine;  me,  mantle  edge;  n,  nephridiuni;  no,  nephrichal 
opening;  op,  operculum;  os,  osphradium;  ov,  ovary;  p,  penis;  pap,  papilla;  pr,  proboscis;  re,  rectum;  s,  siphon;  se,  siphonal  edge; 
sgr,  seminal  groove;  st,  stomach;  tn,  cephalic  tentacle. 


Y.  I.  Kantor  and  M.  G.  Harasewvch,  2()()() 


Page  109 


5  mm 


Figures  32-35.  Obscitranella  papijrodes  new  species.  Anterior  alimentary  system,  Paratype  1,  USNM  896131  32.  Proboscis 
(opened  ventrallv)  and  organs  of  cephalic  haeniocoel.  33.  Anterior  part  of  the  proboscis,  opened  ventrally.  34.  Transverse  section 
through  anterior  oesophagus,  viewed  from  anterior  35.  Esophageal  gland,  opened  dorsally.  Dashed  line  indicates  \entral  midline. 

alsg,  anterior  lobe  of  the  salivary  gland;  ao,  anterior  aorta;  aoe,  anterior  oesophagus;  bm,  buccal  mass;  bt,  buccal  tube;  cm,  circular 
muscle  layer  of  the  proboscis  wall;  j,  jaw;  If,  longitudinal  fold  of  anterior  oesophagus;  !m,  longitudinal  nuiscle  layer  of  the  proboscis 
wall;  mo,  mouth  opening;  nr,  nerve  ring;  ode,  odontophore  caitilages;  oeg,  oesophageal  gland;  plsg,  posterior  lobe  of  the  salivary 
gland;  pn,  proboscis  nerve;  poe,  posterior  oesophagus;  pw,  proboscis  wall;  rs,  radiilar  sack;  sd,  salivary  duct;  tm,  tensor  muscles. 


7,  9  pecimen,  anatomical  de.scription.s  based  on  diis 
specimen),  paratype  2,  USNM  896139  {6  specimen, 
shell  not  present,  male  reproductive  system  based  on 
this  specimen),  both  irom  south  of  Hjort  Seamount, 
RA'  Eltanin,  stn.  1964,  59°5H'S.  ISS^arW.  in  29S5-2992 
m,  10  February  1967;  paratype  3,  USNM  886105  (fig- 
ures 11-12,  sex  undetermined)  Victoria  Land,  south  of 
Scott  Island  Bank,  IW  Eltanin.  stn.  1939,  69203'S, 
I7904IE,  in  3519-3.596  m,  1  February  1967. 

Other  material  examined:  USNM  901317  (Figs  13- 
14,    1    specimen,    sex    undetermined)    Victoria    Land, 


Moubray  Bay,  RA^  Eltanin.  stn.  2002,  72218'S,  177e:35'E, 
in  2005-2010  m,  11  January  1968;  USNM  870610  (fig- 
ures 8-10,  2  dead  shells)  Antarctic  Peninsula,  IW  £/- 
tanin,  stn.  1003,  62^41S,  54243W,  in  210-220  m,  15 
March  1964. 

Etymology:  papijrodes — made  from  papyrus,  refer- 
ring to  the  thinness  of  the  shell. 

Remarks:  The  type  series  oi  ObsciirancUa  papijrodes, 
n.  sp.,  consists  of  lour  specimens,  incluchng  one  paratvpe 
that  lacks  a  shell.  All  were  collected  hving  on  the  abyssal 


Page  110 


THE  NAUTILUS,  Vol.  114,  No.  3 


Figure  36.     Geographic  and  bath\nietric  distribution  of  Obsairnnelln  papijrodes  new  species. 


plain  off  the  Ross  Sea.  We  are  provisionally  attributing 
three  additional  specimens  to  this  species.  One  live-col- 
lected specimen  (figure  13;  USNM  901317),  also  from 
the  abyssal  plain  off  the  Ross  Sea,  was  considered  by 
Dell  (1990:199)  to  be  congeneric  but  not  conspecific 
with  Obsctiranella  papijrodes  n.  sp.  (which  he  identified 
as  Bathi/domiis  ohtcctus  Thiele,  1912)  because  of  its 
more  elongated  shell,  longer  siphon;il  canal,  and  angular 
shoulder.  Even  the  earliest  teleoconch  whorls  of  this 
specimen  appear  angular  because  of  a  prev;ilent  spiral 
cord  along  the  periphery.  This  is  exaggerated  by  a  break 
in  the  shell  and  subsequent  repair,  evidenced  bv  a  thick- 
ened scar  on  the  posterior  part  of  the  last  whorl.  The 
elongated  shell  and  prominent  siphonal  canal  are  fea- 
tures shared  with  an  immature  paratvpe  (paratype  3,  fig- 
ures 11-12)  of  O.  papi/rodes.  This  is  the  only  specimen 
of  Obscuranella  with  some  portion  of  the  earlv  whorls 
intact  (figure  14).  The  protoconch  (2.0  mm  estimated 
diameter)  is  eroded  and  replaced  by  a  plug,  but  the  ear- 
hest  teleoconch  whorls  are  well  preserved  and  clearly 
show  spiral  cords. 

Two  dead  collected  shells  (figures  8-10;  USNM 
870610),  labeled  as  coming  from  upper  slope  depths 
(210-220  m)  off  the  Antarctic  Peninsula,  closely  match 
the  moiphology  of  this  new  species.  We  regard  these 
specimens  to  represent  O  papijrodes,  but  are  skeptical 
of  the  accuracy  of  the  locality  data.  Not  only  is  this  lo- 
cation on  the  opposite  side  of  the  Antarctic  continent 
from  all  records  of  live  collected  O.  papijrodes,  it  is  also 
from  much  shallower  depths  (220  m  vs.  2000-1-  m). 

DISCUSSION 

Ohscuraiudla  can  readily  be  attributed  to  the  superfam- 
ily  Tonnoidea  on  the  basis  of  its  pyriform  shell  with  large 


aperture  and  conspicuous,  if  short  siphonal  canal;  its  ex- 
tensible proboscis;  its  taenioglossan  radula;  its  large  sal- 
ivarv  glands  composed  of  moqihologicallv  differentiated 
anterior  and  posterior  lobes  and  sahvarv  ducts  that  pass 
through  the  nen'e  ring,  as  well  as  its  undifferentiated 
oesophageal  gland.  It  can  be  excluded  from  Ficidae, 
which  was  removed  from  Tonnoidea  and  elevated  to  su- 
perfamily  status  by  Riedel  (1994),  by  its  high  spire,  lack 
of  long  siphonal  canal,  and  also  because  Ficidae  is  char- 
acterized by  small,  tubular  sali\arv  glands.  Similarly,  it 
can  be  excluded  from  Laubierinidae,  a  family  diagnosed 
by  its  monopectinate  osphradium  and  excluded  from 
Tonnoidea  by  Bandel  and  Riedel  (1994),  by  its  nearly 
symmetrical,  bipectinate  osphradium. 

The  shell  of  Obscuranella  suggests  an  affinitv' with  the 
deep-sea  family  Pisanianuridae  (originall)'  proposed  as  a 
subfamily  of  RanelHdae  by  Waren  and  Bouchet,  1990, 
transferred  to  Laubierinidae  by  Bandel  and  Riedel, 
1994,  and  elevated  to  family  status  by  Beu,  1998)  by 
\irtue  of  its  smooth  shell  lacking  regular  varices  and 
weakly  defined  anterior  canal.  The  operculum  of  Pis- 
anianiira  is  shghtly  coiled  but  has  a  terminal  nucleus,  as 
does  Obscuranella.  Howe\er,  the  rachidian  teeth  of  Ob- 
scuranella lack  the  lateral  basal  denticles  present  in  Pis- 
anianuridae (e.g.  Waren  and  Bouchet,  1990:figs.  25-27), 
Bursidae  (e.g.  Waren  and  Bouchet,  1990:figs.  6,  8),  Ton- 
nidae  (e.g.  Waren  and  Bouchet,  1990:figs.  9-14),  and 
Laubierinidae  (e.g.  Waren  and  Bouchet,  1990:figs.  41- 
44),  but  absent  in  Cassidae  (e.g.  Waren  and  Bouchet, 
1990:figs.  15,  16,  18),  Personidae  (e.g.  Beu,  1998:fig. 
15.140  E),  and  Ranellidae  (e.g.  Waren  and  Bouchet, 
1990;figs.  28,  30,  32,  40). 

The  shell  of  Obscuranella  somewhat  resembles  that  of 
Oocorijs  sulcata  Fischer,  1883  (Oocorythinae,  Cassidae) 


Y.  I.  Kantor  and  M.  G.  Harasewvch,  2000 


Page  111 


(see,  eg.  Bouchet  aiu!  W'aivii,  19y3:fig,s.  1936-1943),  and 
some  ranellids,  such  as  Ar^obticcinmn  pnstulosuin 
(Ughtfoot,  1786)  (.see  e.^.  Beii,  1998:fig.  15.12  D).  The 
operculum  of  Obscuranella  has  a  shaqjly  pointed,  ter- 
minal nucleus,  and  differs  from  the  spiralK'  coiled  oper- 
culum of  Oocon/s  (W'aren  and  Bouciu't.  1990:  fig.  66). 
In  adult  Ai-fiobuccinuin  the  nucleus  ot  the  operculum  is 
subcentnJ,  hut  in  ver\'  young  specimens  of  A.  pustulo- 
siim  (Waren  and  Bouchet,  I990:fig.  69)  the  nucleus  is 
terminal.  Howexer,  the  operculmn  of  Ohscuraiwlla  is 
distincti\e  in  its  veiy  small  size,  relatixe  to  the  aperture, 
and  in  having  straight,  anteriorly  converging  margins. 

The  anatomv  of  Obscuranella  is  tvqiicallv  tonnoidean, 
most  closelv  resembling  that  of  the  ranelliil  Ci/inatiinn 
(Houhrick  and  Fretter,  1969).  Obscuranella  can  be  dis- 
tinguished anatomically  from  the  Tonnidae  by  its  lack  of 
a  buccal  gland,  and  from  Pisaniamiridae  and  Laubierin- 
idae  hv  its  lack  of  long  proboscis  retractor  muscles  that 
pass  through  the  nen'e  ring. 

We  assign  this  genus  to  the  family  Ranellidae  because 
of  it  general  similaritx'  to  Arfiobucchuun  in  shell  form, 
radular  moi-pholog\,  and  gross  anatomy.  Moreover,  Ra- 
nellidae is  the  onlv  tonnoidean  familv  to  occur  in  Ant- 
arctic waters — Fusitriton  ma^icllanicus  (Roding,  1798),  a 
species  with  a  wide  geographic  range,  has  been  taken 
from  several  stations  in  the  Weddell  and  Ross  Quadrants 
(e.g.  USNM  S96058,  USNM  896103,  USNM  896277, 
USNM  898520— see  Polar  Invertebrate  Catalog  http:// 
www.nmnh.si.edu/cgi-bin/wdlViz/pci/form).  Obscuranel- 
la papi/rodcs  represents  the  first  record  of  Ranelfidae 
from  abyssal  depths. 


ACKNOWLEDGMENTS 

This  study  was  supported  by  a  USAP  Research  Program 
Award  from  the  Biological  Collections  from  Polar  Re- 
gions at  the  National  Museum  of  National  History  [Na- 
tional Science  Foundation  (Office  of  Polar  Programs) 
Cooperative  Agreement  OPP-9509761].  We  are  grateful 
to  Drs.  Alan  Beu  and  Klaus  Bandel  for  their  constructive 
reviews  of  this  manuscript. 


LITERATURE  CITED 

Aii(lrew.s,  K.  B.,  A.  M.  Page  and  ],  13.  Taylor  1999.  Tfie  fine 
.stnictiire  and  function  of  the  anterior  foregnt  glands  of 
Ci/uuitiiim  iiitenneclius  (Cassoidea:  Ranellidae).  Journal  of 
\iolluscan  Studies  6.5:1-19. 

Bandel,  K.  and  F.  Riedel,  1994.  Classification  of  fossil  and  Re- 
cent CaKptraeoidea  (Caenogastropoda)  with  a  discussion 
on  neoniesogastropod  phylogeny.  Berliner  geowissen- 
schafdiche  Abhandlungen  (E)  13:329-367. 

Beu,  A.  G.  1981.  Australian  gastropods  of  the  family  Bursidae. 
Part  1.  The  families  of  Tonnacea,  the  genera  of  Bursidae, 
and  revision  of  species  prexiously  assigned  to  Tutufa  Jous- 
seaume,  1881.  Records  of  the  Australian  Museinn  33:248- 
324. 

Beu,  A.G.  1998.  Superfamily  Tonnoidea.  In:  Beesley,  P  L.,  G. 
J.  B.  Ross  and  A.  Wells  (eds).  MoUusca:  The  Southern 
Synthesis.  Fauna  of  Australia,  volume  .5,  part  B.  CSIRO 
Publishing,  Melboum,  pp.  792-803. 

Bouchet,  P.  and  A.  Waren.  1993.  Revision  of  the  northeast 
Atlantic  hathyal  and  abyssal  Mesogastropoda,  Bolletino 
Malacologico,  Supplemento  3:.580-840. 

Day,  J.  A.  1969.  Feeding  of  the  cvniatiid  gastropod  Argofci/c- 
cintim  argus,  in  relation  to  the  stRicture  and  secretion  of 
the  proboscis  glands.  American  Zoologist  9:909-916. 

Dell,  R.  K.  1990.  Antarctic  MoUusca.  Bulletin  of  the  Royal 
Society  of  New  Zealand  27:1-311. 

Houhrick,  J.  R.  and  V.  Fretter  1969.  Some  aspects  of  the  func- 
tional anatomy  and  hiologv'  of  Cipnatium  and  Bursa.  Pro- 
ceedings of  the  Malacological  Society  of  London  .38:41.5- 
429. 

Hughes,  R.  N.  and  H.  R  I.  Hughes.  1981.  Moiphological  and 
behavioural  aspects  of  feeding  in  the  Cassidae  (Tonnacea, 
Mesogastropoda).  Malacologia  20:385—102. 

Riedel,  F.  1994.  Recognition  of  the  superfamily  Ficoidea  Meek 
1864  and  definiton  of  the  Thalassocvniidae  fam.  nov.  (Gas- 
tropoda). Zoologische  Jahrbiicher  121:457-474, 

Riedel,  F.  1995.  An  outline  of  cassoidean  phylogeny  (MoUusca, 
Gastropoda).  Contributions  to  Tertiary  and  Quateniary 
Geolog)'  .32:97-1.32. 

Waren,  A.  and  P.  Bouchet.  1990.  Laubierinidae  and  Pisanianu- 
rinae  (Ranellidae),  two  new  deep-sea  taxa  of  the  Tonno- 
idea (Gastropoda:  Prosobranchia).  The  Veliger  33:56-102. 

Weber,  H.  1927.  Der  Darm  von  Dolium  galea  L.,  eine  ver- 
gleichend  anatomische  Untersuchung  unter  besonderer 
Beriicksichtigung  der  rn^()»(i/»i-Arten.  Zeitschrift  fiir 
Morphologic  tmd  Okologie  der  Tiere  8:66.3-804. 


THE  NAUTILUS  114(3):112-116,  2000 


Page  112 


Host-tree  selection  by  Florida  tree  snails,  Ligiiiis  fasciatus 
(Miiller,  1774),  in  Big  Cypress  National  Preserve,  Florida,  USA 


Robert  E.  Bennetts' 

Department  of  Wildlife  Ecology  and 

Conservation 
Florida  Cooperative  Fish  and  Wildlife 

Research  Unit 
P.O.  Box  110450,  University  of  Florida 
GainesN-ille,  FL  32611-0450  USA 
bennetts@tour-du-valat.com 


Steven  A.  Sparks 
Deborah  Jansen 

Big  Cvpress  National  Preserve 
HCR'61,  Box  110 
Ochopee.  FL  34141  USA 


ABSTRACT 

It  has  frequently  been  suggested  that  the  Florida  tree  snail, 
Liguiis  fasciatus,  exhibit  preferences  for  specific  host  trees,  al- 
though most  accounts  of  host-tree  selection  are  based  on  an- 
ecdotal observations.  We  assessed  the  relative  use  of  host  trees 
in  eight  hammocks  in  the  Big  C)'press  National  Preserve  for 
which  the  relative  availability  of  each  potential  host-tree  spe- 
cies was  known.  Based  on  a  total  of  1,464  tree  snail  obsei-va- 
tions  on  21  species  of  host  tree  within  our  study  plots,  host 
trees  were  not  used  in  proportion  to  their  availability  when  the 
expected  values  are  derived  either  from  the  number  of  trees 
or  from  basal  area.  Of  the  most  common  tree  species  on  our 
study  area,  wild  tamarind  (Ltjsiloma  latisiliqurn)  was  consis- 
tentlv  used  in  excess  of  its  availabilitv.  Wild  lime  (Zanthoxi/him 
fagarn),  was  not  a  common  tree  species,  but  was  also  used  far 
in  excess  of  its  availabilitv.  Pigeon  plum  {Coccoloba  diversifolia) 
and  gimibo-limbo  {Bursern  simanibti)  were  consistently  used 
less  than  expected  from  their  respective  availabilities.  Based  on 
Akaike's  Information  Criterion  (AIC),  the  most  parsimonious 
loglinear  model,  after  accounting  for  availabilitv,  was  one  that 
included  only  tree  species  effects.  In  contrast  to  the  model 
selected  using  AIC,  likelihood  ratio  tests  indicated  that  there 
may  also  be  differences  among  hammocks,  but  not  in  relation 
to  a  linear  gradient  of  hurricane  damage.  We  obsers'ed  differ- 
ences in  length  among  3-year-old  snails  on  different  host  trees, 
and  these  differences  were  consistent  with  the  hypothesis  that 
host-tree  selection  reflects  food  availabilitv'.  Snails  found  on 
host-tree  species  that  were  used  in  greater  than  expected  fre- 
quency had  the  greatest  length,  those  found  on  trees  used  less 
than  expected  frequency  were  the  smallest,  and  those  found  in 
proportion  to  their  availability  were  intermediate  in  length. 

Additional  key  iiords:  Big  Cypress  National  Preserve,  host 
tree,  humcane,  Liguus  fasciatus.  tree  snail,  Florida,  hammock. 


INTRODUCTION 

Snails  of  the  genus  Liguus  are  native  to  Cuba,  Hispan- 
iola,  and  Florida  (Pilsbry,  1912).  The  Florida  tree  snail 

'  Author  for  correspondence.  Present  address:  Station  Biolo- 
glque  de  la  Tour  du  Valat,  Le  Sambuc.  F-I320()  Aries,  France 


Liguus  fasciatus  (Mtiller,  1774)  is  found  within  a  restrict- 
ed part  of  this  range,  primarily  some  islands  within  the 
Florida  Keys,  the  Atlantic  coastal  ridge.  Everglades  Na- 
tional Park,  and  Big  Cypress  National  Preserve.  Primar- 
ily because  of  agricultural  and  urban  expansion,  only  a 
few  isolated  populations  remain  outside  of  protected 
public  lands  and  a  few  isolated  islands  in  the  Keys.  Con- 
sequently, they  are  hsted  as  a  species  of  special  concern 
by  the  state  of  Florida.  Thus,  understanding  the  rela- 
tionship between  snails  and  their  habitat  is  essential  for 
the  conservation  and  management  of  remaining  popu- 
lations. 

In  the  Everglades,  L.  fasciatus  is  found  on  islands  of 
subtropical  hardwood  trees  and  scrub  known  as  ham- 
mocks. Hammocks  are  widely  scattered  throughout 
South  Florida  and  may  be  separated  by  water,  sawgrass, 
or  other  habitat  types  such  as  pines.  It  is  widely  beUeved 
that  Florida  tree  snails  exliibit  strong  preferences  for 
host  trees,  particularly  smooth-barked  trees  upon  which 
their  algal  food  source  can  easily  be  grazed  (Pilsbry, 
1912).  In  particular,  snails  are  suggested  to  exhibit  pref- 
erence for  Jamaica  dogwood  (Piscidia  piscipula)  in  the 
Florida  Keys  and  wild  tamarind  (Li/siloma  latisiliqurn) 
on  the  mainland  (Pilsbry,  1912),  although  most  accounts 
of  host-tree  selection  are  based  on  anecdotal  observa- 
tions, and  few  quantitative  data  are  available.  Voss  (1976) 
reported  numbers  of  snails  found  on  each  tree  species 
in  2  hammocks  on  the  coastal  ridge.  However,  he  did 
not  account  for  the  aviulabihty  of  these  host  trees;  thus, 
cUfferential  selection  can  not  be  reliably  assessed.  Brown 
(1978)  is  the  only  study  of  which  we  are  aware  that  com- 
pared the  use  and  availabihty  of  host  trees,  although  this 
was  restricted  to  1  hammock  in  Everglades  National 
Park.  Here  we  assess  the  relative  use  of  host  trees  in 
eight  hammocks  in  the  Big  Cvpress  National  Preserve 
for  which  the  relative  availability  of  each  tree  species  was 
knowni. 

Brown  (1978)  also  hypothesized  that  differences  in 
(juantitv'  and  qualitv'  of  food  resources  of  different  host- 


R.  E.  Bennetts  ct  al,  2000 


Page  113 


tree  species  mav  result  in  size  ditferences  among  snails 
using  olifferent  host  trees,  although  she  was  unable  to 
statisticallv  confirm  such  clifferences.  We  tested  this  hy- 
pothesis h\  comparing  size  cbfferences  among  snails 
found  on  host-tree  species  that  were  used  (1)  in  greater 
than  expected  fretjuencies  (selected),  (2)  did  not  differ 
from  expected  frequencies  (neutral),  and  (3)  in  less  than 
expected  frequencies  (avoided). 

MATERIALS  AND  METHODS 

Our  studv  population  was  in  Big  Cypress  National  Pre- 
sene  (BCNP).  The  Pinecrest  area  of  the  BCNP  contains 
one  of  the  largest  remaining  populations  of  Florida  tree 
snails.  Within  BCNP,  136  hanunocks  have  been  indi\'id- 
ually  identified  and  mapped  (Pilsbiy,  1946);  a  few  others 
may  e.xist  that  have  not  been  identified.  We  selected  8 
of  the  136  hammocks  from  this  pof)l  for  our  study  sites. 
Eight  hanunocks  proxaded  a  reasonable  representation 
of  the  potential  variability  and  represented  a  logistically 
feasible  number  of  sites.  This  study  was  initiated  to  in- 
vestigate the  survival  and  movements  of  Florida  tree 
sn;iils  following  Hurricane  Andrew.  Hurricane  Andrew 
crossed  southern  Florida  from  east  to  west  on  24  August 
1992.  The  eye  of  the  storm  passed  along  the  southern 
boundarv  of  our  study  area  creating  a  decreasing  lati- 
tudinal gradient  of  damage  toward  the  north  with  in- 
creasing distance  from  the  eye.  Because  we  were  inter- 
ested in  the  potential  effects  of  Hurricane  Andrew,  we 
also  wanted  our  study  sites  to  reflect  the  latitudinal  gra- 
dient of  hurricane  impacts.  Consequently,  we  divided 
the  initial  pool  of  hamiuocks  into  4  zones  reflecting  clus- 
ters of  hammocks  positioned  along  a  gradient  from  little 
or  no  hurricane  impact  to  severe  impact  (Bennetts  ct  al., 
in  press).  There  also  exists  considerable  variabifity  in  the 
size  of  these  hammocks,  which  we  beheved  could  ha\'e 
greatly  influenced  their  susceptibihty  to  damage  by  Hur- 
ricane Andrew  and  consequently  affected  tree  snails. 
Therefore,  we  digitized  each  hammock  from  USGS  7.5- 
minute  ortho  photos  and  used  a  Geographic  Information 
System  (CIS)  to  estimate  the  appro.ximate  size  of  each 
hammock.  We  then  used  stratified  random  sampling  to 
select  1  hammock  of  greater  size  than  the  median  size 
and  1  hammock  of  lesser  size  from  each  of  the  4  impact 
zones.  Thus,  our  sample  reflected  the  latitudinal  gradi- 
ent of  hurricane  damage  and  was  balanced  among  larger 
and  smaller  hammocks. 

We  estabhshed  two  80  m-  study  plots  (sub-samples) 
in  each  hammock.  These  plots  were  randomly  located  in 
parts  of  the  hammock  where  tree  snails  occurred.  The 
center  of  each  plot  was  marked  with  a  steel  rod  and  we 
subsequently  marked  all  trees  with  a  diameter  at  breast 
height  (DBH)  >  5  cm  within  a  5  m  radius  of  the  plot 
center.  We  recorded  the  species  and  DBH  of  each  live 
tree  within  these  plots. 

During  each  of  seven  sampling  occasions,  we  searched 
for  all  tree  snails  within  each  plot,  and  as  part  of  a  con- 
current study  on  movements  (Bennetts  et  al..  in  press), 
for  marked  snails  within  an  extended  radius  of  approxi- 


mately 20  HI  around  the  plot.  Sampling  was  conducted 
each  autumn  and  spring  from  autumn  1993  through 
spring  1996.  Our  sampfing  occasions  were  intended  to 
reflect  the  biology  of  the  animal  and  to  minimize  the 
potential  for  handling  to  influence  a  snail's  survival.  Our 
spring  sampling  occasion  was  conducted  as  soon  as  pos- 
sible after  emergence  of  the  snails  from  aestivation.  This 
coincided  with  the  onset  of  the  annual  rainy  season.  At 
this  time  snails  begin  to  put  on  new  growth,  which  be- 
comes yerv'  fragile  as  it  extends  as  a  thin  layer  from  the 
previous  growth.  Thus,  our  sampling  was  intended  to 
precede  the  period  during  which  time  shells  are  fragile, 
although  some  growth  had  occurred  on  some  snails.  Our 
autumn  sampfing  occasion  coincided  with  the  onset  of 
the  dry  season  just  prior  to  aestivation.  At  this  time  the 
summer's  growth  had  been  terminated,  the  shells  had 
thickened,  but  the  animals  had  not  yet  entered  aestiva- 
tion. Animals  that  were  visually  determined  to  be  in  aes- 
tivation (<1%)  by  presence  of  an  epiphragm  (dried  mu- 
cus membrane  across  the  aperture)  were  not  disturbed. 
For  each  snail  we  determined  its  age  from  annual 
growth  scars  (Tuskes,  1981),  measured  its  length  from 
tip  to  tip  parallel  to  the  axis  and  width  perpentficular  to 
the  axis  at  its  widest  point.  We  also  recorded  the  host 
tree  from  which  it  was  collected.  We  then  returned  each 
snail  to  the  same  host  tree  by  placing  them  in  a  conical 
paper  cup  attached  to  the  trunk  of  the  tree. 

Statistical  Analyses:  As  a  prefiminary  analysis  we  test- 
ed for  an  association  between  use  and  availabiliK'  of  all 
host-tree  species  within  our  study  plots  using  a  chi- 
square  goodness  of  fit  test.  This  was  done  for  a  descrip- 
tive account  of  the  relative  selection  of  all  host  species; 
however,  cells  having  expected  values  <5  could  produce 
unreliable  test  statistics  (Cochran,  1954).  Thus,  for  fur- 
ther analyses  including  adcbtional  effects  of  hammock  of 
hurricane  influence,  we  pooled  cells  with  expected  val- 
ues <5  into  an  "other "  category. 

We  tested  whether  host  trees  were  used  in  proportion 
to  their  availabihty  using  a  loghnear  model.  To  account 
for  host  tree  availability  we  used  either  the  log  number 
of  trees,  or  the  log  basal  area  as  an  offset  (Agresti,  1990; 
McCullagh  and  Nelder,  1989).  Thus,  the  frequency  of 
snails  observed  was  modeled  per  available  number  and 
basal  area  of  each  species.  We  used  basal  area  in  addition 
to  the  number  of  trees  because  the  surface  area  of  trees 
are  generally  related  to  its  bas;il  area  (Whittaker  and 
WoodweU,  1967;  Brown,  1978).  We  then  tested  whether 
the  number  of  snails  was  influenced  by  tree  species,  and 
then  whether  this  association  cfiffered  among  hammock 
or  hurricane  zone. 

Model  selection  was  based  on  Akaike's  Information 
Criterion  (AlC)  (Akaike,  1973;  Shibata,  1989),  which  is 
defined  as:  —llni'/)  +  2np,  where  —2  ln(l''),  repre- 
sents the  Ukehhood  ratio  between  the  given  model  and 
a  corresponding  saturated  model;  thus,  represents  a 
measure  of  model  fit.  The  second  term,  2np.  is  the  num- 
ber of  parameters  estimated  in  the  model  antl  can  be 
viewed  as  a  cost  for  adding  excessive  parameters  that  do 


Page  114 


THE  NAUTILUS,  Vol.  114,  No.  3 


Table  1.  Common  and  scientific  name  of  the  host-tree  species  within  our  study  plots.  Also  shown  are  the  total  number  of  trees, 
total  basal  area,  and  total  number  of  snails  observed  for  each  species. 


Common  name 

Scientific  name 

Acronvin 

No.  trees 

Basal  area 

No.  snails 

Cocoplum 

Chnjsobalanus  icaco 

Ci 

1 

174 

1 

Dahoon  holly 

Ilex  cassine 

Ic 

2 

365 

21 

Gumbo  Umbo 

Bursera  simaniha 

Gl 

26 

5423 

85 

Hackberry' 

Celtis  laevigflta 

CI 

2 

212 

15 

Inkwood 

Exothea  paniculata 

Ep 

2 

93 

3 

Lancewood 

Nectandra  coriaceae 

No 

25 

2362 

52 

Wild  lime 

Zanthoxi/him  faopra 

Zf 

1 

79 

20 

Li\e  oak 

Quercus  virainiana 

Qv 

2 

480 

2 

Wild  tamarind 

Li/siloina  latisiqua 

LI 

128 

41820 

983 

Mastic 

Mastichodendron  foetidissimii  m 

Mf 

2 

711 

8 

Paradise  tree 

Simarouba  gjmica 

Sg 

2 

240 

11 

Pigeon  plum 

Coccoloba  diversifolia 

Cd 

80 

9489 

145 

Poisonwood 

Metopium  taxifera 

Mt 

2 

252 

5 

Pond  apple 

Annona  glabra 

Ag 

1 

47 

1 

Red  bay 

Persea  borbonia 

Pb 

1 

620 

9 

Red  stopper 

Eugenia  rhombea 

Er 

1 

22 

1 

Satinleaf 

Ch  n/.wphijUu  m  olivifomie 

Co 

1 

85 

1 

Simpson  stopper 

Mtjrcianthcs  fraarans 

Mf 

6 

723 

18 

Strangler  fio; 

Fiats  aurea 

Fa 

1 

90 

2 

White  stopper 

Eugenia  axillaris 

Ea 

7 

273 

9 

\\'illow  bustic 

Biimelia  salicifolia 

Wb 

19 

2646 

72 

not  contribute  substantially  to  the  overall  model  fit. 
Thus,  AIC  indicates  an  appropriate  balance  between 
precision  and  bias  (i.e.,  over  and  under-fitting  the  model) 
from  a  set  of  candidate  models  (Bumham  and  Anderson, 
1998).  In  contrast  to  AIC,  which  is  used  as  an  optimi- 
zation tool  for  comparison  among  models,  we  also  used 
Ukelihood-ratio  tests  (LRTs)  to  test  for  specific  effects  of 
interest  (Lebreton  et  al,  1992;  Bumham  and  Anderson, 
1998). 

The  selection  of  host  trees  may  be  influenced  by  the 
quantity  and  quality  of  food  resources  (Brown.  1978), 
which  mav  in  turn  result  in  size  differences  of  snails 
using  different  host  trees.  We  tested  this  hypothesis  by 
comparing  size  differences  among  snails  found  on  host- 
tree  species  that  were  used  (1)  in  greater  than  expected 
frequencies  (selected),  (2)  did  not  differ  from  expected 
frequencies  (neutral),  and  (.3)  in  less  than  expected  fre- 
quencies (avoided).  We  compared  the  size  of  snails  (i.e., 
length  and  width)  within  each  age  class  among  host-tree 
species  using  a  fixed-effects  analysis  of  variance  (ANO- 
VA)  model.  For  this  analysis  we  were  not  concerned  with 
the  avaHabihty  of  host  trees,  only  species.  Consequendy, 
we  included  all  snails  encountered  on  our  studv  ham- 
mocks for  this  analysis,  regardless  of  whether  they  were 
found  within  the  designated  plots  (used  to  compare  se- 
lection with  availabihty)  or  the  extended  radius  (used  for 
analyses  of  movement)  (Bennetts  ct  al,  in  press).  How- 
ever, we  restricted  our  sample  to  snails  collected  during 
the  autumn  sampling  occasion  to  minimize  any  con- 
founding attributable  to  variation  in  the  extent  of  new 
growth  during  spring. 


RESULTS 

We  observed  1464  tree  snails  on  21  host-tree  species 
within  our  study  plots  (Table  1).  Host  trees  were  not 
used  in  proportion  to  their  availabiUtv  when  the  expect- 
ed values  are  derived  either  from  the  number  of  trees 
(X'  =  554.18,  20  df  P  <  0.001)  or  from  basal  area  (x' 
=  296.14,  20  df  P  <  0.001).  Of  the  common  tree  spe- 
cies on  our  study  area,  wild  tamarind  was  consistently 
used  in  excess  of  its  availability  (figure  1).  Wild  Ume 
(Zant]uixijlum  fagara).  was  not  common,  but  our  obser- 
vations indicated  that  it  was  used  far  in  excess  of  its 
availabilitv',  regardless  of  which  measure  of  availabihty 
was  used.  Although  the  sample  for  this  species  used  in 
our  analysis  included  only  1  tree,  we  observed  others  on 
our  study  area  that  had  similar  high  use.  In  contrast, 
pigeon  plum  [Coccoloba  diversifolia)  and  gumbo-hmbo 
(Bursera  simaniha)  were  relatively  common,  but  were 
consistentlv  used  less  than  expected.  Willow  bustic  (Bu- 
melia  salicifolia)  and  lancewood  (Nectandra  coriaceae) 
also  were  relatively  common  in  our  study  plots,  but  our 
results  were  conflicting  for  these  species.  In  relation  to 
the  number  of  trees,  willow  bustic  was  used  slightly  less 
than  expected  whereas,  based  on  basal  area,  it  was  used 
shghtlv  more  than  expected.  Lancewood  was  used  less 
than  expected  in  reference  to  the  number  of  trees,  but 
in  proportion  with  its  availabihty-  in  relation  to  basal  area. 
After  pooling  cells  with  low  expected  values,  the  most 
parsimonious  model  based  on  AIC  included  only  tree 
species  effects  (x"  =  101.05,  5  df,  P  <  0.001).  Depar- 
tures from  ex-pected  values  were  consistent  vdth  our  pre- 
hminary  analysis  in  that  wild  tamarind  was  used  in  great- 
er proportion  than  expected  from  its  avaflabifit)-;  pigeon 


R.  E.  Bennetts  et  al,  2000 


Page  115 


LI    Zf    Ic    Pb   CI    Sg   Mf  Fa   Ml  Ag    Ci    Er  Co  Wb  Mf  Ep  Qv  Gl   Ea  Nc  Cd 

Host-Tree  Species 


llll 


■  ■ 


Zf    LI    CI    Ic  Sg  Wb  Ea  Er  Ep  Mf  Fa  Ag  Nc  Mt  Co  Pb  CI  Mf  Qv  Gl  Cd 

Host-tree  Species 

Figure  1.  Adjusted  residuals  (Haberinan  1973)  from  good- 
ness-of-fit  test  between  the  number  of  snails  found  on  different 
host-tree  species  in  relation  to  their  availability.  Host-tree  spe- 
cies are  displayed  in  rank  order  from  selected  in  greatest  pro- 
portion relative  to  its  availabilit)'  to  least.  Expected  values  were 
derived  based  on  the  number  of  trees  of  a  given  species  (top) 
and  the  total  basal  area  for  a  given  species  (bottom).  Positive 
residuals  >1.96  indicate  use  greater  than  expected  from  avail- 
ability and  residuals  <  — 1.96  indicate  use  less  than  expected 
from  a\ailability. 


plum  and  gumbo  limbo  were  used  less  than  expected. 
Lancewood  and  willow  bustic  were  intermediate  as  was 
our  "other"  class.  In  contrast  to  the  model  selected  using 
AIC,  LRTs  indicated  that  there  may  have  been  a  species 
X  hammock  interaction  (x'  =  110.36,  31  df,  P  <  0.001), 
but  not  a  species  X  hurricane  zone  interaction  (x"  = 
9.32,  5  df,  P  =  0.097).  Thus,  our  data  indicated  that 
selection  of  host-tree  species  may  have  differed  among 
hammocks,  but  this  difference  was  not  in  relation  to  a 
gradient  of  hurricane  damage. 

We  found  size  (length)  differences  among  snails  found 
on  host  tree  species  onlv  for  3-year-old  snails  (F^  j,^  = 
3.42,  P  =  0.034)  (figure  2).  Of  the.se,  the  length  of  sniiils 
was  highest  for  snails  found  on  host  trees  that  were  used 
in  greater  frequency  than  their  availabibty  (selected)  and 
lowest  for  snails  found  on  host  trees  that  were  used  less 
than  their  availabihtv'  (avoided).  Sn;uis  found  on  host 
trees  that  were  not  out  of  proportion  to  their  availabihty 


E 
E 


o> 

c 


40  - 

263 

53 

- 

39 

- 

35 

38  - 

37  - 

Selected 


Neutral 


Avoided 


Host-tree  Selection  Class 

Figure  2.  Mean  length  (±SE)  of  .3-vear-old  tree  snails  found 
on  host-tree  species  that  were  used  in  excess  ot  their  avail- 
ability (Selected),  less  than  their  availabilitv'  (Avoided)  and  did 
not  differ  from  availabilitv  (Neutral).  Sample  sizes  are  shown 
for  each  group. 

(neutral)  were  intermediate  in  length,  but  did  not  differ 
from  either  of  the  other  groups  (Fisher's  Least  Signifi- 
cant Difference,  P>0.05).  We  did  not  find  differences 
in  width  among  any  host-tree  species  for  any  age  class 
at  the  a  =  0.05  level. 

DISCUSSION 

Our  data  are  consistent  with  previous  reports  of  wild 
tamarind  being  used  in  excess  of  availabihty;  although 
the  degree  of  this  association  is  highly  sensitive  to  what 
measure  is  used  to  derive  the  e.xpected  values  for  avail- 
ability. When  the  expected  value  is  derived  from  basal 
area,  the  association  is  much  weaker  than  when  the  ex- 
pected value  is  derixed  from  the  number  of  trees.  Ex- 
pected values  derived  from  basal  area  probablv  better 
reflect  the  surface  area  available  tor  foraging.  The  num- 
ber of  stems  would  be  important  if  the  probabifity  of  a 
dispersing  snail  moving  to  a  given  tree  is  dependent  on 
it  encountering  a  trunk  on  the  ground,  although  most 
inter-tree  movement  on  our  study  area  probably  was  via 
intertwined  foliage.  Brown  (1978)  reported  a  remarkably 
similar  result  in  Everglades  National  Park.  She  reported 
an  overall  significant  test  statistic  based  on  both  number 
of  tree  stems  and  estimates  of  bark  surface  area,  al- 
though the  relationship  was  substantially  weaker  based 
on  bark  surface  area.  Regardless  of  the  statistical  signif- 
icance, 983  of  1464  (67%)  of  all  snails  we  observed  were 
found  on  wild  tamarind.  Voss  (1976)  reported  a  similar 
estimate  of  34  of  50  (65%)  for  1  hammock  in  the  eastern 
Everglades  near  Miami,  and  Brown  (1978)  reported  209 
of  39.5  (52%)  in  another  location  in  Everglades  National 
Park.  Although  other  trees  niav  be  preferentially  select- 
ed if  available,  wild  tamarind  was  the  most  abundant  and 


Pase  116 


THE  NAUTILUS,  Vol.  114,  No.  3 


most  used  host  tree  within  the  Big  Cypress  National  Pre- 
serve, which  is  probably  also  the  case  in  most  of  the 
mainland  habitats.  Thus,  wild  tamarind  is  clearly  an  im- 
portant component  of  tree-snail  habitat  in  this  region. 

Although  our  model  selection  did  not  support  that 
host-tree  selection  differed  among  hammocks,  an  LRT 
chd  indicate  a  species  by  hammock  interaction  effect. 
Hammocks  differed  with  respect  to  which  host  tree  spe- 
cies were  present,  which  Ukely  reflects  inter-hammock 
differences  in  attributes  such  as  size,  soils,  and  hydro- 
logic  regimes.  Thus,  the  interaction  effect  we  observed 
could  have  been  attributed  to  these  differences  in  spe- 
cies composition,  but  our  data  are  insufficient  to  evaluate 
such  effects  with  much  reliability. 

Bias  due  to  \isibilit\-  of  snails  was  not  accounted  for 
by  our  approach.  This  effect  would  have  been  most  in- 
fluenced by  dense  foliage  obscuring  snails  from  view. 
For  most  host-tree  species,  we  believe  that  this  bias  was 
negligible,  especially  because  of  reduced  foliage  as  a  re- 
sult of  Hurricane  Andrew.  However,  one  notable  excep- 
tion was  pigeon  plum,  which  often  had  ver)'  dense  fo- 
liage making  it  difficult  to  detect  snails.  Thus,  we  suspect 
that  our  data  indicating  that  this  host  tree  species  was 
underutilized  may  have  been,  at  least  in  part,  an  artifact 
of  detection  bias. 

Although  our  data  are  far  from  conclusive,  they  are 
consistent  with  Browni's  h\-pothesis  that  there  mav  be  an 
association  between  selection  of  chfferent  host  trees  and 
size  of  the  snails.  We  found  size  differences  only  among 
3-year-old  snails.  However,  even  our  relatively  large  sam- 
ple sizes  tend  to  become  quite  small  once  partitioned  by 
age  and  host  tree  species.  Florida  tree  snails  exliibit  most 
growth  during  their  first  2-3  vears,  after  which  growth 
slows  dramatically  (Tuskes,  1981).  Thus,  3-year-old  snails 
probably  represent  a  peak  of  cumulative  growth,  and  are 
a  hkely  age  class  to  observe  size  differences.  For  this  age 
class,  snails  found  on  host-tree  species  used  less  than 
expected  were  smallest  in  length,  those  found  on  trees 
used  in  excess  of  availabihty  were  the  largest  in  length, 
and  those  found  in  proportion  to  their  availability  were 
intermediate  in  length. 

ACKNOWLEDGMENTS 

We  thank  Jimmy  Conner,  Phil  Darby,  Sue  Davis,  Vicky 
Dreitz,  Guy  Fischer,  Paul  Hinchcliff,  Amy  Kazmier, 
Steve  McGehee,  Katie  Golden,  Teresa  Johnson,  Jeff 
Ripple,  Tim  Towles,  Patty  Valentine-Darby  for  field  as- 
sistance,   and    George    Francioni    for    safe    helicopter 


flights.  We  also  appreciate  the  helpful  comments  from  2 
anonymous  reviewers.  Funding  was  provided  by  the  Na- 
tional Park  Service  through  the  South  Florida  Research 
Center,  and  Big  Cypress  National  Preserve  provided  ad- 
ditional logistic  support. 

LITERATURE  CITED 

Agresti,  A.  1990,  Categorical  data  analysis.  John  Wiley  and 
Sons,  New  York.  5.5Spp. 

Akaike,  H.  197.3.  Information  theory'  and  an  e.xtension  of  the 
madmuni  likelihood  principle.  In:  B.  Petrov  and  F.  Czaldl 
(eds.)  Proceedings  of  the  2'"'  International  Symposium  In- 
formation Theory,  Akademiai  Kiado  Budapest,  pp.  267- 
281. 

Bennetts.  R.  E.,  S.  A.  Sparks,  and  D.  Jansen.  In  press.  Factors 
influencing  movement  probabilities  of  Florida  tree  snails 
Liguiis  fasciatiis  (Miiller)  in  Big  Cvpress  National  Pre- 
serve following  Hurricane  Andrew.  Malacologia. 

Brown,  C.  A.  1978.  Demography,  dispersal,  and  microdistri- 
bution  of  a  population  of  the  Florida  tree  snail  Liguus 
fasciatiis.  M.S.  Thesis,  Uni\ersitv  of  Florida,  Gainesville. 

Buniham  K.  P.  and  D.  R.  Anderson.  1998.  Model  selection 
and  inference:  a  practical  infoniiation-dieoretic  approach. 
Springer-Verlag,  New  York. 

Cochran,  W.  G.  19.54.  Some  methods  for  strengthening  the 
common  chi-square  tests.  Biometrics  10:417-451. 

Haberman,  S.  J.  1973.  The  analysis  of  residuals  in  cross-clas- 
sified tables.  Biometrics  29:205-220. 

Lebreton,  J.  D.,  K.  P  Bumham.  J.  Clobert,  and  D  R  Ander- 
son. 1992.  Modeling  survival  and  testing  biological  hy- 
potheses using  marked  animals;  a  unified  approach  with 
case  studies.  Ecological  Monographs  62:67-118. 

McCuUagh.  P.  and  J.  A.  Nelder  1989.  Generalized  hnear  mod- 
els, 2nd  edition.  Chapman  and  HaU,  New  York,  511  pp. 

Pilsbry,  H.  A.  1912.  A  study  of  the  variation  and  zoogeography 
of  Liguus  in  Florida.  Journal  of  the  Academy  of  Sciences 
of  Philadelphia  15:429-470. 

Pilsbry,  H.  A.  1946.  Land  Mollusca  of  North  America.  Acad- 
emy of  Natural  Sciences  Philadelphia,  Monograph  32:1- 
520. 

Shibata,  R.  1989.  Statistical  aspects  of  model  selection.  In:  J. 
C.  Williams  (ed.)  From  data  to  model.  Springer-Verlag, 
New  York,  pp.  21.5-240. 

Tuskes,  P.  M.  1981.  Population  structure  and  biology  of  Liguus 
tree  snails  on  Lignumxitae  Key,  Florida,  The  Nautilus  95: 
162-169. 

Voss,  R.  S.  1976,  Obserxations  of  the  biology  of  the  Florida 
tree  snail,  Liguus  fasciatiis  (Miiller).  The  Nautilus  90:6.5- 
69. 

Whittaker,  R.  H.  and  G.  M.  Woodwell.  1967.  Surface  area  re- 
lations of  woody  plants  and  forest  communities.  American 
Journal  of  Botany  54:931-9.39. 


THE  NAUTILUS  114(3):  117-1 19,  2000 


Page  117 


The  development  of  three  heterobranch  mollusks 
from  California,  USA 


Rachel  Collin 

Committee  on  E\olutionar)'  Biologv' 

Uni\ersits'  of  Chicago 

1025  E.  57th  St. 

Chicago,  IL  60637  USA 

and 

Department  of  Zoology 

Field  Museum  of  Natural  Histor\' 

1400  S,  Lake  Shore  Drive 

Chicago,  IL  60605  USA 

rcoHinCSmidwav.  uchicago.edu 


ABSTRACT 

The  de\elopnient  of  Odostomin  nitinn.  Turbonilla  sp.,  and  Wil- 
hamia  peUoides  from  Cahtoniia,  is  t)pical  ot  heterobranch  mol- 
lusks. Thev  all  produce  small  eggs  (about  60  |jLni)  that  are  in- 
di\idually  encapsulated  and  embedded  in  gelatinous  egg  mas- 
ses. The  capsules  are  connected  by  chalazae.  Cleavage  is  un- 
equal and  gastrulation  occurs  by  inxagination.  Plankiotrophic 
larvae  hatch  at  shell  lengths  of  120-150  |xni. 

Additional  key  words:  Pyramidellidae,  Tiirbonilln.  Odostomia, 
gastropod  development,  Williamin 


INTRODUCTION 

Although  lower  heterobranchs  and  marine  pulmonates 
are  common  in  shallow  marine  waters,  their  develop- 
ment remains  largely  undescribed.  Knowledge  of  their 
development  may  be  useful  and  important  because  de- 
velopmental features  such  as  the  structure  of  egg  mas- 
ses, cleavage  type,  presence  of  the  larval  pigmented 
mantle  organ  (PMO),  and  hydrophobic  larval  shells  may- 
be important  characters  for  phylogenetic  stucbes  (Rob- 
ertson, 1985;  van  den  Biggelaar,  1996;  van  den  Biggelaar 
and  Haszprunar,  1996;  Collin,  1997).  Additionally,  there 
are  a  variety  of  interesting  trends  in  the  evolution  of 
cleavage  patterns  (Freeman  and  Lundelius,  1992;  van 
den  Biggelaar  and  Haszprunar  1996)  and  heterochro- 
nies in  larval  morphologies  (Page,  1994)  whose  docu- 
mentation could  benefit  from  more  extensive  phyloge- 
netic sampling. 

Herein  I  describe  the  development  of  two  pyranii- 
dellid  and  one  siphonariid  pulmonate  species  from  the 
Califomian  coast.  All  animals  were  collected  by  hand  in 
the  summer  of  1997  and  kept  in  small  dishes  in  the 
laboratory  where  thev  laid  egg  masses.  Egg  masses  were 
observed  daily  until  hatching,  but  no  attempt  was  made 
to  raise  the  larvae  to  metamorphosis. 


RESULTS 

Odostomia  altina  Dall  and  Bartsch,  1909 
(Table  1) 

Large  numbers  of  Odostomia  altina  were  found  on  the 
shells  oi  HaliotLs  cornigflta  collected  at  a  depth  of  10  m 
from  Point  Loma,  California  (29°40'N,  117°20'W). 
Specimens  are  deposited  at  the  Field  Museum  of  Nat- 
ural History  (FMNH  282369  and  282370)  and  were 
identified  by  comparison  with  the  original  species  de- 
scription and  illustrations  in  (Dall  and  Bartsch,  1909). 
There  were  as  many  as  50  adult  pvTamidellids  and  nu- 
merous egg  masses  on  one  abalone.  Egg  masses  were 
typical  of  other  pvTamideOids:  Single  eggs  are  enclosed 
in  albumin-filled  oval  capsules  that  are  embedded  in  a 
clear  sticky  gelatinous  mass.  The  capsules  are  connected 
by  thin  extensions  of  the  capsular  covering  called  cha- 
lazae (see  Robertson,  1985;  Collin  and  Wise,  1997  for 
detailed  description).  A  timetable  of  development  is  giv- 
en in  Table  1. 

The  round,  white  eggs  are  60  |jLm  in  diameter  (mean 
=  60.83  |jLm;  sd  =  1.39  |xm;  n  =  31;  eggs  from  3  egg 
masses)  the  inner  capsule  diameter  is  150  (xm  (sd  =  8.11 
|xm;  n  =  19)  and  the  outer  capsule  diameter  is  176  \x.m 
(sd  =  6.992  |xm;  n  =  10).  First  cleavage  is  unequal  and 
at  the  4-cell  stage  the  two  largest  cells  are  adjacent,  as 
are  the  two  small  ones.  Several  embryos  were  observed 
in  a  3-cell  stage,  which  suggests  that  second  cleavage  is 
not  synchronous.  There  is  no  polar  lobe  and  third  cleav- 
age is  unequal.  The  round  blastula  gradually  flattens  and 
invaginates  to  form  a  horseshoe  shaped  gastrula.  During 
subsequent  development  the  embryo  grows  to  fill  the 
capsule  and  the  larval  organs  differentiate.  Plank-totroph- 
ic  larvae  hatch  after  about  7  days  (Table  1)  with  130  |xm 
(mean  =  130.33  |xm;  sd  =  8.55  [xni;  n  =  15;  egg  masses 
from  two  females)  smooth,  left-handed,  hydrophobic 
shells.  Each  larva  has  a  well-developed  metapodial  ten- 


Page  118 


THE  NAUTILUS,  Vol.  114,  No.  3 


Table  1.   De\elopnient  schedule  for  Odostomiti  nltinn  at  22- 
24°C. 


Age 


Stage 


>3  hours  2-cells 

5  hours  4-cells 

12  hours  64-cells 

1  dav  blastula 

1.5  days  gastnilation  by  invagination 

3  days  just  moving,  foot  and  velum  anlangen  visible 

5  days  red-yellow  PMO  visible,  shell  operculum  and 

statocvsts 

~7  days  hatchmg  at  130  (jim 


tacle  and  statocysts,  but  no  detectable  heartbeat,  and  no 
eyes.  The  larval  PMO  consists  of  two  parts:  A  round  red 
central  part  and  an  additional  semi-transparent  yeUow 
lobe  on  one  side.  Observation  of  the  yellow  lobe  under 
the  dissecting  microscope  is  difficult,  but  the  structure 
can  be  seen  easilv'  under  a  compound  microscope. 

Turbonilla  sp. 
(Figures  1,  2,  table  2) 

One  adult  of  an  unknown  species  of  Turbonilla  was  col- 
lected on  the  high  mid-intertidal  zone  at  Alegria,  Cali- 
fornia (34°28'N,^120°17'VV)  in  July  1997.  The  species 
identification  is  uncertain  because  Turbonilla  taxonomy 
is  especially  difficult  and  the  single  specimen  could  not 
be  unambiguously  assigned  to  any  Californian  species. 
The  current  taxonomy  of  Californian  pyramidellids  as- 
signs all  species  with  axial-ribbed  shells  to  the  genus  Tur- 
bonilla (Dall  and  Bartsch,  1909).  This  snail  laid  one  egg 


Figure  2.  Earlv  developmental  stages  of  Turbonilla  .sp.  .\.  2- 
cell  stage  at  the  beginning  of  second  cleavage,  the  cells  are 
clearly  not  equal  in  size,  B.  4-Cell  stage,  C.  blastula,  D.  gas- 
trula,  showing  the  invagination  typical  of  heterobranch  devel- 
opment. Scale  bar  =  50  \x.m. 


mass  of  about  100  eggs  in  the  laboratory  and  develop- 
ment was  followed  until  hatching  (table  2  and  figure  2). 
The  egg  mass  consisted  of  capsules  containing  one  white 
egg  surrounded  by  clear  albumin,  connected  together  by 
chalazae  and  embedded  in  a  clear  sticky  gel.  The  egg 
diameter  was  65.5  |jLm  (n  =  10,  sd  =  1.97  |jLm)  and  the 
capsule  length  was  187.8  |xm  (n  =  10,  sd  =  6.67  (im). 
The  chalazae  are  thicker  and  less  tv\dsted  than  in  O.  col- 
umbiana  Dall  and  Bartsch  1907  (Colhn  and  Wise,  1997). 
First  cleavage  is  slightly  unequal  but  bv  the  4-celI  stage 
the  blastomere  inequalit)'  is  yery  subtle.  The  early  cleav- 
ages do  not  produce  a  polar  lobe.  The  polar  bodies  are 
clearK  visible  throughout  development  because  the  al- 
bumin is  transparent.  After  24  hours  the  embrvos  form 
flattened  blastulas,  which  begin  to  invaginate  at  about  40 
hours.  Gastnilation  continues  by  invagination  for  the 
next  dav.  Bv  the  fourth  dav  the  foot  and  velum  anlagen 
are  visible.  On  the  fifth  dav  the  embrvos  begin  to  move 


Table  2.    Developmental  schedule  for  Turbonilla  sp.  at  16- 
18°C. 


Figure  1.     Adult  Turbonithi  sp.  Shell  length  is  6  mm. 


Age 

Stage 

7  hours 

2-cells 

24  hours 

blastula 

2-3  days 

gastnilation  by  invagination 

4  days 

just  moving,  foot  and  velum  anlangen  visible 

5  days 

shell  and  velum  differentiated.  Embrvo  fills 

1/4  of  the  capsule 

6  davs 

black  PMO  visible 

9  days 

embno  fills  the  capsule 

11  days 

hatching  at  152  |j.m 

R.  Clolhn,  2(M)() 


Page  119 


and  thev  fill  alidiit  a  (juarter  of  each  (.apsiilf.  B\'  thf  next 
cla\'  the  black  PMO  is  \isihle  on  the  right  side  just  pos- 
terior to  the  velum.  The  enibno  grows  to  completely  fill 
the  capsule,  before  hatching.  Shell  length  at  hatching  is 
1.52  jjLni  (n  =  14;  sd  =  2. .37  p,m;  from  a  single  egg  mass). 
The  shell  is  hydrophobic,  left-handed  and  the  lana  has 
no  heart  or  eyes  but  has  a  well-developed  metapodial 
tentacle  on  the  foot.  The  planktotrophic  lanae  swim  ac- 
tively. 


Williamia  pchoides  (Carpenter,  1864) 

Several  adult  W.  peltoidcs  (Carpenter,  1864)  were  col- 
lected in  the  subtidal  zone  (  —  10  ni),  on  hard  substrates, 
in  the  vicinity  of  Santa  Barbara,  California  (.34°28'N, 
120°17'W).  Voucher  specimens  are  deposited  at  the  Bai- 
ley-Matthews Shell  Museum,  Sanibel,  Florida  (BMSM 
4999).  Williamia  species  are  unusual  among  marine  pul- 
monates  because  they  are  predominately  subtidal.  Adults 
produced  several  egg  masses  in  chshes  in  the  laboratory. 
The  structure  of  the  egg  masses  is  verv  similar  to 
those  of  the  pvTamidellids.  The  eggs  are  each  individu- 
ally enclosed  in  a  coating  of  albumin  inside  an  oval  cap- 
sule. The  capsules  are  connected  with  chalazae  and  are 
embedded  in  a  gelatinous  mass.  The  capsules  are  147 
jjim  in  length  (mean  =  147.7  ji-m;  sd  =  3.4  |xm;  n  = 
11).  Unfortunately  uncleaved  eggs  and  early  cleavage 
stages  were  not  observed.  Gastrulation  occurs  via  invag- 
ination forming  a  horseshoe-shaped  gastnila  that  is  69 
(xm  in  diameter  (d  =  69.28  |xm;  sd  =  3.4  |jLm;  n  =  7; 
from  one  egg  mass).  Further  development  progressed  as 
in  the  pyramidellids.  However,  no  pigmented  PMO  de- 
velops: I  could  not  determine  if  there  was  no  PMO  or 
if  the  structure  was  present  but  not  pigmented.  There 
is  some  reddish  pigmentation  along  the  suture  of  the 
larval  shell,  but  the  larval  body  was  pigmentless.  After  9 
days  at  18-22  °C  the  larvae  hatch  at  a  length  of  126  |xm 
(length  =  126.0  jjuni;  sd  =  ■5. .58  |jLm;  n  =  10).  The  plank- 
totrophic lar\ae  have  a  hydrophobic  smooth  left-handed 
shell.  There  is  still  no  pigmented  PMO,  no  eyes,  and  no 
metapodial  tentacle  but  the  statocysts  are  clearly  visible. 

DISCUSSION 

These  observations  are  the  first  descriptions  of  intracap- 
sular development  for  species  in  the  genera  Williamia 
and  Tttrhonilla. 

The  features  of  pyramidellid  development  described 
here  generally  agree  with  prexaous  descriptions  of  pyr- 
amidellid development.  Because  Tnrhoiiilla  and  Odos- 
tomia  are  distantly  related  genera  within  the  Pyramidel- 
lidae  (Wise,  1996),  characters  shared  by  the  species  de- 
scribed here  and  O.  cohtmhiana  (Colhn  and  Wise,  1997) 
may  be  typical  of  pyramidellid  development  in  general. 
All  three  species  have  small  eggs,  unequal  cleavage,  gas- 
tnilation  bv  invagination,  and  hatch  with  smooth,  left- 
handed,  hydrophobic  shells,  a  metapodial  tentacle,  dis- 
tinct PMO,  but  without  eyes  or  a  larxal  heart.  The  color 


oi  the  larval  PMOs  varies  among  species  as  does  the 
cok)r  of  the  adult  PMO  (Robertson,  1985). 

The  development  of  Williamia  peltoidcs  is  strikingly 
similar  to  pyramidellid  development.  The  structure  of 
the  egg  masses  is  indistinguishable  from  that  of  the  pyr- 
amidellids. Hatching  lar\ae  are  distinct  in  their  lack  of 
a  metapodial  tentacle  and  pigmented  PMO  and  the 
small  area  of  shell  pigment.  These  results  are  in  agree- 
ment with  Robertson's  (1985)  conclusion  that  larval 
PMOs  are  not  present  in  pulmonates  while  chalazae  and 
heterostrophy  are  common.  The  retldish  pigment  along 
the  shell  suture  is  also  found  in  larvae  of  Siphouaria  sp. 
from  New  Zealand  (pers.  obs.). 

ACKNOWLEDGMENTS 

This  research  was  conducted  during  a  visit  to  the  labo- 
ratories of  Dr  S.  Gaines  (University  of  California  at  San- 
ta Barbara)  and  Dr  K.  Roy  (University  of  CaUfornia  at 
San  Diego).  I  thank  Ron  McConnaughev  and  Shane  An- 
dersen for  helping  collect  animals  and  two  anon\iiious 
reviewers  for  helpful  comments  on  previous  versions  of 
this  manuscript.  This  work  was  supported  financially  by 
a  NSF  Predoctoral  Fellowship,  and  grants  from  the 
Western  Society  of  Malacologists,  Sigma  Xi,  the  Lemer 
Gray  Fund  (American  Museum  of  Natural  Historv)  and 
the  Hinds  Fund  (University  of  Chicago). 

LITERATURE  CITED 

Biggelaar,  J.  A.  M.  van  den.  1996.  The  significance  of  the  early 
cleavage  pattern  for  the  reconstruction  of  gastropod  phy- 
logeny.  In:  Taylor,  J.  D.  (ed.)  Origin  and  Evolutionary  Ra- 
diation of  the  Molliisca.  Oxford  University  Press,  Oxford, 
UK. 

Biggelaar,  J.  A.  M.  van  den  and  G.  Hazsprunar  1996,  Cleavage 
and  niesentoblast  formation  in  the  Gastropoda:  An  evo- 
lutionarv  perspective.  Evolution  .50:1.520-1.540. 

Collin.  R.  1997.  Hydrophohic  larval  shells:  Another  character 
for  higher  level  systematics  of  gastropods.  Journal  of  Mol- 
luscan  Studies  6.3:425-430. 

Collin,  R.  and  J.  B.  Wise.  1997.  Morphology  and  development 
of  Odostomin  cohnubiann  (Gastropoda:  PyTamidellidae). 
Biological  Bulletin  192:24:3-252. 

Dall,  W.  H.  and  P.  Bartsch.  1909.  A  monograph  of  west  Amer- 
ican pvTamidellid  mollusks.  Bulletin  of  the  United  States 
National  Museum  68:1-258. 

Freeman,  G.  and  ].  W.  Lundelius.  1992.  Evolutionaiy'  impli- 
cations of  the  mode  of  D  quadrant  specification  in  coe- 
lomates  with  spiral  cleavage.  Journal  of  Evolutionary  Bi- 
ology 5:20.5-247. 

Page,  L.  R.  1994.  The  ancestral  gastropod  larval  torm  is  best 
approximated  by  hatching-stage  opisthohranch  larvae:  ev- 
idence from  comparative  developmental  stuthes.  In:  Wil- 
son, W.  H.,  S.  A.  Strieker,  and  G.  L.  Shinn  (eds.)  Repro- 
duction and  Development  of  Manne  Invertebrates.  Johns 
Hopkins  University  Press,  Baltimore. 

Robertson,  R.  1985.  Four  characters  and  the  higher  category 
systematics  of  gastropods,  American  Malacologica!  Bulle- 
tin, Special  Edition  1:1-22. 

Wise,  J,  B,  1996.  Moqiliologv  and  phylogenetic  relationships 
of  certain  pyramidellid  taxa  (Heterobranchia).  Malacologia 
.37:44.3-51  r. 


THE  NAUTILUS  114(3):  120-126,  2000 


Page  120 


Diminishing  species  richness  of  mollusks  in  Oneida  Lake, 
New  York  State,  USA 


Willard  N.  Harman 

State  Universitv'  of  New  York  College  at  Oneonta 
Biological  Field  Station  RD2,  Box  1066 
Cooperstown,  NY  13326  USA 
harmamvn@oneonta.edu 


ABSTRACT 

Between  1915  and  1917,  F.  C.  Baker  studied  the  molluscan 
fauna  of  Oneida  Lake,  a  shallow,  eutrophic  lake  in  central  New 
York  State.  He  hsted  41  living  taxa,  of  which  two  gastropods, 
Bithynia  tentacitlata  (Linnaeus,  1758)  and  Pleurocera  aaita 
(Rafinesque,  1S29),  were  then  recent  introductions.  In  1967- 
68,  John  Foniey  and  I  (Harman  and  Foniey,  1970)  surveyed 
Oneida  Lake,  \'isiting  Baker's  original  collecting  sites.  W'e  tal- 
lied 35  Uving  taxa  dominated  by  the  introduced  European  B 
tentacitlata.  In  1992-95,  I  again  made  extensive  collections  of 
mollusks  finding  a  total  of  24  h\ing  taxa  dominated  bv  Dreis- 
sena  pohjmorpha  (Pallas,  1771),  the  then  recently  introduced 
Eurasian  zebra  mussel.  Fifty  percent  of  Baker's  original  euht- 
toral  collection  sites  could  not  be  found.  An  average  1  m-  area 
in  1992-95  included  6708  individuals  of  D.  pohjmorpha.  60  of 
B.  tcntactilata  and  2  of  the  unionid  bivalve  EUiptio  complanata 
(Lightfoot,  1786).  Molluscan  species  richness  was  reduced  by 
15%  between  1917  and  1968,  a  trend  concurrent  with  the  in- 
crease in  abundance  of  Bithynia.  Species  richness  was  hirther 
reduced  by  31%  between  1968  and  1995,  as  human  acti\-ities 
impacted  euhttoral  habitats  and  D.  polymorpha  colonized 
Oneida.  The  total  decrease  in  species  richness  between  1917 
and  1995  was  of  42%'.  Since  1996  no  li\ing  miionids  ha\e  been 
observed  in  the  lake. 

Additional  key  words:  Exotic  introductions,  habitat  destruc- 
tion, freshwater,  gastropods,  bivalves,  Bithynia  tentacitlata. 
Dreissena  pohjmorpha. 


INTRODUCTION 

Oneida  Lake  is  a  eutrophic  lake  in  central  New  York 
State,  with  about  233  kTn-  in  surface  area  and  a  maxi- 
mum depth  of  16.8  m.  It  is  located  in  a  depression  on 
the  low,  open  rehef  of  the  Ontario  Lake  Plain  in  the 
Oswego  River  drainage  basin  (figure  1).  The  lake  is  ori- 
ented with  its  length  parallel  to  that  of  the  prevailing 
westerly  winds.  It  mixes  throughout  the  ice-free  period, 
with  thermal  stratification  occurring  only  ephemerally 
during  wind-free  periods  (Harman  and  Forney,  1970). 
Tributaries  to  Oneida  drain  densely  populated  areas  of 
Onondaga  Limestone  and  fertile  soils.  As  a  result,  the 
lake  has  been  considered  one  of  the  most  naturally  pro- 
ductive in  the  world  (Mozley,  1954).  Historically,  popu- 


lations of  algae  and  aquatic  vascular  plants  have  thrived 
in  the  shallow  waters  along  the  shoals,  beaches,  and  is- 
lands, on  diverse  eulittoral  substrates  providing  food  and 
cover  for  a  varied  and  abundant  molluscan  fauna  (Har- 
man and  Forney,  1970). 

Central  New  York  has  a  diverse  molluscan  fauna  de- 
rived from  species  immigrating  from  Atlantic  coastal  riv- 
ers, the  Mississippi  River  drainage  (Interior  Basin),  and 
the  Great  Lakes,  soon  after  the  retreat  of  the  last  Pleis- 
tocene glaciers  (Clarke  and  Berg,  1959).  Oneida  Lake  is 
unique  due  to  its  central  position  on  the  New  York  Barge 
Canal  system  and  natural  waterways  (figure  1),  which 
have  continued  to  provide  access  to  immigrating  organ- 
isms from  the  early  1800s.  Since  then,  mollusks  from 
Lake  Erie  and  the  Ohio  River  basins  in  the  west,  and 
from  the  Atlantic  coastal  drainage  in  the  southeast  (via 
the  Mohawk  and  Hudson  drainage  basins)  have  found 
their  ways  to  Oneida  Lake.  Headwaters  of  several  wa- 
tercourses flowing  from  the  Appalachian  highlands  in 
the  southern  Oswego  system  join  the  Susquehanna 
drainage  via  through-viilleys,  which  facilitates  dispersal 
of  aquatic  organisms.  The  Oswego  River  drains  Oneida 
flowing  to  Lake  Ontario  and,  via  the  St.  Lawrence  River, 
joins  Lake  Champlain  and  other  northern  waterways, 
thus  providing  access  to  and  Irom  North  Atlantic  drain- 
age systems. 

During  1915-17,  F.  C.  Baker  studied  the  macroben- 
thos  in  Oneida  Lake,  concentrating  on  the  molluscan 
fauna  excepting  the  fingernail  clams  (Sphaeriidae)  (Bak- 
er, 1916a,  b,  c;  1918a,  b,  c,  d).  His  research  on  Oneida's 
Lower  South  Bay  is  among  the  earliest  quantitative  stud- 
ies of  freshwater  macrobenthos  in  North  America  (Bak- 
er, 1918c).  Baker  studied  Oneida  because  of  its  great 
diversity  of  mollusks.  In  1967  and  1968  I  (Harman  and 
Forney,  1970)  conducted,  as  part  of  a  survey  of  the  gas- 
tropods of  the  Oswego  River  drainage  basin,  a  qualitative 
study  of  the  molluscan  fauna  of  Oneida  Lake.  In  the 
summer  of  1967,  Forney  (Harman  and  Forney,  1970) 
repeated  Baker's  (1918c)  quantitative  work  in  Lower 
South  Bav  as  part  of  a  macrobenthic  monitoring  pro- 
gram carried  out  by  Cornell  University. 

The  introduction  of  zebra  mussels  {Dreissena  poly- 
mor-f)ha)  into  Oneida  Lake  in  1990  (Mitchell  et  al.  1996) 


W.  N.  Hai-man,  2000 


Page  121 


Figure  1.     Central  New  York  State  showing  Oneida  Lake  and 
routes  of  immigrating  mollusks. 


has  had  important  impacts  on  water  clarity  by  reducing 
phvtoplankton  standing  crops.  It  also  appears  to  be  af- 
fecting the  distribution  and  abinidance  ot  aquatic  vas- 
cular macroph\1:es  (E.  L.  Mills,  pers.  comm.).  The  ir- 
ruption of  the  Drcisscna  population  was  expected  to 
have  tremendous  impacts  on  the  molluscan  fauna  (e.g., 
Nalepa  ct  al.  1991;  Ricciardi  ct  al,  1995;  Schloesser  ei 
al,  1996).  That  concern  prompted  my  return  to  Oneida 
during  1992-95  to  determine  the  status  of  the  molluscan 
communitv.  I  effectively  repeated  my  (Harman  and  For- 
ney, 1970)  1967-68  qualitative  work  and  in  1993  did  a 
quantitati\'e  studv  in  Lower  South  Bay  in  an  attempt  to 
repeat  Baker's  191.5-lS  studies  (1918c)  as  well  as  For- 
ney's 1967  work  (Harman  and  Forney,  1970). 

This  paper  describes  the  changes  in  the  molluscan 
fauna  since  1915,  showing  evidence  of  a  constant  reduc- 
tion of  species  richness  concurrent  with  destruction  of 
euhttoral  habitats  and  invasions  of  immigrating  mollusks. 

MATERIAL  AND  METHODS 

In  1915-17,  during  the  summer  months.  Baker  obtained 
(juantitative  samples  in  Lower  South  Bay  using  a  sieve, 
flattened  on  one  edge,  attached  to  a  wooden  pole,  to 
collect  bottom  samples  of  about  10-  cm-.  A  variety  of 
larger  dredges  were  used  in  deeper  water  A  square  met- 
al frame,  viewed  through  a  glass-bottomed  bucket,  was 
used  to  estimate  densities  of  large  snails  and  bivalves. 
Collections  were  made  while  wading,  or  from  a  boat 
(1918c).  Water  depths  in  Lower  South  Bay  are  such  that 
these  methods  allowed  him  to  sample  all  areas  of  the 
bay.  Baker's  samples  at  each  collecting  site  approximated 
10-  cm-.  He  used  varying  numbers  of  replicates,  typically 


2-6.  All  his  computations  were  presented  using  10-  cm- 
(16  in-)  as  the  basic  unit. 

My  1967-68  qualitative  survey  was  made  throughout 
the  lake  in  an  attempt  to  sample  every  habitat  suitable 
for  mollusks.  Samples  were  retrieved  from  inorganic 
substrates  str;iined  through  a  sieve  or  handpicked  vsatli 
forceps  in  shallow  water  Vegetation  was  collected  and 
dried  over  containers  for  retrieval  of  desiccating  organ- 
isms. Collections  from  deeper  water  were  done  similarly 
using  SCL'BA.  Fornev  contributed  several  Ekman  grab 
samples  to  that  study.  Forney's  replication  of  Baker's 
quantitative  study  in  Lower  South  Bay,  again  during  the 
summer  months,  used  Baker's  methods  as  mentioned 
aiio\e.  However,  he  relied  more  on  standard  Ekman 
grabs.  Calculations  were  based  on  10-  cm-  sample  areas 
to  facilitate  comparison  with  Baker's  work.  Eleven  of 
Baker's  original  sample  sites  (out  of  a  total  of  140)  were 
lost  due  to  artificial  modifications  of  the  shoreline. 

In  1992-95  the  same  methodologies  were  used  for  the 
qualitative  work,  over  the  same  time  period  that  Forney 
(Harman  and  Forney  1970)  had  covered  in  1967-68. 
The  1992-95  (juantitative  work  in  Lower  South  Bay  in- 
voked Ekman  samples  in  deep  water  and  square  metal 
frames  in  eulittoral  waters.  Calculations  were  based  on 
1  m-  sample  size.  Information  is  presented  using  this 
size,  as  well  as  10-  cm-  for  comparison  with  the  earlier 
studies.  It  should  be  noted  that  when  converting  to  1  m- 
from  the  earlier  10-  cm-  sample  size,  diversity  is  poten- 
tially underestimated  since  widely  separated  large  indi- 
viduals such  as  unionids  often  encountered  in  the  former 
are  rarely  foimd  in  the  latter  samples,  and  thus  never 
appear  in  the  data  set.  Only  71  of  Baker's  140  original 
sites  were  still  in  existence  in  the  recent  survey.  All  titx- 
onomic  nomenclature  herein  has  been  updated  accord- 
ing to  Turgeon  ci  al  (1998). 

RESULTS  AND  DISCUSSION 

Table  1  indicates  the  taxa  that  Baker  encountered  at  the 
turn  of  the  century.  Baker's  revised  list  includes  41  fiving 
molluscan  taxa  inclutling  the  then  recently  introduced 
European  snail  Bithijnia  tent  acid  at  a,  which  had  ap- 
peared around  1880  in  Lake  Ontario  (Beauchamp, 
1888).  Bithi/nia  tcntavtilata  was  assumed  to  have  been 
introduced  via  ballast  water  in  commercial  seagoing  ves- 
sels plying  the  Great  Lakes  (Baker,  1916b).  The  species 
was  not  abundant  in  Oneida  Lake  in  1915-17,  but  oc- 
curred in  dense  populations  near  the  outlet  at  the  west 
end.  An  average  10-  cm-  bottom  sample  made  in  1917 
contained  7  taxa,  all  consisting  of  native  species  {Pyr- 
gulopsis  lustrica  (Pilsbry,  1890)  (Amnicola  htstiica,  A. 
lustrica  (var)  and  A.  oncida  in  Baker's  original  lists), 
Stagnicola  catcscopiiiin  (Say,  1867)  (Galba  catascopium 
of  Baker),  Gi/ratilus  panii.s  (Say,  1817)  {Planorhis  par- 
vus of  Baker),  FhijscUa  gyrina  (Say,  1821)  (Physa  gyiina 
of  Baker),  P.  Integra  (Haldeman,  1841)  (Physa  intcgra 
of  Baker),  Prmncnetus  cxacuoiis  (Say,  1821)  [Planorhis 
exacuous  of  Baker),  and  Valvata  tiicarinata  (Say,  1817)) 
(figure   2).    A    second    introduction,    Plcnroccra   acuta. 


Page  122 


THE  NAUTILUS,  Vol.  114,  No.  .3 


Table  1.  The  moUu.scan  fauna  of  Oneida  Lake;  reference  conditions,  191.5-17  (modified  from  Baker,  1918). 
Taxa 


Bivalvia:  Palaeohetcrodonta 

Margaritifera  maraarilifera  (Linnaeus,  17.58) 
Elliptio  complanata  (Lightfoot,  1786) 
Alasmidonta  undidata  (Say,  1817) 
Pijganodon  grandis  (Say,  1829) 
Pijganodon  cataracta  (Say,  1817) 
Strophifus  undulatus  (Say,  1812) 
Villosa  iris  (I.  Lea,  1829) 
Lampsilis  radiata  radiata  (Gmelin,  1791) 
Actinonaias  ligmcntina  (Lamark,  1819) 


Total  number  of  species:  41 


Gastropoda:  Basominatophora 

Phi/si'lla  gtjrina  (Sav,  1821) 
Phi/si'lla  Integra  (Haldeman,  1841) 
Phi/sella  heterostropha  (Say,  1817) 
Lijmnaea  stagnalis  (Linnaeus,  1758) 
Fossaria  obrussa  (Say,  1825)* 
Stagnicola  elodes  (Say,  1821) 
Stagnicola  enwrginata  (Say,  1821) 
Stagnicola  catescopium  (Say,  1867) 
Pseudosiiccinea  columella  (Say,  1817) 
Acella  haldniani  (W.  G.  Binney,  1867) 
Helisomn  anceps  (Menke,  1830) 
Phanorbella  trivolvis  (Say,  1817) 
Planorbella  campanulata  (Say,  1821) 
Gyraulus  parvus  (Say,  1817) 
Gyratdus  deflectus  (Say,  1824) 
Promenetus  exacuous  (Say,  1821) 
Laevapex  fuscus  (C,  B.  Adams,  1841) 
Ferrissia  parallelus  (Haldeman,  1841) 
Ferrissia  rivularis  (Sav,  1817) 


Gastropoda:  Caenogastropoda 

Campeloma  decisum  (Say,  1817) 
Vivipams  georgianus  (I.  Lea,  1834) 
Pleurocera  acuta  (Rafinesque,  1820)'' 
Amnicola  limosus  (Say,  1817) 
Pijrgulopsis  lustrica  (Pilsbry,  1890) 
Pomatiopsis  cincinnatiensis  (I.  Lea,  1840) 
Frobijthinella  enwrginata  (Kuster,  1852) 
Soinatogynis  depressus  (Tryon,  1862) 
Bithynia  tentaculata  (Linnaeus,  1758)' 
Valvata  tricarinata  (Say,  1817) 
Valvata  sincera  (Sav,  1824) 
Valvata  lewisi  (Currier,  1868) 
Valvata  bicarinata  (I.  Lea,  1841) 


'  Fossaria  obntssa.  as  used  here  includes  F.  modicelta,  (Say,  1825),  F.  nistica  (I.  Lea,  1841)  and  may  include  F.  parva  (I.  Lea,  1841). 
'•pleurocera  acuta  (Rafinesque,  1820)  had  invaded  Oneida  Lake  via  the  Erie  Canal  system  before  Baker's  1915  work  (Dazo,  1965). 
c  Bithynia  tentaculata  was  apparently  invading  at  the  time  of  collection.  Baker  noted  high  densities  in  the  outlet  (Oneida  River) 
very  few  in  the  Lake  proper 


1 


Q 

5 
8 

a 
B 


3* 


5 
o 
I. 

8. 
I 


5  ^^  I  I 


Figures  2-5. 

Figure  2.     Dominant  mollusks  (individuals  10"-  cnr)  in  Lower  South  Bay,  Oneida  Lake,  in  191.5-17  (Baker,  1918c).  In  this,  and 

figures  3  and  4,  species  occurring  at  a  density  of  less  than  0.5  individuals  10"-  cm-  are  not  considered. 

Figure  3.     Dominant  mollusks  (individuals  10-  cm-)  in  Lower  South  Bay,  Oneida  Lake  in  1967-68  (Harman  and  Forney,  1970). 

Figure  4.     Dominant  mollusks  (incfividuals  10"-  cm-)  in  Lower  South  Bay,  Oneida  Lake  in  1992-95. 

Figure  5.     Dominant  mollusks  (individuals  m"-)  in  Lower  South  Bay,  Oneida  Lake  in  1992-95.  Species  occurring  at  a  density  of 
less  than  0.5  indi\iduals  m"-  are  not  considered. 


W.  N.  Harman,  2000 


Page  123 


Table  2.   Molluscan  species  inxadins;  Oneitla  Lake  followed  by  the  approximate  dates  of  iiitnufnctioii  and  origins. 


Tixxa 


Approximate  dates 
of  introduction 


Origin 


Bivalvia:  Ileterodonta 

Dreissenti  polyinoi-plta  (Pallas.  1771) 
Bivalvia:  Palaeoheterodonta 

Potamihis  alatiis  (Sav,  1817) 
Li'ptodcii  fraailis  (Rafinesque,  1820) 
Liffiinia  recta  (Laniark.  1819) 
Gastropoda:  Caenogastropoda 

Bithi/nia  tentaculata  (Linnaeus,  1758) 
Pleiirocera  acuta  (Rafinesque,  1820) 
Elimia  livescens  (Menke,  1830) 
Eliriiin  virainica  (Say,  1817) 

Total  number  of  species:  8 


1986  Eurasia,  Great  Lakes,  Oswego  River 

1920-1950  Interior  basin,  Erie  Canal  system 

1920-1950  Interior  basin,  Erie  Canal  system 

1920-1950  Interior  basin,  Erie  Canal  system 

1910-1918  Eurasia,  Great  Lakes,  Oswego  River 

1825-1915  Interior  basin,  Erie  Canal  system 

1920-1950  Interior  basin  antl/or  Hvidson  river,  Erie  Canal  system 

1920-1950  Susquehanna  Basin,  through-valleys,  Oswego  Basin 
and/or  Interior  basin,  Erie  Canal  system 


which  had  arrived  via  the  Erie  Canal  from  the  west,  was 
well  established  by  the  time  Baker  completed  his  studies 
(Dazo,  1965;  Baker,  191Sc). 

When  John  Fomey  and  I  (Harman  and  Fomey,  1970) 
collected  in  the  1960s,  35  taxa  were  found.  Three  gas- 
tropods, Elimia  livescens  (Menke,  1830),  E.  virginica 
(Say,  1817),  and  Plcuroccra  acuta,  all  in  the  family  Pleu- 
roceridae,  were  represented  only  bv  dead  specimens. 
Three  pearly  freshwater  mussels  ((Unionidae),  Potamihis 
alatus  (Say,  1817),  Leptodca  fragiUs  (Rafinesque,  1820), 
Ligumia  recta  (Lamark,  1819))  had  immigrated  from  the 
Interior  Basin  (Clarke  and  Berg,  1959;  Dazo,  1965).  Eli- 
mia virginica  was  introduced  into  the  Oswego  watershed 
before  the  turn  of  the  century  (DeKay,  1843;  Beau- 
champ,  1888).  It  is  assumed  that  this  species  crossed  the 
divide  between  the  Susquehanna  and  Oswego  River  wa- 
tersheds via  through-valleys  after  small  headwater  lakes 
and  wetlands  on  the  divide  were  disturbed  to  provide 
water  for  the  Erie  Canal  after  its  opening  in  1825  (Har- 
man, 1970).  It  also  could  have  moved  eastward  along 
the  Erie  Can;il  from  populations  in  the  Interior  Basin 
(Goodrich,  1942).  Bithi/nia  tentaculata,  present  in  1917, 
had  become  abundant  by  the  1960s.  An  average  10-  cm- 
bottom  sample  in  1967  contained  4  individuals  of  B.  ten- 
taculata (table  2,  figure  3). 

Table  3  lists  the  taxa  collected  in  1915-17  (Baker, 
1916a,  b,  c;  1918a,  b,  c,  d)  and  in  1969  (Harman  and 
Fomey,  1970)  that  were  not  collected  in  the  1992-95 
survey.  Only  23  species  of  mollusks  were  present  in 
1992-95,  two  of  them,  Ligumia  recta,  and  Valvata  sin- 
cera  Say,  1824  (Valvatidae),  represented  by  dead  speci- 
mens only.  That  represents  a  decrease  of  species  rich- 
ness of  42%  since  1917.  Collections  made  in  1996  and 
1997  by  faculty  and  students  at  the  Cornell  Biological 
Field  Station  on  Oneida  failed  to  find  anv  living  unionid 
bivalves  (E.  L.  MiUs,  pers.  comm.).  Four  of  the  species 
of  mollusks  found  in  1993-95  were  introductions.  Of 
those,  the  population  of  B.  tentaculata  had  declined 
greatly  since  the  1960s  but  was  exceeded  only  by  the 
exotic  zebra  mussel,  Drcissena  polijmorpha.  This  Eur- 


asian dreissenid  was  apparently  carried  through  the  Os- 
wego River  and  New  York  State  Barge  Canal  svstem  by 
commercial  and/or  recreational  navigation  from  the 
Great  Lakes,  where  it  was  found  in  1988  (Nalepa  and 
Schloesser,  1993).  Like  B.  tentaculata,  D.  polymorpha 
was  supposedly  transported  in  ballast  water  of  ocean- 
crossing  ships  coming  from  Europe.  An  average  10-  cm- 
bottom  sample  in  1993  contained  12  zebra  mussels  (fig- 
ure 4).  An  average  1  m-  sample  collected  at  the  same 
time  and  location  included  6,708  specimens  of  D.  polij- 
morpha, 60  of  B.  tentaculata,  and  2  of  Elliptio  complan- 
ata  (Lightfoot,  1786)  (£.  complanatus  of  Baker)  (figure 
5).  The  tremendous  increase  results  from  large  colonies 
of  D.  poh/morpha  initially  associated  with  specimens  of 
E.  complanata,  which  were  not  encountered  in  the  10- 
cm-  samples.  These  substrates  are  fine  shifting  sands 
practicallv  devoid  of  zebra  mussels,  except  where  the 
comparatively  widely  disbursed  unionids  support  their 
colonies. 

Harman  (1968a,  b)  proposed  that  the  introduced  B. 
tentaculata  was  able  to  displace  the  local  representatives 
of  the  North  American  family  Pleuroceridae  in  eutrophic 
environments.  These  observations  were  based  on  the 
distributions  of  B.  tentaculata,  P.  acuta,  E.  livescens 
(Goniobasis  livescens  of  Baker)  and  E.  virginica  in  the 
Oswego  River  drainage  basin  in  central  New  York  (Har- 
man and  Berg,  1971).  Like  most  caenogastropod  snails 
of  inland  waters,  the  pleurocerids  are  periphyton  grazers 
that  rely  on  the  benthic  algal  community  as  their  main 
energy  source.  They  typically  inhabit  waters  supporting 
comparatively  low  populations  ot  phytoplankton  (Dazo, 
1965).  Bithijnia  tentaculata  also  is  a  grazer  but  in  addi- 
tion it  can  use  planktonic  algae  which  it  filters  from  the 
water  on  specialized  ctenidia  (Jorgensen,  1966)  poten- 
tially giving  it  an  advantage  in  eutrophic  waters  such  as 
Oneida  Lake. 

Observations  in  Oneida  in  the  1960s  regarding  the 
density  of  Bithi/nia  on  rocky  shoreLnes,  and  on  the  ob- 
served behavior  of  pleurocerid  snails  associated  with  B. 
tentaculata    in    aquaria,    substantiated   these   ideas.    In 


Page  124 

THE  NAUTILUS, 

Vol. 

114, 

No. 

3 

Table  3.  Ta\a 

lost  (not 

collected  ali\'e 

since 

dates 

indicated)  followed  by 

assumed 

major  contributing  factors. 

Taxa 

Last  date 
collected 

Contributing  factors  to  decimation 

Bivalvia:  Palaeoheterodonta 

Miir^iaritifcra  margaritifcra  (Linnaeus,  17.58) 
Elliptio  complanata  (Lightfoot,  1786) 
Alasmidonta  undulata  (Say,  1817) 
Pijganodon  grandis  (Say,  1829) 
Fijganodon  cataracta  (Say,  1817) 
Strophitus  undidatus  (Say,  1812) 
Leptodea  fragilis  (Rafinesque,  1820) 
Ligumia  recta  (Lamark,  1819) 
ViUosa  iris  (1.  Lea,  1829) 
Lampsilis  rndiata  radiata  (GmeUn,  1791) 
Actinonaias  ligmentina  (Lamark,  1819) 
Gastropoda:  Basommatophora 
Plu/sella  gyrina  {Sax.  1821) 
Physelln  hcterostrophn  (Say,  1817) 
Li/mnaea  stagnalis  Linnaeus,  1758 
Stagnicola  cmarginata  (Say,  1821) 
Stagnicola  catescopium  (Say,  1867) 
Acelln  haldmani  (W.  G.  Binney  1867) 
Qyranulus  deflectus  (Say,  1824) 
Promenetus  exacuous  (Say,  1821) 
Fenissia  rivularis  (Say  1817) 
Gastropoda:  Caenogastropoda 
Campctoma  decisum  (Say,  1817) 
Pleurocera  acuta  (Rafinesque,  1820) 
Elimia  livenscens  (Menke,  18.30) 
Elimia  virginica  (Say,  1817) 
Pomatiopsis  cincinnatiensis  (L  Lea,  1840) 
Pwbi/thineUa  emarginata  (Kuster,  1852) 
Somatogtjnis  depnisstis  (Tr)-on,  1862) 
Valvata  sincera  (Say,  1824) 
Valiata  lewisi  (Currier,  1868) 
Vakata  bicnrinata  (I.  Lea,  1841) 
Total  number  of  species:  30 


1968 
1995 
1917 
1995 
1917 
1917 
1968 
1968 
1917 
1993 
1917 

1968 
1968 
1917 
1968 
1968 
1968 
1968 
1968 
1917 

1968 
1917 


1968 
1917 
1917 
1968 
1917 
1917 


Direct  competition  with  Dreissena 

Direct  competition  with  Dreissena 

Trophic  alteration 

Direct  competition  with  Dreissena 

Trophic  alterations 

Trophic  alterations 

Direct  competition  with  Dreissena 

Direct  competition  with  Dreissena 

Trophic  alterations 

Direct  competition  with  Dreissena  polymorph 

Trophic  alterations 


polymorpha 
polynwrpha 

'  polymotyha 


polymorfiha 
polymoiyha 


Habitat 
Habitat 
Habitat 
Habitat 
Habitat 
Habitat 
Habitat 
Habitat 
Habitat 


destniction, 
destniction, 
destniction, 
destruction, 
destniction, 
destruction, 
destruction, 
destniction, 
destniction. 


trophic 
trophic 
trophic 
trophic 
trophic 
trophic 
trophic 
trophic 
trophic 


alterations 
alterations 
alterations 
alterations 
alterations 
alterations 
alterations 
alterations 
alterations 


Habitat  destniction 
Direct  competition 
Direct  competition 
Direct  competition 
Habitat  destruction 
Habitat  destruction 
Habitat  destniction 
Habitat  destniction 
Habitat  destniction 
Habitat  destniction 


,  trophic  alterations 
with  Bitht/nia  tcntaculata 
with  Bithynia  tentacxdata 
with  Bithynia  tcntaculata 
,  trophic  alterations 
,  trophic  alterations 
,  trophic  alterations 
,  trophic  alterations 
,  trophic  alterations 
,  trophic  alterations 


'  Empy  shells  collected  in  1968,  not  collected  in  1915;  assumed  introduction  between  1920  and  1950. 


aquaria  enriched  with  planktonic  algae,  pleurocericLs  ac- 
tively grazed  over  the  substrate.  Individuals  of  B.  tcn- 
taculata remained  in  one  spot,  filtering  algae  from  the 
water.  Upon  encountering  specimens  of  B.  tcntaculata, 
pleurocerids  were  rebuffed  by  a  violent  twisting  of  the 
shell  of  that  filter-feeder,  which  invariably  caused  the 
grazing  snails  to  retract  into  their  shells.  They  remained 
in  that  position  for  several  minutes  before  resuming 
grazing.  In  situations  with  high  densities  of  B.  tcntacu- 
lata, indixidual  pleurocerids  spent  a  large  fraction  of  the 
time  in  defensive  positions  rather  than  feeding.  In  Onei- 
da Lake  in  the  1960s,  euhttoral  cobble  substrates  were, 
in  many  areas,  a  solid  mass  of  B.  tcntaculata  (Harman 
and  Forney,  1970).  Under  those  conditions,  all  the  spe- 
cies representing  the  family  Pleuroceridae  chsappeared 
from  the  lake  (Wisenden  and  Bailey,  1995).  Nowadays, 
decreasing  algal  populations  and  increased  water  clarity 
have  occurred  concurrently  with  irrupting  populations  of 
D.  polymorpha,  while  populations  of  B.  tcntaculata  have 
diminished  drastically. 


A  large  number  of  the  pulmonate  gastropod  species 
not  found  in  the  recent  sur^'ev  (e.g.,  one  species  of  the 
planorbid  genus  Gyraulus,  three  in  the  family  Lymnae- 
idae,  and  two  in  the  Physidae)  are  typical  of  eulittoral 
substrates  and  often  associated  with  emergent  vegeta- 
tion. It  is  unlikely  that  these  pulmonates  are  completely 
absent  from  Oneida,  but  their  absence  in  the  recent  sur- 
vey suggests  drastic  reductions  in  their  populations  cor- 
responding with  losses  of  that  habitat  due  to  shorehne 
development.  Most  of  the  shoreline  has  been  modified 
as  a  result  of  the  construction  of  break'waters  or  rip-rap. 
Many  of  Baker's  euhttoral  sites  appear  to  be  buried  un- 
der yards  and  beaches  created  bv  back-fiUing,  which,  bv 
1993-95,  had  destroyed  considerable  areas  of  emergent 
vegetation. 

Other  species  of  gastropods  that  were  not  found  in 
the  1990s  include  several  small  prosobranchs  in  the  fam- 
ily Hydrobiidae,  which  were  typically  abundant  on  sub- 
merged plants  and  nearbv  sub-httoral  and  deeper-water 
substrates.  These  and  other  gastropods  that  historically 


W.  N.  Harman,  2000 


Page  125 


have  been  present  in  abundance  in  Oneida  Lake,  were 
found  in  very  low  numbers  in  the  recent  survey.  An  ex- 
ception, Amnicola  limo.su.s  (Say,  1817)  {Aiunicohi  liinosa 
of  Baker),  occurs  in  high  densities  in  inanv  littoral  and 
sub-littoral  areas.  Changes  in  httoral  vegetation  and  as 
vet  unrecognized  trophic  alterations  associated  at  least 
indirecth'  with  colonization  by  D.  pohjmorfiha  may  be 
affecting  these  organisms  (Stewart  and  Haynes,  1994; 
Howells  ct  al..  1996;  Ricciardi  ct  al,  1997).  ' 

Seven  of  the  12  species  of  freshwater  mussels  found 
in  Oneida  since  1915  had  been  lost  by  1993,  and  no 
li\ing  unionids  ha\'e  been  obser\'ed  since  1995  (table  3). 
These  losses  correspond  to  the  introduction  and  estab- 
lishment of  Drcissena  pohjuunyha.  The  literature  doc- 
umenting the  loss  of  unionid  species  as  a  result  of  com- 
petition with  D.  pohjiiwiyha  (Nalepa  et  al,  1991;  Grif- 
fiths, 1993;  Ricciarchrffl/.,  1995;  Schloesser  ef  a/.,  1996) 
and  observations  of  the  interaction  behA'een  unionids 
and  D.  polyinorpha  in  Oneida  (E.  L.  Mills,  pers.  comm.) 
indicate  that  a  large  percent  of  this  loss  can  be  assumed 
to  be  attributed  to  interactions  with  D.  poJijmotyha. 

Since  1915-17,  the  endemic  molluscan  community  in 
Oneida  Lake  has  been  greatly  modified.  Species  richness 
has  declined  and  the  dominant  faunal  elements  are  now 
Eurasian  in  origin  (figure  5).  With  the  exception  of  re- 
cent changes  since  the  introduction  of  zebra  mussels, 
water  qualit)-  in  Oneida  Lake  has  been  consistent  over 
the  years  (E.  L.  Mills,  pers.  comm.).  It  is  assumed  the 
observed  changes  are  a  result  of  biotic  interactions  with 
introduced  taxa  and  loss  of  habitat  due  to  human  mod- 
ifications of  the  euhttoral  emdronment. 

LITERATURE  CITED 

Baker,  F.  C.  1916a.  The  fresh-water  Mollusca  of  Oneida  Lake, 
New  York.  The  Nautilus  .30:.5-9. 

Baker,  F.  C.  1916b.  The  relation  of  mollusks  to  fish  in  Oneida 
Lake,  New  York.  N.Y  State  College  of  Forestrv.  Technical 
Publication  No.  4.,  .366  p. 

Baker,  F.  C.  1916c.  Description  of  a  new  varietv  of  Lampsilis 
from  Oneida  Lake  ^A^th  notes  on  the  L.  lutcola  group.  The 
Nautilus  .30:74-77, 

Baker,  F.  C.  1918a.  Further  notes  on  the  Mollusca  of  Oneida 
Lake,  New  York:  the  mollusks  of  Lower  South  Bay.  The 
Nautilus  31:81-93. 

Baker,  F.  C.  1918b.  Description  of  a  new  variety  of  fresh-water 
mussel  from  Oneida  lake,  N.Y.  N.Y.  (Appendix)  State  Col- 
lege of  Forestry,  Technical  Fuhlication  9,  43(2):247-248. 

Baker,  F  C.  1918c.  The  productivit)'  of  invertebrate  fish  food 
on  the  bottom  of  Oneida  Lake,  with  special  reference  to 
mollusks.  N.Y.  State  College  of  Forestry,  Technical  Pub- 
lication 9,  43(2):l-2.33  +  235-264  (appendix). 

Baker,  F  C.  1918d.  The  relation  of  shellfish  to  fish  in  Oneida 
Lake,  New  York.  New  York  State  College  of  Forestry,  Cir- 
cular 21,  17(27):l-34. 

Beauchamp,  W.  M.  1888.  Notes  on  American  shells.  The  Con- 
chologists  Exchange  2:114-115. 

Berry,  E.  G.  1943.  The  Amnicolidae  of  Michigan:  distribution, 
ecology,  and  taxonomy.  Miscellaneous  Publications.  Mu- 
seum of  Zoology,  University  of  Michigan.  57:1-68. 

Clarke,  A.  H.  and  C.  O.  Berg.  1959.  The  freshwater  mussels 


of  central  New  York  with  an  illustrated  key  to  the  species 
of  northeastern  North  America.  Cornell  University  Agri- 
cultural Experiment  Station.  Memoir  .367:1-79. 

Dazo,  B.  C.  1965.  The  morphology  and  natural  history  of  P/eu- 
rocera  andn  and  Coniohasis  livescens  (Gastropoda:  Ceri- 
thiacea:  Pleuroceridae).  Malacologia  3:1-80. 

DeKay  J.  E.  1843.  Zoolog\'  of  New  York.  Part  5.  Mollusca. 
Albany,  271  pp..  40  pis. 

Goodrich,  C.  1942.  The  Pleuroceridae  of  the  Atlantic  coastal 
plain.  Museum  of  Zoolog\',  Universit)'  of  Michigan,  Oc- 
casional Papers  456:1-6. 

Griffiths,  R,  W  1993,  Effects  of  zebra  mussels  (Drcissena  poly- 
uioiylui)  on  tiie  benthic  fauna  of  Lake  St.  Clair  In:  T  F 
Nalepa  and  D.  W,  Schlosser  (eds.)  Zebra  mussels:  biology, 
nnpacts,  and  control.  Lewis  Pubhshers,  Boca  Raton,  pp. 
414-437. 

Harman,  W.  N.  1968a.  Replacement  of  pleurocerids  by  Bi- 
thynin  in  polluted  waters  of  central  New  York.  The  Nau- 
tilus 81:77-83. 

Harman,  W.  N.  1968b.  Interspecific  competition  between  Bi- 
thynia  and  pleurocerids.  The  Nautilus  82:72-73. 

Harman,  VV.  N.  1970.  Alterations  in  the  molluscan  fauna  of  a 
meromictic,  marl  lake.  The  Nautilus  84:21-.30. 

Harman,  W.  N.  and  C.  O.  Berg.  1971.  The  freshwater  snails 
of  Central  New  York  with  illustrated  keys  to  the  Genera 
and  Species.  Cornell  Universitv  Agricultural  Ex-periment 
Station.  Search;  Agriculture  Entomolog)-  (Ithaca)  2,  1(4): 
1-68. 

Harman,  W.  N.  and  ].  L.  Forney  1970.  Fifty  years  of  change 
in  the  molluscan  fauna  of  Oneida  Lake,  New  York.  Lim- 
nology and  Oceanography  15:454—460. 

Howells,  E.  T,  M.  H.  Christopher,  R.  W.  Bilyea,  P  B.  Kauss, 
and  K.  Somers.  1996.  Changes  in  environmental  condi- 
tions during  Drcissena  colonization  of  a  monitoring  station 
in  eastern  Lake  Erie,  Journal  of  Great  Lakes  Research  22; 
744-756. 

Jorgensen,  C.  B.  1966.  Biology  of  suspension  feeding.  Perga- 
mon,  Oxford,  357  pp. 

Mitchell,  M.  J.,  E.  L.  Mills,  N.  Idrisi,  and  R.  Michener  1996. 
Stable  isotopes  of  nitrogen  and  carbon  in  an  aquatic  food 
web  recently  inxaded  bv  Drcissena  polymorpha  (Pallas). 
Canadian  Journal  of  Fisheries  and  Aquatic  Sciences  53: 
144.5-1450. 

Mozley,  A.  1954.  An  introduction  to  molluscan  ecology.  H.  K. 
Lewis,  London,  71  pp. 

Nalepa,  T  F,  B.  A.  Manny  J.  C.  Roth,  S.  C.  Mosley  and  D. 
W.  Schloesser  1991.  Long-tenn  decline  in  freshwater 
mussels  (BivaKia:  Unionidae)  of  the  western  basin  of  Lake 
Erie.  Journal  of  Great  Lakes  Research  17(2):214-219. 

Nalepa,  T  F  and  D.  W.  Schloesser  (eds.).  1993.  Zebra  mussels: 
Biology,  impacts  and  control.  Lewis  Publishers,  Boca  Ra- 
ton, 810  pp. 

Ricciardi,  A.,  F  G.  Whoriskev,  and  J.  B,  Rasmussen,  1995. 
Predicting  the  intensitv  and  impact  of  Drcissena  infesta- 
tion on  native  unionid  bivakes  from  Drcissena  field  den- 
sity. Canadian  Journal  of  Fisheries  and  Aquatic  Science 
52:1449-1461. 

Ricciardi,  A.,  F.  G.  Whoriskey,  and  J  B.  Rasmussen  1997  The 
role  of  the  zebra  mussel  (Drcissena  polymorpha)  in  struc- 
turing macroinvertebrate  communities  on  hard  substrata. 
Canadian  Journal  of  Fisheries  and  Aquatic  Sciences  54: 
2596-2608. 

Robertson,  I.  C.  S.  and  C.  L.  Blakeslee.  1948.  The  Mollusca 


Pare  126 


THE  NAUTILUS,  Vol.  114,  No.  3 


of  the  Niagara  frontier  region.  Bulletin  of  the  Bulletin  of 
the  Buffalo  Society  of  Natural  Sciences  19(3):1-191. 

Schloesser,  D.  W.,  T.  R  Nalepa,  and  G.  I.  Maclde.  1996.  Zebra 
mussel  infestation  of  unionid  bivalves  (Unionidae)  in 
North  America.  American  Zoologist  .36:.300-310. 

Stewart,  T.  W  and  J.  M.  Haynes.  1994.  Benthic  macroinver- 
tebrate  communities  of  southwestern  Lake  Ontario  fol- 
lowing invasion  of  Dreissena.  Journal  of  Great  Lakes  Re- 
search 20:479-493. 

Turgeon,  D.  D.,  ].  F.  Quinn,  Jr,  A.  E.  Bogan,  E.  V.  Coan,  F. 


G.  Hochberg,  VV.  G.  Lyons.  P.  M.  Mikkelsen,  R.  ].  Neves, 
C.  F.  E.  Roper,  G.  Rosenberg,  B.  Roth,  A.  Scheltema.  M. 
Vecchione,  and  ].  D.  Wilhams.  199S.  Common  and  sci- 
entific names  of  aquatic  invertebrates  from  the  United 
States  and  Canada:  MoUusks.  2'"'  edition.  American  Fish- 
eries Society  Special  Publication  26,  .526  pp. 
Wisenden,  R  A.  and  R.  C.  Bailey.  199.5.  Development  of  mac- 
roinvertebrate  community  structure  associated  with  zebra 
mussel  {Dreissena  pohjmorpha)  colonization  of  artificial 
substrates.  Canadian  Journal  of  Zoology  73:14.38-1443. 


XtJ"^ — 


Florida  Department  of  State 

Katherlne  Harris 

Secretary  of  State 

Florida  Arts  Council 

Division  of  Cultural  Affairs 

This  program  is  sponsored 
in  part  by  the  State  of  Florida, 

Florida  Department  of  State. 

Division  of  Cultural  Affairs,  and 

the  Florida  Arts  Council 


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THE  NAUTILUS 


Volume  114,  Number  4 
December  15,  2000 
ISSN  0028-1344 

A  quorterUj  devoted 
to  malacology. 


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CONTENTS 


Volume  114,  Number  4 

December  15,  2000 

ISSN  0028-1344 


Luiz  Ricardo  L.  Simone 
Cuido  Pastorino 
Pablo  E.  Penchaszadeh 

Richard  E.  Petit 
M.  G.  Harasewych 

Bruce  A.  Marshall 


Douglas  G.  Smith 


Martin  Avery  Snyder 


Crepiclula  arg^entina  (Gastropoda:  Calyptraeidae),  a  new 

species  from  the  littoral  of  Argentina 127 

Three  new  species  of  the  genus  Merica  ( Neogastropoda: 

Cancellariidae)  from  South  Africa  and  the  Philippines 142 

Systematics  of  the  genus  Infundibuhiin  Montfort,  1810 

(Gastropoda:  Trochidae) 149 

On  the  taxonomic  placement  of  Unio  ochraceus  Say,  1817 

in  the  genus  Ligiimia  (Bivalvia:  Unionidae) 155 

Latints  beckijae,  a  new  species  of  Fasciolariidae 

(Neogastropoda)  from  Brazil 161 


DEC  ^  8  2000 


STATEMENT  OF  OWNERSHIP,  MANAGEMENT  AND  CIRCULATION 

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Sanibel,  FL  33957 

Editor,  Dr.  Jose  H.  Leal,  address  as  above. 
Managing  Editor,  Dr.  Jose  H.  Leal,  address  as  above. 

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not  changed  during  the  preceding  1 2  months. 

13.  Publication  Title,  THE  NAUTILUS 

14.  Issue  Date  for  Circulation  Data  Below,  June  6,  2000 

15.  Extent  and  Nature  of  Circulation 


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B.  Paid  Circulation 

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Average 

Single 

1 2  months 

Issue 

545 

550 

425 

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425 

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22 

25 

2 

2 

24 

27 

449 

433 

96 

117 

545 

550 

95% 

94% 

THE  NAUTILUS  114(4):127-141,  2000 


Page  127 


Crepidiila  argentina  (Gastropoda:  Calyptraeidae),  a  new  species 
from  the  littoral  of  Argentina 


Luiz  Ricardo  L.  Simone 

Museu  de  Zoologia  da  Universidade 

de  Sao  Paulo 
Caixa  Postal  42694 
04299-970  Sao  Paulo 
BRAZIL 
IrsinioneC&'usp.br 


Cuido  Pastorino' 

Museo  Argentino  de  Ciencias 

Naturales 
k\.  Angel  Gallardo  470,  3°  piso,  lab. 

57 
C1405  D]R  Buenos  Aires 
ARGENTINA 
r\pastor@criba.edu.ar 


Pablo  E.  Penchaszadeh 

FCEvN-UBA-CONICET 
Museo  Argentino  de  Ciencias 

Naturales 
Av.  Angel  Gallardo  470,  3°  piso,  lab. 

57 
C1405  DJR  Buenos  Aires 
ARGENTINA 
pablop@mail.retina.ar 


ABSTRACT 

Crepidiila  argentina.  a  new  species  of  gastropod  of  the  famil)' 
Calyptraeidae,  is  described  from  the  littoral  of  Mar  del  Plata, 
proxince  of  Buenos  Aires,  Argentina.  The  new  species  is  con- 
chologicalK  similar  to  C.  protea  d'Orbignv  from  southern  Bra- 
zil. Crepidula  argentina  has  a  larger,  subcircular  shell  and  low- 
er convexitv'  than  C.  protea.  Anatomical  characters  that  allow 
differentiation  from  this  latter  species  are;  larger  lateral  shell 
muscle  and  very  weak  dorsal  shell  muscle;  kidney  proportion- 
ally smaller,  with  a  characteristic  arrangement  of  inner  folds  of 
dorsal  lobe;  presence  of  a  renal  vessel  edging  rectum  and  work- 
ing as  adrectal  sinus;  connection  between  odontophore  muscle 
pairs  m7  and  mil;  longer  salivary  gland;  four  ducts  to  digestive 
gland  in  stomach  (instead  of  two);  distinctive  arrangement  of 
folds  in  inner  surface  of  stomach  between  esophageal  aperture 
and  posterior  pair  of  ducts  to  digestive  gland;  seminal  vesicle 
of  males  broad  and  few  coiled;  penis  and  papilla  long  and  nar- 
row; vaginal  tube  nmning  closely  attached  to  capsule  gland. 
The  reproductive  biology  is  also  distinctive.  Crepidula  argen- 
tina new  species  has  a  very  well  defined  seasonal  reproductive 
cycle.  Females  brood  between  1  and  46  egg  capsules  per 
spawn.  The  average  total  number  of  embrv'os  per  spawn  is 
5600.  The  uncleaved,  laid  egg  diameter  is  170  (xm  and  the 
number  of  eggs  per  egg  capsule  is  about  320.  All  eggs  develop, 
there  are  no  nurse  eggs.  Larvae  hatch  as  planktotrophic  veli- 
gers.  The  new  species  is  compared  with  other  members  of  the 
genus  Crepidula  from  the  southern  Atlantic  coast  of  South 
America. 

Additional  key  word.?:  Crepidula  protea,  anatomy,  reproduc- 
tion, littoral. 


INTRODUCTION 

Most  species  of  moUusks  described  from  the  southern 
Atlantic  coast  of  South  America  are  based  on  specimens 
collected  by  nineteenth  century  expeditions.  Crepidula 
species  are  no  exception.  Alcide  d'Orbigny  (1841)  de- 


Author  for  correspondence. 


scribed  Crepidula  protea  for  the  first  time  on  his  large 
work  "Voyage  dans  I'Amerique  Meridionale".  This  book 
was  published  in  several  parts  in  different  dates.  Ac- 
cording to  Sherboni  and  Woodward  (1901),  Sherbom 
and  Griffin  (1934),  and  Keen  (1966)  the  page  (465)  with 
the  C.  protea  description  (with  no  illustration)  was  pub- 
fished  on  1841.  A  year  later  in  the  French  edition  ol  the 
Atlas  of  R.  de  la  Sagra's  "Histoire  ...  de  la  File  de 
Cuba",  d'Orbigny  pubfished  a  plate  (pi.  24,  figs.  30-32) 
with  three  illustrations  of  C.  protea  and  no  description 
(A  description  was  published  in  1846  in  the  Spanish  edi- 
tion of  the  same  work.)  It  seems  clear  that  the  intention 
of  d'Orbigny  was  to  publish  the  chapter  of  mollusks  in 
Sagra's  work  before  his  'Voyage ",  because  he  cited  that 
other  book  despite  its  later  pubfication.  Crepidula  protea 
was  described  without  mention  of  a  distinct  t\pe  locality. 
A  label  at  The  Natural  Histor)'  Museum,  London 
(BMNH)  states  only  "Patagonia"  without  further  geo- 
graphic details.  The  examination  of  the  type  specimens 
did  not  suggest  any  precise  locafitv  nor  were  there  shell 
characters  that  could  reveal  which  population  they  had 
came  from. 

D'Orbigny's  description  and  illustration  of  Crepidula 
protea  clearly  groups  at  least  two  different  species.  First 
Dall  (1889),  and  later  Hoagland  (1977),  pointed  out  that 
C.  plana  Say,  1822,  and  C.  iin^uiformis  Lamarck,  1822, 
might  be  included  within  his  concept  of  C.  protea.  Ac- 
cording to  d'Orbigny's  illustrations  and  description  this 
seems  to  be  possible. 

Parodiz  (1939)  studied  the  species  oi  Crepidula  from 
Argentina  with  particular  emphasis  on  the  shell  and,  for 
the  first  time,  on  the  radula.  Based  on  a  large  number 
of  lots  from  different  localities  Parodiz  described  and 
illustrated  the  radula  and  shell  of  five  previously  known 
species  and  two  new  subspecies.  The  subspecies  are  now 
considered  to  be  geographical  forms.  Despite  that,  Par- 
odiz (1939)  is  still  the  most  authoritative  paper  on  the 
genus  Crcpidida  from  southern  Atlantic  coast  of  South 


Page  128 


THE  NAUTILUS,  Vol.  114,  No.  4 


America.  Unfortunately,  no  anatomical  descriptions  or 
reproductive  features  were  described  in  that  work.  The 
specimens  examined  by  Parodiz  are  stiU  housed  at  the 
Museo  Argentino  de  Ciencias  Naturales  (MACN)  and 
were  re-examined  in  this  paper  Since  Parodiz's  pio- 
neering work  no  further  articles  on  Crepidula  from  Ar- 
gentina have  been  published. 

Hoagland  (1977)  published  a  comprehensive  study  of 
living  and  fossO  species  of  Crepidula.  Although  her  work 
was  basically  restricted  to  North  America  and  the  east- 
ern Pacific,  she  included  descriptions  and  pictures  of  C. 
protea  and  C.  dilatata  as  well  as  a  few  other  southern 
species.  Hoagland  (1983)  described  several  specimens 
from  BrazU  of  what  she  concluded  was  C  protea.  She 
recognized  C.  protea  as  a  species  distinct  from  C.  plana 
and  C.  ungitifonni.s  and  studied  d'Orbigny's  type  mate- 
rial of  C.  protea.  In  the  same  paper,  Hoagland  described 
the  larval  development  of  C.  protea  based  on  specimens 
from  the  southeastern  Brazihan  coast.  Here  we  consider 
the  specimens  she  studied  to  represent  actual  C.  protea 
mainly  because  they  are  conchologically  similar  to  the 
lectotype  and  paralectotypes. 

In  an  annotated  Ust  of  several  type  specimens  from 
the  coast  of  Argentina,  Aguirre  (1993)  designated  a  lec- 
totype for  C.  protea.  Unfortunately,  she  selected  the  only 
complete  specimen  in  d'Orbigny's  collection  at  BMNH. 
She  seems  to  have  been  unaware  of  the  differences 
among  the  southern  species  of  this  genus.  Aguirre 
(1993)  mentioned  only  4  syn types  when  the  count  is  ac- 
tually 16,  arranged  in  two  lots  of  5  and  11  specimens 
respectivelv  (BMNH  1854.12.4.573  and  574)  (Hoagland, 
1983). 

Gallardo  (1977,  1979)  and  Brown  and  Olivares  (1996) 
described  several  new  species  of  Crepidula  from  Chile 
with  distinctive  reproductive  patterns  as  diagnostic  char- 
acters. These  authors  led  the  way  in  the  search  for  new 
characters  in  this  variable  genus,  showing  that  in  several 
cases  the  shell  lacks  real  ta.\onomic  value  at  the  specific 
level. 

Hoagland  (1983)  and  Rios  (1985)  pointed  out  that 
Crepidula  protea  occurs  from  Rio  de  Janeiro,  Brazil, 
south  to  Miramar,  in  the  province  of  Buenos  Aires.  How- 
ever, studies  on  different  population  samples  along  this 
range  revealed  that  more  than  one  pattern  of  reproduc- 
tive strategy  and  anatomical  characters  are  found,  indi- 
cating the  presence  of  an  unnamed  species. 

In  this  paper  we  describe  this  new  species  of  Crepi- 
dula and  provide  the  groundwork  for  a  future  revision 
of  the  group  in  the  southwestern  Atlantic.  The  new  spe- 
cies was  originally  recognized  first  by  its  distinctive  re- 
productive pattern  and  second  through  detailed  anatom- 
ical studies.  Comparison  with  several  kiiown  species 
from  South  America  is  presented.  The  study  of  repro- 
ductive biology  in  the  new  species  is  the  main  topic  of 
another  paper  (Cledon  and  Penchaszadeh,  submitted.). 

MATERIALS  AND  METHODS 

Fresh  specimens  of  Crepidula  argentina  new  species 
used  in  this  studv  were  collected  by  commercial  fish 


trawlers  at  Mar  del  Plata  (38°00'  S,  57°33'  "W)  and 
Puerto  Quequen  (38°35'  S,  58°42'  'W)  along  the  coast 
of  the  province  of  Buenos  Aires,  Argentina.  Tvpes  and 
material  examined  are  deposited  at:  Museu  Oceanograf- 
ico  de  Rio  Grande,  Rio  Grande  do  Sul,  Brazil  (FURG); 
Museo  Argentino  de  Ciencias  Naturales  "Bernardino  Ri- 
vadavia",  Buenos  Aires  (MACN);  Museo  Nacional  de 
Historia  Natural,  Montevideo,  Uruguay  (MHNM);  Mu- 
seu de  Zoologia,  Universidade  de  Sao  Paulo  (MZSP); 
National  Museum  of  Natural  History,  Smithsonian  In- 
stitution, Washington  (USNM).  The  type  material  of  C. 
protea  was  studied  at  The  Natural  History  Museum, 
London  (BMNH). 

Specimens  were  fixed  in  fomiahn  for  at  least  24  hours 
and  preserved  in  70%  ethanol.  They  were  not  relaxed. 
All  observations  were  made  on  preserved  material.  Dis- 
sections were  perfonned  on  3  males  and  13  females  us- 
ing standard  techniques  under  a  stereomicroscope  with 
the  specimens  immersed  on  70%  ethanol.  Radulae  were 
prepared  (two  specimens  of  C  argentina  new  species  of 
32.2  and  35  mm  length  and  two  of  C.  protea  of  18.5  and 
17.4  mm)  according  to  the  method  described  by  Solem 
(1972)  and  observed  and  illustrated  using  a  LEO  440 
scanning  electron  microscope  (SEM)  at  the  National 
Museum  oi  Natural  History,  Washington. 

Shell  measurements  including  convexity  index,  were 
done  using  the  parameters  established  by  Hoagland 
(1977).  Most  photographs  were  taken  using  a  Nikon  N70 
with  a  60  mm  Nikkor  macro  lens.  These  images  were 
latter  scanned  from  black  and  white  35  mm  negatives 
using  a  Nikon  Coolscan  III  slide  scanner  All  images  and 
plates  were  processed  with  the  software  Photoshop  5.02. 

In  the  figures  and  text,  the  following  abbreviations  are 
used:  aa,  anterior  aorta;  ab,  auricle  region  beyond  ven- 
tricle connection;  ac,  anterior  extremity  of  gill  on  mantle 
border;  ag,  albumen  gland;  an,  anus;  au,  auricle;  bb, 
bulged  part  of  br;  bg,  buccal  ganglion;  br,  subradular 
membrane;  bv,  mantle  blood  vessel  inserting  in  kidney; 
eg,  capsule  gland;  cv,  ctenidial  vein;  da,  aperture  of  duct 
to  digestive  gland;  dd,  duct  to  digestive  gland;  dg,  di- 
gestive gland;  dm,  dorsal  shell  muscle;  dp,  posterior 
duct  to  digestive  gland;  ea,  esophageal  aperture;  en,  en- 
dostv'le;  es,  esophagus;  fd,  dorsal  surface  of  foot;  fg, 
food  groove;  fl,  female  papilla;  fp,  female  pore;  ft,  foot; 
gd,  gono-pericardial  duct;  gi,  gill;  gs,  gastric  shield;  bg, 
hypobranchial  gland;  in,  intestine;  ir,  insertion  ol  m4  in 
tissue  on  radula  preceding  its  exposed  (in  use)  portion; 
iu,  U-shaped  loop  of  intestine  on  palhal  roof;  kd,  dorsal 
lobe  of  kidney;  Id,  kidney;  kv,  ventral  lobe  of  kidney 
attached  to  intestine;  11,  left  lateral  ex-pansion  (Hap)  of 
neck;  hn,  lateral  membrane  restricting  paUial  cavity;  ml 
to  ml4,  odontophore  muscles;  mb,  mantle  border;  mj, 
muscles  of  jaws  and  mouth;  ml,  mantle  region  restricting 
palhal  cavity;  mo,  mouth;  mr,  mantle  reinforcement; 
mt,  mantle;  ne,  nephrostome;  nr,  nerve  ring;  ns,  neck 
"sole";  oc,  odontophore  cartilage;  os,  osphradium;  ov, 
paUial  oviduct;  oy,  ovary;  pc,  pericardium;  pd,  penis 
sperm  groove;  pe,  penis;  pg,  pedal  gland  anterior  fur- 
row; pp,  penis  papilla;  pr,  propodiuni;  pt,  palfial  sperm 


L.  R.  L.  Simone  et  ai,  2000 


Page  129 


Table   1.    Shell  morphometries  of  Crepidula  argentina  new 
species.  Linear  measurements  in  mm. 


Speci- 

Septum 

Convex- 

men 

Total 

Diam- 

Septum 

free 

ity 

# 

length 

Height  Width 

eter 

length 

area 

D/L 

1 

31.5 

6.3 

25 

37 

13.4 

17.9 

1.17 

2 

35.7 

7.7 

24.7 

41 

15.1 

19.8 

1.14 

3 

32.8 

8.6 

23.1 

39 

17.6 

17.5 

4.18 

4 

35.8 

8.9 

25 

42 

17 

19.4 

1.17 

5 

33.8 

8.8 

27 

43 

13.7 

21.8 

1.27 

6 

28.1 

7.1 

21.7 

36 

11.8 

16.4 

1.28 

( 

32.6 

7.3 

22.5 

39 

13.3 

17.6 

1.23 

8 

30 

7.3 

23.4 

37 

13.3 

16.8 

1.14 

9 

29.8 

7.1 

23.9 

34 

12.3 

16.3 

1.14 

10 

33 

9.3 

28 

41 

14.4 

19.3 

1.24 

groove;  ra,  radiila;  rl,  right  lateral  expansion  (flap)  of 
neck;  m,  radular  nucleus;  rs,  radular  sac;  sc,  subradular 
cartilage;  sg,  salivary  gland;  sn,  snout-proboscis;  sp,  ap- 
erture of  vas  deferens  into  pallial  cavity;  ss,  style  sac;  st, 
stomach;  sv,  seminal  vesicle;  te,  cephalic  tentacle;  tm, 
net  of  transversal  muscles  of  haemocoel;  to,  tissue  cov- 
ering middle  region  of  radula  before  its  exposed  part; 
ts,  testis,  vc,  visceral  connection  with  haemocoel;  ve, 
ventricle;  vg,  vaginal  duct;  vm,  visceral  mass;  vo,  visceral 
oviduct;  vs,  vesicles  of  pallial  oviduct. 

SYSTEMATICS 

Class  Gastropoda  Cuvier,  1797 

Subclass  Orthogastropoda  Ponder  and  Lindberg,  1996 

Superorder  Caenogastropoda  Cox,  1960 

Order  Sorbeoconcha  Ponder  and  Lindberg,  1997 

Family  Calyptraeidae  Lamarck,  1809 

Genus  Crepidula  Lamarck,  1799 


Crepidula  argentina  new  species 
Figures  1-21,  28^6 

Crepidula  protea:  Parodiz,  1939:  702,  fig.  8,  pi.  1,  fig.  6  (not 
d'Orbigny,  1841). 

Diagnosis:  Shell:  Large  and  wide,  slightly  convex.  Pro- 
toconch  smooth,  with  1%  whorls.  Aperture  elhptical. 
Beak  solid,  very  small.  Septum  planar,  septum  margin 
with  sulcus  on  left  side  and  clear  notch  in  center.  Muscle 
scars  absent.  Shell  externally  opaque  white,  internally 
white  porcelanaceus.  Periostracum  absent. 

Anatomij:  Large  lateral  shell  muscle.  Dorsal  shell  mus- 
cle weak.  Kidney  proportionally  smaU,  with  characteristic 
arrangement  of  inner  folds  of  dorsal  lobe,  renal  vessel 
edging  rectum.  Connection  between  odontophore  mus- 
cle pairs  m7  and  mil.  Long  salivary  glands.  Four  gastric 
ducts  to  digestive  gland,  distinctive  arrangement  of  folds 
(sorting  area)  in  inner  surface  of  stomach  between 
esophageal  aperture  and  posterior  pair  of  ducts  to  di- 
gestive gland.  Seminal  vesicle  broad  and  few  coiled.  Pe- 


nis and  its  apical  papilla  long  and  narrow.  Vaginal  tube 
running  attached  to  capsule  gland. 

Description:  Shell  (Figures  1-15,  20,  21):  Large  (up 
to  40  nun  in  length)  and  wide,  thin  (0.35—0.45  mm  in 
thickness),  slightly  convex,  convexity  =  1.20  (see  table  1 
for  other  measurements),  male  specimens  very  thin, 
brittle,  transparent,  planar  shells.  Protoconch  smooth, 
with  1%  whorls,  transition  to  teleoconch  clearly  defined. 
Aperture  elliptical  or  subcircular.  Beak  .soUd,  very  small, 
turned  to  right  on  females,  almost  central  on  males,  at 
level  of  or  only  slightly  above  margin,  in  males  never 
reaching  margin,  in  females  never  extended  beyond  it. 
Septum  planar  (never  convex),  with  central  ridge  almost 
imperceptible  but  present,  margin  of  septum  with  sulcus 
on  left  side  and  clear  notch  in  center,  covering  less  than 
half  of  aperture,  color  bright  white  with  translucent 
edge.  Muscle  scars  absent.  Growth  lines  covering  entire 
shell.  Shell  externally  opaque-white,  internally  porcehiin- 
white;  some  specimens  externally  with  diffuse  radial  or- 
ange lines  and/or  internally  with  radial  brownish  lines. 
Males  always  translucent-white  externally  and  bright- 
white  internally.  Periostracum  absent. 

Head-foot  (Figures  28,  30,  34,  43):  Head  protruded,  at 
the  end  of  long  (about  same  length  as  foot),  dorso-ven- 
trally  flattened,  neck  region.  Snout  short  and  cylindrical, 
able  to  retract  and  partially  invaginate  for  about  half  of 
its  length  within  haemocoelic  cavity.  Tentacles  long, 
stubby,  tip  somewhat  rounded.  Eyes  dark,  situated  on 
low  ommatophores  about  midway  on  lateral  margin  of 
tentacles.  Neck  region  with  pair  of  lateral,  flattened  ex- 
pansions (nuchal  lobes);  right  nuchal  lobe  bears  shallow 
food  groove  along  its  limit  with  head  (figure  28).  Sperm 
groove  of  males  (described  below)  running  externally 
along  food  groove  (figure  43).  Ventral  surface  of  neck 
region  forming  an  additional,  anterior  "sole"  (figure  30). 
Foot  veiy  ample  (occupies  about  %  of  shell  aperture), 
dorso-ventrally  flattened.  Shell  septum  defining  dorsal 
limit  of  foot.  Mantle  fusing  with  dorsal  surface  of  foot 
and  protruding  beyond  its  borders.  Furrow  of  pedal 
glands  transversal,  located  on  central  region  of  anterior 
margin  of  foot  (about  1/3  of  its  width);  a  small,  pointed 
expansion  present  on  each  side  of  furrow  (figure  30). 
Anterior  margin  of  foot  covering  ventrally  posterior  re- 
gion of  neck  "sole".  Columellar  muscle  very  reduced, 
small  flap  contouring  anterior  border  of  shell  septum, 
only  evident  on  right  side.  Inner  haemocoelic  cavity  nar- 
row, running  approximately  along  center  of  neck  region, 
almost  entirely  filled  by  mass  of  salivary  glands  (de- 
scribed below)  and  numerous,  transversal,  very  slender 
muscle  fibers  (figure  34);  these  fibers  connect  ventral 
surface  of  dorsal  haemocoelic  wall  with  dorsal  surface  of 
its  ventral  wall. 

Mantle  organs  (Figures  29,  31-33,  44):  Mantle  border 
very  thick,  edging  entire  ventral  inargin  of  sheU,  free  on 
its  anterior  half  and  attached  to  foot  edge  on  its  posterior 
half.  Mantle  border  without  appendages,  but  entirely 
covered  by  a  series  of  minute  glands.  Mantle  border  with 


Page  130 


THE  NAUTILUS,  Vol.  114,  No.  4 


Figures  1-15.  Shells  of  Crepidula  argentina  new  species.  1-3.  Holotvpe,  MACN  .3450S,  Mar  del  Plata,  Buenos  Aires,  Argentina. 
4-5.  Paratope,  MACN  .34.509,  Mar  del  Plata,  Buenos  Aires,  Argentina.  6-7.  Paratope,  MACN  .34509.  Puerto  Quequen.  Buenos 
Aires,  Argentina.  8-9.  Paratype.  10-11.  Paratype,  MACN  .34.509,  male  specimen,  scale  bar  on  right  side  of  figure  11  =  0.5  mm 
(for  figures  10  and  11).  12-13.  Paratjpe,  MACN  .34509  14-15.  Parat>pe,  MACN  .34509.  Scale  bar  under  figure  8  =  1  cm  for  all 
specimens,  except  figures  10  and  11. 


L.  R.  L.  Simone  ci  ai,  2()()() 


Page  131 


Figures  16-21.  Crepidula  argentina  new  species.  16-17.  Radula  16.  Radula,  frontal  view.  Scale  bar  =  30  \x.m.  17.  Radula, 
marginal  teeth,  scale  bar  =  30  jxni.  18-19.  Penis,  critical-point  dried.  18.  Dorsal  view.  Scale  bar  =  200  (xni.  19.  Ventral  \iew. 
Scale  bar  =  200  \Lm.  20-21.  Protoconch.  20.  Dorsal  view.  Scale  bar  =  300  ^.m.  21.  Detail  of  figure  20.  Scale  bar  =  100  p-ni. 


characteristic  arrangement  of  folds  along  central  region 
of  aperture  of  pallial  cavity  (figure  33),  a  broad  furrow 
beginning  on  anterior  extremity  of  gill,  running  toward 
left  side,  and  ending  at  about  right  third  of  osphradium; 
this  thick  fold  presents  broad  central  furrow.  Dorsal  shell 
muscle  apparently  lacking  in  most  specimens,  very  re- 
duced and  difficult  to  observe  in  others  (figure  29).  Lat- 
eral shell  muscle  well  developed,  inserting  broadly  in  left 
lateral  region  of  mande  border  and  region  posterior  to 
it,  originating  shortly  in  inner  shell  surface  in  region 
close  to  left  anterior  edge  of  shell  septum.  Pallial  cavity 
aperture  occupving  about  2/3  of  anterior  half  of  shell 
border,  turned  to  right  (drawing  an  analogy  between 
shell  in  dorsal  view  and  a  clock,  with  head  occupying 
12:00,  pallial  aperture  occupies  a  sector  beginning  at 
9:00  and  ending  at  2:00).  Pallial  cavity  deep,  broad,  tri- 
angular, arched  and  flattened  dorso-ventrally  (figure  29). 


Anterior  extremity  of  pallial  cavity  slightly  larger  than  its 
aperture  due  to  constriction  on  left  and  right  extremities 
produced  by  fusion  of  mantle  and  foot  (figures  31,  44). 
Pallial  ca\ity  narrows  gradually  in  posterior  direction, 
penetrating  left  side  of  visceral  mass  (described  below); 
cavity  length  about  3/4  of  total  length  of  animal.  Os- 
phradium (figure  33)  long,  monopectinate,  located  be- 
tween anterior  region  of  gill  and  mantle  border,  occu- 
pying about  middle  region  of  pallial  aperture,  somewhat 
perpenthcular  to  longitudinal  axis  of  animal,  compressed 
between  gill  and  mantle  border.  Osphradium  length 
slightlv  exceeding  1/5  of  palfial  aperture  length.  Osphra- 
dium leaflets  long,  somewhat  thick,  closely  packed,  with 
rounded  tip.  Gill  very  large,  with  somewhat  narrow  base, 
edging  anterior  and  left  margin  oi  pallial  cavity  along 
almost  its  entire  length;  anterior  extremity  of  gill  in  an- 
terior and  left  region  of  pallial  cavity  aperture,  near  its 


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THE  NAUTILUS,  Vol.  114,  No.  4 


Figures  22-27.  Crepidula  protea  dOrbigiiy,  1841.  22-24.  Shell,  MACN  34511,  off  Ubatuba,  Sao  Paulo,  Brazil,  2.3°30'  S,  44°54' 
W,  42  m,  dorsal,  ventral,  and  lateral  \iews  of  the  same  specimen.  25.  Shell,  MACN  34511,  other  specimen  in  lot,  ventral  view. 
Scale  bar  =  1  cm  for  all  shells.  26.  Radula,  frontal  view.  Scale  bar  =  100  |j.m.  27.  Protoconch,  dorsal  view.  Scale  bar  =  300  (Am. 


right  limit,  on  thick  mantle  border;  posterior  extremity 
of  gill  in  posterior  end  of  palhal  cavaty  (figure  32).  Base 
of  gill  filaments  triangular.  Gill  filament  with  very  long, 
almost  straight,  stiff  rod  extending  to  right;  rods  extend 
for  about  twice  as  long  as  their  triangular,  membranous 
base;  these  rods  begin  in  region  of  ctenidial,  in  left  mar- 
gin of  cavity  roof,  touching  food  groove  of  head-foot,  in 
right  margin  of  cavit)'  floor  Rod  apex  rounded  and  pre- 
ceded by  a  thicker  region.  Gill  filaments  connected  to 
each  other  by  cUia,  mainly  along  their  thicker  apical  re- 
gion, which  helps  maintain  somewhat  gill  structure.  Gill 
filaments  longer  in  central  region  of  gill,  shortening 
gradually  toward  both  extremities.  Anterior  extremity  of 
gill  with  short  filaments,  abruptly  turning  foiAvards,  end- 
ing at  mantle  border  (figure  33).  Ctenitliiil  vein  cylin- 


drical. Endostyle  well  developed,  yellowish  (a  somewhat 
narrow  glandular  ridge  located  on  ventral  surface  of 
ctenidial  vein  and  present  along  its  entire  length)  (figure 
33).  Hvpobranchial  gland  whitish,  low,  slightly  devel- 
oped, occupying  surface  between  gill  and  visceral  mass 
(figures  1,  5).  About  1/3  of  visceral  mass  encroaches  on 
pallial  cavity  roof,  occupying  about  1/3  of  posterior  and 
right  sectors  of  this  region;  pericardium  and  kidney  lo- 
cated in  posterior  part;  long  intestinal  loop,  anus,  and 
palhal  oviduct  in  anterior  part  (described  below). 

Visceral  mms  (Fiffircs  2.9,  31,  32,  44):  A  dorso-ventrally 
flattened  cone  lying  in  shell  chamber  produced  by  sep- 
tum. Thin  calcareous  septum  separating  visceral  mass 
from  dorsal  surface  of  foot.  Left  and  anterior  region  of 


L.  R.  L.  Simone  ct  at.,  2()()() 


Page  133 


30 


31 


Figures  28-31.  Anatomy  of  Crepklula  argentina  new  species.  28.  Female  animal  with  shell,  visceral  mass,  and  pallia!  cavity- 
removed,  dorsal  view,  29.  Same  animal,  whole,  dorsal  view.  30.  Same,  ventral  view.  31.  Isolated  visceral  mass  and  pallial  cavity, 
ventral  view.  Scale  bar  =  5  mm. 


visceral  ma.ss  occupied  by  palLal  cavity  (figures  31,  44). 
Remaining  region  of  visceral  mass  with  stomach  as  central 
structure,  immetliately  and  iilmost  completely  surrounded 
by  digestive  gland  (except  in  some  ventral  and  dorsal 
parts).  Gonad  surrounding  digestive  gland  externally. 


Visceral  mass  encroaching  on  right  and  posterior  re- 
gions of  palli;il  cavitv  roof.  Anterior  extremity  of  visceral 
mass  (ventral  to  posterior  pallial  cavity  region)  covering 
columellar  muscle  just  posterior  to  anterior  border  of 
shell  septum. 


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THE  NAUTILUS,  Vol.  114,  No.  4 


L.  R.  L.  Simone  et  a/.,  2000 


Page  135 


Circidatani  and  cxcrctoni  si/.stciii.s  (Figures  29,  35); 
Peric-archiini  \en  loiiij,  soiiiewhat  peqieiidicular  to  loTi- 
gitialinal  avis  ot  animal  (figure  29);  verv'  narrow  in  pos- 
terior extreniit)'  of  gill  (posterior  and  left  end  of  pallial 
cavity);  running  along  anterior  margin  of  xasteral  mass 
where  it  encroaches  on  piillial  roof,  enlarging  gradually 
and  ending  at  about  middle  level  of  this  region  of  vis- 
ceral mass,  near  its  mechan  line.  Pericartlium  circum- 
scribes (1)  palhal  cavit)'  in  anterior  and  ventral  direction, 

(2)  visceral  mass  (mostlv  gonad)  in  posterior  direction, 

(3)  mantle  in  dorsal  direction,  and  (4)  kidnev  to  right. 
Auricle  thin-walled  and  vew  long,  running  long  entire 
pericardium  length,  attached  to  its  anterior  and  dorsal 
inner  surfaces;  auricle  connects  with  ventricle  approxi- 
matelv  along  its  middle  and  right  thirds;  auricle  portion 
beyond  connection  with  ventricle  represented  by  broad 
binil  sac.  Ventricle  elliptical,  veiT  muscular,  its  connec- 
tion with  auricle  located  about  middle  region  of  its  an- 
terior surface;  aortas  originating  in  opposite  side.  Ante- 
rior aorta  broad,  directed  awav  from  posterior  aorta.  An- 
terior aorta  directed  toward  right,  edging  posterior  inner 
surface  of  pericardium.  Anterior  aorta  penetrates  head 
haemocoel.  Kidney  small,  occupying  about  1/4  of  area 
of  visceral  mass  when  projected  on  inner  surface  of  pal- 
lial cavity.  Kidnev  circumscribes  (1)  mantle  in  dorsal  ch- 
rection,  (2)  pallial  ca\ity  in  ventral  and  left  lateral  direc- 
tion, (3)  visceral  mass  (mostly  gonad)  in  posterior  and 
right  direction,  (4)  pericardium  in  posterior  and  left  di- 
rection, (5)  an  intestinal  loop  in  anterior  direction,  (6) 
intestine  and  oviduct  (when  present)  in  lateral  right  di- 
rection. Kidney  mostly  hollow,  with  pair  of  veiy  irregu- 
larly shaped  lobes.  Ventral  lobe  with  several,  slight  nar- 
row transverse  folds  attached  to  posterior  surface  of  ad- 
jacent intestine.  Dorsal  lobe  occupies  most  of  dorsal  and 
lateral  surfaces,  bearing  several  irregular  folds  in  left  di- 
rection; part  of  lobe  covering  ventral  surface  around 
nephrostome.  Nephridial  gland  thin,  present  along 
boundary  between  kidney  and  pericardium,  bearing  se- 
ries of  transverse,  narrow  folds  connected  with  dorsal 
renal  lobe.  Nephrostome  a  very  small  sht  in  left  region 
of  ventral  wall.  A  broad  vessel  connected  with  left  ex- 
tremity of  kidney  and  ninning  along  external  pallial  loop 
of  intestine  for  about  %  of  its  length. 

Digestive  system  (Figures  29,  31,  32,  34-42):  Mouth 
longitudinal,  in  center  of  anterior  surface  of  snout  (figure 
34).  Buccal  mass  very  large,  occupying  most  of  inner 
space  of  snout.  Buccal  mass  capable  of  some  protraction 
and  invagination.  Dorsal  wall  of  buccal  mass  with  pair 
of  broad  and  thin  jaw  plates.  Pair  of  df)rsal  folds  broad 
and  low,  connected  to  jaws  in  posterior  direction.  Odon- 


tophore  large,  consisting  of  most  of  volume  of  buccal 
mass.  Odontophore  muscles  (figures  36-38,  40,  41): 
(ml)  jugai  muscles,  several  verv  narrow  muscles  con- 
necting buccal  ma.ss  to  adjacent  wall  of  snout,  more  con- 
centrated anteriorly  around  mouth;  (mla)  pair  of  dorsal 
protractor  muscles,  narrow,  thin  and  superficial,  with  or- 
igin in  antcro-dorsal  region  of  mouth,  close  to  its  median 
line,  insertion  in  posterio-dorsal  and  lateral  region  of 
odontophore;  (mj)  jaws  and  peribuccal  muscles,  some- 
what thick,  surrounding  lateral  and  dorsal  wall  of  buccal 
mass,  with  origin  around  mouth,  insertion  in  middle  re- 
gion of  lateral  and  dorsal  wall  of  odontophore;  (m2)  pair 
of  retractor  muscles  of  buccal  mass  (retractor  of  phar- 
ynx), broad,  with  origin  in  latero-ventral  region  of  hae- 
mocoel just  posterior  to  snout,  nmning  in  anterior  di- 
rection, with  insertion  in  postero-Iateral  and  dorsal  re- 
gion of  odontophore  cartilages;  (m2a)  pair  of  dorsal  ten- 
sor muscles  of  radula,  continuation  of  m2  after  insertion 
in  cartilages,  nmning  in  anterit)r  direction,  with  insertion 
in  subradular  cartilage  in  middle  region  of  its  dorsal  in- 
ner surface;  (mt)  dorsal  transversal  muscle  or  ventral  ap- 
proximator muscle  of  cartilages,  connecting  dorsally  pos- 
terior-dorsal and  lateral  surface  of  both  cartilages,  lying 
between  superficial  membrane  that  covers  odontophore 
and  tissue  on  middle  region  of  radula  (to);  (m4)  pair  of 
median  dorsal  tensor  muscles  of  radula,  very  large  and 
thick,  with  origin  in  ventral-central  and  posterior  region 
of  odontophore  cartilages,  running  along  their  middle 
region,  contouring  meso-ventral  surface  of  cartilages, 
running  along  their  dorsal  surface,  with  insertion  in  dor- 
sal-posterior and  medial  extremity  of  subradular  carti- 
lage; (mSf)  pair  of  median  radular  tensor  muscles,  thick, 
with  origin  in  meso-posterior  and  dorsal  regions  of  odon- 
tophore cartilages,  just  adjacent  to  m2  insertion  and  m2a 
origin,  crossing  middle  region  of  m4,  running  toward 
medial  region  of  m4,  with  insertion  along  radular  sac  on 
both  sides  (each  branch  of  m5  nms  along  a  side  of  rad- 
ular sac,  medially  and  dorsally);  (m6)  horizontal  muscle, 
very'  thin,  uniting  anterior  half  of  odontophore  cartilages, 
with  insertion  on  their  dorsal  margin;  (m7)  pair  of  ven- 
tral tensor  muscles  of  radula,  thin  and  narrow,  with  or- 
igin in  meso-anterior  margin  of  m4,  running  in  posterior 
direction  adjacent  to  subradular  membrane,  bifurcating 
in  posterior  region  of  odontophore,  median  branch  con- 
necting with  that  of  other  member  of  pair  and  inserted 
in  meso-posterior  region  of  radular  sac,  lateral  branch 
connecting  with  mil;  (m8)  pair  of  strong  muscles  with 
origin  in  postero-dorsal  and  lateral  region  of  odonto- 
phore cartilages  just  adjacent  to  insertion  of  m2,  nmning 
along  and  attached  to  dorsal  margin  of  odontophore  car- 


Figures  32-36.  Anatomy  oi  Crepidula  ar^entina  new  species.  32.  Pallial  cavity,  ventral  view,  part  of  visceral  mass  deflected,  some 
gill  filaments  of  central  part  of  gill  removed.  33.  Detail  of  mantle  border  and  pallial  cavity  at  anterior  extremity  of  gill.  34.  Head 
and  haemocoel,  ventral  view,  foot  and  neck  "sole"  removed.  35.  Detail  of  visceral  mass  at  border  of  pallial  cavit)-,  adjacent  to 
pericardial  structures,  ventral  view,  ventral  surface  of  pericardium  and  kidney  membranes  removed.  36.  Buccal  mass,  dorsal  view. 
All  scale  bars  =  1  nun. 


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THE  NAUTILUS,  Vol.  114,  No.  4 


mo 


ra 


to      \ 


37 


mil     /  . 

m         rnl4 


40 


L.  R.  L.  Simone  et  al.,  2000 


Page  137 


tilages,  with  insfrtioii  in  their  antero-tlorsal  region  ad- 
jacent to  horizontal  muscle  (m6);  (ni9)  pair  of  dorso- 
medial  tensor  muscle  of  radula,  broad  and  thin,  with 
origin  along  ineso-dorsal  surface  of  radular  sac  (in  its 
region  internal  to  odontophore),  crossiTig  to  dorsal  sur- 
face, with  insertion  in  dorso-ventral  margin  of  suhradn- 
lar  cartilage;  (mil)  small,  narrow  pair  of  muscles,  with 
origin  in  meso-ventnd  region  of  mouth,  nmning  in  pos- 
terior direction  along  its  median  line,  penetrating  in  me- 
dian region  of  odontophore,  rumiing  between  m7  and 
m4,  with  insertion  in  anterior  region  of  subradular  mem- 
brane; (ml4)  pair  of  broad  and  thin  muscles,  with  origin 
in  postero-dorsal  region  of  otlontophore,  close  to  origins 
of  m2  and  m5,  nmning  in  antero-ventral  direction,  with 
insertion  in  inner  ventral  surface  of  snout  at  about  mid- 
dle region  of  odontophore;  to)  tissue  covering  mitldle 
region  of  radida  within  odontophore,  along  its  dorsal  sur- 
face; br)  subradular  membrane,  covering  ventral  surface 
of  subradular  cartilage  and  some  neighboring  areas. 

Radula  taenioglossate,  short,  measuring  little  more 
than  odontophore  length  (figures  37,  38).  Rachidian 
tooth  tall,  narrow  with  cin^ved,  convex  base,  central  cusp 
ver)  large  and  shaqi,  at  least  two  weak  denticles  on  each 
side  decreasing  in  size  toward  lateral  teedi,  no  basal 
cusps  but  pair  of  lateral  reinforcements  present  along 
borders.  Lateral  tooth  broad  (about  3  times  rachidian 
width),  curved  inward,  with  conspicuous  apical  cusp 
turned  toward  rachidian  line  and  5-7  short,  triangular 
denticles  along  edge  on  marginal  side  and  2—3  very 
weak  denticles  on  edge  on  rachidian  side,  denticles  de- 
creasing in  size  in  both  directions,  disappearing  at  about 
middle  region  of  tooth,  only  a  thickened  border  remain- 
ing. Marginal  teeth  long,  curved,  tall,  tip  sharply  pointed, 
with  serrate  inner  margin  (at  least  7  denticles);  inner 
marginal  tooth  broad,  about  twice  as  wide  as  outer  mar- 
ginal tooth. 

Pair  of  buccal  ganglia  large,  close  together  near  me- 
dian line  (figure  37),  situated  between  buccal  mass  and 
adjacent  esophagus.  Salivary  glands  very  long,  tubular, 
coiled  (about  3  times  longer  than  haemocoel  length 
when  straightened)  (figure  34).  Several  narrow  trans- 
verse muscles  unite  internally  dorsal  and  ventral  surfaces 
of  haemocoel,  passing  both  sides  of  salivary  glands, 
esophagus,  and  aorta  (figure  34).  Salivary  glands  not 
passing  through  nerve  ring.  Ducts  of  salivary  glands 
thick,  running  to  dorsal  surface  of  buccal  mass,  pene- 
trating adjacent  wall  of  buccal  mass;  apertures  small, 
close  to  site  of  penetration,  located  in  anterior  region  of 
dorsal  folds  of  buccal  mass  (figure  36). 

Esophagus  narrow  and  long  (figure  39).  Inner  surface 
of  anterior  esophagus  with  a  pair  of  broad  folds.  Middle 


esophagus  with  pair  of  narrow  folds  (continuation  from 
those  of  anterior  esophagus)  and  moderately  spacious 
glandular  chamber.  Iimer  surface  of  posterior  esophagus 
with  4—5  longitudinal,  narrow,  folds  of  similar  (hmen- 
sions.  Stomach  (figures  .39,  42)  large,  slightly  conical,  oc- 
cupying about  half  of  visceral  mass;  esophagus  inserting 
in  left  side  of  its  po.sterior  region  of  stomach,  adjacent 
to  region  of  shell  apex.  Four  ducts  to  digestive  gland 
present:  anterior  pair  narrow,  inserted  in  ventral  surface, 
between  insertion  of  esophagus  and  posterior  gastric 
end,  one  turned  in  anterior  direction  and  other  to  op- 
posite side;  posterior  pair  of  ducts  thick,  located  in  mid- 
dle region  of  ventral  and  left  surface  of  stomach,  one  at 
considerable  distance  from  other.  Stomach  graduallv 
narrows  in  anterior  and  left  direction,  close  to  left  and 
posterior  extremity  of  paUial  cavity.  Stomach  inner  sur- 
face (figure  42)  with  p;ur  of  narrow  and  long  folds,  both 
with  origin  adjacent  to  insertion  of  esophagus,  lining 
posterior  surface  of  stomach,  gradually  disappearing  in 
direction  posterior  to  apertures  of  anterior  ducts  to  di- 
gestive gland.  Gastric  shield  thin,  presenting  transversal 
folds  marking  a  somewhat  elfiptical,  ccjucave  area.  An- 
terior half  of  stomach  with  a  pair  of  slight  tall,  longitu- 
dinal folds;  posterior  region  between  both  folds  smaller 
than  anterior  region;  smaller  region  as  intestinal  branch 
of  stomach;  broader  (anterior)  region  as  style  sac.  Di- 
gestive gland  pale-brown  in  color,  surrounding  stomach 
except  for  some  spots  in  dorsal  and  ventral  surfaces. 

Intestine  very  narrow  and  sinuous  (figure  39);  running 
from  left  to  right  adjacent  to  anterior  border  of  stomach, 
running  up  near  median  fine  to  right  and  anterior  ex- 
tremity of  visceral  mass;  in  this  region  and  toward  left, 
it  surrounds  right  and  anterior  border  of  kidney,  turning 
suddenly  to  right  and  nmning  pandlel  to  preceding  loop; 
both  loops  exposed  along  pallial  roof  (figures  29,  32,  35, 
39).  Anus  small,  sfightly  siphoned,  situated  in  right  re- 
gion of  pallial  cavity  close  to  mande  border  (figures  31, 
32,  39).  Last  loops  of  intestine  contiiin  numerous,  small, 
somewhat  elliptical  fecal  pellets. 

Male  genital  sifstem  {Figures  43.  44,  46):  Only  small 
specimens  (up  to  9  mm)  are  functional  males.  Testis  sit- 
uated in  anterior  region  of  visceral  mass,  color  orange. 
Seminal  vesicle  convoluted,  very  thickened,  color  cream, 
situated  in  anterior  and  right  region  of  visceral  mass, 
where  it  abruptly  narrows,  becoming  very  slender  and 
opening  in  postero-ventral  and  right  region  of  pallial  cav- 
itv.  A  shallow  groove  runs  from  this  aperture  to  penis 
liase,  on  pallial  floor  near  right  margin  of  head.  Sperm 
groove  better  defined  and  deeper  in  anterior  direction. 
Penis  long  (about  3  times  tentacle  length),  curved,  orig- 


Figures  37-41.  Anatomy  oi  Crcpidiila  nruentina  new  species.  37.  Buc'c;il  mass,  ventral  \iew.  38.  Odontophore,  dorsal  view,  some 
muscles  sectioned  and  deflected,  part  of  right  region  of  subradular  curtilage  remo\ed  to  show  muscular  insertions  in  it.  39.  Digestive 
tubes  seen  in  situ  with  visceral  mass  as  a  transparent  structure,  ventral  view.  40.  Odontophore,  ventral  view,  superficial  membrane 
and  muscles  removed,  nght  mj  (left  in  figure)  also  removed.  4L  Odontophore,  ventral  view,  radular  sac  deflected  and  only  partially 
shown,  odontophore  cartilages  deflected  from  each  other,  right  m4  (left  in  figure)  deflected  downward.  All  scale  bars  =  1  mm. 


Page  138 


THE  NAUTILUS,  Vol.  114,  No.  4 


L.  R.  L.  Sinione  et  al,  2000 


Page  139 


inating  dorsallv  to  right  tentacle.  Papilla  on  penis  tip, 
very  long,  about  half  of  penis  length.  Penis  duct  opened, 
running  along  middle  region  of  ventral  surface  of  penis 
to  tip  of  papilla. 

Female  genital  st/stem  (Figures  29,  31,  32,  35,  45): 
Ovarv  pale  brown,  surrounding  digestive  gland,  denser 
in  anterior  region  of  visceral  mass  (figures  29,  31).  Vis- 
ceral oviduct  \er\'  narrow,  ninning  from  left  to  right  in 
anterior  border  of  \asceral  mass.  Gonopericardial  duct 
well  developed,  slightlv  thicker  than  visceral  oviduct, 
with  origin  in  ventral  and  right  extremitv  of  pericardium 
(figure  35);  nmning  along  visceral  glands  encroached  in 
pallial  cavity;  inserted  in  posterior  extremity  of  pallial 
oviduct  where  it  joins  insertion  of  visceral  oviduct.  Al- 
bumen gland  long,  moderately  thick,  whitish;  walls  thick, 
glandular;  situated  in  anterior  and  right  extremity  of  vis- 
ceral mass;  3  to  4  seminal  receptacles  inserted  along 
right  surface  of  albumen  gland.  Capsule  gland  a  contin- 
uation of  albumen  gland,  marked  by  sudden  increase  of 
secretorv  tissue  in  wall  and  bv  turn  toward  left;  walls 
irregular,  thick,  glandular.  Vaginal  tube  moderately  nar- 
row, originating  in  posterior  region  of  capsule  gland,  run- 
ning attached  to  this  latter  up  to  its  right  limit,  where  it 
abniptlv  turns  in  ventral  direction  to  form  tall  genital 
papilla.  Papilla  with  p;ur  of  low  folds  iimning  along  its 
posterior  side,  from  base  almost  to  tip;  folds  close  to 
each  other  with  narrow  furrow  in  between.  Female  gen- 
ital pore  situated  in  tip  of  papilla  (figures  31,  32,  45)  sfit- 
hke,  transversal,  with  posterior  and  anterior  edges  slight- 
ly projected. 

Etymology:  The  specific  epithet,  a  noun  in  apposition, 
refers  to  the  name  of  the  countiy  where  the  species  oc- 
curs, Argentina. 

Type  locality:  Mar  del  Plata,  Buenos  Aires,  Argentina 
in  35-50  m,  on  shells  of  Mytiliis  cdiilis  platcnsis 
d'Orbigny,  1846. 

Type  material:  Holotyjie,  MACN  34508;  20  paratv- 
pes,  MACN  .34509  (10  dn,'  specimens)  and  MACN 
34510  (10  specimens  preserved  in  ethanol);  6  paratypes, 
MLP  5578;  16  paratypes  (3  males,  13  females)  MZSP 
32152;  6  paratypes  USNM  2016009  (1  specimen  pre- 
served in  ethanol);  MHNM,  6  specimens  without  num- 
ber; 6  paratypes  MHNM  15105;  all  from  type  locality. 

Additional  material  examined:  MACN  18504,  39 
specimens,  Puerto  Quequen,  Buenos  Aires;  MACN 
8887,  9  specimens,  off  Mar  del  Plata,  in  46  m;  MACN 
11367,  36  specimens;  MACN  9361^9,  5  specimens; 
MACN  18374,  2  specimens;  MACN  8653,  4  specimens; 
MACN  11586,  27  specimens;  all  from  Mar  del  Plata; 
MACN  20529-1  more  than  60  specimens,  mouth  of  Rio 
Negro. 


Literature  record.s:  Due  to  similarities  with  C  pro- 
tea,  most  hteratnre  records  should  be  re-checked  from 
now  on.  Records  cited  by  Parodiz  (1939)  were  revised 
and  are  included  in  the  material  examined. 

Di.stribution:  Province  of  Buenos  Aires,  Argentina, 
from  Mar  del  Plata  to  the  mouth  of  Rio  Negro,  on  banks 
of  Mytiltis  edulis  phiten.sis,  35-50  m  depth.  Records 
from  Uiiiguav  and  Brazil  need  to  be  confirmed. 

Reproductive  pattern:  Crepidula  argentina  new  spe- 
cies is  a  protandrous  hermaphrodite  that  undergoes 
complete  sex  change.  Males  mature  at  about  4  mm 
length.  Sex  change  begins  when  individuals  reach  ap- 
proximately 9  mm  and  is  completed  when  the  first  pre- 
vitelogenic  ovocites  appear,  usually  at  around  11  mm 
length.  The  smallest  broochng  female  we  found  was  15.0 
mm  length  and  the  largest  39.0  mm. 

The  new  species  has  a  well-defined  seasonal  repro- 
ductive cycle.  Between  September  and  March  more 
than  30%  of  the  females  are  brooding,  with  maximum 
brooding  in  January  (57%).  In  June  and  July  brooding  is 
uncommon  (0-10%).  Females  l)rood  1—46  egg  capsules 
per  spawn.  The  average  number  of  embryos  per  spawn 
is  5600.  There  is  no  correlation  between  size  of  the  fe- 
male and  number  of  egg  capsules  or  eggs  per  spawn. 
However,  female  size,  capsule  size,  and  the  average 
number  of  embryos  per  capsule  are  positively  correlated 
(Cledon  and  Penchaszadeh,  submitted).  Uncleaved  egg 
diameter  is  170  |xm  and  there  are  about  320  eggs  per 
egg  capsule.  All  eggs  develop,  there  are  no  nurse  eggs, 
and  the  onlv  extra-embi"vonic  nutritit)nal  source  is  the 
intracapsular  liquid.  There  is  no  record  of  cannibalism 
in  early  or  advanced  stages  of  development.  The  embry- 
os hatch  as  plauktotrophic  veliger  lanae. 

DISCUSSION 

Crepidula  argentina  new  species  is  usually  foimd  on  the 
posterior  edge  of  fiviug  shells  of  Mytiliis  edulis  platensis 
d'Orbigny,  1846,  typically  in  association  with  Calyp- 
traeotheres  gaiihi  (Fenucci,  1975)  (Crustacea:  Brachi- 
ura).  Mvtihd  banks,  distributed  all  around  the  coast  of 
Argentina  (Penchaszadeh,  1971a)  are  a  suitable  habitat 
for  Crepidula  argentina. 

Crepidula  argentina  was  included  by  different  authors 
(Parodiz,  1939;  Hoagland,  1977;  1983,' in  part)  in  C.  pro- 
tea  and  referred  to  as  C.  ungiiifonnis  bv  Penchaszadeh 
(1971b:  480).  The  t\pe  specimens  of  C.  protea  were  ex- 
amined by  Hoagland  (1983).  At  BMNH,  there  are  two 
lots  with  5  and  11  syntypes  under  the  numbers 
18.54.12.4.573  and  574  respectively.  Aguirre  (1993)  des- 
ignated and  illustrated  the  only  whole  specimen  as  lec- 
totype   plus   tvvo   paralectotypes,   with   no   mention   to 


Figures  42-46.  Anatomy  of  Crepidula  an^cntina  new  species.  42.  Stomach,  ventral  view,  inner  surface  exposed  by  means  of  a 
longitudinal  incision.  43.  Head  of  male,  dorsal  view.  44.  Visceral  mass  and  adjacent  part  of  pallial  cavity,  male,  ventral  view;  45. 
Pallial  oviduct,  ventral  view.  46.  Penis,  ventral  view.  All  scale  bars  =  1  mm. 


Page  140 


THE  NAUTILUS,  Vol.  114.  No.  4 


Hoagland's  paper  or  d'Orbigny's  (in  Sagra)  illustrations. 
Most  of  the  type  specimens  are  smaller  than  30  mm  and 
have  the  conchological  attributes  of  what  Hoagland  de- 
scribed for  C.  protea  from  Brazil.  In  terms  of  shell  char- 
acters, C.  protea  and  C.  argcntino  new  species  are  sim- 
ilar. Measurements  of  C.  argentina  in  Table  1  are  iden- 
tical to  those  given  to  C.  protea  by  Hoagland  (1977). 
However,  in  a  later  paper,  Hoagland  (1983)  mentioned 
that  the  largest  specimen  of  C.  protea  was  20  mm  long. 
Therefore,  we  suspect  that  both  species  were  combined 
under  the  name  C.  protea  in  her  1977  paper  Crepidida 
argentina  is  larger  and  wider,  particularly  in  young  spec- 
imens. Crepidula  protea  has  an  elliptical  aperture  with 
a  thicker  and  more  convex  shell,  while  the  new  species 
has  a  subcircular  aperture  and  a  thinner  and  flatter  shell. 
Both  species  differ  primarily  in  their  anatomical  features 
and  reproductive  strategy'.  Further  anatomical  study  of 
C.  protea  is  provided  in  another  paper  (Simone,  sub- 
mitted); some  data  from  that  paper  are  here  discussed 
for  comparative  purposes.  Crepidula  argentina  differs 
moq^hologically  from  C.  protea  in  that  C.  argentina  has: 
1)  a  more  developed  lateral  shell  muscle,  2)  a  poorly 
developed  dorsal  shell  muscle,  3)  the  kidney  proportion- 
ally smaller  and  with  a  different  arrangement  of  inner 
folds  of  dorsal  lobe,  4)  presence  of  a  renal  vessel  edging 
rectum  and  working  as  adrectal  sinus,  5)  a  connection 
between  odontophore  muscle  p<iirs  m7  and  mil,  6)  a 
longer  salivary  gland,  7)  four  ducts  to  digestive  gland  in 
stomach,  instead  of  two,  8)  different  arrangement  of 
folds  in  inner  surface  of  stomach  between  esophageal 
aperture  and  posterior  pair  of  ducts  to  digestive  gland, 
9)  seminal  vesicles  of  males  broader  and  less  coiled,  10) 
penis  and  papilla  longer  and  narrower,  and  11)  vaginal 
tube  running  closely  attached  to  capsule  gland. 

Embyological  features  of  Crepidula  argentina  are  very- 
distinct  from  those  of  C.  protea  (fide  Hoagland,  1983). 
The  total  number  of  embryos  per  spawn  and  per  egg 
capsule  and  the  size  range  of  brooding  females  are  die 
most  remarkable  differences.  Furthermore,  Hoagland 
(1986)  described  later  stages  of  developing  embryos  (ve- 
liger  stage)  as  pinkish  in  color  and  embedded  in  a  sticky 
gelatinous  matrix  in  which  the  embryos  did  not  move 
freely.  We  never  observed  this  in  pre-hatching  stages  of 
C.  argentina. 

It  is  possible  to  differentiate  several  species  of  Cre- 
pidula along  die  southern  Atlantic  coast  of  South  Amer- 
ica. Parodiz  (1939)  cited  five  species  of  Crepidula  for 
the  Argentine  coasts:  C.  ddatata  Lamarck;  C.  aculcata 
(Gmelin);  C  protea  d'Orbignv;  C  onyx  Sowerby  and  C. 
tinguiformis  Lamarck.  Specimens  of  Crepidula  ungui- 
fonnis  were  described  by  Parodiz  as  usually  associated 
with  hermit  crabs.  The  distribution  of  C.  unguifonnis  is 
given  by  Hoagland  (1977)  as  restricted  to  the  Mediter- 
ranean Sea  and  Northern  Africa.  It  is  probable  that  "C. 
unguifonnis"  sensu  Parodiz  is  a  distinct,  perhaps  un- 
named, species.  Crepidula  oni/x  from  the  northern  Pa- 
cific was  cited  by  Parodiz  (1939)  as  Uving  in  northern 
Patagonia.  However,  Hoagland  (1977)  based  on  Parodiz's 
drawing  of  the  radula  of  C.  onyx  presumed  that    "C. 


onyx"  sensu  Parodiz  is  a  different  species:  C  aph/sioides 
Reeve,  1859.  The  latter  species  was  cited  as  a  sviionyin 
of  C.  onyx  by  Parodiz  (1939).  Hoagland  (1977)  described 
C.  aphjsioides  as  a  different  species  ranging  from  Gre- 
nada south  to  Brazil  and  Argentina.  Reeve's  Upes  were 
not  examined.  This  type  material  includes  four  speci- 
mens and  is  housed  at  the  BMNH  (Reg.  Number; 
1977137).  As  far  as  we  know,  there  is  no  species  similar 
to  C.  aphjsioides  Reeve  on  the  coast  of  Argentina. 

The  study  of  the  family  Calyptraeidae  and  the  genus 
Crepidula  in  particular  is  less  than  finished  for  the  re- 
gion in  study.  It  is  expected  that  additional  new  species 
will  be  detected  and  described  following  detailed  studies 
on  anatomy  and  reproductive  biology  of  these  moUusks. 

ACKNOWLEDGMENTS 

We  thank  the  following  people  for  access  to  material  in 
their  collections:  K.  M.  Way  (BMNH);  A.  Tablado 
(MACN);  and  C.  Ituarte  (MLP).  We  are  grateful  to  two 
anonymous  reviewers  for  thorough  revision  and  helpful 
suggestions  that  considerably  improved  the  clarity  of  the 
manuscript. 

Several  SEM  pictures  were  taken  during  a  short-term 
visit  to  the  National  Museum  of  Natural  History,  Smith- 
sonian Institution,  Washington,  DC.  This  study  was  sup- 
ported in  part  b\'  a  Cooperative  Argentina-Brazil-Chile 
Research  Grant  from  Fundacion  Antorchas,  Argentina, 
a  grant  to  L.  R.  L.  Simone  from  Funda^ao  de  Amparo 
a  Pesquisa  do  Estado  de  Sao  Paulo  (FAPESP  grant  #  96- 
06756-2)  and  the  project  BID  802/OC-AR-PICT  No.  01- 
04321  from  the  National  Agency  for  Scientific  and  Tech- 
nique Promotion,  Argentina. 

LITERATURE  CITED 

Aguirre,  M.  L.  199.3.  Type  specimens  of  Quaternary  marine 
gastropods  from  Argentina.  Ameghiniana  30:2.3-.38. 

Brown,  D.  I.  and  C.  A.  Olivares.  1996.  A  new  species  o£  Cre- 
pidula (Mollusca:  Mesogastropoda:  CaKptraeidae)  from 
Chile:  additional  cliaracters  for  the  identification  of  east- 
ern Pacific  planar  Crepidula  group.  Journal  of  Natural 
Historv- .30: 144:3-1458. 

Cledon,  M.  and  P.  E.  Penchaszadeh.  Submitted.  Reproduction 
and  brooding  of  Crepidula  argentina  Simone,  Pastorino 
and  Penchaszadeh  (Gastropoda:  Calyptraeidae).  The  Nau- 
tilus. 

Cox,  L.  R  1960  Thoughts  on  the  classification  of  the  Gastro- 
poda. Proceedings  of  the  Malacological  Societv  of  London 
33:239-261. 

Dall,  W.  H.  1S89.  Reports  on  the  results  of  dredging  by  the 
U.  S.  Coast  SuA'ey  Steamer  "Blake",  report  on  the  mol- 
lusca part  2:  Gastropoda  and  Scaphopoda.  Bulletin  of  the 
Museum  of  Comparative  Zoolog\'  18:1—492. 

Gallardo,  C.  S.  1979.  Especies  gemelas  del  genero  Crepidida 
(Gastropoda,  Calyptraeidae)  en  la  costa  de  Chile;  una  re- 
descripcion  de  C.  dilatata  Lamarck  y  descripcion  de  C. 
fecunda  n.  sp.  Studies  on  Neotropical  Fauna  and  En\n- 
ronment  14:215-226. 

Gallardo,  C.  S.  1977.  Crepidula  philippiana  n.  sp.,  nuevo  gas- 
tropodo  Calyptraeidae  de  Chile  con  especial  referencia  al 


L.  R.  L.  Simone  et  ai,  2000 


Page  141 


patron  de  desanollo.  Stiiilies  on  Neotropical  Fanna  an<! 
Enxironnient  12:177-185. 

Hoagland,  K.  E.  1977.  Systematic  review  of  fossil  and  Recent 
Crepidula  and  discussion  of  e\'olution  of  the  Calvptraei- 
dae.  Malacologia  16;353-420. 

Hoagland,  K.  E.  1983.  EcologN'  and  lanal  development  of  Crc- 
piilula  protea  (Prosoliranchia:  Orepidiiiidae)  from  south- 
ern Brasil:  a  new  t)pe  of  egg  capsule  for  the  gemis  The 
Nautilus  97:105-109. 

Hoagland,  K.  E.  1986.  Patterns  of  encapsulation  and  hrooding 
in  the  Calvptraeidae  (Prosobranciiia:  Mesogastropoda). 
American  Malacological  Bulletin  4:173-183. 

Keen,  A.  M.  1966.  West  American  mollusk  types  at  the  British 
Museum  (Natural  History)  HI.  Alcide  d'Orbigny's  South 
American  collection.  The  Vehger  9:1-7. 

d'Orbigny,  A.  D.  1834-1847.  Voyage  dans  i'Amerique  Meri- 
dionale.  Mollusques.  Volume  5,  partie  3,  atlas,  85  pis.  Par- 
is, Chez  Ve.  Levrault,  758  pp.  |  Publication  dates  in  Keen 
(1966,)] 

d'Orbigny.  A.  D.  1841-1853.  Mollusques  In:  Sagra,  R.de  la 
(ed.)  Histoire  phvsique,  poHtique  et  naturelle  de  I'lle  de 
Cuba.  Volumes  1  and  2,  380  pp.,  atlas,  28  pis,  Paris.  [Puli- 
lication  dates  in  Sherboni  and  Griffin  (19.34)] 

Parodiz,  J.  J.  1939.  Las  especies  de  Crepidula  de  las  costas 
Argentinas.  Physis  17:685-709. 

Penchaszadeh,  P.  E.  1971a.  Estudios  sobre  el  mejillon  {Mt/tiliis 
platensis  d'Orb.),  en  ex|3lotaci6n  comercial  del  sector  bon 
aerense.  Mar  Argentino.  I.  Reproduccion,  crecimiento  y 
estructura  de  la  poblacion.  FAO.  CARPAS  5/D  Tc.  12:  1- 
15. 

Penchaszadeh,  P.  E.  1971b.  Aspectos  de  la  embriogenesis  de 
algunos  gasteropodos  del  genero  Bticcinanops  d'Orbigny, 
1841  (Gastropoda,  Prosobranchiata,  Buccinidae).  Physis 
30  (81):475-^82. 


Ponder,  'W'.  E  and  D.  R.  I.intlberg  1996.  Giistropod  Phvloge- 
ny — Challenges  for  tlie  90s.  In:  Taylor,  J.  D.  (ed.)  Origin 
and  evolutionary  radiation  of  the  Moliusca.  Oxford  Uni- 
versity Press,  Oxford,  pp.  135-154  pp. 

Ponder,  W.  F.  and  D.  R.  Lindberg.  1997.  Towarils  a  phylogeny 
of  gastropod  molluscs:  an  analysis  using  morphological 
characters.  Zoological  Journal  of  the  Linnean  Society  119: 
83-265. 

Reeve,  L.  A.  1859.  Conchologia  Iconica:  or,  illustrations  of  the 
shells  ot  molu.scous  animals. Monograph  of  the  genus  Cre- 
pidula V.  11,  5  pis. 

Rios,  E.  de  C.  1985.  Seashells  of  Brazil.  Editora  da  Fundafao 
Universidade  do  Rio  Grande,  Rio  Grande,  328  pp. 

Say,  T.  1822.  An  account  of  the  marine  shells  of  the  United 
States.  Journal  of  the  Academy  of  Natural  Sciences  of 
Philadelphia  2:221-227. 

Sherboni,  C.  D.  and  B.  B.  Woodward.  1901  Notes  on  the 
dates  of  publication  of  the  natural  history  portions  of  some 
French  voyages.  Part  1.  'Amerique  meridionale";  Indes  or- 
ientales';  'Pole  Sud';  I'Astrolabe  and  'Zele';  La  Bonite';  La 
Coquilie';  and  "PUranie  et  Physicienne'.  Annals  and  Mag- 
azine of  Natural  History,  series  7,  7:  388-.392. 

Sherboni,  C.  D.  and  F  J.  Griffin.  1934.  On  the  dates  of  pub- 
lication of  the  natural  history  portions  of  Alcide 
d'Orbigny's  Voyage  Amerique  Meridionale'.  Annals  and 
Magazine  of  Natural  History,  series  10,  13:130-134. 

Simone,  L.  R.  L.  Submitted,  Morphological  comparative  study 
of  representatives  of  the  Superfamilies  Caivptraeoidea  and 
Hipponicoidea,  (Moliusca,  Caenogastropoda)  with  ac- 
counts on  their  phylogeny.  Journal  of  the  Linnean  Society. 

Soleni,  A.  1972.  Malacological  application  of  scanning  electron 
microscopy.  II.  Radular  structure  and  functioning.  The 
Veliger  14:327-336. 


THE  NAUTILUS  114(4):  142-148,  2000 


Page  142 


Three  new  species  of  the  genus  Merica  (Neogastropoda: 
Cancellariidae)  from  South  Africa  and  the  PhiUppines 


Richard  E.  Petit' 
M.  G.  Harasewych 

Department  of  Invertebrate  Zoology 
National  Museum  of  Natural  History 
Smithsonian  Institution 
Washington,  DC  20560-0118  USA 


ABSTRACT 

Three  new  species  of  the  cancellariid  genus  Merica  are  de- 
scribed: Merica  lussii  from  South  Africa,  and  M.  deijnzeri  and 
M.  ektijphos  from  the  southern  Philippines.  The  South  African 
Merica  lussii  has  previously  been  misidentified  in  the  literature 
as  Cancellaria  bifasciata  Deshayes,  1830,  a  junior  subjective 
synon\Tn  of  C  oblonga  Sowerby,  1825,  from  which  it  differs  in 
having  a  broader,  stepped  spire,  a  proportionally  shorter  but 
broader  aperture,  and  a  deeply  impressed  sutural  canal.  Merica 
deijnzeri  differs  from  other  Philippine  species  of  Merica  by  its 
deeply  impressed  sutural  canal,  its  much  stronger  and  coarser 
axial  and  spiral  sculpture,  more  rounded  apert\ire,  and  its  lack 
of  banding  pattern.  Merica  ektt/phos  can  be  distinguished  from 
all  other  described  species  of  Merica  in  its  distinctively  heavy, 
inflated  shell,  as  well  as  by  its  extremely  coarse  axial  and  spiral 
sculpture. 

Additional  key  words:  Gastropoda,  Caenogastropoda,  Recent, 
western  Pacific. 


INTRODUCTION 

Merica  was  originally  proposed  as  a  subgenus  of  Can- 
cellaria  by  H.  and  A.  Adams  (1S54:  277)  to  distinguish 
those  species  with  reticulate  sculpture,  oblique  colu- 
mellar  folds  and  an  internally  lirate  outer  Up,  but  lacking 
a  prominent  emargination  or  notch  on  the  outer  lip  at 
the  base  of  the  siphonal  canal.  This  taxon  has  been  ac- 
corded generic  status  in  most  twentieth  century  works 
(e.g.,  Habe,  1961;  Petit,  1974;  Noda,  1980;  Verhecken, 
1986a,  1986b),  while  retained  as  a  subgenus  of  Cancel- 
laria in  several  faunistic  surveys  (e.g.,  Springsteen  and 
Leobrera,  1986;  Wilson,  1994;  Higo  ct  ai,  1999).  Merica 
has  been  reported  from  the  later  Tertiaiy  of  Europe 
(Cossmann,  1899;  Sacco,  1904;  Janssen,  1984)  and  the 
Indo-Pacific  region  (Shuto,  1969;  Noda,  1908;  Ladd, 
1982).  In  the  Recent  fauna,  Merica  appears  hmited  to 
the  Indo-West  Pacific,  ranging  from  South  Africa,  along 

'  Research  Associate;  mailing  address:  806  St.  Charles  Road, 
North  M>Ttle  Beach,  SC  29852-2846  USA. 


the  northern  Indian  Ocean,  through  Indonesia  to  Aus- 
traha  and  northward  to  Japan,  usually  at  subUttoriil 
depths.  As  pointed  out  by  Verhecken  (1986a:  38—40), 
there  is  considerable  confusion  in  the  literature  as  to  the 
correct  nomenclature  for  many  of  the  currently  recog- 
nized species  of  Merica. 

This  paper  describes  three  new  Recent  species  ot 
Merica,  one  from  Soudi  Africa  and  two  from  the  Phil- 
ippines. The  assignment  of  Cancellaria  oblonga  Sower- 
bv,  1825,  to  the  genus  Merica  is  reaffirmed,  and  the  spe- 
cies is  illustrated  for  comparative  purposes.  Holofypes  of 
the  new  species  are  housed  in  the  collections  of  the  Na- 
tal Museum  (NM),  Pietermaritzburg,  Repubhc  of  South 
Africa,  and  the  National  Museum  of  Natural  History 
(USNM) 
USA. 


Smithsonian    Institution,    Washington,    DC, 


SYSTEMATICS 

Family  Cancellariidae  Forbes  and  Hanley,  1851 
Subfamily  Cancellariinae  Forbes  and  Hanley,  1851 
Genus  Merica  H.  and  A.  Adams,  1854 

Merica  H.  and  A.  Adams,  1854:277.  Tvpe  species:  Cancellaria 
melanostoma  Sowerbv,  1849,  by  subsequent  designation  of 
Cossmann,  1899. 

Diagnosis:  Shell  with  conical  spire,  large  body  whorl, 
paucispiral  protoconch  weakly  to  strongly  deflected  from 
teleoconch  axis,  prominent  suture,  generally  weak  si- 
phonal  fasciole,  narrow  pseudo-umbihcus,  and  large, 
ovate  aperture.  Columella  short,  thick,  weakly  concave, 
with  2  sharply  keeled  columellar  folds  and  strong  si- 
phonal  fold.  Columellar  Up  with  pustules  and  secondary 
folds.  Parietal  callus  generally  present.  Emargination 
along  outer  lip  at  base  of  siphonal  canal  poorly  defined 
or  absent. 

Remarks:  Merica  appears  to  be  the  Indo-Pacific  stem 
group  for  the  genus  Cancellaria,  which  is  hmited  to  the 
eastern  Pacific  and  the  western  Adantic.  Merica  differs 
in  lacking  a  pronounced  emargination  along  the  outer 
lip  of  the  shell  as  well  as  a  heavy,  bifurcated  posterior 


R.  E.  Petit  and  M.  G.  Harasewvch,  2000 


Page  143 


columellar  fold,  both  characteristic  of  CanccUnria.  ('oss- 
inann  (1899:  13)  accorded  generic  status  to  Mcrica,  and 
most  authors  have  followed  this. 

Kuroda  and  Habe  (1971:  310,  202)  proposed  the  ge- 
nus Momocbora  based  primarily  on  the  presence  of  a 
strongly  dexiated  protoconch  in  its  tvpe  species,  Can- 
ccllaria  sinensis  Reeve,  1856.  Mouiorhora  has  been  con- 
sidered a  spionxTii  of  Mcrica  by  most  subsecjuent  au- 
thors (Garrard,  1975:  3;  Verhecken,  1986a:  44). 

Mcrica  oblonga  (Sowerbv,  1825) 
(Figures  1,  5,  6) 

Cancellaria  oblonga  Sowerby,  1825:  Appendix:  15;  1S32:  fig. 
19. 

Cancellaria  bifasciata  Deshayes,  1830:  181;  Lobbecke,  1885: 
30,  pi.  9,  figs.  1,2. 

Cancellaria  oblonga  Sowerby:  Kieiier,  1841:  6,  pi.  3,  fig.  3;  Ab- 
bott and  Dance,  1982:  225;  Springsteen  and  Leobrera, 
1986:  78,  pi.  18,  fig.  26. 

Cancellaria  (Merica)  oblonga  Kiener  [sic]:  Chenu,  1859:  277, 
fig.  1847. 

Not  Cancellaria  (Merica)  bifasciata  Deshayes:  Chenu,  1858: 
277,  fig.  1845  [?=M.  sinensis  (Reeve,  1856)]. 

Not  Cancellaria  bifasciata  Deshayes:  Bamard,  1959:  13-14; 
Kenslev,  1973:  194,  fig.  749  [  =  M.  hissii  new  species]. 

Merica  bifasciata  (Deshayes):  Habe,  1961:  pi.  24,  fig.  27. 

Cancellaria  (Merica)  bifasciata  Deshayes:  Oyama  and  Take- 
mura,  1963:  Cancellaria  plate,  fig.  6. 

Merica  oblonga  (Sowerbv):  Petit,  1974:  112,  fig.  5;  Verhecken, 
1986a:  41,  figs.  7-8;  Verhecken  and  Wranik,  1991:  60;  Ver- 
hecken, 1997;  308,  fig.  37. 

Momoebora  bifasciata  (Deshayes):  Higo.  1973:  179. 

Momocbora  oblonga  (Sowerby):  Higo  and  Goto,  1993:  276; 
Higoe/fl/.,  1999:  294. 

Remarks:  For  many  years  there  was  confusion  about 
priority  of  the  names  apphed  to  this  species,  with  Can- 
cellaria bifasciata  Deshayes,  1830,  generally  given  pre- 
cedence over  Cancellaria  oblonga  Sowerby,  1825.  These 
two  available  names  were  correctly  dated  by  Perit 
(1974).  Merica  oblonga  (as  Merica  bifasciata)  was  attri- 
buted to  the  genus  Momoebora  by  Higo  (1973:179),  and 
this  placement  remained  unchanged  in  the  two  later  re- 
visions and  enlargements  of  that  work  (Higo  and  Goto, 
1993;  Higo  et  al,  1999).  As  M.  oblonga  has  a  protoconch 
that  is  not  strongly  deflected  from  the  coihng  axis  of  the 
teleoconch  (Figs.  5,6),  we  see  bttle  justification  for  the 
usage  of  Momoebora. 

There  were,  and  to  a  lesser  degree  still  remain,  ques- 
tions about  the  geographic  range  of  this  species.  Its  oc- 
currence in  Panama,  as  reported  by  Kiener  (1841),  was 
shown  to  be  incorrect  by  Keen  (1971:649).  Verhecken 
(1997:309)  gave  the  distribution  of  Merica  oblonga  as 
"Japan  to  Indonesia;  Northern  Indian  Ocean  to  Aden; 
Eastern  South  Africa  (?)."  His  query  regarchng  the  South 
African  records  is  appropriate,  as  the  South  African 
specimens  previously  identified  as  Mcrica  bifasciata  rep- 
resent a  new  species,  described  herein  as  Merica  hissii. 

There  are  no  verifiable  Japanese  records  for  Mcrica 
oblonga  (Kazunori  Hasegawa,  personal  communication). 


The  figme  of  M.  oblonga  pubUshed  by  Habe  (1961)  and 
ri'produced  b\'  Petit  (1974)  is  of  a  specimen  from  Taiwan 
that  was  inchulcd  b\'  Habe  for  comparison  with  Merica 
asprella  [sic]  Lamarck.  The  specimen  figured  by  Oyama 
and  Takemura  (1963)  is  also  from  Taiwan.  Verhecken's 
(1986a:  41)  record  for  Japan  is  based  on  Habe's  usage. 
Records  of  M.  oblonga  are  noticeably  absent  in  most 
standard  Japanese  compendia  and  fists.  Its  inclusion  in 
the  exhaustive  faunal  fists  of  Higo  (1973),  Higo  and  Goto 
(1993)  and  Higo,  Callomon  and  Goto  (1999)  was  based 
on  a  Japanese  publication  of  uncertain  authorship. 

Mcrica  oblonga  ranges  as  far  north  as  Taiwan.  It  is 
uncommon  in  the  Phifippines.  Springsteen  and  Leobrera 
(1986:  78)  record  this  species  as  "sporadically  found 
throughout  the  Philippines  in  limited  quantity-."  The  col- 
or figure  in  Abbott  and  Dance  (1982:225)  is  of  a  Phil- 
ippine specimen  taken  from  over  30  m  depth  in  tangle 
nets  (Petit  collection).  Verhecken  and  Wranik  (1991:  60) 
state  that  M.  oblonga  'seems  to  have  its  distribution  cen- 
ter in  the  eastern  Inthan  Ocean  (Gulf  of  Bengal). '  While 
this  species  is  uncommon  in  other  areas,  it  appears  to 
be  common  in  India.  We  have  seen  commercial  lots 
from  India  containing  hundreds  of  specimens  of  M.  ob- 
longa. 

Merica  litssii  new  species 
(Figures  2,  7,  8) 

Description:  Shell  (figure  2)  to  35  mm,  thin,  biconic, 
elongate.  Spire  high  (spire  angle  53-55°)  comprising 
one-half  of  total  shell  length.  Protoconch  (figures  7,  8) 
paucispiral,  nearly  coaxial  with  teleoconch  (deviated 
from  coifing  axis  by  up  to  4°),  increasing  in  diameter 
from  300  p,m  to  2.15  mm  in  2%  elongated  whorls.  Tran- 
sition to  teleoconch  marked  by  onset  of  spiral  then  axial 
sculpture.  Teleoconch  of  about  5  evenly  rounded  whorls. 
Shoulder  rounded.  Suture  broadly  impressed,  forming 
conspicuous  canal.  Axial  sculpture  of  30-34  very  weak, 
rounded,  sfightly  prosocline  ribs,  as  broad  as  inter\'ening 
spaces.  Axial  ribs  become  prominent,  irregular,  crowded 
on  final  V4  of  body  whorl.  Spiral  sculpture  more  pro- 
nounced than  axial,  of  regularly  spaced,  flat,  primary 
cords  (8-10  on  penultimate  whorl,  2.5-32  on  body  whorl) 
with  weaker  secondaiy  cords  present  irregularly,  but 
most  often  on  posterior  portion  of  whorls.  Aperture 
large,  broadly  ovate,  deflected  from  coifing  axis  by  16- 
17°.  Outer  lip  thin,  weakly  flared  outwarcUv,  without  cfis- 
cemable  emargination.  Inner  fip  smooth  or  with  weak 
spiral  firae  reflecting  spiral  cords,  especially  in  the  an- 
terior portion.  Inner  lip  with  2  columellar,  1  siphonal 
fold.  Siphonal  fold  flat,  bifid  on  holot^pe  but  sharp  on 
other  specimens.  Columellar  lip  thick,  forming  induc- 
tural  shield  that  obscures  chink-fike  umbificus.  Siphonal 
canal  short,  broad,  inconspicuous.  Base  color  fight  yel- 
low-brown, irregularly  marked  with  white  and  various 
shades  or  redcfish  brown.  Two  bands  of  brown  some- 
times present,  visible  inside  aperture.  Wider  band  just 
below  shoulder,  narrower  band  just  posterior  to  base  of 


Page  144 


THE  NAUTILUS,  Vol.  114,  No.  4 


R.  E.  Petit  and  M.  G.  Harasewvch,  2000 


Page  145 


siphonal  canal.  Axial  flammules  of  browii,  \t>ll()\v,  white, 
irregular  in  outline  and  intensity 

Tjpe  locality:  Off  Park  Rviiie,  about  SO  km  S  ol  Dur- 
ban, on  the  KwaZulu-Natal  south  coast.  South  Africa, 
appro>dmatelv  30°17'  S,  3()°45'  E,  dredged  in  150  ni  on 
nibble  and  sponge  substrate. 

Type  material:  Holot\pe,  NM  ¥7648/11312,  height 
34.4  mm;  Paratvpe  1,  Marcus  Lussi  Collection,  both 
from  the  t\pe  locality-;  Paratope  2,  Dawni  Brink  Collec- 
tion, near  type  localit\'  in  140  m,  nibble  and  sponge  bot- 
tom; Paratvpe  3,  NM  E8794,  Zululand,  off  Matigulu 
River  mouth,  29°21.4'  S,  3r56.5'  E,  in  145  m,  mud, 
shell  rubble,  NMDP  16  Apr  1989,  station  ZR7. 

Etymology:  Named  for  Mr.  Markus  Lussi  of  Durban 
North,  SoutJi  Africa,  who  has  made  numerous  contri- 
butions to  the  Uterature  on  South  African  Mollusca,  and 
who  provided  specimens  for  study. 

Comparative  remarks:  Merica  lussii  differs  from  M. 
ohlonga  in  having  a  broader  shell  with  a  stepped  spire, 
a  proportioniilly  shorter  but  broader  aperture,  and  a 
deeply  impressed  sutural  canal.  Merica  lussii  lacks  the 
closely  packed  strong  axial  ribs  that  produce  the  finely 
beaded  surface  characteristic  of  M.  ohlonga.  Barnard 
(1959:  14)  and  Verhecken  (1986a:  42)  have  previously 
suggested  that  South  African  records  of  M.  bifasciata  (  = 
M.  oblonga)  might  represent  a  distinct  species. 

Merica  detjnzeri  new  species 
(Figures  3,  9,  10) 

Description:  Shell  (figure  3)  to  35  mm,  thick,  biconic, 
weakly  pseudo-umbilicate.  Spire  high  (spire  angle  55- 
58°).  Protoconch  (figures  9,  10)  paucispiral,  deviated 
from  coiling  a.xis  by  about  14°,  increasing  in  diameter 
from  300  jim  to  2.15  mm  in  2V4  evenly  rounded,  glassy 
whorls.  Transition  to  teleoconch  marked  by  onset  of  spi- 
ral cords  followed  almost  immediately  by  axial  ribs.  Te- 
leoconch of  up  to  6  whorls.  Shoulder  inconspicuous.  Su- 
ture deeply  impressed,  forming  narrow  canal.  Axial 
sculpture  of  27-33  low,  evenly  spaced,  prosochne  ribs, 
rounded  in  profile,  as  wide  as  intervening  spaces.  Peri- 
odic internal  varices  (see  Harasewych  and  Petit,  1982: 
111)  barely  discernable  on  outer  surface  of  shell  by 
thickened  axial  rib  slightly  expanded  along  edge  of  su- 
tural canal  followed  closely  by  thin  axial  rib  and  slightly 
reflected  fip,  begin  after  about  2%  whorls,  continue  at 
intervals  of  about  240°  thereafter.  Spiral  sculpture  of 
strong,  broad  cords  (20-26  on  body  whorl,  8-9  on  pen- 


ultiiriatc  whorl)  that  are  narrower,  more  closely  spaced 
near  suture  and  siphcjn.  Spiral  cords  overlav  axial  ribs  to 
produce  a  cancellate  surface  on  early  whorls.  Single,  sec- 
ondarv'  cords  occur  between  primary  cords  along  central 
region  of  whorls.  Axial  ribs  Iteconie  progressivi'ly  broad- 
er, irregular,  more  widely  spaced  on  last  half  of  body 
whorl,  obscuring  cancellate  appearance.  Aperture  large, 
broad,  nearl\'  hemi-elliptical,  deflecteil  from  coifing  axis 
by  14-15°.  Outer  lip  weakly  recurved  in  profile,  with 
finely  .serrated  edge,  lacking  emargination.  Inner  surface 
of  outer  Up  with  16  recessed  spiral  Urae  that  diminish  1/ 
3  whorl  into  the  aperture.  Inner  Up  with  2  columellar,  1 
siphonal  fold,  columellar  lip  nearly  covering  pseudo-um- 
bificus.  Posterior  columellar  fold  strongest,  overlying 
weak  siphonal  fasciole.  Anterior  columellar  fold  extends 
to  edge  of  columellar  fip.  Spiral  ridge  parallels  anterior 
columellar  ff)ld  adapically,  but  is  confined  to  the  colu- 
mellar hp  without  extending  into  aperture.  Surface  of 
siphonal  fold  with  single  pustule  in  some  specimens.  Si- 
phonal canal  short,  narrow,  axial.  Color  uniform  yellow- 
brown,  sometimes  with  narrow  darker  bands  above  and 
below  mid-whorl.  Portions  of  spire  whorls  intficate  that 
juveniles  may  be  mottled. 

Type  locality:  Off  Baficasag  Island,  Bohol,  Phifip- 
pines.  Taken  in  tangle  nets  at  approximately  200  meters. 

Type  material:  Holotvpe,  USNM  880645,  height  35.2 
mm;  Paratype  1,  R.  Petit  Collection  2465,  both  from  the 
type  locahty;  Paratypes  2-A,  USNM  242316,  off  Pt.  Du- 
murug,  Ma.sbate,  Philippines  (12°00'30"  N,  124°05'.36" 
E),  dredged  in  280  m,  green  mud,  USES  Albatross,  sta. 
5394,  l3'Mar  1909;  Paratvpes  5-8,  USNM  242321,  off 
Adyagan  Island,  E  Masbate,  PhiUppines  (12°12'35"  N, 
124°02'48"  E),  dredged  in  247  m,  green  mud,  sand. 
USES  Albatross,  sta.  5392,  13  Mar  1909;  Paratype  9, 
USNM  277485,  off  Sibugav  Island,  E  of  Masbate,  Phil- 
ippines (12°04'15"  N,  124°()4'36"  E)  in  198  m,  grav  sand, 
mud,  15.5°C.  USES  Albatross,  sta.  5212,  20  Apr'l908. 

Etymology:  We  take  pleasure  in  naming  this  species 
for  Mr.  Al  Deynzer  of  Sanibel,  Elorida,  who  obtained 
these  and  other  uncommon  Cancellariidae  in  the  Phil- 
ippines and  made  them  available  for  study. 

Comparative  remarks:  This  new  species  is  distinc- 
tive in  having  a  well-developed  sutural  margin  strongly 
rounded  to  form  a  deeply  impressed  sutural  canal.  Mer- 
ica deynzeri  is  similar  to,  but  stockier  and  broader  than 
M.  oblonga,  from  which  it  can  also  be  distinguished  by 
its  much  stronger  and  coarser  axial  and  spiral  sculpture, 
and  by  its  more  rounded  aperture.  A  notable  feature  of 


<- 

Figures  1—4.  Apertural,  lateral  and  dorsal  views  of  the  shells  of.  1.  Merica  oblonga  (Sowerby,  1825),  USNM  845168,  taken  by 
fishing  trawler  in  Manila  Bay,  Luzon,  Philippines,  1969  2.  Merica  lussii  new  species,  holotype,  NM  V7648Ari.312,  off  Park  Ryiiie, 
about  80  km  south  of  Durban  on  the  KwaZulu-Natal  south  coast.  South  Africa,  approximately  30°17'S,  .30°45'E,  dredged  in  150 
meters,  rubble  and  sponge  substrate.  3.  Merica  deijnzeri  new  species,  holotype,  USNM  880645.  off  Balicasag  Island,  Bohol,  Phil- 
ippines, taken  in  tangle  nets  from  about  200  meters.  4.  Merica  ekti/phos  new  species,  holotype,  USNM  880646,  Off  Balicasag 
Island,  Bohol,  Philippines,  taken  in  tangle  nets  from  about  200  meters.  Scale  bar  =  2  cm  for  all  figures. 


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THE  NAUTILUS,  Vol.  114,  No.  4 


Figures  5-12.  Scanning  Electron  Micrographs  of  a\ial  and  lateral  aspects  ot  pnitcicDnclis  nt  spcnini-ns  iii  figures  1—4.  5-6.  Merica 
oblonga  (Sowerby,  1825).  7-8.  Merica  lussii  new  species.  9-10.  Merica  detjnzeri  new  species.  11-12.  Merica  ekttjphos  new  species. 
Arrowheads  indicate  transition  from  protoconch  to  teleoconch.  Scale  bar  =  1  mm  for  all  figures. 


R.  E.  Petit  and  M.  G.  Harasewvch,  2000 


Page  147 


Merica  deynzeri  is  the  lack  of  color  bands  that  arc  char- 
acteristic of  most  species  of  Merica. 

Merica  ektyphos  new  species 
(Figures  4,  11,  12) 

Description:  Shell  (figure  4)  to  33  mm,  very  thick, 
globose,  pseudo-umbilicate.  Spire  relatively  low  (spire 
angle  63°).  Protoconch  (figures  11,  12)  paucispiral,  de- 
viated from  coiling  ;L\is  by  about  8°,  increasing  in  di- 
ameter from  400  |xm  to  2.10  nun  in  2y2  low,  evenlv 
rounded  whorls.  Transition  to  teleoconch  marked  by  on- 
set of  axial  ribs  followed  by  overlying  spiral  cords.  Te- 
leoconch of  up  to  6  rounded  whorls.  Suture  impressed, 
forming  deep,  narrow  sutural  canal.  A.\ial  sculpture  of 
prominent,  prosocline  ribs  (31  on  body  whorl,  31  on 
penultimate  whorl).  Ribs  become  smaller,  more  densely 
spaced  in  narrow  bands  spaced  every  Vi  to  %  whorl,  in- 
dicating position  of  internal  varices.  Spiral  sculpture  of 
strong,  flattened  cords  (15-16  on  bodv  whorl,  6  on  pen- 
ultimate whorl)  that  form  nodes  at  intersections  with  ax- 
ial ribs.  Nodes  especially  pronounced  at  margin  of  su- 
tural canal,  forming  serrated  ridge.  Finer  secondaiy 
threads  present  between  adjacent  cords  above  suture 
line.  Aperture  large,  deflected  from  coifing  axis  by  12°, 
with  broadly  elfiptical  outer  lip,  sharp,  48°  angle  between 
columella  and  parietal  wall.  Outer  fip  prosocline,  sfightly 
crenate,  with  12-13  strong,  sfightly  recessed  firae  that 
become  reduced,  disappearing  about  V4  whorl  within  the 
aperture.  Outer  lip  with  emargination  discernible  only 
by  touch  as  shallow  depression.  Columella  with  2  colu- 
mellar,  1  siphonal  fold.  Posteriormost  fold  strongest, 
overlving  conspicuous  siphonal  fasciole,  does  not  reach 
edge  of  columellar  lip  as  anterior  and  siphonal  folds  do. 
Columellar  fip  thick,  rounded,  nearly  covering  pseudo- 
umbificus.  Four  weak,  short  spiral  ridges  run  along  col- 
umellar fip  parallel  to  and  posterior  to  anterior  colu- 
mellar fold,  but  do  not  extend  into  aperture.  Siphonal 
canal  short,  narrow,  well  demarcated,  deflected  from 
axis.  Color  whitish,  with  ginger  brown  bands  along  su- 
ture (darkest),  above  mid-whorl  (broadest),  below  mid- 
whorl  (narrowest)  and  along  siphonal  fasciole.  Gaps  in 
coloration  of  sutural  band  correlate  with  internal  varices. 

Type  locality:  Off  Baficasag  Island,  Bohol,  Phifip- 
pines.  Taken  in  tangle  nets  at  appro.ximately  200  meters. 

Type  material:  Holotype,  USNM  880646,  32.3  mm; 
Paratypes  1-4,  R.  Petit  Collection  2472;  Paratypes  5-7, 
Deynizer  Collection,  all  from  tyjie  locafity. 

Etymology:  Gr  ektyphos.  puffed  up.  Named  for  its 
inflated  shefl. 

Comparative  remarks:  This  new  species  can  be  dis- 
tinguished from  all  known  species  of  Merica  by  the 
thickness  of  its  shell  and  the  strength  of  its  sculpture.  In 
outfine  it  is  closest  to  the  Australian  M.  wcstralis  (Gar- 
rard, 1975),  from  which  it  differs  in  having  much  coarser 
axial  and  spiral  sculpture,  a  straighter  columella  with 


Tabic  1.  Merica  deynzeri  new  species.  Measurements  of  shell 
characters.  Linear  measiirpnients  in  mm.  n  =  5.  °  n  =  3  for 
no.  whorls,  protoconch. 


(;!iaractcr 

Mean 

SU 

Range 

Shell  length  (SL) 

.35.0 

2.6 

.34.9-38.6 

Aperture  length  (AL) 

22.1 

1.7 

19.6-24.3 

AIVSL 

0.629 

0.003 

0.624-0.632 

No.  whorls,  protoconch" 

1.5 

0 

2.5-2.5 

No.  whorls,  tcleoconcii 

5.53 

0.14 

5..33-5.67 

.Spire  angle 

56.7° 

1..3° 

5.5-58° 

stronger  folds,  and  a  body  whorl  that  is  niuch  less  con- 
stricted beliind  the  siphonal  fasciole. 

ACKNOWLEDGMENTS 

We  thank  Marcus  Lussi  and  Dawn  Brink  of  South  Africa 
for  making  specimens  of  Merica  lussii  available  for  this 
study.  While  on  visits  to  the  Phifippines  Al  Deynzer 
made  special  efforts  to  obtain  specimens  for  this  study. 
We  are  grateful  to  Paul  Callomon  of  Osaka  and  Dr.  Ka- 
zunori  Hasegawa  of  Ibaraki  for  information  about  the 
Japanese  records  of  M.  ohlonga. 

LITERATURE  CITED 

Abbott,  R.  T.  and  S.  P.  Dance,  1982.  Compendium  of  Seashells. 
E.  P  Dntton.  New  York,  x  -I-  411  pp. 

Adams,  H.  and  A.  Adams.  18.5.3-.58.  The  genera  of  Recent  Mol- 
lusca;  arranged  according  to  their  organization.  John  van 
Voorst,  London.  2  vols.  [1:1-256,  pis.  1-.32,  1853;  1:257- 
484,  2:1-92,  pis.  .33-72,  1854;  2:9.3-284,  pis.  7;3-96,  1855; 
2:285-412,  pis.  97-112,  18.56;  2:41.3-540,  pis.  113-128, 
1857;  2:.541-660.  pis.  129-1.38,  18,58.] 

Barnard,  K.  H.  1959.  Contributions  to  the  faiowledge  ot  South 
African  marine  Mollusca.  Part  II,  Gastropoda:  Prosobran- 
chiata:  Rachiglossa.  Annals  of  the  South  African  Museum 
45:1-2.37. 

Chenu,  J.  C.  1859-62.  Manuel  de  conch\liologie  et  de  paleon- 
tologie  conchyliologique.  Librarie  Victor  Masson,  Paris.  2 
vols.  [l:i-vii,  1-508,  18.59;  2:1-327,  1862]. 

Cossmann,  M.  1899.  Essais  de  Paleoconchologie  Comparee. 
Troisieme  hvraison.  Paris.  201  p.,  8  pis. 

Deshayes,  G.  P.  1830.  Encyclopedic  Methodique.  Histoire  na- 
turelle  des  vers  2(l):l-256,  Paris. 

Forbes,  E.  and  S.  Hanley  1848-1853.  A  history  of  British  Mob 
lusca  and  their  shells.  London.  4  volumes.  [Published  in 
parts;  for  dates  of  parts  see  Fischer  and  Tonilin,  19.35, 
Journal  of  Conchology  20(5):  150-151.] 

Garrard,  T.  A.  1975.  A  revision  of  the  Australian  Cancellariidae 
(Gastropoda:  Mollusca).  Records  of  the  Australian  Muse- 
um .30:1-62. 

Habe,  T.  1961.  Description  of  four  new  cancellariid  species, 
with  a  list  of  the  Japanese  species  of  the  familv  Cancel- 
lariidae. Venus  21:431-441.  pis.  2:3-24, 

Harasewvch,  M.  G.  and  R,  E,  Petit.  1982.  Notes  on  the  mor- 
phology of  Cancellaria  reticulata  (Gastropoda:  Cancellar- 
iidae). The  Nautilus  96:104-113. 

Higo,  S.  1973.  A  catalogue  of  molluscan  faima  of  the  Japanese 


Page  148 


THE  NAUTILUS,  Vol.  114,  No.  4 


Islands  and  the  adjacent  area.  S.  Higo.  Isahaya,  Japan.  (58) 
+  397  +  61  pp. 

Higo,  S.  and  Y.  Goto.  1993.  A  systematic  list  of  molluscan  shells 
from  the  Japanese  Is.  and  the  adjacent  area.  Elle  Corpo- 
ration, Osaka,  Japan,3  +  22  +  693  +  13  +  149  pp. 

Higo,  S.,  Callomon,  P.  and  Y.  Goto.  1999.  Catalogue  and  bib- 
liography of  the  marine  shell-bearing  Mollusca  of  Japan. 
Elle  Scientific  Publications,  Osaka,  49  pp. 

Janssen,  A.  W.  1984.  An  account  of  the  Cancellariidae  (Gastro- 
poda) of  Winterswijk-Mi.ste  (Miocene,  Henimoorian),The 
Netherlands.  Scripta  Geologica  68;  1-39,  pis.  1-6. 

Keen,  A.  M.  1971.  Sea  shells  of  tropical  West  America.  Second 
edition.  Stanford  University  Press,  Stanford,  xiv  -t-  1064 
pp.,  22  pis. 

KensJey,  B.  1973.  Sea-Shells  of  southern  Africa,  Gastropods. 
Maskew  Miller  Ltd.,  Cape  Town,  236  pp. 

Kiener,  L.  C.  1841.  Genre  Cancellaire.  Species  general  et  icon- 
ographic  des  coquiUes  vivantes.  Paris,  44p.,  9  pis. 

Kuroda,  T.  and  T.  Habe.  1971.  [Descriptions  of  species]  In: 
Kuroda,  Habe  and  Oyama,  The  sea  shells  of  Sagami  Bay. 
Manizen  Co.,  Tokyo.  Pp.  i-xix,  1-741  (Japanese),  121  pis., 
1-489  (English),  1-51  (index). 

Ladd,  H.  S.  1982.  Cenozoic  fossil  mollusks  from  Western  Pa- 
cific Islands;  gastropods  (Eulimidae  and  Volutidae  through 
Terebridae).  United  States  Geological  Suwey  Professional 
Paper  1171:1-100,  pis.  1-41. 

Lobbecke,  T.  1881-1887a.  Das  genus  Cancellaiia.  Systema- 
tisches  Conchylien-cabinet  von  Martini  und  Chemnitz  4: 
1-96,  pis.  1-23  [issued  in  parts;  pt.  .309;  1-16,  pis.  1-5, 
1881;  pt.  335;17^32,  pis.  6-10,  1885;  pt.  .340;.3;3-56,  pis. 
11-15,  1886;  pt.  343;5780,  pis.  16-20,  1886;  pt.  .346;81- 
96,  pis.  21-23,  1887.] 

Matsukiuua,  A.  1998.  Index  and  collation  of  The  Molluscan 
Shells  by  Katura  Oyama  and  Yoshio  Takemura  (1957- 
1963).  The  Yuriyagai  6;91-122. 

Noda,  H.  1980.  Molluscan  fossils  from  the  Ryukyu  Islands, 
southwestern  Japan.  Part  1.  Gastropoda  and  Pelecypoda 
from  the  Shinzato  Fomiation  in  southeastern  part  of  Old- 
nawa-jinia.  Science  Reports  of  the  Institute  of  Geoscience, 
University  of  Tsuk-uba,  SecHon  B,  l;l-95,  pis.  1-12. 

Oyama,  K.  and  Y.  Takemura.  1963.  The  Molluscan  Shells.  Pt. 


6,  Cancellaria  plate,  [for  collation  of  all  parts  and  plates 
see  Matsuk-imia.  1998]. 

Petit,  R.  E.  1974.  Notes  on  Japanese  Cancellariidae.  Venus  33: 
109-115. 

Reeve,  L.  A.  1856.  Monograph  of  the  genus  Cancellaria  Con- 
chologia  Iconica  10;  [unpaginated  text],  pis.  1-18. 

Sacco,  F.  1904.  I  molluschi  dei  terreni  terziarii  del  Piemonte 
e  deila  Liguria.  Pt.  .30  (Aggiunte  e  Correzione).  C.  Clau- 
sen, Torino,  xxxvi  +  203  pp.,  31  pis. 

Shuto,  T.  1969.  Neogene  gastropods  from  Panay  Island,  the 
Philippines  (Contributions  to  the  Geology  and  Paleontol- 
ogy of  Southeast  Asia,  LXVIII).  Memoirs  of  the  Faculty 
of  Science,  Kyushu  University,  Series  D,  Geology  19(1): 
1-250,  pis.  1-24. 

Sowerby,  G.  B.,  I.  1825.  A  catalogue  of  the  shells  contained  in 
the  collection  of  the  late  Earl  of  Tankerville,  arranged  ac- 
cording to  the  Lamarcldan  conchological  system;  together 
with  an  Appendix,  containing  descriptions  of  many  new 
species.  London,  vii  -1-92-1-  xxxi\-  pp.  [Appendix],  9  pis. 

Sowerby,  G.  B.,  II.  1849.  Descriptions  of  some  new  species  of 
Cancellaria  m  the  collection  of  Mr  H.  Cuming,  Proceed- 
ings of  the  Zoological  Society  of  London  for  1848:1.36- 
138. 

Springsteen,  F  J,  and  F  M.  Leobrera.  1986,  Shells  of  the  Phil- 
ippines. Carfel  Seashell  Museum,  Manila,  377  pp. 

Verhecken,  A.  1986a.  The  Recent  Cancellariidae  of  Indonesia 
(Neogastropoda,  Cancellariacea).  Gloria  Maris  25(2):29- 
66. 

Verhecken,  A.  1986b.  A  revision  of  the  Cancellariidae  (Neo- 
gastropoda: Cancellariacea)  of  the  Red  Sea  and  the  Gulf 
of  Aden.  Gloria  Maris  25(4):  13.3-153. 

Verhecken,  A.  1997.  Mollusca  Gastropoda:  Arafura  Sea  Can- 
cellariidae collected  during  the  KARUBAR  Cmise;  7m.  A. 
Crosnier  and  P.  Bouchet  (eds),  Resultats  des  Campagnes 
MUSORSTOM,  Vol.  16.  Memoirs  du  Museum  d'Histoire 
Naturelle  172:29.5^23. 

Verhecken,  A,  and  W.  Wranik.  1991.  Additional  data  on  the 
Cancellariidae  of  the  Gulf  of  Aden.  Gloria  Maris  .30(4); 
59-63. 

Wilson,  B.  R.  1994.  AustraUan  Marine  Shells.  Prosobranch 
Gastropods,  Part  two  (Neogastropods).  Odyssey  Publish- 
ing, Kallaroo,  370  pp. 


I 


THE  NAUTILUS  114(4):149-154,  2000 


Page  149 


Systematics  of  the  genus  Infundihulum  Montfort,  1810 
(Gastropoda:  Trochidae) 


Bruce  A.  Marshall 

Museum  of  New  Zealand  Te  Papa 

Tongarewa 
P.O.  Box  467,  Wellington 
NEW  ZEALAND 
brucem@tepapa.govt.nz 


Two  Recent  Infundibtiluin  species  are  recognized:  /. 
concaviiin  (Gnielin,  1791)  (type  species)  and  /.  tonilini 
(Fulton,  1930).  Their  shells  are  convergent  on  those  of 
calyptraeid  limpets  of  the  genus  Trochita  Schumacher, 
1817,  having  the  most  widely  excavated  bases  and  the 
most  strongly  tangential  apertures  of  any  living  Trochini. 
They  Hve  firmly  attached  to  rocks  on  highly  e.xposed 
shores  from  low  tide  level  to  several  meters  depth  in 
tropical  seas.  Reliable  records  of  /.  concavum  are  from 
Reunion  and  Mauritius.  All  Recent  specimens  of  /.  tom- 
lini  seen  were  from  old  collections,  and  most  have  either 
minimal,  often  dubious  locality  data,  or  none  at  all, 
though  it  would  seem  that  /.  tomlini  has  a  patchy  chs- 
tribution  in  the  tropical  southwest  Pacific. 

Additional  key  words:  Mollusca,  Trochoidea,  fossils. 
Pleistocene,  Kermadec  Islands. 


INTRODUCTION 

The  present  contribution  was  initiated  some  years  ago 
when  I  recognized  a  species  oi  Infundihulum  among 
Pleistocene  fossils  from  Raoul  Island,  Kermadec  Islands, 
northeast  of  New  Zealand.  A  search  of  the  literature  re- 
vealed that  Infundihulum  species  are  sui"prisingly  poorly 
known.  Moreover,  inquiries  revealed  that  most  of  the 
few  specimens  of  Infundihulum  species  in  museum  col- 
lections were  acquired  during  the  nineteenth  century, 
and  have  broad,  often  dubious  locality  data,  or  none  at 
all.  The  fossils  from  Raoul  Island  prove  to  be  the  little 
known  species  /.  tomlini  (Fulton,  1930),  which  is  rep- 
resented in  old  collections  labeled  as  originating  vari- 
ously from  Penang  (Pinang,  Malaysia),  Australia,  Upolu 
(Western  Samoa),  Lifou  (Loyalty  Islands)  or  the  New 
Hebrides  (Vanuatu).  Of  these  locality  records,  only  Upo- 
lu, Lifou,  and  Vanuatu  seem  hkely  to  be  correct.  Mu- 
seum specimens  of  the  type  species  (Trochus  concavus 
Gmehn,  1791)  are  labeled  as  having  originated  from  lo- 
calities in  the  Indian  Ocean,  as  well  as  the  Red  Sea  and 
Cochin  China  (southern  Vietnam),  though  the  only  re- 
hably  locaUzed  specimens  seen  during  the  present  study 


were  from  Reunion.  An  explanation  for  the  poor  rep- 
resentation of  /.  concavum  (and  presumably  /.  tomlini) 
in  collections  is  suggested  by  the  fact  that  specimens 
collected  ahve  were  cryptic  at  and  below  low  tide,  at- 
tached to  exposed,  wave-swept  volcanic  rocks,  which 
tend  to  receive  far  less  attention  from  collectors  than 
more  congenial  and  species-rich  coral  reef  and  lagoon 
environments,  where  they  have  never  been  recorded.  In- 
stitutional acronyms:  ANSP,  Academy  of  Natural  Scienc- 
es, Philadelphia;  AMS,  Australian  Museum,  Sydney; 
RMNH,  The  Natural  Histoiy  Museum,  London;  LACM, 
Los  Angeles  County  Museum  of  Natural  History; 
MNHN,  Museum  National  d'Histoire  Naturelle,  Paris; 
NMNZ,  Museum  of  New  Zealand,  Welfington;  NMP, 
Natal  Museum,  Pietermaritzburg;  USNM,  National  Mu- 
seum of  Natural  History,  Washington,  DC. 

SYSTEMATICS 

Superfamily  Trochoidea  Rafinesque,  1815 
Family  Trochidae  Rafinesque,  1815 
Subfamily  Trochinae  Rafinesque,  1815 
Tribe  Trochini  Rafinesque,  1815 
Genus  Infundihulum  Montfort,  1810 

Infundibuluiii  Montfort,  ISIO:  167.  Type  species  (by  original 
designation):  Infundihulum  tijpus  Montfort,  1810  =  Tro- 
cluis  concavus  Gmelin,  1791;  Recent,  Rennion. 

Carinidea  Swainson,  1S40:  .350.  T^pe  species  (by  nionotypy): 
Trochus  concavus  Gnielin,  1791. 

Description:  Shell  up  to  54  mm  in  diameter,  stout, 
base  excavate,  umbilicus  narrow;  aperture  very  strongly 
tangential,  as  wide  as  shell;  periphery  sharply  angulate, 
not  stellate  at  any  stage  of  growth,  almost  flush  against 
a  flat  surface.  Spire  sculptured  with  strong,  close,  round- 
ed spiral  cords  and  a.xial  costae,  intersections  finely  and 
bluntly  nodular  Base  weakly  convex,  evenly  curving  into 
narrow  umbilicus,  excavated  from  periphery.  ColumeUar 
lip  adapicallv  with  strong,  rounded  denticle,  sharply  re- 
tracted to  insertion  within  umbilicus.  Foot  (/.  concavum) 
very  large,  epipothum  very  well  developed,  edge  finely 
fringed,  well  developed  epipodial  flaps  beside  epipodial 


Page  150 


THE  NAUTILUS.  Vol.  114,  No.  4 


insertion;  cephalic  lappets  large,  papillate.  Radula  (/. 
concavum,  figure  9)  with  the  formula  ii  +  .5+l+5  +  n, 
very  similar  to  that  in  Trochiis  (scnsu  stricto). 

Remarks:  Infundibiilum  species  are  referable  to  sub- 
family Trochinae,  tribe  Trochini  because  of  similaritv'  to 
species  oiTrochtis  Linnaeus,  175S,  and  Tectiis  Montfort, 
1810,  in  shell  and  radular  morphology,  and  external  anat- 
omy. External  anatomy  is  close  to  that  of  Tcctiis  fenes- 
trattis  (Gniehn,  1791)  (Hickman  and  McLean,  1990:  fig. 
55b).  lufiindibtihtm  species  differ  irom  Trochiis  and  Tcc- 
tus  in  the  combination  of  low,  evenly  conical  spire,  weak- 
ly convex  whorls,  low  sculpture,  lack  of  peripheral  nod- 
ules at  any  stage  of  growth,  and  widely  excavated  base, 
which  fits  almost  flush  with  a  flat  surface.  All  of  these 
character  states  (apomoq^hies)  evidentlv  enhance  attach- 
ment to  rocks  exposed  to  strong  wave  action.  Infundib- 
iilum concavum  (Gmehn,  1791)  and  /.  tomlini  (Fulton, 
19.30)  have  the  most  strongly  tangential  apertures  and 
the  most  widely  excavated  bases  of  any  li\ing  Trochini, 
and  they  resemble  limpets  in  ventral  view  when  the  foot 
is  fully  extended  (shdes  of  aquarium  specimens  of  7.  con- 
coviim  provided  by  D.  G.  Herbert).  Gross  shell  mor- 
phology strikingly  parallels  calyptraeid  fimpets  of  the  ge- 
nus Trochita  Schumacher,  1817. 

Whereas  Infiindibulum  has  been  interpreted  as  a  sub- 
genus of  Trochus  by  many  authors,  I  prefer  to  treat  In- 
fiindibulum as  a  distinct  genus  because  of  current  lack 
of  clear  exddence  for  degrees  of  relatedness. 

Trochus  hochii  Phihppi,  1844,  from  Oman  (figures  8, 
10)  has  a  similar  shape  and  sculpture  to  Infundibiihtm 
species,  but  differs  in  having  the  base  considerably  less 
excavated,  and  a  much  more  weaklv  tangential  aperture 
(and  thus  presumably  a  more  weakly  expanding  foot). 
The  relationships  of  this  species  are  unclear,  and  I  refer 
it  to  Trochus,  subgenus  Infundibidops  Pilsbry,  1889  (type 
species  Trochus  cri/thracus  Brocchi,  1821)  with  some 
hesitation. 

Inftindibiihim  concavum  (Gmelin,  1791) 
(Figures  1-.3,  9) 

Trochus  ptjramidalis  rarissimus  .  .  .  Chemnitz,   1781:  86,  pi. 

168,  figs.  1620,  1621  (not  binominal). 
Trochus  concavus  Gmelin,   1791:  .3.570  (refers  to  Chemnitz, 

1781.  figs.   1620,   1621);  Dillw>ii.  1817:  763;  Blaimille. 

1825:  425.  pi.  .32  bis.  fig.  1;  Wood.  1825:134.  pi.  28.  fig. 

7;  Sganzin.  1843:  22;  Philippi.  1848:  .38.  pi.  8.  figs.  7.  8; 

Deshayes.   1863:   71;  Fischer.   1876:   105.  pi.  33,  fig.  2; 

Kaicher.  1979:  card  2087. 
Infundibulum  tijpus  Montfort,  1810:  167,  text  fig. 


Carinidea  concavus:  Swainson,  1840:  .350. 

Pohjdonta  (Carinidea)  concavus:  Chenu.  1859:  357.  fig.  2646 

Trochus  (Pohjdonta)  concavus:  Martens.  1880:  295. 

Trochus  (Infundibulum)  concavus:  Pilsbry,  1889:  40,  pi.  43,  fig. 

13;  VVenz,  1938:  311,  fig.  693. 
Infundibulum  concavum:  Cossmann.  1918:  188.  text  fig.  65. 
Tectus  concavus:  Dri\'as  and  Jay.  1988.  pi.  2,  fig.  3. 
Not  Poh/donta  (Infundibulum)  concava:  MeMlI  and  Standen, 

1895:  125  =  /.  tomUni. 

Description:  Shell  (figures  1-3)  up  to  56  mm  in  di- 
ameter, stout,  broadly  conical,  wider  than  high,  weakly 
cyrtoconoid,  base  excavate;  aperture  strongly  tangential, 
as  wide  as  shell;  peripherv  sharplv  angulate,  almost  flush 
against  a  flat  sin4ace,  narrow  umbihcus.  Spire  dull  green 
with  subsutur;il  band  of  irregular,  duU,  red  and  white 
maculations,  last  few  whorls  pinkish  milky  green.  Base 
white,  inner  half  with  porcellanous  glaze.  Protoconch 
and  first  few  teleoconch  whorls  unknown  (eroded).  Spire 
whorls  moderately  convex  at  first,  becoming  weakly  con- 
vex, sculptured  with  low,  rounded  spiral  cords  and  op- 
isthocline,  non-coUabral  axial  costae,  interspaces  narrow- 
er; intersections  with  strong,  bluntlv  rounded  nodules; 
axial  costae  on  last  2  or  .3  whorls  traversing  rounded  axial 
folds  that  are  broader  than  the  costae  and  tend  to  be 
more  strongly  opisthochne.  Spiral  cords  7,  strong  and 
similar  on  early  whorls,  weakening  after  shell  attains 
about  8  mm  diameter,  obsolete  on  last  2  or  3  adult 
whorls.  Excavate  base  convex,  smoothly  cuning  into  nar- 
row umbihcus.  Outer  part  of  base  with  6  or  7  smooth, 
simflar  spiral  cords,  becoming  obsolete  on  last  whorl  at 
maturit\';  inner  (porceUanous)  part  of  base  with  6  or  7 
spiral  cords  that  become  obscured  bv  indiictura  at  ma- 
turity. Aperture  ovate.  Basal  Mp  strongly  tluckened, 
smoothly  continuous  with  columellar  lip,  which  is 
stronglv  thickened  adapicallv  then  sharplv  retracted  to 
insertion  within  umbihcus. 

Animal  (retracted,  ex  ethanol).  Foot  extremely  large 
and  spreachng.  Epipodium  very  weU  developed,  edge 
finely  fringed  throughout;  inner  surface  finely  papillate, 
tapered  macropapillae  over  posterior  half  neck  lobes 
convoluted,  finely  fringed,  left  lobe  considerably  larger 
and  more  deeply  convoluted  than  right.  Epipodial  ten- 
tacles slender,  narrowly  tapering,  4  pairs.  Epipodial  flaps 
well-developed,  bases  close  to  epipodial  insertion,  edges 
finely  fringed,  5  on  right,  5  or  6  on  left.  Cephahc  ten- 
tacles slender,  narrowly  tapered,  subcyhndrical.  right 
tentacle  base  adjacent  to  tip  of  left  tentacle  due  to 
obhque  asymmetry  of  head.  Eye  stalks  large,  dorsoven- 
trally  flattened,  well  developed  eyes  in  tips.  Cephahc  lap- 
pets well  developed,  edge  and  adjacent  ventral  surface 


Figures  1-10.  Infundibulum  and  Trochus  (Infiindibulops)  species.  1-3.  /.  concavum  (Gmelin.  1791).  St.  Paul  Bav.  Reunion,  .56 
X  40  mm  (NMNZ  M. 270521).  4-6.  I  tomlini  (Fulton.  19.30).  s\Titype,  locality  unhiown  (possibly  Western  Samoa  or  Vanuatu). 
19.0  X  26.6  mm  (BMNH  19.30.4.2.3).  7.  7  tomlini.  Boat  Cove.  Raoul  Island.  Kemiadec  Islands  (Pleistocene).  25.5  X  .33  (est.)  mm 
(NMNZ  M.214580);  8,  10.  Trochus  (Infundibidops)  kochii  (Philippi.  1844).  SW  coast  of  As  Sawda.  Juzor  Al  Halaaniyaat.  Oman, 
32  X  ,38  mm  (Zoological  Museum,  Amsterdam).  9.  Radula  of  7  concavum.  north  of  Cap  la  Houssaye,  St.  Paul  Bav.  Reunion  (NMP 
K4898). 


B.  A.  Marshall,  2000 


Page  151 


Page  152 


THE  NAUTILUS,  Vol.  114,  No.  4 


papillate.  Cephalic  tentacles  blackish  brown,  sides  of 
foot  darker,  elsewhere  cream. 

Radula  (figure  9)  with  the  formula  n+5+1+5+n, 
very  similar  to  that  in  Trochtis  {scnsit  stricto)  (Hickman 
and  McLean,  1990:  fig.  57D). 

Type  data:  Holotype,  the  specimen  illustrated  by 
Chemnitz  (1781:  figs.  1620,  1621),  ex  Spengler  collec- 
tion, in  Zoological  Museum,  Copenhagen  (T.  Schi0tte, 
pers.  comm.);  "k-uste  von  Coromandel". 

Material  examined:  North  of  Cap  la  Houssaye,  St. 
Paul  Ba\-,  Reunion,  exposed  rocky  shore,  extreme  low 
tide  to  2  m,  24  September  1988,  R.  N.  Kilbum  and  D. 
G,  Herbert  (3,  NMP);  St  Paul  Bay,  Reunion,  breaker 
zone  on  exposed  rock-\'  shore  at  low  tide,  M.  Jay,  1994 
(.3  NMNZ);  St  Paul,  Reunion,  M.  Jav,  1985  (3  ANSP); 
"Zanzibar"  (8  in  3  lots,  MNHN):  Seychelles  (4,  MNHN); 
"Seychelles  [and]  Madagascar",  H.  Cuming  collection  (2, 
BMNH);  "Madagascar(?)",  M.  Larrey,  1874  (1,  MNHN); 
"Madagascar",  M.  Ballot  (1,  MNHN);  "Madagascar"  (1, 
MNHN);  "Indian  Ocean",  H.  Fischer  (1,  MNHN);  "In- 
dian Ocean",  Staadt  colln  (2  MNHN);  "Ceylon",  ex- 
Gould  coUn  (1  USNM);  "Cochin  China",  M.  Harmand, 
1876  (2,  MNHN);  "Penang  (?)",  ex  Marie  and  Sowerby 
collections  (7,  BMNH);  "Penang"  (2,  LACM);  "Austra- 
lia". Steams  colln  (1,  USNM). 

Distribution:  Reunion  and  (based  on  Sganzin,  1843) 
Mauritius.  Living  on  faces  of  lava  rock  and  boulders  ex- 
posed to  tlie  open  sea  from  low  tide  to  2  m  depth  (M. 
Jay,  pers.  comm.).  Records  from  Madagascar  and  the 
Seychelles  require  confirmation.  Specimens  in  old  col- 
lections reputedly  from  Zanzibar,  Coromandel  Coast, 
Ceylon  (Sri  Lanka)  are  of  uncertain  provenance,  whereas 
Penang,  Cochin  China  (southern  Vietnam)  and  Austrafia 
are  extremely  doubtful. 

Remarks:  The  shell  oi  Infundibuhim  concavum  is  dis- 
tinctive among  Trochoidea  in  the  combination  of  broadly 
conical,  weakly  cyrtoconoid  spire;  flat,  inwardly  sloping 
base,  conical  umbilicus,  and  very  strongly  prosocline  out- 
er Up.  Among  trochid  animals  examined  or  that  have 
been  recorded  in  the  literature,  the  external  anatomy  of 
this  species  seems  most  similar  to  that  of  Tcctiis  fcncs- 
tratiis  (Gmehn,  1791)  (Hickman  and  McLean,  1990:  fig. 
55A). 

Infundibulum  tomlini  (Fulton,  1930) 
(Figures  4-7) 

Pobjdonta  {Infundibidum)  concava:  MeMl  and  Standen,  1895: 

125  (not  Gmelin,  1791). 
Trochtts  (Infundibulum)  tomlini  Fulton,  19.30:  16,  pi.  .3,  fig.  5. 
Trochus  tomlini:  Kaicher,  1979:  card  2162. 
Trochus  mnculatus:  Marshall,  1979:  535;  Marshall,  1981:  90 

(not  Linnaeus,  1758). 
Trochus  (Infundibulum)  sp.:  Brook,  1998:  256. 

Description:  Shell  (figures  4-7)  up  to  36.0  mm  wide, 
stout,  broadly  conical,  wider  than  high,  weakly  cyrtocon- 
oid, base  excavate;  aperture  strongly  tangential,  as  wide 


as  shell,  narrow  umbilicus;  periphery  sharply  angulate, 
not  stellate,  almost  flush  against  a  flat  surface.  Proto- 
conch  white.  First  3.5  teleoconch  whorls  CTeen  with  red 
spiral  bands;  2  or  3  continuous  median  bands  and  1  su- 
prasutural  band  on  1st  2  whorls;  spiral  cords  spotted  and 
streaked  with  red  on  3rd  whorl.  Subsequent  spire  whorls 
deep  duU  greenish  or  reddish  grey  with  subsutural  band 
of  white  spots  or  streaks.  Base  considerably  paler  than 
spire,  inner  third  (approximately)  porcelain-white.  Pro- 
toconch  200  fjLm  wide,  tip  narrowly  tapered,  sculpture 
unknown  (eroded).  Teleoconch  of  up  to  7.5  whorls.  Ear- 
ly spire  whorls  moderately  convex,  later  whorls  moder- 
ately or  weakly  convex;  sculptured  with  rounded  spiral 
cords  and  rounded,  prosocUne,  non-coUabral  axial  cos- 
tae,  interspaces  narrower;  intersections  with  strong, 
bluntly  rounded  nodules;  fine,  crowded  coUabral  growth 
lines  throughout.  Spiral  cords  numbering  7  per  spire 
whorl;  adapical  spiral  broadest  in  most  specimens,  al- 
most fusing  with  adjacent  spiral  in  some  specimens;  pe- 
ripheral and  adjacent  spiral  narrowest,  similar;  other  spi- 
rals similar,  peripheral  spiral  partly  covered  by  succeed- 
ing whorls.  Excavated  base  broad,  weakly  convex,  evenly 
curving  into  narrow  steep-sided  umbilicus,  base  of  which 
is  fiUed  with  callus.  Basal  spirals  much  finer  than  spire 
spirals;  outer  (pigmented)  part  of  base  of  most  speci- 
mens with  7  or  8  weakly  nodular  spirals,  typically  1  sec- 
ondary spiral  in  each  interspace,  some  interspaces  in 
some  specimens  with  2  secondary  spirals;  6-8  smooth 
spirals  on  inner  (white)  part  of  base,  smooth,  similar. 
Aperture  ovate-trapezoidal.  Columellar  and  basal  fips 
smoothly  continuous,  thickened  adapicaUy,  sharply  re- 
tracted to  insertion  within  umbilicus.  Rounded  spiral 
ridge  at  base  of  outer  lip. 
Animal  unk-nown. 

Type  data:  Originally  figured  syntype  BMNH 
1930.4.2.3  (height  19.0  mm,  diameter  26.6  mm);  syntype 
National  Museum  of  Wales  Cardiff  (Trew,  1984).  Fulton 
(1930)  stated  that  he  had  seen  five  specimens  and  did 
not  segregate  a  holotype.  The  specimens  could  not  be 
locahzed  and  evidently  originated  from  somewhere  in 
the  tropical  southwest  Pacific. 

Other  material  examined:  Titi  Knob  (Trig  Sentinel), 
Boat  Cove,  Raoul  Island,  Kermadec  Islands,  cemented 
tuffaceous  conglomerate  (Pleistocene),  W.  R.  B.  Oliver 
(1,  NMNZ);  DavTell  Islet.  Raoul  Island,  hard  sandy  tuff 
(Pleistocene),  F.'j.  Brook,  June  1991  (2,  Auckland  Insti- 
tute and  Museum);  Upolu,  Western  Samoa,  R.  W  Tate 
(7,  NMNZ);  Lifou,  Loyalty-  Islands,  J.  Brazier,  1873  (4, 
BMNH);  Vanuatu,  C.  Bertie,  1895  (1,  AMS);  Erroman- 
go,  Vanuatu,  H.  A.  Robertson,  1898  (3  in  2  lots,  AMS); 
"Red  Sea",  E.  C.  Freeman  collection  (3,  BMNH);  "Red 
Sea"  (4,  LACM);  "Penang"  (1,  NMNZ);  "Penang,  ex 
Sowerby"  (1,  NMP). 

Distribution:  Raoul  Island,  Kermadec  Islands  (Pleis- 
tocene): Upolu,  Western  Samoa  (record  plausible  be- 
cause R.W.  Tate  coUected  extensively  there);  Aneiteum 
and    Erromango,   Vanuatu    (records   plausible  because 


B.  A.  Marshall.  2000 


Page  153 


there  are  extensive  stretches  of  exposed  rockv'  shore), 
and  possibly  Lifou.  Specimens  in  old  collections  reput- 
edl\  from  the  Red  Sea  and  Penang  are  of  extremely 
doubtful  provenance. 

Remarks:  Compared  with  Infundihiiliim  concamiin.  I. 
tonilini  differs  in  attaining  smaller  size  (nia.\imum  di- 
ameter 36.0  mm  instead  of  56.0  nnn).  in  being  more 
broadly  conical,  in  ha\'ing  more  strongly  convex  whorls, 
and  in  having  much  stronger  axial  costae  and  nodules  on 
the  spire,  especially  after  the  shell  is  about  6  mm  wide. 
In  /.  concaium  the  spiral  cords  and  nodules  progres- 
sivelyweaken  and  become  obsolete  when  the  shell  is  12- 
15  mm  wide,  while  in  /,  touiliui  the  spiral  cords  pro- 
gressively enlarge  throughout. 

One  lot  (NMP)  labelled  "Penang,  ex  Sowerby",  sug- 
gests origin  from  the  London  shell  dealers  Messrs.  Sow- 
erby and  Fulton,  who  advertised  Trochus  concavus  in 
their  1908  price  list.  Other  "Penang"  specimens  of  I. 
coucavtiin  and  /.  totulini  in  old  collections  mav  well  have 
originated  from  this  source. 

The  Pleistocene  specimens  from  Raoul  Island  (figure 
7)  are  indistinguishable  from  the  syntype  (figures  4-6) 
and  other  Recent  specimens.  /.  tomlini  is  absent  from 
the  Recent  fauna  of  the  Kermadec  Islands  (Brook  and 
Marshall,  1998). 

ACKNOWLEDGMENTS 

For  loan  or  gift  of  specimens  I  thank  Philippe  Bouchet 
(Museum  National  d'Histoire  Naturelle,  Paris),  Dai  Her- 
bert (Natal  Museum,  Pietermaritzburg),  Maurice  Jay  (La 
Reunion),  Alan  Kabat  (National  Museum  of  Natural  His- 
tory, Washington  D.C.),  Ian  Loch  (Australian  Museum, 
Sydney),  Jim  McLean  (Los  Angeles  County  Museum  of 
Natural  History),  Robert  Moolenbeek  (Zoological  Mu- 
seum, Amsterdam),  David  Reid  and  Kathie  Way  (The 
Natural  Historv^  Museum,  London),  and  Gary  Rosenberg 
(Academy  of  Natural  Sciences  of  Philadelphia).  Thanks 
to  Dai  Herbert  and  Jim  McLean  for  comments  on  the 
manuscript,  Norman  Heke  (Museum  of  New  Zealand) 
for  the  photography,  and  to  Robert  Thompson  (Victoria 
Universitv',  Wellington)  for  access  to  the  scanning  elec- 
tron microscope. 

LITERATURE  CITED 

Blainville,  H.  M.  D.  de  1825-1827.  Manuel  de  malacologie  et 
de  conchy liologie.  .  ,  LevTaiilt,  Strasbourg,  664  pp. 

Brook,  F.  J.  199S.  Stratigraphy  and  Paleontology  of  Pleistocene 
submarine  volcanic-sedimentarv  sequences  at  the  north- 
em  Kermadec  Islands.  Journal  of  the  Royal  Society  of 
New  Zealand  2S;235-257. 

Brook,  F.  J.  and  B.  A.  Marshall.  1998.  Checklist  of  benthic 
coastal  marine  Mollusca  from  the  northern  Kermadec  Is- 
lands, pp. 210-233  (Appendix)  In:  Brook,  F.  J.  The  coastal 
molluscan  fauna  of  northern  Keniiadec  Islands,  southwest 
Pacific  Ocean.  Journal  of  the  Royal  Society  of  New  Zea- 
land 28:185-2.33. 

Chemnitz,  J.  H.  1781.  Neues  Systematisches  Conchylien  Cab- 
inet 5.  Raspe,  Nuniberg,  324  pp. 


Chenu,  J.  C.  1859.  Manuel  de  conchvliologie  et  de  paleonto- 
logie  conchyliologique  1.  Masson,  Paris,  508  pp. 

Cossmann,  M.  1918.  Essais  de  paleoconchologie  comparee  11. 
Cossmann,  Paris,  388  pp. 

Deshayes,  G.  P.  1863.  Catalogue  des  Mollusques  de  I'ile  de  la 
Reunion  (Bourbon)  7m.  Maillard,  L.  (ed.)  Notes  sur  I'ile 
de  la  Reunion.  Annexe  E,  Dentu,  Paris,  pp. 1-144. 

Dillwyn,  L.  W.  1817.  A  descriptive  catalogue  of  Recent  shells, 
arranged  according  to  the  Linnaean  method  2.  Arch,  Lon- 
don, pp.  581-1092. 

Drivas.  J.  and  M.  Jay,  1988.  Coquillages  de  la  Reunion  et  de 
I'ile  Maurice.  Delachaux  et  Niestle,  Paris,  159  pp. 

Fi.scher,  P.  1875-1880.  Species  general  et  Iconographie  des  co- 
quilles  vivantes,  comprenant  la  collection  du  Museum 
d'Histoire  naturelle  de  Paris,  la  collection  Lamarck,  celle 
du  Prince  Massena.    .11.  Bailliere,  Paris,  480  pp. 

Fulton,  H.  C.  1930.  Descriptions  of  new  species  of  Fiisinus. 
Biplex,  Trochus.  and  Biisliia.  Proceedings  of  the  Malaco- 
logical  Society  of  London  19:16-17. 

Gmelin,  J.  F.  1791.  Caroli  a  Linne  .  .  .  Systema  naturae  per 
regna  tria  naturae,  secundum  classes  .  .  .  Editio  decima 
tertia,  aucta,  reformata.  1  (6).  Vermes  testacea.  Beer,  Lip- 
siae,  pp.  .'3021-4120. 

Kaicher,  S.  D.  1979.  Card  catalogue  of  world-wide  shells.  Pack 
21.  Trochidae  1.  Kaicher,  St.  Petersburg,  Florida,  cards 
2072-2177. 

Keen,  A.  M.  and  L.  R.  Cox.  1960.  7/i.  Moore,  R.  C.  (ed)  Trea- 
tise on  invertebrate  paleontology.  Part  1.  Mollusca  1.  Geo- 
logical Society  of  America  and  University  of  Kansa.s, 
Lawrence,  pp.  246-275. 

Marshall,  B.  A.  1979.  The  Trochidae  and  Turbinidae  of  the 
Kermadec  Ridge.  New  Zealand  Journal  of  Zoology  6:521- 
552. 

Marshall,  B.  A.  1981.  Boat  Cove  Formation:  macrofossils  hi: 
Lloyd,  E.  F.  and  S.  Nathan  (eds.)  Geology  and  Tephro- 
chronology  of  Raoul  Island,  Kemiadec  Group,  New  Zea- 
land. New  Zealand  Geological  Sur\'ey  Bulletin  95,  pp.  90- 
91. 

Martens,  E.  von  1880.  Die  Mollusken  der  Maskarenen  und 
Seychellen  auf  Gniud  der  von  Professor  Karl  Mohius  ges- 
ammelten  Mollusken  In:  Mohius,  K.  (ed.)  Beitriige  zur 
Meeresfauna  der  Insel  Mauritius  und  der  Seychellen. 
Gutmann'schen,  Berlin,  pp.  179-.343. 

Melvill,  J.  C.  and  R.  Standen  1895.  Notes  on  a  collection  of 
shells  from  Lifu  and  Uvea,  Loyalt)'  Islands,  formed  by  the 
Rev.  James  and  Mrs.  Hadfield,  with  list  of  species.  Journal 
of  Conchology  8:84-1.32. 

Montfbrt,  D.  de  1810.  Conchyliologie  systematique,  et  classifica- 
tion mediodique  des  coquilles.  .  .  2.  Schoell,  Paris,  676  pp. 

Oliver,  W.  R.  B.  1915.  The  Mollusca  of  the  Kermadec  Islands. 
Transactions  of  the  New  Zealand  Institute  47:509-568. 

Philippi,  R.  A.  1846-1855.  Systematisches  Conchylien-Cabinet 
von  Martini  und  Chemnitz.  Neu  herausgegeben  von  ver- 
voUstaeudigt  \on  H.  C.  Kuester  (iiach  dessen  Tode  fort- 
gesetzt  von  W.  Kobelt)  2  (3).  Bauer  and  Raspe,  Nurem- 
berg, 372  pp. 

Pilsbry,  H.  A.  1889-1890.  Manual  of  Conchology;  structural 
and  systematic.  With  illustrations  of  the  species.  Volume 
11.  Academy  of  Natural  Sciences,  Philadelphia,  519  pp. 

Sganzin,  V.  1843.  Catalogue  des  cotjuilles  troiuees  aux  ties  de 
France,  de  Bourbon  et  de  Madagascar  Memoires  de  la 
Societe  d'Histoire  Naturelle  de  Strasbourg  3  (2):l-.'30. 

Swainson,  W.   1840.  A  treatise  on  malacology,  or  shells  and 


Page  154  THE  NAUTILUS,  Vol.  114,  No.  4 

shellfish.  Longman,  Omie,  Brown.  Green,  Longmans  and  Prosobranchia.  In:  Schindewolf,  O.  H.  (ed.)  Handbuch 

Taylor,  London,  419  pp.  der  Palaozoologie  6.  Bonitraeger,  Berlin,  pp.  24-^80. 

Thiele,  J.  1929.  Handbuch  der  Systematischen  Weichtierkunde  Wood,  W.  1825.  Index  testaceologicus;  or  a  catalogue  of  shells, 

1(1).  Fischer,  Jena,  pp.  1-.376.  British  and  foreign,  arranged  according  to  the  Linnean 

Trew,  A.  1984.  The  Melvill-Tomlin  Collection.  Part  30.  Tro-  system;  with  the  Latin  and  Enghsh  names,  references  to 

chacea.  Handlists  of  the  moUuscan  collections  in  the  De-  authors,  and  places  where  found.  Illustrated  with  2300 

partment  of  Zoology,  National  Museum  of  Wales,  94  pp.  figures.  Taylor,  London,  188  pp. 

Wenz,  W  1938.  Gastropoda.  Teil  1  (2):  Allgemeiner  Tell  und 


THE  NAUTILUS  114(4):155-160,  2000 


Page  155 


On  the  tcLxononiic  placement  of  Uiiio  ochraceus  Say,  1817  in  the 
genus  Ligumia  (Bivalvia:  Unionidae) 


Douglas  G.  Smith 

Department  of  Bi()log^  and 
Graduate  Program  in  Organisniic  and 

Exolutionan'  BioloijN 
Uni\ersitv  of  Massachusetts 
Amherst,' MA  01003-5810  USA 
d<rsmith@bio. imiass.edu 


ABSTRACT 

Since  1975,  the  freshwater  unionid  mussel  orig;inally  described 
as  Vnio  ochraceus  Sav,  1S17,  has  been  \ariouslv  assigned  to  the 
genera  LampsiUs  or  Leptodca.  The  taxonomic  stabilit\'  of  tliis 
species  has  been  comphcated  h\  incomplete  biowledge  of  cer- 
tain features  of  its  anatomy.  Study  of  the  mantle  of  this  taxon 
leads  to  the  conclusion  that  V.  ochraceus  does  not  belong  to 
either  Lampsilis  or  Leptodea.  The  species  is  assigned  to  the 
genus  Ligumia  according  to  established  taxonomic  criteria  to 
differentiate  lampsiline  genera  that  include  characters  of  the 
anatomy  and  lar\a.  The  mantle  edge  contains  distinct,  regular 
papillae  along  an  luiinternipted,  lamellate  border  and  the  glo- 
chidia  are  of  the  large  t\pe. 

Additional  kiij  words:  LampsiUs,  Leptodea.  Unionoidea,  fresh- 
water nuissels,  new  combination. 


INTRODUCTION 

The  unionid  mu.ssel  originally  described  as  Unio  ochra- 
ceus Sav,  1817,  is  discontinuouslv  distributed  east  of  the 
Appalachian  divide  in  drainages  veiy  near  the  Atlantic 
coast  of  North  America.  The  northernmost  populations 
are  found  in  Nova  Scotia  (Atheam  and  Clarke,  1962)  and 
the  southernmost  in  Georgia  (Johnson,  1947). 

The  generic  position  of  this  species  has  remained  an 
open  question  since  Morrison  (1975)  and  Bereza  and 
Fuller  (1975)  challenged  the  placement  of  U.  ochraceus 
in  Lampsilis.  The  present  paper  attempts  to  resoK'e  the 
issue  in  light  of  widely  accepted  anatomical  definitions 
of  the  various  genera  comprising  the  Lanipsihnae  (sen.su 
Heard  and  Guckert,  1971)  or  Lampsihni  (sensu  Davis 
and  Fuller,  19S1).  It  is  proposed  that  the  species  be 
placed  in  the  genus  Ligumia  and  is  treated  as  such  in 
the  remaining  portion  of  the  paper. 

HISTORY  OF  THE  PROBLEM 

The  question  of  generic  allocation  of  this  species  was 
raised  by  Morrison  (1975)  and  Bereza  and  Fuller  (1975). 
Morrison's  (1975)  discussion  was  the  most  comprehen- 


sive of  the  two;  he  argued  that  the  taxon  Mijtilus  fluvia- 
tihs  Gmelin,  1791,  took  precedence  over  Unio  ochraceus 
and  that  M.  fluviatilis,  for  which  he  elected  a  neotype, 
belonged  in  Leptodea  because  of  its  "smaller  glochidia, 
and  a  complete  lack  of  'mantle  flaps'  or  papillae  on  the 
mantle  margins  of  the  females  ".  Although  his  conclusion 
regarding  the  generic  placement  of  U.  ochracea  has 
been  widely  adopted,  his  argument  for  M.  fluviatilis  has 
not  gained  acceptance.  One  major  issue  regarding  the 
position  of  L.  ochracea  concerns  the  lack  of  the  "mantle 
flap "  (Morrison,  1975),  a  character  included  in  the  pre- 
vailing definition  of  Lampsilis  (sensu  stricto),  the  genus 
to  which  the  species  has  been  assigned  bv  manv  authors 
(e.g..  Ortmann,  1919;  Reardon,  1929;  Johnson,  1947, 
1970;  Burch,  1973).  Ortmann  (1912)  reassessed  the 
characters  defining  lampsiline  genera.  This  author  used 
primarily  anatomical  and  lanal  characters  in  the  diag- 
nosis of  the  various  genera,  including:  ( 1 )  features  of  the 
mantle  margin  anterior  to  the  inhalant  aperture,  (2)  the 
size  and  shape  of  the  glochidia,  (3)  the  nature  of  the 
marsupial  portion  of  the  female  gill  (four  "types"),  and 
(4)  the  degree  of  attachment  of  the  inner  gill  to  the 
abdomen.  Ortmann  (1912)  defined  the  mantle  flap,  a 
feature  of  Lampsilis,  as  a  highly  pigmented  free  lobe 
extending  from  the  mantle  margin  anterior  to  the  inhal- 
ant aperture  with  a  'lacerated  appearance. "  Papillae  are 
not  present  in  this  region  of  die  mantle.  The  mantle  flap 
is  best  represented  in  the  female,  reduced  in  the  male. 
Ortmann  (1912)  was  apparentlv  unable  to  examine  an- 
atomical material  of  Ligumia  ochracea  and  consequentlv 
omitted  the  species  in  his  discussion.  Later,  Ortmann 
(1919),  still  unfamiliar  with  the  anatomv,  placed  the  spe- 
cies in  Lampsilis,  presumably  on  the  basis  of  shefl  char- 
acters. Subsequent  study  (Bereza  and  Fuller,  1975;  Mor- 
rison, 1975;  Kat,  19S3;  Smith,  1995)  has  shown  that  Lig- 
umia ochracea  clearly  does  not  possess  a  mantle  flap. 
Nevertheless,  L.  ochracea  continued  to  be  included  in 
Lampsilis,  at  least  proxdsionallv,  by  several  in\'estigators 
(Fuller,  1977;  Johnson,  19S0;  Clarke,  1981a;  Kat,  1983; 
Porter,  1985;  Strayer,  1987).  Certainly  part  of  the  reason 


Page  156 


THE  NAUTILUS,  Vol.  114,  No.  4 


for  maintaining  L.  ochracca  in  Lampsilis  has  been  the 
strong  resemblance  of  its  shell  to  other  .species  oi  Lamp- 
silis sensii  stiicto  and  conservatism  in  the  absence  of  a 
definitive  description  of  the  species. 

Otherwise,  following  the  suggestion  of  Morrison 
(1975),  the  species  has  been  placed  within  Lcpfodea  (D. 
Stan.sberrv',  in  Porter,  1985;  Smith,  1991;  Stiven  and  Al- 
derman, 1992;  Strayer  and  Jirka,  1997;  Turgeon  ct  al., 
1998).  Ortmann  (1912)  stated  that  the  mande  margin  of 
Paraptcra  (=  Lepiodea),  based  on  the  type,  L.  fragilis, 
was  lamellate  with  at  most  marginal  crenulations,  but 
without  papillae  or  a  flap.  There  were  little  or  no  ob- 
served differences  behveen  the  sexes.  The  glochidia 
were  described  as  "very  small"  and  "suboval." 

Descriptions  of  the  mantle  o{  Ligumia  ochracca  from 
North  Carohna  and  Nova  Scotia  (Porter,  1985,  and  Kat, 
1983,  respectively)  have  clearly  shown,  in  conjunction 
with  characters  discussed  below,  that  L.  ochracca  does 
not  belong  within  Leptodea  either.  Porter  (1985)  noticed 
in  his  studv  of  this  species  the  occurrence  of  some  pa- 
pillae along  the  margin  of  the  mantle  of  females  and 
males.  Whereas  Lea  (1863),  Reardon  (1929),  Bereza  and 
Fuller  (1975)  and  Morrison  (1975)  did  not  report  these 
papillae,  Kat  (1983)  mentioned  observing  a  few  papillae. 

At  various  jimctures,  and  with  apparent  awareness  of 
the  difficulties  regarding  the  placement  of  Ligumia 
ochracca,  both  Fuller  (1977)  and  Clarke  (in  Porter, 
1985)  indicated  that  a  new  genus  name  was  forthcoming. 
To  date,  however,  no  such  name  has  been  pubhshed. 


Figure  1.  Mantle  region  of  an  adult  female  Ligumia  ochra- 
cca posterior  to  the  foot.  Arrows  denote  papillae  along  inner 
margin.  F  =  foot,  lA  =  inhalant  aperture,  M  =  mantle.  Scale 
line  =  15  mm. 


MATERIALS  AND  METHODS 

Materia]  cited  in  the  discussion  of  Ligumia  ochracca  is 
cataloged  in  the  Museum  of  Zoology,  In\ertebrate  Di- 
vision, University  of  Massachusetts  at  Amherst.  The  cat- 
alog numbers  are  as  follows: 

Leptodea  fragilis  (Rafinesque,  1820)  UMA  MO.  1270,  1.381, 
1.38.3,  Vermont; 

Lioionia  nasuta  (Say.  1817)  UMA  MO.  769,  8.32,  1129,  1810, 
1849,  1904,  Connecticut,  Massachusetts,  New  York; 

Lioimia  ochracea  (Say,  1817)  UMA  MO.  1053,  1234,  1241, 
1809,  1822,  1823,  1850  Connecticut,  Maine,  Massachu- 
setts, and  North  Carolina; 

Ligumia  recta  (Lamarck,  1819)  UMA  MO.  1137.  1379.  1902. 
\'ermont.  New  York; 

Ligumia  subrostrata  (Say  1831)  UMA  MO.  1903,  Illinois. 

Specific  locahHes  wiU  be  furnished  upon  request. 

All  specimens  used  in  this  studv  were  prepared  fol- 
lowing the  method  described  by  Smith  (1996).  For  his- 
tological studies,  a  portion  of  the  mantle  edge  measuring 
appro.ximatelv  5  mm-  was  excised,  cleared,  and  infiltrat- 
ed with  paraplast®.  Embedded  specimens  v\'ere  sec- 
tioned at  5  (xm  inten'als  and  stained  with  hematoxvhn 
and  eosin.  Bright  field  micrography  was  accomphshed 
with  a  Wild  M20  microscope.  Glochidia  were  either 
whole  mounted  on  slides  or  mounted  on  Cambridge 
SEM  stubs  and  sputter-coated  with  gold.  SEM  microg- 
raphy used  a  JEOL  JSM-5200  scanning  electron  micro- 
scope. 


RESULTS 

Close  examination  of  the  mantle  margin  of  carefully  re- 
la.\ed  preserved  male  and  female  specimens  of  L.  ochra- 
cca revealed  a  row  of  small  but  regular  papillae  extend- 
ing along  the  mantle  margin  from  the  base  of  the  in- 
halant aperture  anteriorlv  to  the  gape  of  the  foot.  The 
papillae  are  regularlv  arranged  (figure  I),  rather  closelv 
set,  and  average  about  100  (jim  in  length  (figures  2,  3). 
They  are  less  developed  than  those  of  Ligumia  nasuta, 
L.  subrostrata,  or  L.  recta,  which  mav  reach  >1  mm  in 
length.  The  mantle  edge  bearing  these  papillae  is  slightly 
lamellate  and  there  is  no  indication  of  a  flap  or  lacera- 
tions, as  is  also  the  case  with  L.  nasuta,  L.  subrostrata, 
and  L.  recta.  The  papillae  of  L.  ochracea  appear  at  a 
young  age,  about  two  years  as  evidenced  by  prominent 
grov\4h  marks  on  shells  of  20-25  mm  in  length.  At  this 
age  (shell  size),  the  papillae  are  not  as  closeK-  set  as  in 
the  adult  and  there  is  no  evidence  of  crenulation,  a  flap, 
or  a  "lacerated"  structure  (sensu  Ortmann,  1912). 

The  mantle  edge  of  L.  ochracca  is  pigmented  with  a 
cream,  orange-ochre,  or  hght-grev  background  and  var- 
iegated with  darker  grev.  Although  most  of  the  darker 
pigment  is  distributed  near  the  posterior  portion  of  the 
mantle  edge  (figure  1),  no  specific  pattern  is  e\ident.  In 
this  regard,  L.  ochracea  is  intermediate  between  L.  na- 
suta, which  has  verv  dark  pigment  throughout  and  a  dis- 
cernible pattern,  and  L    recta  and  L.  subrostrata.  in 


D.  G.  Smith,  2000 


Page  157 


3f^' 


Figure  2.  Section  of  mantle  edge  of  female  Lii^iimia  ocJira- 
cca  (sliell  lengtli  =  55  mm)  from  the  Connecticut  Ri\er,  Con- 
necticut. Scale  line  =  50  ni. 

Figure  3.  Same  as  figure  2,  different  specimen  (shell  length 
=  72  mm).  Scale  hne  =  50  ni. 

Figure  4.  Glochidia  of  Liotania  ochracea  (left)  and  Leptoclea 
Jrajiilis  (right).  Scale  line  =  200  m. 


which  the  mantle  edge  is  a  siightK"  variegated,  pattern- 
less,  light-  to  dark-gray.  The  mantle  margins  o(  Ligiimia 
subrosirata  and  L.  recta  differ  in  that  the  inner  edge  of 
the  mantle  of  L.  suhrostrata  has  a  very  dark-brown  bor- 
der, a  feature  absent  in  L.  recta.  Both  species  possess  a 
densely  pigmented  "eye  spot"  at  the  base  of  the  inhahint 
aperture.  In  L.  recta,  the  spot  is  small  and  not  swollen; 
in  L.  suhrostrata,  however,  the  spot  is  actually  a  shghtly 
thickened  pad,  almost  lobe-like  in  a  few  specimens.  The 


function  of  these  spots  is  as  yet  unclear,  although  it  is 
supposed  that  thev  act  as  some  sort  of  light  rece]itor.  No 
such  spots  are  appari'Ht  externalK  in  either  L  iiasulti  or 
L.  ochracea. 

E.\aniined  specimens  ol  Leptodea  fragiUs  comply  with 
Ortmann's  (1912)  (hagnosis  for  the  most  part.  In  one 
female,  a  few  small  papillae  extend  slightly  anteriorly 
from  the  inhalant  apertural  papillae  but  subsequently 
disappear  completely.  In  the  remaining  specimens,  no 
extra-apertural  papillae  are  evident.  The  crenulation  re- 
ported by  Ortmann  (1912)  is  actually  no  more  than  a 
shght  sinuation  of  the  margin,  the  edge  being  uninter- 
rupted whatsoever  for  the  length  of  the  mantle.  The 
mantle  margin  is  slightly  more  lamellate  in  the  female 
than  in  the  male,  but  there  is  httle  difference  between 
the  sexes.  Although  the  specimens  have  been  in  alcohol 
for  several  years,  there  is  no  indication  of  concentration 
of  pigment  in  the  posterior  region  <jf  the  mantle,  and  no 
specific  pattern  was  observed  during  the  original  prep- 
aration of  the  specimens  (D.  G.  Smith,  unpublished  ob- 
servations). 

The  glocliidia  of  L.  ochracea  are  over  twice  the  size 
of  those  of  Leptodea  fragiUs  (figiu-e  4)  (see  also  Porter, 
1985).  The  glochidia  of  L.  fragilis  from  Vermont  mea- 
sure on  the  average  7.3  |xm  (length)  by  94  fim  (height) 
and  possess  a  narrow  hinge.  This  compares  well  with 
observations  on  glochicha  of  this  species  by  Surber 
(1912).  Glochidia  of  L.  ochracea  from  Massachusetts 
have  a  mean  length  ot  222  fxm  and  a  mean  height  of 
287  (xm  (see  Porter  and  Horn,  1980,  for  similar  values 
of  North  Carohna  populations).  These  values  are:  shghtly 
smaller  than  those  for  glochidia  from  L.  nasuta  from 
Massachusetts  and  New  York,  which  have  a  combined 
mean  length  of  256  ^xm  and  mean  height  of  .318  |xm; 
shghtlv  larger  than  L.  suhrostrata  v\'ith  a  mean  length  of 
205  fjLm  and  mean  height  of  261  [xm;  and  nearest  to 
values  reported  for  L.  recta,  with  mean  values  of  220 
|xm  by  270  (xm  (Ortmann,  1912;  Baker,  1928).  The  ratio 
of  glochidia  length:height  among  the  species  is  rather 
consistent,  0.77  for  Ligumia  ochracea,  0.78  for  L.  recta 
and  L.  suhrostrata.  and  0.80  for  L.  na.suta.  The  ratio  of 
glochidial  length:height  in  Leptodea  fragihs  is  similar, 
0.78,  however,  as  noted  above,  the  glochidia  of  this  spe- 
cies are  less  than  one  third  the  size  of  Ligumia  ochracea. 
The  glochidia  of  L.  ochracea  (figure  5)  are  most  similar 
to  those  of  L.  suhrostrata.  which  also  has  a  narrow  hinge: 
those  of  L.  nasuta  (figure  7)  possess  a  wider  hinge.  The 
surfaces  of  the  glochidial  shell  of  each  species  are  pro- 
vided with  minute  pores  less  than  5  |xni  in  diameter 
(figures  6,  8)  and  the  margin  opposite  the  hinge  is  evenly 
cur\'ed  and  smooth. 

DISCUSSION 

As  concluded  earlier  by  Bereza  and  Fuller  (1975),  Lig- 
umia ochracea  can  not  be  assigned  to  either  Lampsilis 
or  Leptodea  as  these  genera  are  currenth'  defined.  In 
the  case  of  Lampsilis,  this  is  further  supported  by  studies 
using  biochemical  techniques  (Kat,  1983;  Stiven  and  Al- 


Page  158 


THE  NAUTILUS,  Vol.  114,  No.  4 


Figure  5.  Glochidiuni  of  Li ffania  ochracea  Scale  line  =  100  [jliii.  Figure  6.  Enlargement  of  glochidial  shell  surface  ofLigumia 
ochracea.  Scale  line  =  10  [j.m  Figure  7.  Glochidiuin  of  Ligiiinin  nasuta.  Scale  line  =  100  (jini.  Figure  8.  Enlargement  of 
glochidial  shell  surface  ot  Ligumia  nasuta.  Scale  hne  =  10  [jim. 


derman,  1992).  which  lia\'e  .showii  that  Liaumia  ochra- 
cea is  quite  distantly  related  to  two  s)inpatric  species  of 
Lampsilis  with  mantle  flaps.  The  only  recognized  lamp- 
sihne  genus  to  which  the  species  ochracea  can  be  allo- 
cated is  Ligtitnia.  In  re\aew,  Ortmann  (1912),  defined  a 
large  genus,  Eiinjnia,  which  was  distinguished  from  oth- 
er lampsihne  genera  principally  by  the  presence  ol  pa- 
pillae, variously  developed,  along  the  mantle  margin  and 
exddent  in  both  the  male  and  female.  Glochidia  of  Eii- 
njnia species  were  hsted  as  "subovate,  oi  mediimi  size, 
or  rather  large."  Other  characters,  including  the  nature 
of  the  marsupial  gill  (Ortmann  "type  4")  and  degree  of 
attachment  of  the  inner  gill  were  essentially  as  in  Lamp- 
silis. The  three  subgenera  oi  Eiin/nia  listed  by  Ortmann 
(1912)  [Micronuja,  Canincitlina,  Eiinjnia]  were  even- 
tually raised  to  genus  rank  and  their  nomenclature  sta- 
bihzed  (Ortmann  and  Walker,  1922).  The  nominotypical 
genus  sen.su  Ortmann  (1912)  is  Ligumia  Swainson,  1840 
[type-species:  Unio  recta  Lamarck,  1819]  and  is  defined 
(Ortmann,  1912,  as  subgenus  Etirynia)  by  having  "quite 
regular,  uniform,  smaller  or  larger  papillae,  reaching 
about  the  middle  of  the  lower  margin."  Ligumia  ochra- 


cea fits  this  description  and,  though  less  dex'eloped,  its 
papillae  are  similar  in  their  external  gross  morjiholog)-  to 
those  of  L.  nasuta,  less  so  to  L.  recta,  in  which  the  pa- 
pillae are  somewhat  crowded  and  slightly  irregular. 

At  present,  erection  of  a  new  genus  for  this  species 
seems  unwarranted,  in  the  absence  of  urgently  needed 
critical  anatomical  studies  of  the  various  nominal  lamp- 
sine  genera. 


ACKNOWLEDGMENTS 


1  thank  Richard  I.  Johnson  whom,  though  not  in  com- 
plete agreement  with  some  of  what  has  iieen  presented, 
provided  useful  comments  on  an  earher  draft  of  this  ar- 
ticle. David  Strayer,  Catherine  Corey,  Kevin  Cummings, 
Ethan  Nedeau,  and  Sarah  Riseman  donated  many  spec- 
imens used  in  this  studv.  The  study  was  funded  in  part 
by  a  grant  from  The  Nature  Conservancy,  Connecticut 
Chapter. 


D.  G.  Smith,  2000 


Page  159 


Figure  9.     Mantle  margin  of  female  Ligumia  nasuta.  Arrow 

denotes  papilla.  Scale  line  =  5  mm. 

Figure   10.      Mantle  margin  of  female  Ligtnuiti  stihrostrnta. 

Arrov\'  denote.s  papilla.  Scale  line  =  5  mm. 

Figure  11.     Mantle  margin  of  female  Ligumia  recta.  Arrow 

denotes  papilla.  Scale  line  =  5  mm. 


LITERATURE  CITED 

Atheani.  H.  D.  and  A.  H.  Clarke,  Jr  1962.  The  freshwater 
mussels  of  No\'a  Scotia.  Bulletin  National  Museum  Can- 
ada 183:11-41. 

Baker  F.  C.  1928.  The  freshwater  Mollnsca  of  Wisconsin.  Part 
II.  Pelec\poda.  Bulletin  \Vi.sconsin  Geological  and  Natural 
History  Survey  70:1—482. 

Bereza,  D.  J.  and  S.  L.  H.  Fuller  1975.  Notes  on  "Lainpsilis" 
ochracea  (Say)  (Mollusca:  Bi\alvia).  Association  of  South- 
easteni  Biologists  Bulletin  (Abstract)  22:42. 

Burch,  J.  B.  1973.  Freshwater  Unionacean  Clams  (Mollusca: 
Pelecypoda)  of  North  America.  Biota  of  Freshwater  Eco- 
systems. Identification  Manual  11,  United  States  Environ- 
mental Protection  Agency,  Washington,  D.C.,  176  pp. 

Clarke,  A.  H.  1981.  The  Freshwater  Molluscs  of  Canada.  Na- 
tional Museum  of  Natural  Sciences,  National  Museums  of 
Canada,  Ottawa,  446  pp. 

Davis,  G.  M.  and  S.  L.  H.  Fuller  1981.  Genetic  relationships 
among  Recent  Unionacea  (Bivalvia)  of  North  America. 
Malacologia  20:217-253. 


Fuller,  S.  L.  H.  1977.  Freshwater  and  terrestrial  mollusks.  In: 
J.  E.  Cooper,  S.  S.  Robin.son,  J.  B.  Fimderbnrg  (eds.)  En- 
dangered and  Threatened  Plants  and  Animals  of  Nortli 
Carolina.  North  Carolina  State  .Museinn  Natural  History, 
Raleigh,  pp.  14.3-191. 

Johnson,  R.  I.  1947.  Laiitpsilis  rariosa  Say  and  Lainpsilis 
ochracea  Say.  Occasional  I'apers  on  Mollusks  1:145-156. 

Johnson,  R.  I.  1970.  The  .systematics  and  zoogeography  of  the 
Unionidae  (Mollusca:  Bivalvia)  of  the  southern  Atlantic 
Slope  region.  Bulletin  of  the  Museum  of  Comparative  Zo- 
ology 140:26.3-449. 

Johnson,  R.  I.  1980.  Zoogeography  of  North  American  Union- 
acea (Mollusca:  BivaKia)  north  of  the  maximum  Pleisto- 
cene glaciation.  Bulletin  ot  the  Museum  of  Comparative 
Zoology  149:77-189. 

Kat,  P.  W.  1983.  Morphologic  divergence,  genetics,  and  spe- 
ciation  among  LampsiUs  (Bivalvia:  Unionidae).  Journal 
Molluscan  Studies  49:1.3:3-145. 

Lea,  I.  1863.  Unionidae,  their  soft  parts  and  embryonic  fonns. 
Obsenations  on  the  genus  Unio  10:10-92. 

Morrison,  J.  P.  E.  1975.  Maryland  and  Mrginia  mussels  of  List- 
er. Bulletin  of  the  American  Malacological  Union  for 
I974:36-.39. 

Ortmann,  A.  E.  1912.  Notes  upon  the  families  and  genera  of 
the  najades.  Annals  of  the  Carnegie  Museum  8:222-.365. 

Ortmann,  A.  E.  1919.  Monograph  of  the  Naiades  of  Pennsyl- 
vania. Part  III.  Systematic  account  of  the  genera  and  spe- 
cies. Memoirs  of  the  Carnegie  Museum  8:1-385. 

Ortmann,  A.  E.  and  B.  \\'alker  1922.  On  the  nomenclature  of 
certain  North  American  naiades.  Occasional  Papers  of  the 
Museum  ot  Zoology,  University  Michigan  112:1-75. 

Porter,  H.  J.  1985.  Molluscan  census  and  ecological  interrela- 
tionships. Vol.  1  Text.  Rare  and  Endangered  Fauna  of 
Lake  Waccamaw.  Final  Report,  North  Carolina  Endan- 
gered Species  Restoration.  North  Carolina  Wildlife  Re- 
sources Connnission,  Raleigh,  170  pp. 

Porter  H.  J.  and  K  J.  Honi.  1980.  Freshwater  nmssel  glochidia 
from  Lake  Waccamaw,  Colinnbus  County,  North  Carolina. 
Bulletin  American  Malacological  Union  for  1980:1.3-17. 

Reardon,  L.  1929.  A  contribution  to  our  knowledge  of  the  anat- 
omy of  the  fresh-water  mussels  of  the  District  of  Colimi- 
bia.  Proceedings  of  the  United  States  National  Museum 
75:1-12. 

Smith,  D.  G.  1991.  Keys  to  the  Freshwater  Macroin\ertebrates 
of  Massachusetts.  Author's  edition,  Sunderland,  236  pp. 

Smith,  D.  G.  1995.  Keys  to  the  Freshwater  Macroinvertebrates 
of  Massachusetts.  Second  Edition.  Author's  edition,  Sun- 
derland, 243  pp. 

Smith,  D.  G.  1996.  A  method  ot  preparing  freshwater  mussels 
(Mollusca:  Unionoida)  tor  anatomical  study.  American 
Malacological  Bulletin  13:12.5-128. 

Stiven,  A.  E.  and  J.  Alderman.  1992. 
among  certain  freshwater  mussel 
Lampsilis  genus  in  North  Carolina, 
369. 

Strayer  D.  1987.  Ecology  and  zoogeography  of  the  freshwater 
mollusks  of  the  Hudson  River  Basin.  Malacological  Re- 
view 20:1-68. 

Strayer  D,  L,  and  K.  J.  Jirka.  1997.  The  Pearly  Mussels  of 
New  York  State.  New  York  State  Museinn  Memoir  26:1- 
113. 

Surber,  T  1912.  Identification  of  the  glochidia  of  freshwater 
mussels.  Report  of  the  Commissioner  of  Fisheries  for 
1912  and  Special  Papers,  Document  771:1-10. 


Genetic  similarities 
populations  of  the 
Malacologia  :34:355- 


Page  160  THE  NAUTILUS,  Vol.  114,  No.  4 

Turgeon,  D.  D.,  J.  F.  Quinn,  Jr., A.  E.  Bogan,  E.  V.  Coan,  F.  Common  and  Scientific  Names  of  Aquatic  Invertebrates 

G.  Hochberg,  W.  G.  Lyons,  P.  M.  Mikkelsen,  R.  J.  Neves,  from  the  United  States  and  Canada;  Mollusks,  2nd.  Edi- 

C.  F.  E.  Roper,  G.  Rosenberg,  B.  Roth,  A.  Scheltema,  F.  tion.  American  Fisheries  Society  Special  Publication  26. 

G.  Thompson,  M.  Vecchione,  and  J.  D.  Williams.  1998.  5.36  pp. 


THE  NAUTILUS  114{4);161-163,  2000 


Page  161 


Latims  beckijae,  a  new  species  of  Fasciolariidae 
(Neogastropoda)  from  Brazil 


Martin  Avery  Snyder 

745  Newtown  Road 
Villanova,  PA  190S5  USA 


ABSTRACT 

Latirus  becktjae  new  species  is  described  from  off  Vitoria,  Es- 
pirito  Santo  State,  and  off  Santos,  Sao  Paulo  State,  Brazil.  The 
new  species  is  readily  distinguished  from  conchologically  sim- 
ilar species  in  the  same  genus  by  its  caramel  color,  white  spiral 
banding,  and  angular  profile. 

Additional  key  words:  Southwestern  Atlantic  Ocean,  south- 
eastern Brazil,  Peristeniiinae. 


INTRODUCTION 

In  May,  1994,  local  .shrimpers  trawling  in  30-35  ni  off 
Vitoria,  Espi'rito  Santo  State,  and  in  190  in  off  Santos, 
Sao  Paulo,  Brazil,  collected  specimens  of  Latims  that 
were  tentatively  identified  as  Latims  cf.  varai  Bullock, 
1970.  Although  superficially  similar  to  L.  varai,  these 
specimens  are  readily  distinguished  from  all  other  spe- 
cies of  western  Atlantic  Latims.  Despite  having  been 
collected  alive,  no  soft  parts  ot  the  new  species  were 
available  for  study,  and  the  species  is  herein  described 
solely  on  the  basis  of  shell  morjihology.  Institutional  ab- 
breviations are  as  follows:  ANSP,  The  Academy  of  Nat- 
ural Sciences,  Philadelphia;  IBUFRJ,  Instituto  de  Bio- 
logia,  Universidade  Federal  do  Rio  Janeiro,  Brazil; 
MNRJ,  Museu  Nacional,  Rio  de  Janeiro,  Brazil;  MORG, 
Museu  Oceanografico,  Funda^iio  Universidade  do  Rio 
Grande,  Brazil;  UNSM,  National  Museum  of  Natural 
History,  Smithsonian  Institution,  Washington,  DC. 

SYSTEM  ATICS 

Family  Fasciolariidae  Gray,  1853 

Subfamily  Peristeniiinae  Tryon,  1880 

Genus  Latims  Montfort,  1810 

Type  species:  Latims  aurantiacus  Montfort,  1810,  by 

monotypy  [=  L.  gibhulus  (Gmelin,  1791)]. 

Latims  beckijac  new  species 
(Figures  1,  2) 

Description:  Shell  length  to  54  mm.  Shell  broadly  fu- 
siform, biconic,  moderately  heavy,  with  prominent  axial 
and  spiral  sculpture.  Protf)conch  with  1  3/4—2  smooth. 


bulbous,  white,  translucent  whorls,  with  axial  sculpture 
and  shoulder  developing  in  last  1/4  whorl.  Transition  to 
teleoconch  abiTipt,  distinguished  by  coarser  axial  sculp- 
ture, onset  of  spiral  sculpture,  and  by  change  of  color 
from  white  to  orange-tan.  Teleoconch  with  up  to  8  1/2 
convex,  strongly  sculptured  whorls.  Axial  sculpture  dom- 
inant, consisting  of  7-8  strong,  broad  ribs  per  whorl, 
ahgned  with  spaces  between  ribs  of  previous  whorl.  Ribs 
rounded  on  early  whorls,  becoming  increasingly  angular 
with  size.  Growth  lines  forming  widely  spaced  lamellae, 
eroded  on  early  whorls  and  on  spiral  cords,  most  proin- 
inent  along  adpressed  suture  and  between  spiral  cords 
on  body  whorl.  Spiral  sculpture  of  strong,  cream-colored 
cords  (3-5  on  early  whorls,  5-7  between  suture  and  pe- 
ripheiy,  6-7  between  peripherv  and  siphonal  canal,  6-9 
on  siphonal  canal)  that  are  most  pronounced  along  axial 
ribs.  Weak  threads  (1-2)  may  be  present  between  adja- 
cent cords.  Aperture  ovate,  with  weakly  developed  pos- 
terior canal.  Golumella  with  1  weak,  narrow  fold  below 
mid-aperture,  and  one  weak,  broad  fold  over  the  fasci- 
ole,  which  may  represent  two  partially  fused  folds.  Outer 
lip  crenulated,  with  12-15  irregular  hrae  running  into 
the  aperture.  Siphonal  fasciole  well  developed.  Pseudo- 
umbilicus  slit-like,  prominent,  open.  Siphonal  canal  as 
long  as  aperture,  broad,  open,  proximal  end  demarcated 
by  tubercles  on  columella  and  outer  hp.  Shell  color  a 
dense  caramel-tan,  with  cream  spiral  cords  and  threads. 
Operculum  medium  dark-brown,  claw-like,  with  termi- 
nal nucleus.  Periostracum  unknown. 


Type  locality: 

in  30-50  m. 


Off  Vitoria,  Espirito  Santo  State,  Brazil, 


Type  material:  Holotype,  USNM  880231  (51.3  mm); 
Parat\pe  1,  USNM  880232  (48.8  mm);  Paratypes  2-3, 
Snyder  Collection  (43.2  mm  and  53.4  mm);  Paratypes 
4-5,  IBUFRJ  9121  (55.3  and  51.2  mm);  Paratvi:)e  6. 
MORG  39008  (50.4  mm);  Paraty^ie  7,  MNRJ  7696  (52.3 
mm),  all  from  the  type  localit}'.  Paratvpe  8,  ANSP  (52.7 
mm),  off  Santos,  Sao  Paulo  State,  Brazil,  in  190  ni. 

Etymology:  The  species  is  named  for  the  author's  el- 
dest daughter,  Becky. 

Discussion:  Latims  hecki/ae  resembles  L.  varoi  (figure 
3;  see  also  Bullock,  1970)  but  is  reachlv  distinguished 


Page  162 


THE  NAUTILUS,  Vol.  114,  No.  4 


Figures  1-2.     Latinis  becktjae  new  species  1.  Holotype,  USNM  880231  (51.3  mm),  2.  Paratype  1,  USNM  880232  (48.8  mm), 

both  from  off  Vitoria,  Espirito  Santo  State,  Brazil,  in  30-50  m. 

Figure  3.     Latinis  varai  Bullock,  1970.  Holotype,  MCZ  262589  (70.0  mm),  from  Off  Gibara,  Oriente  Province,  Cuba,  in  183  m. 


from  it  b\'  its  smaller,  differently  colored  shell.  Latinis 
bcckyae  has  roughK'  half  as  many  strong  spiral  cords  as 
L  varai.  The  axial  ribs  in  L.  heckijae  are  whitish  whereas 
in  L.  varai  the  axial  ribs  are  light  chestnut  brown.  Larger 
(80+  mm),  whitish  specimens  of  Latinis  have  recently 
been  taken  off  San  Salvador,  Bahamas,  in  depths  of  490 
m  by  research  submersibles  (D.  Dan,  pers.  comm.). 
These  closely  resemble  L.  varai,  but  have  far  fewer  spi- 
ral cords  (R.  Bullock,  pers.  comm.).  Not  enough  material 
is  presently  available  to  determine  whether  these  spec- 
imens represent  a  deep-water  form  of  L.  varai  or  an- 
other new  species  of  deep-water  Latinis.  Latinis  beck- 
yae  is  easilv  differentiated  from  the  San  Salvador  spec- 
imens bv  its  smaller  size,  darker  color,  and  coarser,  more 
prominent  sculpture. 

Latinis  beckyae  might  also  be  confused  with  some 
specimens  of  Latinis  ctina  Petuch,  1990,  from  Honduras 
and  east  Panama  (Petuch,  1990).  This  latter  species  is 
usuallv  bright  orange  or  yellow  and  broader  overall  with 
a  proportionally  shorter  canal  than  L.  bcckyae.  Latinis 
cuna  has  pairs  of  white  cords  on  the  body  whorl  and 
sometimes  on  earher  whorls  as  well  as  on  the  terminal 
part  of  the  body  whorl.  Because  of  these  white  spiral 
cords,  an  orange-brown  color  form  of  L.  cuna  superfi- 
cially resembles  L.  beckyae.  Separation  of  the  two  spe- 
cies is  easy:  L.  beckyae  is  more  elongate  and  angular, 
with  a  relatively  larger  teleoconch  than  L.  cuna. 

There  are  four  previously  known  species  of  Latinis 
endemic  to  Brazil,  all  of  which  are  very  distinct  from 
Latinis  becki/ae.  Latinis  devyanac  Rios,  Costa  and  Cal- 


vo,  1994.  from  southeastern  Brazil,  has  a  distinctive 
curved  siphonal  canal  and  grows  to  just  35  mm  (Rios, 
Costa  and  Calvo,  1994).  Latinis  lacteum  Matthews-Gas- 
con. Matthews  and  Rocha.  1991,  from  northern  Brazil. 
is  glossy-white  and  grows  to  just  32  mm  (Matthews-Gas- 
con, Matthews  and  Rocha,  1991).  Latinis  ogum  Petuch, 
1979  (Petuch,  1979;  Rios,  1994:  pi.  42,  fig.  574),  from 
eastern  Brazil  has  few  large  smooth  axial  nodules  and 
attains  43  mm.  Latinis  vennciji  Petuch,  19S6,  from  the 
northern  coast  of  Brazil,  is  very  small  with  a  short 
stumpv  canal,  growing  to  just  26  mm  (see  Petuch,  1986). 

There  are  five  other  Caribbean  species  of  Latinis  also 
found  in  northern  Brazil.  Four  of  them  differ  in  shape 
and  color  from  Latinis  bcckyae:  Latinis  carinifer  (La- 
marck, 1822),  (Rios,  1994:  pi.  42,  fig.  572),  Latinis  an- 
giilatus  (Roding,  1798)  (Rios,  1994:  pi.  42,  fig.  572),  La- 
tinis infundibuhim  (Gmehn,  1791)  (not  Rios,  1994:  pi. 
42,  fig.  573;  see  Abbott,  1974:  color  pi.  11,  fig.  2491), 
and  Latinis  virginensis  Abbott.  1958  (Rios.  1994:  pi.  42. 
fig.  575).  Latinis  beniadensis  Bullock,  1974,  is  uniformly 
light-cream-orange,  a  color  that  sets  its  apart  from  La- 
tinis beckyae  (see  Bullock,  1974). 

There  are  six  species  of  Fu.sinus  known  from  Brazil, 
of  which  two  are  superficially  similar  to  Latinis  bcckyae. 
The  other  four  species  that  cannot  possibly  be  confused 
with  L.  beckyae  are  large  Ftisiniis,  all  with  more  or  less 
dark  browm  markings.  These  are  Fusinus  brasilicnsis 
(Grabau,  1904)  (Rios,  1994:  pi.  42,  fig.  566),  F  closter 
(Phihppi,  1850)  (Rios,  1994:  pi.  42,  fig.  567),  F  mar- 
moratus  (Phihppi,  1846)  (Rios,  1994:  pi.  42,  fig.  569), 


M.  A.  Snyder,  2000 


Page  163 


and  F.  stri^atu.s  (Pliilippi,  1850)  (Rios,  1994:  pi.  42,  fig. 
570).  The  supei-ficiallv  similar  species  are  Fusiuiis  Jtart- 
vigii  (Shiittleworth,  1S56)  (Ahl)()tt,  1974:  color  pi.  11,  fig. 
2494)  and  Fnsinn.s  frcngiwllii  (Carcelles,  1953)  (Rios, 
1994:  pi.  42,  fig.  568).  Fusiiius  haiivigii,  a  species  er- 
roneously placed  in  Latirus  by  Kaicher  (1978:  card 
1816),  exliibits  a  different  color  pattern  and  more  nu- 
merous ittia]  ribs  than  Latim.s  hcckijac.  (Fnsinus  pactcli 
(Dunker,  1867)  is  a  .s\non\in  of  F  hartvigii  named  from 
Brazilian  material.)  Finally,  young  specimens  ol  Fusinus 
frcngucUii  could  be  confused  with  Latinis  bech/ae  (the 
species  was  originally  named  in  the  genus  Lathi/ nis  [  = 
Latinis]).  The  axial  ribs  of  this  species  are  more  nu- 
merous and  its  coloration  ranges  from  uniform  white  to 
pale  orange. 

LITERATURE  CITED 

Abbott,  R.  T.  19.58.  The  nianiie  niollusks  of  Grand  Cayman 
Island,  British  West  Indies.  Monographs  of  the  Academy 
of  Natural  Sciences  of  Philadelphia  11:138  pp.,  5  pis.,  7 
figs- 
Abbott,  R.  T.  1974.  American  seashells.  2nd  ed.  Van  Nostrand- 

Reinhold,  New  York,  663  pp.,  24  pis. 
Bullock,  R.  C.  1970.  Latinis  varai.  a  new  fasciolariid  gastropod 
from  the  Caribbean.  The  Nautilus  83:13:3-1.3.5. 


Bullock,  R.  C.  1974.  A  contribution  to  the  systematics  of  some 
West  Indian  Latinis  (Gastropoda:  Fasciolariidae).  The 
Nautilus  88:69-79. 

Lamarck,  J.  B.  P.  A.  1822.  Hi.stoire  naturelie  des  animau.x  .sans 
vertebres  .  .  .  Vol.  7,  (2),  Paris,  711  pp. 

Matthews-Cascon,  H.,  H.  R.  Matthews  and  C.  A.  Rocha.  1991. 
Nova  especie  de  Latinis  Montfort.  1810  (Mollusca:Gas- 
tropoda),  Boletim  do  Museu  Nacional,  no\a  serie.  Zool- 
ogia  349:1-6,  figs.  1-4. 

Petuch,  E.  ].  1979.  New  gastropods  from  the  Abrolhos  archi- 
pelago and  reef  complex,  Brazil.  Proceedings  of  the  Bio- 
logical Society  of  Washington  92:510-526. 

Petuch,  E.  J.  1986  New  South  American  gastropods  in  the 
genera  Comis  (Conidae)  and  Latinis  (Fasciolariidae).  Pro- 
ceedings of  the  Biological  Societ\-  of  Washington  99:8-14. 

Petuch,  E.  J.  1990.  A  new  molluscan  faunule  from  the  Carib- 
bean coast  of  Panama.  The  Nautilus  104:  57-71. 

Rios,  E.  C.  1994.  Seashells  of  Brazil.  2nd  ed.  Funda^ao  Cidade 
[and]  Funda^ao  Universidade  do  Rio  Grande,  Museu 
Oceanografico  "Prof  Eliezer  de  Carxalho  Rios",  Rio 
Grande,  368  pp.,  113  pis. 

Rios,  E.  C.,  R  M.  Costa  and  I.  S.  Calvo.  1994.  From  off  south- 
em  Brazil  a  new  species  of  Latinis.  La  Conchiglia 
26(273):3.3-36. 

Roding,  P.  F.  1798.  Museum  Boltenianus  sive  Catalogus  ci- 
nielioruni  e  tribus  regnis  naturae  quae  olim  collegerat. 
Pars  seciuida  continens  Conchylia  sive  Testacea  univalvia, 
bivalvia  &  nuiltivahia.  Johan  Christi  Trappii,  Hamburg, 
viii  -I-  199  pp. 


THE  NAUTILUS 


Volume  114 
2000 


AUTHOR  INDEX 


Bennetts,  R.  E. 
BURCH,  K.  W.    .. 

CiAiN,  A.  ] 

Collin,  R 

cordeiro,  ] 

Darrigran,  G.  . 


112 
,  18 
,   93 

117 
.  80 
,   69 


deMaintenon,  M.  J 14 

EZCURRA  DE  DrAGO,  1 69 

Gittenberger,  a 1 

GiTTENBERGER,  E 1 

GOUD,  J 1 

Harasewtch,  M.  G 38,  103,  142 

Hakman,  W.  N 120 

Jansen,  D 112 

K,WANO,  T.  74 


k-antor,  y.  i 

Kim,  W.  

Leal,  J.  H 

Marshall,  B.  A 

MgLean,  J.  H 

monteiro,  w.    

Pastorino,  G 

Penchaszadeh,  p.  E. 

Petit,  R.  E 

Rex,  M.  a 

SiMONE,  L.   R.   L 

Smith,  D.  G 

Snyder,  M.  A 

Sparks,  S.  A 

Yoon,  S.  H 


103 

85 

59 

18,  150 

99 

74 

38,  127 

127 

142 

93 

59,  127 

31,  156 

162 

112 

85 


NEW  TAXA  PROPOSED  IN  VOLUME  114  (2000) 

GASTROPODA 

Columbella  moinensis  deMaintenon,  2000,  new  species  (Columbellidae) 15 

Copulabvssia  riosi  Leal  and  Simone,  2000,  new  species  (Pseudococculinidae) 60 

Crepidula  argentina  Simone,  Pastorino,  and  Penchaszadeh,  2000,  new  species  (Ceilyptraeidae) 129 

Epitonium  hoeksemai  A.  Gittenberger  and  Goud,  2000,  new  species  (Epitoniidae) 4 

Epitonitim  ingridae  A.  Gittenberger  and  Goud,  2000,  new  species  (Epitoniidae) 7 

Epitonium  lochi  A.  Gittenberger  and  Goud.  2000,  new  species  (Epitoniidae)  9 

Epitonium  twilae  A.  Gittenberger  and  Goud,  2000,  new  species  (Epitoniidae) 10 

Latinis  becht/ae  Sn)der,  2000,  new  species  ( Fasciolariidae) 162 

Mericn  deynzeri  Petit  and  Harasewych.  2000,  new  species  (Cancellariidae) 145 

Merica  ekti/phos  Petit  and  Harasewych,  2000.  new  species  (Cancellariidae) 148 

Merica  lussii  Petit  and  Harasewych,  2000,  new  species  (Cancellariidae) 143 

Muricopsis  {Murexstd)  profunda  Marshall  and  Burch,  2000,  new  species  (Muricidae) 24 

Muricopsis  (Rolandiella )  .scoff i  Marshall  and  Burch,  2000,  new  species  (Muricidae) 26 

Obsctiranella  Kantor  and  Harasewych,  2000,  new  genus  (Ranellidae) 103 

Ohscuranella  papijrodes  Kantor  and  Harasewych,  2000,  new  species  (Ranellidae) 103 

Perimnngelia  McLean,  2000,  new  genus  (Turridae) 101 

RetidriUia  McLean,  2000,  new  genus  (Turridae)  100 

Rolandiella  Marshall  and  Burch,  2000,  new  subgenus  (Muricidae) 25 

Subniso  McLean,  2000,  new  genus  (Eulimidae) 100 

Torellivelutina  .McLean.  2000,  new  genus  (Velutinidae) 99 


REVIEWERS  FOR  VOLUME  114 


L.  Adamkewicz 
K.  Bandel 
A.  G.  Beu 
R  Bouchet 
H.  W.  Chaney 
R.  Collin 
R.  H.  Cowie 
K.  S.  Cummings 
D.  J.  Eemisse 
K.  C.  Emberton 
C.  S.  Gallardo 


M.  G.  Harasewych 
G.  Haszprunar 
J.  M.  Healv 

D.  G.  Herisert 

E.  Jokinen 

R.  N.  Kilbum 
C.  M.  Lalli 
H.  G.  Lee 
C.  Lydeard 
W,  G.  Lyons 


J.  H.  McLean 
R   E.  Petit 
R.  Robertson 
B.  Roth 
R.  R,  Seapy 
D.  Strayer 
A.  Verhecken 
G.  T.  Walters 
A.  Waren 
J.  B.  Wise 


Florida  Department  of  State 

Katherine  Harris 

Secretary  of  State 

Florida  Arts  Council 

Division  of  Cultural  Affairs 

This  program  is  sponsored 
in  part  by  the  State  of  Florida, 

Florida  Department  of  State, 

Division  of  Cultural  Affairs,  and 

the  Florida  Arts  Council 


't  ^:  W  L 


7 


INSTRUCTIONS  TO  AUTHORS 


THE  NAUTILUS  publishes  papers  on  all  aspects  of  the 
bio!og)'and  systematics  of  mollusks.  Manuscripts  describing 
original,  unpublished  research  as  well  as  review  articles  will 
be  considered.  Brief  articles,  not  exceeding  1000  words,  vvill 
be  published  as  notes  and  do  not  require  an  abstract.  No- 
tices of  meetings  and  other  items  of  interest  to  malacolo- 
gists  will  appear  in  a  news  and  notices  section. 

Manuscripts:  Each  original  manuscript  and  accompanying 
illustrations  should  be  submitted  in  triplicate.  Text  must  be 
typed  on  one  side  of  SV2  X  11  inch  white  paper,  double 
spaced  throughout  (including  literature  cited,  tables  and 
figure  captions),  with  at  least  1  inch  of  margin  on  all  sides. 
^\11  pages  must  be  numbered  consecutively.  If  printed  on  a 
word  processor,  the  right  margin  should  be  ragged  rather 
dian  justified.  Audiors  should  follow  the  recommendations 
of  the  Scientific  Sttjle  and  Format — The  CBE  Manual  for 
Authors,  Editors,  and  Publishers,  which  is  available  from 
die  Council  of  Biolog)-  Editors,  Inc.,  11250  Roger  Bacon 
Drive,  Suite  8,  Reston,VA  20190,  USA  (http://w\vw.cbe.org/ 
cbe).  The  first  mention  of  a  scientific  name  in  the  text 
should  be  accompanied  by  the  taxonomic  authority,  includ- 
ing year.  Latin  names  and  words  to  be  printed  in  italics 
must  be  underlined;  leave  other  indications  to  the  editor. 
Metric  and  Celsius  units  are  to  be  used. 

The  sequence  of  sections  should  be:  tide  page,  abstract 
page,  introduction,  materials  and  methods,  results,  discus- 
sion, acknowledgments,  literature  cited,  tables,  figure  cap- 
tions, figures.  The  title  page  should  include  the  title,  au- 
thor's name(s)  and  address(es).  The  abstract  page  should 
contain  the  title  and  abstract,  which  should  summarize  in 
250  words  or  less  the  scope,  main  results  and  conclusions 
of  the  paper.  The  abstract  may  be  followed  by  a  maximum 
of  S  key  words.  All  references  cited  in  the  text  must  appear 
in  the  Hterature  cited  section  and  vice  versa.  In  the  litera- 
ture cited  section,  all  authors  must  be  fully  identified  and 
listed  alphabetically.  Follow  a  recent  issue  of  THE  NAU- 
TILUS for  bibliographic  style,  noting  that  journal  titles 
must  be  unabbreviated.  Information  on  plates  and  figures 
should  be  cited  only  if  not  included  in  the  pagination.  Ta- 
bles must  be  numbered  and  each  placed  on  a  separate 
sheet.  A  brief  legend  must  accompany  each  table.  Captions 
for  each  group  of  illustrations  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,  clear- 
ly detailed  and  completely  labeled.  Photographs  must  be 
on  glossy,  high  contrast  paper.  All  figures  are  to  be  consec- 
utively numbered  (figs.  1,  2,  3,  ...  ,  NOT  figs,  la,  lb,  Ic, 


.  .  .  NOR  plate  1,  fig.  1  .  .  .).  Illustrations  must  be  arranged 
in  proportions  that  will  conform  widi  the  width  of  a  page 
(6%  inches  or  171  mm)  or  a  column  (.3%  inches  or  82  mm). 
The  ma.ximum  size  of  a  printed  figure  is  6%  by  9  inches  or 
171  by  228  mm.  All  illustrations  must  be  fully  cropped, 
mounted  on  a  firm,  white  backing,  numbered,  labeled  and 
camera  ready.  The  author's  name,  paper  title  and  figure 
number(s)  should  appear  on  the  back.  Original  illustrations 
must  be  between  one  and  two  times  the  desired  final  size. 
It  is  the  author's  responsibilitv'  that  the  hue  weight  and  let- 
tering are  appropriate  for  the  desired  reduction.  Original 
illustrations  will  be  returned  to  die  author  if  requested.  Col- 
or illustrations  can  be  included  at  extra  cost  to  the  author. 

Voucher  Material:  Deposition  of  t)'pe  material  in  a  rec- 
ognized public  museum  is  a  requirement  for  pubhcation  of 
papers  in  which  new  species  are  described.  Deposition  of 
representative  voucher  specimens  in  such  institutions  is 
strongly  encouraged  for  all  other  tvpes  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  ac- 
ceptance or  rejection.  Accepted  manuscripts  are  returned 
to  the  author  for  consideration  of  the  reviewers'  comments. 

Final  Manuscript  Submission:  Audiors  of  accepted 
manuscripts  will  be  required  to  submit  an  electronic  version 
of  the  manuscript  correctly  formatted  for  THE  NAUTI- 
LUS. The  formatted  manuscript  may  be  sent  as  an  e-mail 
attachment  to  nautilus@sliellmuseum.org  or  in  a  diskette, 
preferably  prepared  using  an  IBM  PC-compatible  text  pro- 
cessor. Original  illustrations  may  be  submitted  separately  by 
regular  mail  or  as  digital  files  (zip  disks  or  CDs),  preferably 
in  TIFF  or  BMP  formats. 

Proofs:  After  typesetting,  two  sets  of  proofs  are  sent  to  the 
author  for  corrections.  Changes  other  than  tvpesetting  er- 
rors will  be  charged  to  the  author  at  cost.  One  set  of  cor- 
rected proofs  should  be  sent  to  the  editor  as  soon  as  pos- 
sible. 

Reprints  and  Page  Charges:  An  order  form  for  reprints 
will  accompany  die  proofs.  Reprints  may  be  ordered 
through  the  editor.  Authors  with  institudonal,  grant,  or  oth- 
er research  support  will  be  billed  for  page  charges  at  the 
rate  of  $60  per  printed  page. 

Manuscripts,  corrected  proofs  and  correspondence  re- 
garding editorial  matters  should  be  sent  to:  Dr.  Jose  H. 
Leal,  Editor,  The  Nautilus,  PO.  Box  1580,  Sanibel,  FL 
33957,  USA. 


@  This  paper  meets  the  requirements  of  ANSI/NISO  Z39.48- 1 992  (Permanence  of  Paper). 


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