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THE 

NAUTILUS 


THE  PILSBRY  QUARTERLY 

DEVOTED  TO  THE  INTERESTS 

OF  CONCHOLOGISTS 


VOL.  75 
JULY,  1961  to  APRIL,  1962 


EDITORS    AND    PUBLISHERS 

HORACE  BURRINGTON  BAKER 
Professor  Emeritus  of  Zoology,  University  of  Pennsylvania 

CHARLES  B.  WURTZ 
Consulting  Biologists  Inc.,  Bethlehem  Pike,  Spring  House,  Pa. 

R.  TUCKER  ABBOTT 
H.  A.  Pilsbry  Chair  of  Malacology,  Academy  of  Natural  Sciences 

MRS.  HORACE  B.  BAKER 

Philadelphia,  Pennsylvania 


PONV    PRINTING,    UPPER    DARBY,    PA. 


April,   1962  nautilus  iii 

CONTENTS 

Names  of  new  genera,  species,  etc.  in  italics 

Acteocina  candei  distinct  from  Retusa  canaliculata  87 

Adventitious  dispersal  94,  124 

Alabama  32,  63,  97,  104,  106,  123,  124,  145 

American  Malacological  Union  83 

Ancylidae,   radulae   97,  145 

Anomia  aculeata,  variation  and  sculpture 131 

Arcidens  confragosus  in  Kansas  95 

Arkansas  28 

Atlantic,  eastern 50,  109 

western   1,  7,  21,  40,  50,  55,  85,  87,  94,  108, 

109,  127,  128,  131,  138,  149 

Australorbis  albicans,  anatomy  and  shell  156 

Beckianum  H.  B.  Baker   (genus  of  Achatinidae  or  subgenus 

of  Leptinaria  84 

Brachidontes  recurvus,  gonad  development  and  spawning  149 

Bulimulus  dealbatus  jonesi  166 

California,    inland      84 

marines   19 

Canada    103 

Capulus  sericeus  Burch  &  Burch  19 

Camaenidae,   Puerto  Rican  64 

Caracolus  marginella  mayaguezi  H.  B.  Baker  64 

Catinella  pugilator  Hubricht  31,  pi.     4,  61 

Catinella  texana  Hubricht  31,  pi.     4,  61 

Cerion,  anatomy    33 

Cionella  lubrica,  aggregations  Ill 

Clausilia,  pallial  complex  36 

Corbicula  fluminea  from  Ohio  River  126 

Cuba    156 

Cymatium  caribbaeum,  range  extension  94 

Dates  of  the  Nautilus  39 

Directory  of  conchologists  40 

Engina  zonata  Gray,  type  species 107 

Eupleura  caudata,  sexual  behavior  7 

Europse    36,  39 

European  Malacological  Congress  39 


82208 


iv  NAUTILUS  Vol.  75  (Index) 

Fluorescence,  enhancement  by  glycerin  138 

Florida,  inland  63,  142 

marine  94 

Gaeotis  nigrolineata  121 

Gardner,  Julia  Anna,  obituary  122 

"Gastrondonta"  saludensis  to  Helicodiscus  105 

Georgia    63,  124 

Geotaxis  in  Physa  75 

Glyphyalinia  junalaskana,  anatomy  125 

Gonads  and  spawning  149 

Haiti    71 

Helicodiscus  barri  Hubricht  105 

Helicodiscus  hadenoecus  Hubricht  106 

Helicodiscus  jacksoni  Hubricht   106 

Helicodiscus  multidejis  Hubricht  102 

Helicodiscus  notius  &  H.  n.  specus  Hubricht  104,  105 

Helicodiscus  saludensis  105 

Helicodiscus  shimeki  Hubricht  103 

Helicodiscus,  subgenus  Troglodiscus  105 

Helisoma  anceps  carried  by  bug  124 

Holopodopes,  infraorder  o£  Geophila 116 

Illinois   113,  126 

Ilyanassa  obsoleta,  mass  movement 85 

Indiana  123 

Iowa 103 

Jamaica  142 

Kansas 46,     95 

Kentucky  105,  106 

Lacinaria,  pallial  complex  36 

Leptinaria,  subgenus   (?)   Beckianum  84 

Louisiana    79 

Maryland  62,  107 

Mexico 31,  84,  142 

Michigan 124 

Mississippi 63 

Missouri    104 

Nenia  tridens,  anatomy  35 

Neoplanorbinae,  radulae  145 

New  Guinea  70 


April,  1962  nautilus  v 

New   Mexico  28 

New  York  103 

North  Carolina  62,  124,  126 

Notes  and  news  39,  84,  123 

Nucella  109 

Obeliscus  (Stenogyra)  terebraster  rarisinister  H.  B.  Baker  117 

Oklahoma  46,  104,  124 

Oleacinoids,  Puerto  Rican  142 

Oliva  rejecta  Burch  &  Burch  165 

Oliva  spicata  &  O.  venulata  162,   165 

Olivella  mutica,  reproduction  139 

Pacific,  eastern  19,  162,  165 

Papuina  ferussaci  67 

"Paravitrea"  roundyi  to  Helicodiscus   (?)    107 

Physa,  geotactic  behavior  75 

"Pilsbryna"  tridens  to  Helicodiscus    (?)    107 

Pleistocene   43 

Polydontes  incerta  &  P.  lima  asperula  66 

Polygyra  gracilis  Hubricht  26 

Polygyra   lithica   Hubricht  28 

Pomacea  paludosa  in  Alabama  123 

Practicolella  berlandieri  campi  29 

Proptera  capax  in  Kansas  95 

Publications  received  41,  86,    (3)    iii,    (4)    iii 

Puerto  Rico  33,  64,  116,  142,  156 

Pupillidae  33 

Radulae  of  ancylids  97,  145 

Retusa  canaliculata  distinct  from  Acteocina  candei  87 

Rhodacmeinae,  radulae  97 

Rissoellidae    21 

Sagdidae   142 

Schwengel,  Jeanne  Sanderson,  obituary  36 

Sexual  behavior,  Eupleura  and  Urosalpinx  7 

Smith,  Maxwell,  death  notice  and  portrait 84,    (4)    pi.     18 

South  America  16 

South  Carolina  63 

Spiraxidae  142 

Stenotrema  calvescens  Hubricht  28 

Strobilops  lonsdalei  &  S.  I.  cansasiana  Ho  &:  Leonard 43,     46 


vi  NAUTILUS  Vol.  75  (Index) 

Strombus  canaliculatus  127,  129 

Succinea  Indiana  60,  123 

Succinea  solastra  Hubricht  30 

Succinea  urbana  Hubricht  32 

Tennessee    29,  102,  105,  106 

Thyasira  sarsi,  synonymy  50 

Texas   27,  30,  43,  61 

Troglodiscus,  subgenus  of  Helicodiscus  105 

Urosalpinx  cinerea,  sexual  behavior  7 

Valvata  windhauseni  Parodiz   16 

Vasum  globulus  nuttingi  1 

Vertigo  teskeyae  62 

Virgin  Island  marines  55,  108 

Virginia  104 

Viviparus  subpurpureus  in  Oklahoma  124 

X-ray  diffraction  &  shell  structure  in  Oliva  162 

Xylophaga  atlantica  40 


INDEX  TO  AUTHORS 

Allen,  J.  Frances  149 

Baker,  H.   Burrington  33,  39,  64,  84,  116,  142 

Basch,  Paul  F 97,  145 

Bode,  William  T.   (Shuster  &)  1 

Branson,   Branley  A.   124 

Briggs,  Rev.  H.  E.  J 39 

Burch,  John  Q 40 

Burch  &  Rose  L.  Burch 19,  165 

Deslandes,  Newton    (Paraense  &)    156 

Dexter,  Ralph  W 40,  85 

Donohue,  Jerry  &  Kenneth  Hardcastle  162 

Dundee,  Dee  Saunders  &  Patti  Watt  79 

Editors 36 

Eyerdam,  Walter  Jacob  71 

Fechtner,  Frederick  R 126 

Gregg,  Wendell  0 84 

Hardcastle,  Kenneth  (Donohue  &)   162 

Hargis,  William  J.,  Jr.  &  Clyde  L.  Mackensie,  Jr 7 

Henrard,  J.  B 67 


April,  1962  nautilus  vii 

Ho,  Tong-Yun  &  A.  B.  Leonard  43 

Hubricht,    Leslie    26,  60,  102,  123,  123,  125,  166 

Leonard,  A.  B.   (Ho  &)    43 

Mackensie,  Clyde  L.,  Jr.    (Hargis  &)    7 

McClary,  Andrew  75 

Merrill,  Arthur  S 94,   1 3 1 

Moore,  D.  R.  (Olsson  &)  127 

Murray,  Harold  D 95 

Ockelmann,  K.  W 50 

Olsson,  A.  A.  &  D.  R.  Moore 127 

Orr,  Virginia  107 

Owen,  D.  F 1 24 

Paine,  Robert  T 1 39 

Paraense,  W.  Lobato  8c  Newton  Deslandes 156 

Parodiz,  J.  J.  16 

Rehder,   Harald  A 1 09 

Robertson,  Robert  2L  128 

Roscoe,  Ernest  J.  1 1 1 

Shuster,  Carl  N.,  Jr.  &  William  T.  Bode  1 

Teskey,  Margaret  C 83 

Watt,  Patti   (Dundee  &)   79 

Weber,  J.  A 55 

Wells,  Harry  J.  &  Mary  Jane  Wells  87 

Wilson,  Druid  122 

Woodridge,  Richard  G.,  Ill  138 


THE  NAUTILUS 

Vol.   75  July,    1961  No.   1 


OBSERVATIONS  ON  VASUM  GLOBULUS  NUTTINGI, 
WITH  COMMENTS  ON  OTHER  CARIBBEAN  VASE  SHELLS 

By  carl  N.  SHUSTER,  JR.,  and  WILLIAM  T.  BODE 

Department  of  Biological  Sciences,  University  of  Delaware^  and 

El  Camino  High  School,  Sacramento,  California 

The  information  in  this  article  is  based  upon  the  observations 
by  Mr.  John  B.  Henderson,  Jr.  (Nutting,  1919),  and  from  field 
notes  made  in  connection  with  the  Smithsonian-Bredin  Caribbean 
Expedition,  1958,  and  by  Mr.  Bode  during  two  years,  1957-1958,  as 
a  Fulbright  Fellow  at  Antigua.^ 

Information  on  the  distribution,  ecology,  evolution,  and  sys- 
tematics  of  the  genus  Vasum  in  the  western  Atlantic  is  of  particu- 
lar interest  to  the  writers  of  this  article.  Since  at  least  three  species, 
V.  jnuricatum,  capitelliim,  and  V.  globulus  (Globivasum)  nuttingi 
(Henderson),  have  overlapping  ranges  in  the  Lesser  Antilles,  spe- 
cial care  should  be  made  to  report  the  exact  locality  and  observa- 
tions of  each  find. 

Habitat  notes.  Vasum  muricatum  and  V.  globulus  nuttingi  were 
found  within  a  few  miles  of  each  other  at  Barbuda,  but  in  dis- 
tinctly different  habitats.  Vasum  muricatum  is  generally  found  in 
or  close  to  turtle  grass  beds. 
Vasum  muricatum    (Born) 

Anegada:  from  Pomato  Point  in  shoal  water  on  Turtle  Grass 
flat.  Barbuda:  south  shore  (Gavenor's  Landing) .  5  specimens 
from  shallow  water,  3  to  6  feet  depths,  on  sandy  bottom  in  region 
of  coral  heads  and  nearby  shoreward  expanse  of  turtle  grass.  Ab- 
bott (1950)  noted  that  "It  seems  to  prefer  rather  sheltered,  shal- 
low waters,  but  is  sometimes  found  on  relatively  exposed  reefs." 
Antigua:  Jumby  Bay,  Long  Island. 
Vasum  globulus  nuttingi   (Henderson) 

Barbuda:  windward  side  of  Spanish  Point.  Specimens  found  in 
shallow  water  along  coral   (dead)   rock  ledges  and  on  patches  of 

1  University  of  Delaware  Marine  Laboratories,  Contribution  No.  7. 

2  For  the  opportunities  afforded  him,  as  an  invited  member  of  the  expedi- 
tion, the  senior  author  wishes  to  express  his  gratitude  to  the  sponsors  of  the 
expedition,  Mr.  and  Mrs.  J.  Bruce  Bredin  and  to  the  expedition  leader, 
Dr.  Waldo  L.  Schmitt. 

I 


2  NAUTILUS  Vol.    75     (1) 

sand  among  the  rocks.  Roughly  40%  of  the  specimens  were  within 
a  few  inches  of  Diadema.  Several  Vasum  were  only  an  inch  away 
from  the  periphery  of  the  Diadema.  Depths  were  1  to  3  feet.  In 
only  one  case,  a  specimen  was  collected  in  another  habitat:  turtle 
grass  on  a  sand-covered  ledge  of  coral  rock,  depth  6  inches. 
Antigua:  from  areas  just  inside  points  of  land,  in  semi-protected 
waters.  As  at  Barbuda,  where  an  offshore  coral  reef  broke  the 
direct  force  of  the  surf,  the  localities  in  Antigua  were  areas  with 
well-circulated  seawater  but  not  exposed  to  direct  surf.  We  found 
no  Nutting's  vase  well  within  bays.  Localities  included  are:  Car- 
lisle Bay:  in  3  feet  of  water.  Dead  Sands:  rocky  area  on  the  east 
side,  in  5  feet  of  water.  Ding-a-Dong  Nook:  in  4  to  5  feet  of  water. 
Exchange  Bay  and  Smith  Island  at  Mill  Reef:  in  one  to  21/2  feet. 
Freeman's  Bay:  inside  harbor  mouth.  Galley  Bay:  in  3  to  6  feet 
of  water.  Literally  hundreds  of  Nutting's  Vase  shells  containing 
hermit  crabs  were  observed  on  the  Galley  Bay  side  of  the  point 
separating  Galley  Bay  and  Little  Galley  Bay.  Specimens  can  be 
collected  at  Little  Galley  Bay  when  the  sea  is  calm.  Guana  Island: 
at  the  part  of  the  island  called  the  Headlands,  at  3  to  4  foot 
depths.  Half  Moon  Bay:  inside  rocky  area  on  the  Mill  Reef 
side  of  the  bay  and  in  2  to  6  feet  of  water,  along  western  shore- 
line, midway  between  the  sand  beach  and  the  open  sea  to  the 
south.  Johnson's  Island:  northwest  side,  in  4  to  5  foot  depths. 
Rendezvous  Bay:  in  4  feet  of  water;  this  area  may  be  heavily 
populated,  but  extremely  rough  surge  makes  it  difficult  to  collect; 
it  was  the  only  locality  on  Antigua  where  specimens  were  found 
directly  exposed  to  the  open  sea.  Windward  Bay:  in  tide  pools 
formed  by  a  small  reef  of  dead  coral,  at  the  water's  edge  to  one 
and  a  half  feet;  in  3  to  6  foot  depths  outside  of  the  tide  pools. 

Natural  history  of  species  of  Vasum:  Little  is  known  about 
the  living  animal.  Vasum  muricatum  is  known  to  be  predatory 
upon  worms  and  clams  (Abbott,  1954)  ;  presumably,  the  other 
species  are  also  predatory. 

In  its  habitat,  Vasum  globulus  nuttingi  escapes  notice  from 
all  but  the  experienced  collector.  Nutting  (1919)  reported  from 
the  notes  of  Mr.  Henderson  that,  "Miss  Sykes  seemed  to  be  the 
only  collector  with  eyes  properly  focussed  to  detect  these  turbin- 
ellas  clinging  to  the  rocks,  so  well  concealed  are  they  by  their 
coloration  and  by  the  calcareous  deposit  that  further  hides  them 
from  their  enemy  (but  not  Miss  Sykes).  One  of  the  chief  enemies 
of  these  rock-living  mollusks  is  a  fish  of  the  grouper  family  that  at 
high  tide  swims  about  the  reefs  examining  every  inch  of  their 
surface  for  little  mollusks  that  have  not  hidden  themselves  very 
carefully."   Miss   Esther   Bates,   in   a   mimeographed   newsletter, 


July,   1961  NAUTILUS  3 

commented  upon  collecting  at  Carlisle  Bay,  Antigua,  on  22 
April,  1958:  "The  tide  was  low  and  they  [natives]  led  us  to  a 
coral  reef  at  the  water's  edge.  They  [Nutting's  vase]  are  difficult 
to  see  against  the  rocky  background  but  I  came  away  with  a 
goodly  numbei"  ..." 

Vasiim  muricatum  may  be  nocturnal,  hiding  during  the  bright 
sunlight  hours  among  a  little  species  of  coral  and  turtle  grass; 
traveling  across  the  bottom,  in  2  to  3  feet  of  water,  in  the  late 
afternoon.  This  may  also  be  the  case  for  V.  globulus  nuttingi. 
Most  specimens  appear  to  be,  to  the  collector  swimming  about  in 
search  of  them,  at  rest  among  growths  on  rocks  or  partially  cov- 
ered with  sand.  Possibly  visual,  audible,  or  pressure  stimuli, 
caused  by  the  swimmer,  cause  the  snails  to  "freeze."  Specimens 
removed  from  the  water  frequently  come  part  way  out  of  their 
shells  and  appear  to  be  quite  active. 

Three  of  the  species  have  been  dredged  from  moderate  depths 
{Abbott,  1950;  Rehder  and  Abbott,  1951)  :  V.  latiriforme  (18  to 
20  fathoms) ,  capitellum  (10  fathoms),  and  globulus  nuttingi  (7 
fathoms) .  The  type  locality  of  globulus  nuttingi  was  designated 
as  7  fathoms  (Abbott,  1950) ,  whereas  the  published  account  of 
the  living  animals  (Nutting,  1919)  reported  them  on  the  rocks 
in  shallow  water  about  the  reefs  at  the  entrance  to  Freeman's 
Bay.  There  is  no  doubt  that  shallow  water  areas  are  characteristic 
habitats;  it  remains  to  be  proved  that  moderate  depths  are  also 
usual  or  common.  If  a  range  in  depth  of  habitat  exists,  then  a 
seasonal  migration  pattern,  perhaps  for  feeding  or  breeding,  may 
be  involved.  In  this  respect,  the  collections  by  Bode  and  Shuster 
were  made  only  in  shallow  water,  during  the  period  from  9 
March  through  25  June,  1958.  A  false  distribution  may  be  caused 
by  the  activities  of  predators,  such  as  the  grouper  (Nutting, 
1919) ,  or  by  hermit  crabs. 

V.  globulus  nuttingi  has  been  found  at  depths  ranging  from 
the  water's  edge  to,  as  a  rule,  not  more  than  6  or  8  feet  of  water. 
In  every  case,  except  at  Rendezvous  Bay,  the  snails  were  found  in 
an  area  protected  from  the  direct  waves  of  the  open  sea,  yet  they 
were  always  located  where  there  was  considerable  water  circula- 
tion. As  a  rule,  they  were  not  found  in  sheltered  areas,  as  at  the 
head  of  a  bay.  The  most  sheltered  habitat  observed  by  Mr.  Bode 
was  just  inside  the  entrance  to  Freeman's  Bay,  where  the  snails 


4  NAUTILUS  Vol.   75     (1) 

were  found  in  shallow  water,  not  exceeding  a  foot  in  depth.  Their 
distribution  in  this  particular  area,  as  in  several  others,  was 
restricted  to  a  zone  covered  by  a  moss-like  giowth  on  the  rocks. 
This  growth  was  not  collected  nor  identified,  although  it  can 
be  recognized  easily  again  once  a  person  has  seen  it.  It  appears 
like  a  coarse  moss,  14  inch  to  I1/2  inches  long,  ranging  in  color 
from  a  grey  to  a  brown. 

In  some  areas  the  Nutting's  vase  was  found  in  the  shallow  water 
within  a  short  distance  of  the  beach,  among  rocks  and  dead  coral, 
with  a  higher  ridge  of  rock  and  dead  coral  between  the  open 
sea  and  their  habitat.  Often  a  bank  of  live  coral  would  be  located 
at  the  edge  of  this  "protecting"  reef  and  the  water  depths  quickly 
dropped  off  to  at  least  25  or  30  feet.  Although  examples  of  V. 
globulus  nuttingi  were  found  in  shallow  water,  they  were  within 
a  distance  generally  not  exceeding  50  to  a  100  feet  from  deep 
water. 

Specimens  of  Nutting's  vase  are  frequently  found  partially 
buried  in  pockets  of  coral  sand  in  crevasses  of  dead  coral.  When 
collected  out  in  the  open,  on  the  top  of  rocks,  they  are  always 
beneath  the  water's  surface.  No  examples  were  ever  found  on 
living  coral,  although  some,  as  at  Half  Moon  Bay,  were  almost 
completely  encrusted  with  what  appeared  to  be  a  living  coral. 
This  coral  had  a  very  light  lavender  color  when  first  taken 
from  the  water.  It  completely  covered  and  hid  the  shape  of  the 
shell,  with  the  exception  of  the  aperture. 

Large  numbers  of  immature  V.  globulus  nuttingi  have  not  yet 
been  found.  In  some  areas  only  adult  specimens  occuned.  In 
others,  only  immature  examples  were  found  and  when  one  or 
two  snails  were  found  in  an  area  they  were  most  often  immature. 
On  the  basis  of  these  observations,  since  in  most  areas  the  adult 
specimens  considerably  outnumber  the  immature  ones,  we  believe 
that  either  the  young  stages  are  to  be  found  in  another  habitat, 
perhaps  a  breeding  area,  or  that  the  adult  populations  are  main- 
tained by  a  low  number  of  surviving  immature  snails.  Mr.  Bode 
has  noted  that  the  local  populations  can  be  over-fished.  This 
observation  suggests  that  the  snails  either  have  a  wider  range 
of  distribution,  as  into  greater  depths,  or  more  probably,  there 
is  a  low  level  of  population  replenishment  due  to  predation  and 
other  forms  of  mortality  or  to  a  low  reproductive  rate. 


July,    196 1 


NAUTILUS 


ligmcs  a  lu  d,  J'a.suin  {(.'Aabivaauin)  globulus  nutliugi  (Henderson),  lo 
illnstrate  variation  in  adult  shells:  a,  the  "fossilized"  shell  frim  a  kitchen 
midden,  b,  characteristic  shell,  c.  an  elongate  specimen,  d,  the  largest  shell. 
Scale  block  5  cm.  in  length. 

An  area  that  needs  considerably  more  investigation  comprises 
the  small  islands  off  the  northeast  coast  of  Antigua,  and  in  par- 
ticular, the  windward  side  of  Long  Island  and  Guana  Island. 
Exchange  Island,  Little  Exchange,  Hellgate,  Pelican  Island  and 
Maid  Island  should  also  be  more  closely  investigated.  In  this 
general  area,  dead  shells  have  been  picked  up  that  appear  to  be 
larger  and  longer  than  those  found  on  the  south  side  of  the  island. 

Among  the  several  dozens  of  V.  globulus  nuttingi  collected  by 
Mr.  Bode,  3  are  worthy  of  special  mention:  2  are  atypical,  one 
was  found  in  a  kitchen  midden.  These  3  specimens  (figs  a,  c,  d) 
and  a  more  usual  one  (fig.  b)  from  Barbuda  (Shuster!)  are  de- 
picted on  page  5.  An  elongate  specimen  (fig.  c)  was  collected 
alive  at  Windward  Bay,  Antigua,  on  9  March,  1958.  It  was  found 
at  a  depth  of  about  3  feet  of  water  in  an  area  consisting  of  broken, 
dead  coral  and  coral  rock,  with  small  patches  of  sand.  A  very 
large  shell  was  found  in  the  same  area.  The  third  shell  (fig.  a) 
was  found  in  a  kitchen  midden,  approximately  two  feet  below 
the  surface,  in  conjunction  with  potshard  and  broken  conch  and 
nuuex  shells.  The  midden  site  was  at  the  western  end  of  a  "shell 
beach"  on  tlie  north  side  of  Antigua,  between  the  airjjort  and 


NAUTILUS 


Vol. 


;i) 


■n 


IP  I 


EupU'ura  caudaiu  (Say)  .  Fig.  1 ,  male  and  female  in  ropi.la.  Fig.  2.  two  males 
attempting  copulation  with  one  female. 


July,   1961  NAUTILUS  7 

Long  Island.  This  shell  was  in  a  bank  approximately  7  feet  above 
mean  tide  and  10  feet  back  from  the  water's  edge.  Two  of  these 
specimens  were  pictured  at  slightly  larger  than  actual  size  by 
Shuster  (1959)  :  the  example  with  tlie  elevated  spire  and  the  low- 
spired  shell  from  the  midden. 

References 
Abbott,    R.   T.    1950.   The  genera   Xancus   and    Vasurn   in   the 

western  Atlantic.  Johnsonia,  2  (28)  :  20 1-2 18. 

1954.  American  Seashells.  D.  van  Nostrand,  Princeton,  N.  J. 

Nutting,  C.  C.  1919.  Barbados-Antigua  expedition.  University  of 

Iowa  Studies  in  Natural  History,  8(3)  :  199-203. 
Rehder,  H.  A.  and  R.  T.  Abbott.  1951.  Some  new  and  interesting 

mollusks  from  the  deeper  waters  of  the  Gulf  of  Mexico.  Rev. 

Soc.  Malacologica,  8  (2) -.5 3-66. 
Shuster,  C.  N.  Jr.  1958.  Caribbean  adventure.  Estuarine  Bulletin, 

5(2):7-12. 

SEXUAL  BEHAVIOR  OF  THE  OYSTER  DRILLS: 
EUPLEURA  CAUDATA  AND  UROSALPINX  CINEREA' 

By  WILLIAM  J.  HARGIS,  JR.,  and  CLYDE  L.  MACKENZIE.  JR.2 

Virginia  Fisheries  Laboratoi7 

The  observations  reported  herein  are  part  of  an  extensive  study 
of  the  biology  of  Urosalpinx  cinerea  (Say) ,  the  smooth  oyster 
drill,  and  Eupleura  caudata  (Say) ,  the  rough  oyster  drill,  which 
is  in  progress  at  our  laboratory.  Results  of  studies  of  other  aspects 
of  the  behavior  and  ecology  of  these  predatory  snails  will  be  re- 
ported as  they  become  available. 

Little  is  known  of  the  copulatory  behavior  of  dioecious  marine 
gastropods.  Though  Stauber  (1943)  reported  a  partial  pairing  of 
E.  caudata,  pairing  of  U.  cinerea  has  never  been  described.  Our 
studies  show  that  pairing  of  both  species  is  a  complex  process 
involving  fairly  intricate  behavior  of  both  sexes. 

Copulation  o^  Eupleura  caudata.  According  to  Stauber's  (1943) 
description  of  a  partial  mating  of  E.  caudata  the  male  mounted 
the  right  side  of  the  female  and  formed  the  anterior  part  of  his 
foot  into  a  copulatory  groove  through  which  the  penis  was  ex- 


1  Contributions  from  the  Virginia  Fisheries  Laboratory,  No.  00.  This  re- 
search was  conducted  under  contract  with  the  U.  S.  Fish  and  Wildlife  Service, 
Bureau  of  Commercial  Fisheries,  No.  14-19-008-2372,  Study  of  Oyster  Drills  in 
Chesapeake  Bay. 

-  Present  address.  Bureau  of  Commercial  Fisheries  Biological  Laboratory, 
Milford,  Connecticut. 


8  NAUTILUS  Vol.    75     (1) 

tended  into  her  mantle  cavity.  Even  though  disturbed  repeatedly 
this  pair  copulated  intermittently  for  21  days.  Our  laboratory 
observations  of  pairing  of  captive  E.  caudata  confirm  and  enlarge 
upon  those  of  Stauber.  Eighty-one  separate  pairings  were  observed 
but  because  the  average  time  each  couple  was  together  was  ex- 
tensive and  coitus  occurred  intermittently,  only  two  copulations 
were  seen  from  beginning  to  end. 

Prior  to  coupling,  the  female  usually  moved  about  the  aquar- 
ium walls  and  finally  assumed  a  stationary  position  with  her 
siphonal  tip  upward.  Pairing  also  occurred  in  horizontal  positions 
on  oysters,  or  on  the  aquarium  bottom,  or  rarely,  on  the  walls. 
The  mantle  cavity  was  then  exposed  by  allowing  the  shell  to 
hang  down  and  away  from  the  posterior  part  of  the  foot.  Though 
in  most  cases  a  male  (or  males)  had  already  assumed  a  position 
upon  the  shell  of  the  female,  several  unpaired  females  also  ex- 
posed their  mantle  cavities  in  the  same  fashion  prior  to  copulating 
and  probably  this  behavior  is  normal  precopulatory  activity.  Fol- 
lowing this  the  male  often  moved  about  on  the  female's  shell  for 
a  short  time  but  eventually  assumed  a  stationary  position  on  her 
right  ventral  surface  slightly  posterior  to  the  mantle  cavity  with 
his  siphonal  cavity  pointing  in  the  same  direction  as  hers  (Fig.  1, 
p.  6) .  The  foot  of  the  male  extended  to  the  rim  of  the  mantle 
cavity  and  a  copulatory  groove  formed  in  its  anterior  surface 
through  which  the  penis  was  protruded  into  the  mantle  cavity 
and  presumably  the  vagina,  though  that  orifice  was  always  hidden 
from  view  by  the  shell.  On  termination  of  copulation,  the  female 
often  twisted  from  side  to  side  and  opened  and  closed  the  cavity 
opening.  Similar  twisting  behavior  has  been  observed  in  other 
gastropods  apparently  attempting  to  dislodge  predatory  snails, 
Odostomia,  from  their  shells  (Allen,  1958) .  The  male  then  with- 
drew and  either  remained  in  position  or  moved  about  on  the 
shell  of  the  female  or  to  the  substrate. 

E.  caudata  usually  remained  in  position  and  copulated  inter- 
mittently for  extensive  periods,  often  up  to  48  hours.  Two  pairs 
copulated  continuously  for  2  hours  45  minutes  ±  15  minutes  and 
3  hour  39  minutes  ±  10  minutes,  and  a  third  for  5  hours  ±  10 
minutes,  but  all  were  paired  longer.  Another  pair  copulated 
intermittently  for  a  total  of  at  least  8  hoins  48  minutes  over  a 
period  of  two  days,  during  which  the  male  maintained  his  posi- 


July,  1961  NAUTILUS  9 

tion  on  the  female.  A  marked  pair  copulated  1 1  times  in  five 
months.  During  this  period  the  pair  often  separated  and  the 
female  paired  with  other  males. 

Eupleura  caudata  showed  marked  promiscuity.  Of  29  marked 
pairs,  12  females  copulated  with  more  than  one  male  and  15  males 
with  more  than  one  female.  One  male  copulated  with  6  different 
females  and  one  female  accepted  5  males  throughout  the  season. 
Although  these  are  observations  on  drills  confined  in  running- 
water  aquaria  and  cages,  there  is  little  reason  to  doubt  that, 
depending  on  density  and  movement,  similar  promiscuity  occurs 
in  nature. 

Usually,  males  were  smaller  than  their  consorts  (48  males: 
mean  height  18.7  mm.,  range  14.9-23.5  mm.,  standard  deviation 
2.3;  44  females:  mean  height  22.6  mm.,  range  17.5-28.9  mm., 
standard  deviation  3.2) . 

Seasonal  periodicity  of  copulation  was  determined  in  outside 
cage  experiments  and  running-water  aquarium  observations. 
Though  the  frequency  of  observation  was  not  always  constant 
throughout  the  year,  the  number  of  pairings  seen  in  aquaria  per 
month  roughly  corresponded  to  those  in  more  careful  cage  exper- 
iments. In  1956  pairings  were  recorded  as  follows:  March-6, 
April-14,  May-15,  June-3,  July-9,  August-1,  September-1,  October- 
9,  November- 1,  and  December- 1.  The  first  occurred  on  March  7 

(12.9°)  and  the  first  peak  occurred  in  late  April  and  early  May. 
A  second  peak  came  in  July,  comparatively  few  pairings  occurred 
from  July  through  September,  but  an  increase  took  place  in  the 
last    part    of    October.    Copulation    ceased    after    December    10 

(10.3°C) .  The  lowest  temperature  at  which  pairings  were  ob- 
served was  10.3°C,  the  highest  28.4°C.  Also  in  1956,  30  females 
and  30  males,  caged  in  a  single  large  compartment,  were  examined 
every  two  days  between  1400  and  1700  hours.  Onset  of  the  mating 
period  was  not  observed  because  the  experiment  was  established 
too  late  in  the  season.  Pairing  was  first  seen  in  late  April  at  13.7°C, 
reached  a  peak  on  June  7  at  23.7°C,  and  ceased  on  July  7  at 
26.1  °C,  (Figure  3).  A  late  wave  of  28  pairings,  about  one-third  as 
intense   as   the   spring   wave,    began    at    the   end   of   September 

(21.7°C)  and  ended  November  11  (16.4°C.)  Observations  made 
two  or  three  times  a  week  during  the  winter  of  1956-57  revealed 
no  copulatory  activity  but  this  was  expected  because  all  other 


10 


NAUTILUS 


Vol.  75    (1) 


i 


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1 1 1 1 1 1  1 1 1 


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lOvAuHro        9  1IH01a 


t.joo  to   on 


:.<I03    JO     ON 


overt  activity  had  ceased.  Observations  in  the  first  half  of  the 
summer  of  1957  on  two  cage  compartments  containing  45  speci- 
mens each  (sex  ratio  of  entire  sample,  50  females  to  40  males) 
indicated  that  copulation  was  less  intense  than  in  1956.  Pairing 
was  first  observed  on  April  4  (1 1.1  °C) .  There  was  no  distinct  peak 


July,    1961  NAUTILUS  11 

and  after  the  week  of  May  6  activity  slowed  and  ceased  on  July 
6  (26.1°C).  Most  activity  occurred  slightly  earlier  than  in  1956, 
probably  as  a  result  of  earlier  warming.  No  observations  were 
made  in  late  summer. 

Little  diurnal  periodicity  was  noticed  and  nocturnal  activity 
was  not  investigated.  Of  56  pairings,  23  (48  per  cent)  were  com- 
pleted before  1200  hours  and  29  (52  per  cent)  occurred  after  1200 
hours.  Because  observational  effort  varied  somewhat  these  data 
ajre  not  precise,  though  they  are  probably  indicative  of  general 
conditions  and  comparable  to  those  pertaining  to  U.  cinerea 
gathered  in  similar  fashion,  see  below. 

On  two  occasions,  pairs  of  males  were  observed  in  copula. 
Careful  external  examination  and  studies  of  gonad  smears  showed 
all  four  animals  to  be  normal  males  with  no  detectable  female 
characteristics.  Five  trios  in  copula,  each  composed  of  a  female 
and  a  male  in  the  usual  position  with  an  extra  male  in  copulatory 
position  on  the  first  male,  were  also  observed  in  aquaria.  The 
extended  penes  of  both  males  were  seen  twice.  Usually  the  male 
next  to  the  female  copulated  with  her  while  the  intromittent 
organ  of  the  second  male  extended  into  the  mantle  cavity  of  the 
first,  but  at  times  both  males  attempted  copulation  with  the 
female  (Fig.  2,  p.  6) .  Copulations  involving  two  functional  males, 
the  middle  one  acting  as  both  male  and  female,  and  another 
female  has  been  reported  for  the  hermaphroditic  species, 
Lymnea  stagnalis,  (Crabbe,  1927)  but  never  for  dioecious  gas- 
tropods. Once  a  quartet  with  3  males,  all  situated  chain-fashion 
on  a  female,  was  observed.  The  penes  of  at  least  two  of  these 
males  were  extended  into  the  mantle  cavity  of  the  animal  before 
them.  In  dioecious  animals,  like  drills,  these  abeirrant  copulatory 
groups  have  little  reproductive  significance,  but  seem  to  support 
the  hypothesis  of  exocrine  stimulation  or  attraction  of  males  to 
females.  Possibly  the  males  were  attracted  to  the  females,  or  to 
each  other,  by  release  of  female  exocrine  in  the  vicinity  and, 
without  being  able  to  discriminate  further,  established  and  main- 
tained contact  with  each  other. 

Copulation  of  Urosalpinx  cinerea.  Observations  of  76  pairs  of 
U.  cinerea  showed  their  copulatory  behavior  to  be  generally  the 
same  as  that  of  E.  caudata,  but  with  several  distinct  differences. 
The   female   usually    assumed   a    stationary    position    with    her 


12  NAUTILUS  Vol.    75     (1) 

siphonal  tip  upwards,  allowed  the  shell  to  fall  away  from  the 
substrate,  thus  exposing  the  right  rear  corner  of  the  mantle 
cavity  and  twisted  the  shell  from  side  to  side  several  times  in  a 
sort  of  "precopulatory  dance."  Most  often  a  male  was  already  on 
the  female's  back  when  this  precopulatory  play  was  observed,  but 
several  times  males  appeared  to  be  attracted  from  afar  to  the 
demonstrating  female.  Sometimes  unattached  males,  often  several 
at  a  time,  were  attached  to  a  demonstrating  female  or  copulating 
pair,  possibly  drawn  to  the  receptive  female  or  the  pair  by  some 
stimulus,  probably  an  exocrine.  Following  the  precopulatory  dem- 
onstration by  the  female  the  male  assumed  a  position  on  her  right 
ventral  side,  formed  a  copulatory  groove  of  the  anterior  portion 
of  his  foot  and  inserted  his  penis  through  the  groove  into  the 
female's  mantle  cavity.  On  completion  of  coitus,  the  intromittent 
organ  was  withdrawn,  and  in  contrast  to  the  slow  separation  or 
intermittent  resumption  of  E.  candata,  the  male  moved  quickly 
away.  The  shortest  complete  copulation  observed  lasted  4  min- 
utes, the  longest  32  min.  Including  preplay  time,  one  complete 
pairing  occupied  one  hour  and  25  minutes,  but  actual  copulatory 
contact  required  only  1 1  minutes.  Usually  U.  cinerea  copulation 
occupied  only  3  to  4  minutes.  This  characteristic  short  contact, 
seldom  more  than  4  or  5  minutes,  probably  explains  why  U. 
cinerea  mating  has  not  been  previously  reported. 

According  to  our  observations,  only  one  pair  of  20  marked 
pairs  recopulated  and  the  pairings  were  U/o  months  apart.  Of 
the  marked  pairs  7  males  and  2  females  coupled  with  different 
mates.  Thus,  E.  caudata  seemed  more  promiscuous  than  U. 
cinerea.  However,  the  incidence  of  promiscuity  in  U.  cinerea 
may  be  actually  greater  than  these  data  indicate  because  of  the 
unlikelihood  of  observing  its  brief  sexual  contacts  as  readily  as 
the  much  longer  ones  of  E.  caudata. 

The  male  is  usually  the  smaller  of  the  pair:  41  pairing  Seaside 
(from  the  ocean  coast  of  the  Eastern  Shore  of  Virginia)  males 
averaged  29.8  mm.  in  height,  range  19.5-38.7,  standard  deviation 
5.1;  46  females  averaged  34.4  mm.  in  height,  range  21.9-44.6  mm., 
standard  deviation  4.7;  six  York  River  males  averaged  19.3  mm. 
in  height,  range  16.5-24.6  mm.  and  five  females  averaged  19.5  mm. 
(17.3-21.2  mm.). 

Pairing  was  seen  in  late  October  and  early  November,   1955, 


July,   1961  NAUTILUS  13 

until  the  temperature  dropped  to  13.7°C.  In  1956,  it  began  on 
Februai-y  29  (8.2°C.,  increased  in  March  with  10  pairings,  reached 
a  peak  in  April  with  17  pairings,  and  diminished  in  May  to  three 
pairings  (20,3°C) .  Pairing  was  not  observed  again  until  Oc- 
tober, (20.8°C),  none  in  November  and  2  on  December  7 
(11.5°C).  Of  59  pairings,  nine  (15  per  cent)  were  observed 
before  1200  hours  and  50  (85  per  cent)  after  1200  hours.  In 
contrast  to  E.  caudata,  U.  cinerea  seems  to  exhibit  marked 
preference  for  the  afternoon. 

Two  pairs  of  structurally  normal  males  were  observed  in  copula 
and,  on  another  occasion,  two  males  were  observed  copulating 
with  one  female. 

Sperm  viability  and  storage.  One  U.  cinerea  male,  forcibly 
separated  from  a  female,  exuded  sperm  in  a  continuous,  viscous 
stream,  thus  indicating  that  discrete  spermatophores  are  prob- 
ably not  employed.  Examination  of  seminal  receptacles  of  at 
least  50  females  of  both  species  support  this  conclusion.  Seminal 
receptacle  smears  of  two  females  taken  immediately  after  copula- 
tion contained  immotile  sperm  while  those  in  another  were 
motile;  therefore,  it  is  not  clear  whether  the  sperm  are  immotile 
when  passed.  Microscopic  examinations  of  seminal  receptacles  of 
several  hundied  individuals  of  both  species  revealed  that  many 
mature  females  contained  fully  or  partially  motile  sperm  at  all 
times  of  the  year  though  the  sperm  in  some  were  entirely  immo- 
tile. Whether  they  are  stored  in  the  motile  condition  is  not  clear 
because  the  mechanical  action  of  smearing  may  have  stimulated 
the  sperm  to  activity.  Apparently  spermatozoal  energy  is  sustained 
in  some  way,  either  through  conservation  of  energy  by  immotility 
or  special  nourishment  because  both  species  can  store  viable  sperm 
for  considerable  periods  of  time. 

Stauber  (1943)  reported  that  a  U.  cinerea  female  isolated  from 
April  to  October  of  the  same  year  deposited  e^^  cases  containing 
viable  embryos  through  the  period.  In  our  experiments  4  females 
isolated  in  November  and  December  of  1955  produced  viable 
eggs  the  following  spring  and  summer.  Of  these,  one  deposited  in 
May  after  5  months,  two  spawned  in  August,  after  somewhat  less 
than  9  months,  and  one  in  September  after  9  months.  Two  post 
copula  E.  caudata  females  isolated  in  1955  produced  egg  cases 
through  1956  and  into  1957,  but  the  embryos  produced  in  1957 


14  NAUTILUS  Vol.   75     (1) 

did  not  develop.  Unless  parthenogenesis  occurred — which  is  most 
unlikely — or  the  so-called  "embryos"  included  in  these  1957  egg 
cases  were  merely  unfertilized  ova  and  not  really  embryos  at  all, 
the  sperm  must  have  been  vigorous  enough  to  affect  fertilization 
even  after  a  storage  period  of  over  14  months.  In  any  case,  how- 
ever, the  embryos  (or  unfertilized  ova)  produced  in  1957  failed 
to  develop  even  though  handled  in  the  same  fashion  as  others 
which  survived.  Eight  other  females,  isolated  in  late  fall  or  early 
winter  of  1956  produced  viable  cases  for  periods  of  at  least  six 
months  after  isolation. 

Discussion.  Several  points  of  biological  interest  have  arisen 
during  the  present  study.  Because  of  the  nature  of  the  photo 
receptive  organs  it  is  doubtful,  but  not  certainly  so,  that  the 
precopulatory  "dance"  or  movements  of  the  female  attracts  the 
male.  More  feasible  is  the  chemical  stimulation  of  the  male  by 
the  female.  If  female  oyster  drills  actually  attract  males  during 
their  "premating  ritual"  by  release  of  an  exocrine,  such  a  chemical 
might  be  useful  as  an  experimental  or  control  tool  because  it  is 
probably  highly  specific  and  powerful  enough  to  be  effective  in 
extreme  dilution. 

It  has  been  shown  that  females  of  both  species  are  able  to  store 
sperm  in  their  seminal  receptacles  for  extensive  periods.  The 
mechanisms  by  which  they  are  sustained  should  be  of  interest  in 
studies  of  cell  culture  and  nutrition. 

Oyster  drills  are  promiscuous  and  have  the  facility  for  sperm 
storage;  therefore,  unless  sperm  from  previous  pairings  are  dis- 
carded or  resorbed  completely  before  another  mating  occurs,  a 
female  contains  viable  sperm  from  several  different  males  in  her 
seminal  receptacle.  Under  these  circumstances,  embryos  produced 
therefrom  might  have  different  paternal  hereditary  materials. 
Thus,  differences  in  rate  of  embryonic  development,  time  of 
hatching,  appearance,  etc.  may  be  due  to  varied  paternity  and 
not  the  usual  genetic  difference  found  among  siblings.  Until  the 
precise  nature  of  sperm  storage,  syngamy  and  egg  case  formation 
is  understood,  experimenters  working  with  drill  embryos  cannot 
safely  assume  that  all  embryos  in  a  unimaternal  cluster  or  even  a 
single  egg  case  are  of  the  same  parentage. 

Summary 

1.  Eupleura  caudata  and  U.  cirierea  exhibited  fairly  complex 


July,   1961  NAUTILUS  15 

mating  behavior  involving  definite  precopulatory  movement  pat- 
terns by  the  female,  stimulation  of  nearby  males,  possibly  by 
exocrine  activity,  assumption  of  relatively  constant  copulaiory 
positions  by  both  sexes  and  copulatory  groove  formation  by  males. 
Females  often  twisted  violently  immediately  prior  to  cessation  of 
copulation. 

2.  Urosalpinx  cinerea  completed  copulation  in  a  matter  of 
minutes,  but  E.  caudata  often  persisted  intermittently  for  days, 
remaining  in  position  all  the  while. 

3.  Eupleiira  caudata  was  apparently  more  promiscuous  than 
U.  cinerea  but  possibly  this  disparity  may  have  been  a  function  of 
a  species  different  in  frequency  of  a  pairing,  i.e.,  if  E.  caudata 
normally  pairs  more  often,  its  comparative  incidence  of  promis- 
cuity would  naturally  be  greater.  Or,  it  may  also  have  been  a 
function  of  the  length  of  time  that  pairing  consumes.  For  ex- 
ample, E.  caudata  pairing  takes  much   longer  than   U.  cinerea 

(12  to  20  hours  vs.  3  to  4  minutes  or  up  to  200  times  longer)  ; 
therefore,  U.  cinerea  copulation  would  more  easily  be  overlooked, 
resulting  in  a  numerical  bias  in  favor  of  E.  caudata  in  any  com- 
parison of  frequency  of  pairing. 

4.  Though  U.  cinerea  paired  at  lower  temperatures  than  E. 
caudata  (8.2°C  vs.  10.2°C) ,  mating  activities  of  both  increased 
during  April.  U.  cinerea  ceased  copulatory  activity  in  June  while 
E.  caudata  persisted  through  the  first  week  in  July  before  stopping 
temporarily.  Both  resumed  mating  activities  in  September  which 
increased  in  October,  diminished  in  November  and  ceased  alto- 
gether in  early  December. 

5.  In  our  laboratory  aquaria,  U.  cinerea  copulated  more  fre- 
quently in  the  afternoon  and  evening  while  mating  activities  of 
E.  caudata  were  more  evenly  distributed  throughout  the  day. 

6.  Females  of  both  species  stored  viable  sperm  in  their  seminal 
receptacles  for  periods  of  at  least  6  to  9  months. 

7.  Several  items  of  general  biological  interest,  sperm  nutrition 
and  multiple  paternity  of  embryos  within  a  single  eg^  case,  etc., 
have  been  discussed. 

Literature  cited 
Allen,  J.  Frances.  1958.  Nautilus  72:11-15. 

Bushland,  R.  C,  A.  W.  Lindquist  and  E.  F.  Knipling,  1955.  Sci- 
ence 722:287-288. 


16  NAUTILUS  Vol.    75     (1) 

Crabbe,  E.  D,  1927.  Biol.  Bull.  55:67-109. 

Stauber,  L.  A.  1943.  Ecological  studies  on  the  Oyster  Drill,  Urosal- 
pinx  cinerea  in  Delaware  Bay,  with  notes  on  the  associated 
drill,  Eupleura  and  with  special  consideration  of  control 
methods,  Unpubl.  Typescript,  Oyster  Res.  Lab.,  Rutgers 
Univ.  New  Brunswick,  N.  J.:  1-180. 


NOTES  ON  VALVATIDAE  FROM  EARLY  TERTIARY  OF 

SOUTH  AMERICA,  WITH  A  NEW  SPECIES 

By  J.  J.  PARODIZ 

The  living  Valvatidae  are  widely  distributed  in  the  northern 
hemisphere,  with  only  two  species  known  south  of  the  equator  in 
New  Caledonia.  Fossils  are  known  from  the  Jurassic  and  Tertiary 
of  the  Old  World,  as  well  as  from  the  Pliocene  of  California  and 
Pleistocene  of  central  United  States.  The  Neotropical  region  has 
three  species  described  from  Guatemala  and  one  from  Jamaica. 

However,  from  South  America,  where  living  species  of  Vnhiata 
are  unknown,  C.  H.  Fritzche  in  1924  (1)  described  several  fossil 
forms,  discovered  by  the  geologist  G.  Steinmann  in  very  early  Ter- 
tiary strata  (Puca  Formation)  of  Bolivia  and  N.W.  Argentina. 
These  are: 

Valvata  humilis  Fr.  (op.cit  p. 23,  pl.II,  f.6) ,  limestone  of  Mira- 
flores,  near  Potosi,  Bolivia. 

Valvata  yaviana  Fr.  (p. 23,  pl.II,  f.7) ,  and  V.  satira  Fr.  (p.24, 
pl.II,f.8) ,  limestone  of  Yavi,  N.  of  province  of  Jujuy,  Argentina, 
close  to  the  Bolivian  border. 

According  to  the  original  descriptions  and  figures,  there  are 
very  little  specific  differences  among  Fritzche's  forms,  without 
sculpturing  or  carination,  and  no  indication  to  which  of  the 
present  subgenera  they  may  be  related,  but  all  seem  to  be  of  regu- 
lar planorboid  shape  as  in  Valvata  s.s.,  differing  also  from  V. 
Q-uatemalensis  Morelet  or  strebeli  Cross,  and  Fisch. 

A  very  different  form,  found  in  strata  of  southern  regions  but 
same  relative  age,  presents  the  characteristic  strong  keel  recalling 
the  subgenus  Tropidina.  It  was  previously  an  unknown  taxono- 
mic  element  in  the  extinct  Patagonian  fauna,  and  belongs  to  a 
new  species,  as  follows: 
Valvata  windhauseni  new  species.  PI.  1,  figs.  1-6 

Shell  very  large,  with  almost  5  whorls  (the  last  portion  of  the 


July,  1961  NAUTILUS  17 

body  whoii  not  completely  preserved) ,  increasing  regularly,  and 
very  convex,  but  at  middle  zone  of  the  last  whorl  an  angulosity 
is  marked  by  a  strong  carina  or  marginate  keel,  which  starts  at  the 
end  of  the  suture,  above  the  apertural  angle;  below  the  keel,  the 
body  becomes  convex  again.  The  best  preserved  portions  of  the 
last  two  whorls,  are  conspicuously  marked  with  oblique  striae 
very  regularly  spaced  (approx.  0.4  mm.  apart),  also  visible  on  the 
base  of  the  shell.  Although  partially  covered  with  a  hard  calcar- 
eous matrix,  a  wdde  umbilical  area  is  observed.  The  median  alti- 
tude of  the  last  whorl,  from  suture  to  keel  is  about  the  same  as 
the  rest  of  the  spire,  and  the  altitude  of  the  shell  is  almost  equal  to 
its  width.  The  apex,  well  preserved,  is  rather  obtuse. 

Dimensions:  total  height  6  mm.;  height  from  apex  to  carina 
4.5  mm.;  maj.  diam.  8.5,  min.  diam.  7.5  mm. 

Holotype*,  and  4  paratypes,  from  Nahuel  Niyeu  (25  miles  W. 
of  Valcheta) ,  Rio  Negro  province,  Argentina,  in  lacustrine  de- 
posits of  the  Jahuel  Formation  (Windhausen  1918),  of  Danian 
age  (Paleocene)  ;  collector  Dr.  R.  Wichmann  1923   (2) . 

The  species  is  named  in  memory  of  Dr.  A.  Windhausen,  who 
contributed  greatly  to  the  knowledge  of  the  Cretaceous-Tertiary 
boundary  stratigraphy  in  Patagonia,  and  its  fossils. 

The  smaller  paratype,  is  a  well  preserved  inner  cast,  crystallized 
by  infiltration  of  silica  solution,  a  characteristic  of  many  of  the 
fresh-water  fossils  found  in  these  strata.  This  silicification  is  also 
present  in  the  apex  of  the  holotype.  The  other  paratypes  are 
larger  than  the  holotype,  showing  the  carina,  filled  with  tuffaceous 
matrix,  and  deformed  by  lateral  pressure,  probably  due  to  the 
strong  diastrophism  which  subsequently  affected  the  sediments 
during  the  Tertiary. 

The  area  where  the  specimens  were  found,  south  of  the  Rio 
Negro  River,  corresponds  to  a  great  depression  in  the  territory, 
partially  filled  with  lacustrine  and  tuffaceous  sediments,  parallel, 
and  sometimes  mixed,  or  underlying,  with  the  Roca  marine 
beds  (3) ,  covering  in  unconformity  the  Senonian  substratum. 
From  this  area  also,  an  abundant  fresh-water  fauna  is  known,  with 
Diplodon  bondenbenderi  D-J.,  Physa  doering  D-J.,  Physa  wich- 
manni  Parodiz,  several  pleurocerids  under  the  old  name  of 
"Melania",  Lioplacodes,  and  others  (4) .  This  fauna  shows  resem- 
blance with  that  of  Upper  Cretaceous  and  Lower  Tertiai^  of 
western  United  States. 


*  Type   and   paratypes  in   Museo   Argentine   Nat.   Ciencias   and    Carnegie 
Musuem  respectively. 


18  NAUTILUS  Vol.   75     (1) 

This  new  species  is  the  only  well  carinate  Valvata,  fossil  or 
living,  known  from  the  Neotropical  region.  To  assign  it  to  the 
subgenus  Tropidina,  on  account  of  its  strong  keel,  might  be,  how- 
ever, a  taxonomic  risk.  The  apex  is  not  flattened  but  somewhat 
elevated,  as  in  Pleurovalvata,  although  the  spaced  radial  striae  are 
very  different  from  the  coarse  rib-like  lamellae  of  that  subgenus, 
and  the  general  outline,  on  the  other  hand,  resembles  Cincinna. 
By  all  these  combinations  of  features,  V.  windhauseni  cannot  be 
restricted  to  any  of  the  known  subgenera.  Valvata  is  a  genus  very 
plastic  to  the  enviroment,  and  Haas  (5)  reached  the  conclusion 
that  lacustrine  habits  are  in  someway  related  with  sculpturing  in 
fresh-water  mollusks  and,  in  the  case  of  Tropidina,  the  presence  of 
a  keeled  sculpture  is  probably  due  to  a  genotypic  feature  acquired 
during  prolonged  lacustrine  life  and  become  invariable.  For  very 
variable  living  species,  as  V.  utahensis,  Morrison  (6)  questioned 
if  the  specimens  should  be  distributed  in  thirds  to  the  subgenera 
Cincinna,  Pleurovalvata  and  Tropidina,  or  should  one  disregard 
these  names  as  long  as  they  are  based  on  superficial  characters 
only. 

Notes  and  references 

(1)  C.  H.  Fritzche,  Neues  Jahrb,  fur  Min.  Geol.  und  Palaont., 
50:1-56,  taf.1-4,  Stuttgart  1924.  Paper  continues  on  pages  313-334 
about  older  fauna  which  are  of  no  concern  here,  but  the  explana- 
tions of  plates  all  on  pag.  334. 

(2)  R.  Wichmann,  1927,  published  an  account  of  this  strati- 
graphic  series:  Boletin  Academia  Nacional  Ciencias,  Cordoba, 
50:384-407.  Recent  studies  proved  that  the  moUuscan  fauna  is 
Danian. 

(3)  Roca  (Rocanean  or  "Rocanense")  now  considered  as 
northern  member  of  San  Jorge  Formation  (Montian) ,  and  it  is 
synchronic  with  Puca  Formation  of  Peru  and  Bolivia  (with  Val- 
vata) which  Fritzche  formerly  placed  in  the  Upper  Cretaceous. 
See  also  references  to  other  fresh-water  mollusks  of  the  Puca  For- 
mation in  H.  A,  Pilsbry,  John  Hopkins  Univ.,  Studies  in  Geol. 
ii:  69-72,  1939;  and  F.  Ahlfeld,  Revista  Museo  La  Plata  (Geol.) 
3:1-370,  1946. 

(4)  See  M.  Doello  Jurado,  BoL  Acad.  Nac.  Cienc.  Cordoba, 
50:407-416,  1927. 

(5)  F.  Haas,  Zool.  Ser.  Field  Museum  Nat.  Hist.,  24,  No.  8, 
1939. 

(6)  Naut.  55:140,  1940. 


NAin  ILUS  75    (I) 


PLA  11.    1 


I'ah'ala  windhauseni  Paiodiz,  from  tvpe  lot.  Figs.  3  to  6,  holotvpe. 


NAUTILUS  75    (1) 


PLATE  2 


Capulus  sericeus  Burch  &  Burch.  Figs.  1   &  2,  holotype,  ventral  and  dorsal 
views.  Figs.  3  &  4,  additional  examples,  mentioned  in  text 


July,   19G1  NAUTILUS  19 

A  NEW  CAPULUS 
FROM  GULF  OF  CALIFORNIA 

By  JOHN  Q.  and  ROSE  L.  BURCH 

Among  other  interesting  species  taken  on  the  Ariel  Expedition 
to  the  Gulf  of  California,  Aug.  30  to  Sept.  6,  1960,  were  several 
specimens  of  a  species  of  Capulus  which  appears  to  be  new. 

Capulus  sericeus,  new  species.  Plate  2. 

Shell  capshaped,  oval;  apex  spiral,  turned  towards  the  pos- 
terior side,  curling  downwards,  and  twisted  to  the  left  as  in  some 
species  of  the  genus  Crepidula;  aperture  transversely  oval,  with 
an  irregularly  sinuated  margin,  the  posterior  expanded;  exterior 
surface  with  definite  and  distinct  axial  and  radiating  raised  lines, 
a  velvety  periostracum  extending  beyond  the  margins;  interior 
rose  color  with  darker  rays  extending  from  the  apex  to  the  anter- 
ior margin;  shell  not  symmetrical  but  modified  in  shape  by  its 
sessile  habit  conforming  to  the  surface  on  which  it  is  attached; 
growth  marks  conspicuous  but  irregular.  Length  of  holotype  14.8 
mm.;  width  12.3  mm.;  height  6.3  mm. 

This  new  form  was  taken  commensal  on  Pecten  sericeus  Hinds, 
1845.  It  is  obviously  related  to  Capulus  californicus  Dall  of  Cali- 
fornia, which  is  commensal  on  Pecten  diegensis  Dall,  but  differs 
from  that  species  in  the  sculpture  of  the  exterior  surface  and  the 
color  and  rays  of  the  interior.  The  axial  lines  are  much  more 
pronounced  than  are  those  ©n  Capulus  californicus  Dall,  and  the 
radial  lines  are  much  more  prominent.  The  periostracum  on 
Capulus  sericeus  is  much  more  like  overlapping  shingles  than  on 
Capulus  californicus  Dall,  which  seems  to  be  nearer  to  Capulus 
ungaricoides  (Orbigny)  in  that  it  is  smooth  under  the  periostra- 
cum. On  Capulus  californicus  Dall,  the  radiating  lines  are  not  as 
uniform,  but  seem  to  be  more  as  irregularities  in  the  shell  due  to 
conformation  to  the  host  shell.  A  study  of  the  protoconch  shows 
little  variation  in  species,  all  being  of  the  same  general  form,  but 
a  comparison  indicates  that  Capulus  sericeus  has  a  more  sunken 
protoconch.  Capulus  sericeus  resembles  Capulus  hungaricus  Lin- 
naeus, type  species  of  the  genus,  in  having  radiating  raised  lines, 
but  lacks  the  fine  ribs  that  radiate  from  the  apex  toward  the  mar- 
gin in  the  latter  species.  The  only  other  species  described  from  the 
eastern  Pacific  is  Capulus  ungaricoides  (Orbigny,  1841),  type 
locality,  Payta,  Peru.  Orbigny  desaibed  his  species  under  the 
name  Pileopsis  Lamarack,  1812  which  is  in  the  synonymy  of  Capu- 


20  NAUTILUS  Vol.    75     (1) 

lus  Montford,  1810.  This  species  is  smooth  except  for  radiating 
raised  lines  on  the  periostracum.  We  wish  to  thank  Dr.  S.  P. 
Dance,  British  Museum  (Natural  History)  for  his  advice  that 
Capulus  ungaricoides  (Orbigny)  does  not  seem  to  him  to  be  the 
same  species.  In  addition  to  the  shell  characters  mentioned  above, 
we  are  convinced,  from  a  study  of  ocean  currents,  that  it  is  un- 
likely that  this  occurrence  constitutes  a  range  extension  from 
Peru.  Capulus  chilensis  Dall,  1904,  from  the  Antarctic,  is  possibly 
a  species  related  to  Capulus  ungaricoides  (Orbigny) . 

The  type  specimen  (pi.  2,  figs.  1  &  2)  was  trawled  off  Cabo 
Haro,  near  Guaymas,  Sonora,  Mexico  at  a  depth  of  100  fathoms, 
and  will  be  deposited  in  the  Stanford  University  Type  Collection, 
no.  8519.  This  specimen  is  from  the  collection  of  Dr.  Homer  King. 

Other  specimens  here  named  paratypes  are:  one  specimen 
trawled  off  Cabo  Haro,  Sonora,  Mexico  in  20-40  fathoms,  Dec.  27, 
1959  on  the  ship  General  Yanes,  collection  of  Dr.  Donald  Shasky; 
one  specimen  trawled  off  Cabo  Haro,  Sonora,  Mexico,  collection 
of  Mr.  Mark  Rogers. 

The  following  are  mentioned  as  hypotypes,  all  from  the  Gulf 
of  California,  but  not  the  type  locality;  one  specimen  trawled  off 
Espiritu  Santo  Island  in  40-90  fathoms,  fig.  3,  collection  of  Dr. 
Donald  Shasky;  one  specimen  trawled  off  Monserrate  Island  in 
20-40  fathoms,  Sept.  1,  1960,  fig.  4,  collection  of  Dr.  Donald 
Shasky;  one  specimen  trawled  off  MonseiTate  Island  in  75  fath- 
oms, collection  of  Mr.  Mark  Rogers. 

References 

Carcelles,  Alberto.  1944.  Comm.  Zool.  del  Museo  De  Hist.  Nat.  de 

Montevideo. 
Dall,  W.  H.  1889.  Marine  mollusks  of  the  southeastern  coast.  U.S. 

Nat.  Mus.  Bull.  37:152. 
Dall.  W.  H.  1900.  Naut.  13  (9):  100. 
Dall,  W.  H.  1908.  Mollusca  and  Brachiopoda.  Albatross  Rep.  Bull. 

Mus.  Comp.  Zool.  ^i  (6)  :329. 
Dall,  W.  H.  1927.  In  Eastman-Zittel,  Textbook  of  Paleontology, 

2nd  ed.,  London. 
Orbigny,  A.  d'  1811.  Amer.  Merid.    (Moll.),  p.457. 
Thiele,  Johannes.  1931  Handbuck  der  systematischen  Weichtier- 

kunde,  vol.  1,  p.  246. 
Tryon,  George  W.  1886.  Man.  of  Conch.  (1)  5:105,131. 


July,  1961  NAUTILUS  21 

A  SECOND  WESTERN  ATLANTIC  RISSOELLA 
AND  A  LIST  OF  THE  SPECIES  IN  THE  RISSOELLIDAE 

By  ROBERT  ROBERTSON 
(Concluded  from  April  no.) 

Genera,  subgenera  and  species  in  the  family  Rissoellidae 

The  genera  and  subgenera  Laciinella  (family  Lacunidae) , 
Fairhankia  (Micromelaniidae) ,  Hyala,  "Dardania"  (=:Darda- 
nula),  Tatea  (Rissoidae) ,  and  Diala  (Cerithiidae)  have,  at  one 
time  or  another,  been  wrongly  included  in  the  Rissoellidae  by 
various  malacologists.  Some  of  the  species  listed  below  (notably 
those  named  by  A.  Adams  and  W.  H.  Turton)  probably  are  also 
not  rissoellids.  All  species  originally  named  as  rissoellids  are  in- 
cluded in  the  list,  as  well  as  species  which  have  been  rightly  or 
wrongly  transferred  to  rissoellid  genera. 

alhella,  Rissoa.  Alder  (1844) .  See  also  Thompson  (1844) .  An 
older  name  for  R.  glabra  Alder  non  Brown  than  R.  diaphana. 
The  name  R.  albella  Alder  never  has  been  adopted  and  the  "50 
year  rule"  is  here  invoked  to  suppress  it  (I.C.Z.N.  ruling 
pending) . 

alderi,  fjeffreysia.  Carpenter  (1856).  "Mazatlan"  [W.  coast 
Mexico].  According  to  Bartsch  (1920)  a  Barleeia  (Rissoidae) . 
See  also  Baker,  Hanna  &  Strong   (1930) . 

angulijerens,  Rissoa.  Folin  (1869)  .  "Bale  de  Panama."  Re- 
ferred by  Bartsch    (1920)    to  Rissoella. 

atlantica,  Jeffreysia.  Smith   (1892).  "St.  Helena." 

bakeri,  Rissoella  (?) .  Strong  (1938).  "9  to  15  fms.  off  Guada- 
lupe Island,  Mexico." 

becki,  Jeffreysia.  Turton    (1932) .  "Port  Alfred,  South  Africa." 

bifasciata,  Jeffreysia.  Carpenter  (1856).  "Mazatlan"  [W.  coast 
Mexico].  See  Bartsch   (1920). 

caffra,  Rissoa  (Cingula).  Sowerby  (1897).  "Port  Elizabeth" 
[South  Africa].  Referred  by  Bartsch  (1915)  and  Turton  (1932) 
to  Jeffreysia.  Specimens  collected  by  Turton  at  Port  Alfred 
(M.C.Z.   101515)    are  not  rissoellids. 

californica,  fRissoella.  Bartsch  (1927),  "San  Clemente  Island, 
California." 

capensis,  Rissoa  (Cingula).  Sowerby  (1892) .  "Port  Elizabeth" 
[South  Africa].  Apparently  this  is  the  species  referred  by  Bartsch 
(1915)   io  Jeffreysia.  See  Turton   (1932). 

caribaea,  Rissoella  (Phycodrosus).  Rehder  (1943a) .  "Bonefish 
Key,  Fla."  [One  of  the  Crawl  Keys  between  Grassy  Key  and  V^aca 
Key,  Florida  Keys.] 

cylindrica,  Jeffreysia.  Jeffreys  (1856) .  ".  .  .  in  about  12  fathoms 
at  Spezia"    [Italy].   Referred  by   Monterosato    (1878)    to  Aclis 


22  NAUTILUS  Vol.  75    (1) 

(Aclididae) .  Specimens  from  Sicily  (A.N.S.P.  247781),  which 
presumably  are  correctly  identified,  have  hyperstrophic  proto- 
conchs  and  are  pyramidellids. 

diaphana,  Rissoa?  Alder  (1848.  New  name  for  R.?  glabra 
Alder  non  R.  glabra  (Brown) .  Referred  by  Forbes  &  Hanley 
(1850)  to  Jeffreysia  and  incorrectly  by  Clark  (1850)  to  "Chem- 
nitzia"  (Pyramidellidae) ,  who  claimed,  wrongly,  that  the  apex 
is  reflexed  (i.e.,  hyperstrophic)  ;  see  Jeffreys  (1851)  and  Alder 
(1851a) .  See  albella  and  nitidus. 

duperrei,  Paludestrina.  Velain  (1877).  [Misspelled  'Duperei' 
p.  144.]  "He  Saint-Paul;  dans  I'ouest  du  banc  Roure,  par  les 
fonds  de  35  metres  a  50  metres."  [S.  Indian  Ocean].  Referred  by 
Thiele    (1912)    to  Jeffreysiopsis. 

eburnea,  Rissoa.  Stimpson  (1851a,  b)  .  ".  .  .  in  thirty  fathoms, 
off  Cape  Ann"  [Massachusetts].  Doubtfully  and  wrongly  referred 
by  Gould  (1870)  to  Rissoella.  An  "Odostomia"  (Pyramidellidae); 
see  Bartsch  (1909) . 

edzvardiensis,  Jeffreysia.  Watson  (1880).  [Challenger]  "St.  145a 
.  .  .  Lat.  46°  41'  S.,  long.  38°  10'  E.  Prince  Edward  Island,  between 
Cape  of  Good  Hope  and  Kerguelen.  50  to  150  fms."  Figured  Wat- 
son (1886);  possibly  from  310  fms.,  or  from  Sta.  145.  Referred 
doubtfully  by  Thiele  (1912)  to  Jeffreysiella,  and  definitely  by 
Iredale   (1912)   to  Heterorissoa. 

excolpa,  Rissoella.  Bartsch  (1920) .  "Concepcion  Bay,  Lower 
California"  [Mexico].  See  Baker,  Hanna  &:  Strong    (1930). 

julgida,  Rissoella.  "A.  Ad."  Dunker    (1882).   [Japan]. 

fiiscotincta,  Jeffreysia.  Turton  (1932) .  "Port  Alfred,  South 
Africa." 

galba,  Rissoella.  Robertson,  this  paper. 

glabra,  Rissoa?.  Alder  (1844).  "Dalkey  Island  near  Dublin 
[Ireland],  and  at  Cullercoats,  Northumberland  [England]."  A 
misapplication  of  the  name  Rissoa  glabra  (Brown)  Brown  (1844), 
a  pyramidellid.  R.  glabra  Alder  referred  to  Rissoella  by  Gray 
(1847) .  Weinkauff  (1868)  wrongly  referred  i?.  g/o 6 ra  (Brown) 
to  Jeffreysia.  See  diaphana. 

globularis,  Jeffreysia.  "Jeffreys,  MS."  Forbes  &  Hanley  (1852). 
".  .  .  at  Skye  [Scotland]  and  the  Shetlands  .  .  ."  Figured   (1853) . 

gulsonae,  Chemnitzia.  Clark  (1850).  ".  .  .  prope  ostia  Iscae 
Danmoniorum."  [Exmouth,  England].  Referred  doubtfully  by 
Jeffreys  (1851)  to  Jeffreysia  and  later  (1867)  to  Aclis.  Type 
species  of  Pherusina   (Aclididae) . 

hera,  Jeffreysia.  Turton    (1932).  "Port  Alfred,  South  Africa." 

hertleini,  Rissoella.  Smith  &  Gordon  (1948).  "10  fathoms  .  .  . 
off  Cabrillo  Point,  Monterey  Bay,  California." 

Heterorissoa  Iredale  (1912).  Type  species  (original  designa- 
tion) :  H.  secunda.  Synonym  of  Jeffreysiella  according  to  Thiele 
(1925). 


July,  1961  NAUTILUS  23 

hydrophana,  Rissoella.  A.  Adams  (1860) .  "Tabu-Sima  [Tobi 
Shima];  25  fathoms"  [Japan]. 

indistincta,  Jeffreysia.  Turton  (1932) .  "Port  Alfred,  South 
Africa." 

infiata,  Jeffreysia.  Monterosato  (1878).  Nom.  nudum.  "Pal, 
[ermo]  e  Trap,   [ani]    (Monts.)  ;  Messina    (Granata) ."  [Sicily]. 

Jeffreysia  Alder  in  Forbes  &  Hanley  (1850) .  Type  species  (or- 
iginal designation)  :  /.  diaphana.  Objective  junior  synonym  of 
Rissoella. 

Jeffreysiella  Thiele  (1912) .  Type  species  (original  designation): 
/.  notnbilis. 

Jeffreysilla  Thiele  (1925) .  Type  species  (monotypy)  :  Rissoella 
zebra. 

Jeffreysina  Thiele  (1925) .  Type  species  (subsequent  designa- 
tion, Winckworth,  1932)  :  /.  globularis.  Ranked  as  genus  by 
Rehder    (1943a). 

Jeffreysiopsis  Thiele  (1912) .  Type  species  (here  designated) : 
Paludestrina  duperrei  Veiain.  Synonymized  with  Rissoella,  s.s., 
by  Thiele   (1925). 

joJtnstojii,  Rissoella.  Baker,  Hanna  &:  Strong  (1930) .  "Cape 
San  Lucas,  Lower  California"   [Mexico]. 

nialayensis,  Rissoella?  Thiele  (1925).  "Station  211  .  .  ."  [805 
meters,  7°  48.8'  N.,  93°  7.6'  E.,  near  Nicobar  Islands]. 

minima,  Rissoella.  A.  Adams  (1860) .  "Tsu-Sima;  26  fathoms" 
[Japan]. 

mundula,  Rissoella.  A.  Adams  (1860) .  "Tsu-Sima;  26  fathoms" 
[Japan]. 

nitidus.  Turbo.  J.  Adams  (1797).  Nom.  dubium.  "Pembroke- 
shire" [Wales].  Considered  by  Jeffreys  (1867)  a  possible  older 
name  for  Rissoa?  diaphana  Alder. 

nitida,  Jeffreysia.  "Sars"  Friele  (1876).  Hydrobia  nitida  Sars 
(1859),  nom.  nudum.  "Bergen"  [Norway].  Close  to  "Rissoella" 
eburnea  (Stimpson) ,  a  pyramidellid,  according  to  Sars  (1878) . 
See  eburnea. 

notabilis,  Jeffreysiella.  Thiele  (1912) .  "Observatory  Bay  .  .  . 
Kerguelen."   [S.  Indian  Ocean].  See  also  Thiele    (1925). 

omphalotropis,  Rissoella.  A.  Adams  (1860) .  "Sado;  30  fathoms" 
[Japan]. 

opalina,  Rissoa    (?) .  Jeffreys    (1848).   "Guernsey  and  Sark" 
[Channel  Islands].  Referred  by  Forbes  &  Hanley    (1850)    to  Jej- 
freysia  and  incorrectly  by  Clark  (1851)   to  "Chemnitzia"  (Pyram- 
idellidae;  see  Alder   (1851b). 

pauli,  Rissoella.  See  sancti-pauli. 

Phycodrosus  Rehder  (1943a) .  Type  species  (original  designa- 
tion) :  Rissoella  caribaea.  Subjective  synonym  of  Jeffreysilla  (this 
paper) . 

Rissoella   Gray    (1847).   Type   species    (monotypy):    "Rissoa? 


24  NAUTILUS  Vol.  75    (1) 

glaber,  Alder."   [z=Rissoa?  diaphana  Alder]. 

sancti-pauli,  Rissoella  [  (Jeffreysia)].  Velain  (1877).  Velain 
(1876),  nom.  nudum.  "He  Saint-Paul  .  .  .  au  niveau  de  la  basse 
mer  . . ."  [S.  Indian  Ocean].  Emended  to  R.  paidi  by  Crosse  (1879). 

secunda,  Heterorissoa.  Iredale  (1912).  "Kermadec  Islands." 
[S.  Pacific  Ocean]. 

simoniana,  Jeffreysiopsis.  Thiele  (1912) .  "Simonsbai"  [prob- 
ably Simonstown,  nr.  Cape  Town,  South  Africa]. 

spiralis,  Rissoella.  A.  Adams  (1860).  "Sado;  30  fathoms" 
[Japan]. 

sulcosa,  Phasianella.  Mighels  (1843).  "Casco  Bay  [Mame]. 
Wrongly  referred  by  Gould  (1870)  to  Rissoella.  An  "Odostomia" 
(Pyramidellidae)  ;  see  Johnson    (1915) . 

tenuis,  Jeffreysia.  Turton   (1932)  .  "Port  Alfred,  South  Africa." 

translucens,  ? Jeffreysia.  Carpenter  (1866).  Carpenter  (1864), 
nom.  nudum.  "S.  Diego"  [California].  Referred  by  Bartsch  (1920) 
to  "Syncera"    (=Assiminea,  Assimineidae) . 

tumens,  Jeffreysia.  Carpenter  (1856).  "Mazatlan"  [W.  coast 
Mexico].  Referred  by  Carpenter  (1866)  to  "Cythna"  (PStilifer- 
idae) .  Retained  by  Bartsch  (1920)  in  Rissoella.  See  also  Baker, 
Hanna  &  Strong   (1930) . 

turgidula,  Rissoella.  A.  Adams  (1860).  "Korea  Strait;  46  fath- 
oms." 

vesicalis,  Rissoella.  A.  Adams  (1860) .  "Sado;  30  fathoms" 
[Japan]. 

vitrina,  Rissoella.  A.  Adams  (1860).  "Tabu-Sima  [Tobi  Shima]; 

25  fathoms"  [Japan]. 

wilfredi,  Jeffreysia.  Gatliff  &  Gabriel  (1911).  "Ocean  beach, 
near  Point  Nepean"  [Victoria,  Australia].  Referred  by  Iredale 
(1912)    to  Heterorissoa    (misspelled  'wilfridi') . 

zebra,  Rissoella.  Thiele  (1925) .  "Deutsch-Ostafrika  .  .  .  Dar- 
essalam"   [Tanganyika]. 

References 
Adams,  A.  1860.  Ann.  Mag.  Nat.  Hist.,    (S)  6:  332,  333. 
Adams,  J.  1797.  Trans.  Linn.  Soc.  [London],  3:  65. 
Alder,  J.  1844.  Ann.  Mag.  Nat.  Hist.,  (1)  13:  325,  326,  pi.  8,  figs. 

1-4.  1848.  Trans.  Tyneside  Nat.  Field  Club,  1:  149.  1851a.  Ann. 

Mag.  Nat.  Hist.,   (2)7:  193-196.  1851b.  Ibid.,  pp.  460-465. 
Baker,  F.,  G.  D.  Hanna  Sc  A.  M.  Strong,   1930.  Proc.  California 

Acad.  Sci.,    (i)  19:  36-38,  fig.  3,  pi.  1,  figs.  10,  13,  16. 
Bartsch,  P.  1909.  Proc.  Boston  Soc.  Nat.  Hist.,  34:  109.  1915.  Bull. 

U.S.  Natl.  Mus.,  91:  134,  135,  225.  1920.  Proc.  U.S.  Natl.  Mus., 

^8:  159-164,  175,  176,  pi.  12,  figs.  1-3,  6,  7,  9.  1927.  Ibid.  70(11)  : 

31,  pi.  4,  fig.  2. 
Brown,  T.  1844.  Illustr.  Rec.  Conchology  Gt.  Britain  and  Ireland, 

ed.  2,  p.  13,  pi.  9,  fig.  37. 
Carpenter,  P.  P.   1856.  Cat.   Mazatlan  Shells  British   Mus.,  pp. 


July,  1961  NAUTILUS  25 

361-363.  1864.  Rept.  British  Assoc.  Adv.  Sci.  "1863,"  pp.  613, 

657.  1866.  Proc.  California  Acad.  Sci.,   (1)   3:  219. 
Clark,  W.  1850.  Ann.  Mag.  Nat.  Hist.,   (2)  6:  454,  455,  459.  1851. 

Ibid.,    (2)    7:   292-297.   1855.  British  Mar.  Test  Moll.,  pp.  7, 

387-395. 
Crosse,  H.  1879.  Jour.  Conchyl.,  27:  55. 
Dunker,  G.   1882.  Index  Moll.  Mar.  Japonici,  p.   117. 
Folin,  L.  de.  1869.  Fonds  de  la  Mer,  1:  134,  pi.  20,  fig.  6. 
Forbes,  E.  &  S.  Hanley.  1850.  Hist.  British  Moll.,  3:  151-155,  pi. 

J.}.,  figs,  1,  2,  pi.  76,  figs.  1,  3,  4.  1852.  Ibid.,  4:  267-269,  pi.  133, 

figs.  5,  6.  1853.  Ibid.,  pi.  M.M.,  fig  2. 
Fretter,  V.   1948.  Jour.  Mar.  Biol.  Assoc.  U.K.,    (2)27:   597-632, 

figs.  1-6,  pi.  4.  1956.  Proc.  Zool.  Soc.  London,  126:   380.   1954 

(&  A.  Graham) .  Jour.  Mar.  Biol.  Assoc.  U.K.,    (2)  33:  577-583. 
Friele,  H.  1876.  Forh.  Vid.-Selsk.  Christiania,  "1875,"  p.  61,  pi.  1, 

fig.  6. 
Gatliff,  J.  H.  &  C.  J.  Gabriel.    1911.   Proc.  Roy.   Soc.   Victoria, 

{2)24:  188,  189,  pi.  46,  fig.  3. 
Gould,  A.  A.  1870.  Rept.  Invert.  Massachusetts,  ed.  2,  pp.  296,  297, 

figs.  564,  565. 
Gray,  J.  E.  1847.  Proc.  Zool.  Soc.  London,  15:  159. 
Gray,  M.  E.  1850.  Figs.  Moll.  Anim.,  4:  86. 
Iredale,  T.  1912   [Oct.  30].  Proc.  Malac.  Soc.  London,  10:  221, 

222,  1  fig.  1915.  Ibid.,  11:  332. 
Jeffreys,  J.  G.  1848.  Ann.  Mag.  Nat.  Hist.,   (2)2:  351.  1851.  Ibid., 

(2)7:  465-469,  pi.  15.  1856.  Ibid.,   (2)17:   184,  pi.  2,  figs.  8,  9. 

1867.  British  Conch.,  4:  58-63,  106,  107,  pi.  1,  fig.  3. 
Johnson,  C.  W.  1915.  Occas.  Papers  Boston  Soc.  Nat.  Hist.,  7:  99. 
Lebour,  M.  V.  1936.  Jour.  Mar.  Biol.  Assoc.  U.K.,    (2)20:  552, 

pi.  1,  fig.  17. 
McGinty,  T.  L.  1948.  Mollusca  (Paul  H.  Reed,  Tavares,  Florida) , 

2:  63.  Mimeogr. 
Mighels,  J.  W.  1843.  Boston  Jour.  Nat.  Hist.,  4:  348,  pi.  16,  fig.  4. 
Monterosato,  T.  A.  di.  1878.  Gior.  Sci.  Nat.  Econ.  Palermo,  13: 

87,  91. 
Rehder,  H.  A.  1943a.  Proc.  U.S.  Natl.  Mus.,  93:  194,  pi.  20,  fig.  7. 

1943b.  Naut.,  57:  33. 
Robertson,  R.  1960.  American  Malac.  Union  Ann.  Repts.  "1959," 

pp.  22,  23.  In  press.  Collecting  minute  mollusks  which  live  in 

marine  algae.  In  How  to  collect  shells.  Publ.  American  Malac. 

Union,  ed.  2. 
Sars,  G.  O.   1878.  Moll.  Reg.  Arcticae  Norvegiae,  pp.  206,  347, 

pi.  10,  fig.  13,  pi.  34,  fig.  8,  pi.  vi,  fig.  16. 
Sars,  M.   1859.  Forh.  Vid.-Selsk.  Christiania,  "1858,"  p.  85. 
Smith,  A.  G.  &  M.  Gordon,  Jr.  1948.  Proc.  California  Acad.  Sci., 

(4)26:  224,  225,  pi.  3,  fig.  15. 
Smith,  E.  A.  1892.  Ann.  Mag.  Nat.  Hist.,  (6)  10:  130,  pi.  12,  fig.  7. 
Sowerby,  G.  B.  1892.  Mar.  Shells  South  Africa,  p.  38,  pi.  2,  fig.  41. 


26  NAUTILUS  Vol.   75    (1) 

1897.  Appendix  Mar.  Shells  South  Africa,  p.  17,  pi.  6,  fig.  15. 
Stimpson,  W.   1851a.  Proc.  Boston  Soc.  Nat.  Hist.,  4:   14.  1851b. 

Shells  New  England,  p.  34,  pi.  1,  fig.  1. 
Strong,  A.  M.  1938.  Proc.  California  Acad.  Sci.,    (4)25:  211,  212, 

pi.  15,  fig.  5 
Thiele,  J.  1912  [Aug.].  Deutsche  Sudpolar-Exped.  1901-1903,  13, 

Zool.  5:  239,  240,  260,  272,  276-278,  fig.  13,  pi.  15,  fig.  5,  pi.  16, 

figs.  3,  4,  pi.  19,  figs.  15,  21.  1925.  Wiss.  Erg.  deutschen  Tiefsee- 

Exped.  "Valdivia,"  17:  87,  91,  92,  363,  pi.  16,  figs.  13-15,  pi.  19, 

figs.  13-15.  1929.  Handb.  syst.  Weichtierkunde,  /:  178,  179,  figs. 

162,  163. 
Thompson,  W.   1844.  Rept.  British  Assoc.  Adv.  Sci.,  "1843,"  p. 

256. 
Turton,  W.  H.  1932.  Mar.  Shells  Port  Alfred  S.  Africa,  pp.  151- 

153,  pi.  34,  figs.  1080,  1083-1087. 
Velain,  C.  1876.  C.  R.  Acad.  Sci.  [Paris],  83:  285.  1877.  Arch.  Zool. 

Exper.  Gen.,  6:  115,  116,  144,  pi.  3,  figs.  18-20. 
Watson,  R.  B.  1880.  Jour.  Linn.  Soc.  [London],  Zool.,  15:  99. 

1886.  Rept.  Voy.  Challenger.  Zool.,  15:  584,  701,  pi.  43,  fig.  5. 
Weinkauff,  H.  C.  1868.  Conch.  Mittelmeeres,  2:   275. 
Winckworth,  R.  1932.  Jour.  Conchol.,  19:  217,  223. 

EIGHT  NEW  SPECIES  OF  LAND  SNAILS  FROM 
THE  SOUTHERN  UNITED  STATES 

By  LESLIE  HUBRICHT 

PoLYGYRA  GRACILIS,  new  species.  PI.  4,  figs.  N  &  O. 

Shell  pale  brown,  base  lighter,  translucent,  shining,  of  5  to  5.5 
whorls.  Spire  slightly  convex,  of  slowly  increasing  whorls,  the  last 
shortly  descending  in  front;  periphery  above  the  middle,  rounded. 
First  whorl  smooth,  following  whorls  becoming  inaeasingly  stri- 
ate; last  two  whorls  weakly  ribbed  above,  base  smooth  with 
growth  lines  only,  or  with  weak  ribs  behind  the  lip.  Umbilicus 
well-like,  expanding  in  the  last  whorl  to  about  4  times  its  earlier 
diameter,  contained  about  3.5  times  in  the  diameter  of  the  shell; 
last  whorl  with  a  shallow  furrow  parallel  to  the  umbilical  suture. 
Aperture  very  oblique,  deeply  furrowed  behind,  outer  and  basal 
margins  of  lip  strongly  reflected,  thickened  within,  its  inner  edge 
bearing  two  teeth;  one  on  the  basal  margin  and  one  on  the  outer 
margin.  Ends  of  the  lip  joined  by  a  parietal  callus  bearing  a 
V-shaped  tooth.  A  short  distance  within  there  is  a  callous  tubercle 
on  the  columellar  axis. 


Height 

Diameter 

Whorls 

H./D. 

3.9  mm. 

8.3  mm. 

5.5 

0.47  Holotype. 

4.1  mm. 

9.1  mm. 

5.5 

0.45  Paratype. 

3.6  mm. 

7.8  mm. 

5.0 

0.46  Paratype. 

3.2  mm. 

6.7  mm. 

5.0 

0.48  Medina  River  drift 

2.7  mm. 

6.7  mm. 

5.0 

0.40  Nueces  River  drift. 

NAUTILUS  75    (1) 


PLATE  3 


Figs.  1  &  2,  Xotia  tridcns.  1,  internal  view  of  pallial  complex.  2.  penis  and 
accessories;  optical  longitudinal  .section  of  flattened  mount  in  glycerin  jelly; 
semi-diagrammatic.  Figs.  3  &  4,  Cerion  uva.  3,  penis,  cut  open  and  pinned 
out.  4,  terminal  genitalia,  dissected  apart.  .Scales  represent  1  mm.  Drawn  with 
aid  of  camera  lucida. 


NAUTILUS  75    (I) 


PUATF  4 


Holotypcs:  A-C,  Succinca  solaslra  Hubricht.  D  &  E,  .S'.  urbana  Hubricht. 
F-H,  Calinella  lexana  Hubricht.  I-K,  C.  pugilntor  Hubricht.  L  &  M,  Pol\g\ra 
litliira  Hubricht.  N  &  O,  P.  gracilis  Hubricht.  P  &  Q,  Stciiotreiua  calvescens 
Hubricht.  Pliotographs  by  John  B.  Burch,  University  of  Michigan. 


July,  1961  NAUTILUS  27 

Distribution. — Texas:  Comal  Co.:  Guadalupe  River  bluff,  2 
miles  south  of  Sattler,  holotype  205894  and  paratypes  205893, 
University  of  Michigan  Museum  of  Zoology,  other  paratypes 
14834,  collection  of  the  author;  drift,  Guadalupe  River,  New 
Braunfels.  Kerr  Co.:  12  miles  southwest  of  Kerrville;  river  bluff, 
5  miles  southwest  of  Hunt.  Bandera  Co.:  drift,  Medina  River,  4.7 
miles  northwest  of  Medina.  Medina  Co.:  Hondo.  Real  Co.:  Frio 
River  bluff,  5.5  miles  north  of  Leakey.  Uvalde  Co.:  drift,  Nueces 
River,  7  miles  west  of  Uvalde;  drift,  Nueces  River,  1.5  miles  south 
of  Laguna;  Garner  State  Park. 

Polygyra  gracilis  is  a  species  of  river  bluffs  and  ravines  in  the 
Edwards  Plateau  of  Texas.  It  is  most  closely  related  to  Polygyra 
mooreana  (W.  G.  Binney) ,  which  is  found  in  the  same  region.  It 
differs  in  its  more  slender  whorls,  more  depressed  shape,  and  more 
glossy  and  translucent  shell.  Of  the  specimens  measured  P.  gracilis 
had  a  height  diameter  index  below  0.50  with  an  average  of  0.45, 
while  in  P.  mooreana  the  measured  shells  were  all  above  0.50  with 
an  average  of  0.57.  The  lip  teeth  are  usually  smaller  and  more 
widely  spaced  and  are  not  as  laterally  compressed.  The  parietal 
tooth  usually  does  not  have  the  distinct  lower  angle  found  in  P. 
mooreana,  being  more  rounded.  Despite  its  close  relationship  to 
P.  mooreana,  the  two  species  could  be  readily  separated  in  river 
drift  material. 
Polygyra  tamaulipasensis  Lea. 

Polygyra  texasiana  texasensis  Pilsbry,  1940,  Land  Moll.  N. 
Amer.  /,  p.  619,  fig.  394g,  h. 

Polygyra  texasensis  Pilsbry,  Pilsbry  &  Hubricht,  1956,  Naut. 
69:  94. 

Polygyra  texasiana  tamaulipasensis  Lea,  Pilsbry  &  Hubricht, 
1956.  Naut.  69:  95,  PI.  5,  figs.,  1,1a. 

This  species  has  been  considered  to  be  a  subspecies  of  Polygyra 
texasiana  (Moricand) ,  but  is  quite  distinct.  I  have  seen  no  inter- 
gradation  in  the  beach  drift  material,  nor  in  any  material  collec- 
ted in  west  Texas.  It  differs  from  P.  texasiana  in  having  a  more 
depressed  shell  with  narrower  whorls  and  a  larger  umbilicus.  The 
shell  is  never  banded.  The  parietal  tooth  is  longer,  higher  and 
straighter,  the  outer  end  extending  a  little  beyond  the  lower  end 
of  the  lip.  The  outer  end  of  the  upper  branch  is  enlarged  into  a 
low  tooth.  The  animal  is  pale  gray  rather  than  brownish-yellow. 
It  lives  in  rock  piles  and  under  prostrate  yuccas  on  the  tops  of 
the  mesas,  while  P.  texasiana  in  west  Texas  lives  in  low  ground 
near  streams  and  ponds. 


28  NAUTILUS  Vol.   75    (1) 

Polygyra  texasiana  is  quite  variable  in  its  sculpture.  Those 
from  near  the  coast  are  often  strongly  ribbed  above  and  below. 
Away  from  the  coast  the  ribbing  on  the  base  disappears  and  they 
are  only  ribbed  above.  Farther  west  the  upper  surface  becomes 
smooth.  Within  the  range  of  P.  tamaulipasensis,  P.  texasiana  can 
not  be  distinguished  by  the  sculpture.  The  presence  of  the  tooth 
on  the  end  of  the  upper  branch  of  the  parietal  tooth  will  always 
separate  P.  tamaulipasensis  from  P.  texasiana. 

P.  tamaulipasensis  is  found  living  from  near  Junction,  Kimble 
Co.,  west  to  Alpine,  Brewster  Co.  and  north  to  Colorado  City, 
Mitchell  Co.,  Texas.  The  record  from  Lyford,  Cameron  Co., 
Texas  is  based  on  P.  scintilla  Pilsbry  &  Hubricht.  The  record 
from  near  Roswell,  New  Mexico  is  based  on  a  smooth  P.  texasiana. 
Polygyra  lithica,  new  species.  PI.  4,  figs.  L  &  M 

Shell  depressed,  pale  brown,  of  5  to  6  whorls.  Spire  weakly  con- 
vex, of  slowly  increasing  whorls,  periphery  rounded.  Embryonic 
whorls  smooth,  later  whorls  becoming  increasingly  striate,  last 
two  whorls  weakly  rib-striate,  rib-striae  becoming  much  weaker 
below  the  periphery,  base  with  distinct  spiral  striae.  Umbilicus 
deep,  well-like,  expanding  in  the  last  whorl  to  about  one-third 
the  diameter  of  the  shell.  Aperture  oblique,  deeply  furrowed 
behind,  lip,  reflected  and  strongly  thickened,  with  two  teeth  on 
the  inner  margin,  the  basal  tooth  laterally  compressed,  not  im- 
mersed; the  outer  tooth  only  slightly  immersed,  of  about  the  same 
size  as  the  basal  tooth.  Ends  of  the  lip  joined  by  a  heavy  parietal 
callus  bearing  a  heavy  tooth  which  is  rounded  in  front  view.  A 
short  distance  within  there  is  a  callous  tubercle  on  the  columellar 
axis. 


Height 

Diameter 

Whorls 

H./D. 

3.2  mm. 

7.4  mm. 

5.8 

0.43  Holotype. 

3.7  mm. 

8.1  mm. 

6.0 

0.46  Paratype. 

3.3  mm. 

7.0  mm. 

5.5 

0.49  Paratype. 

2.7  mm. 

7.9  mm. 

5.2 

0.46  Rushing. 

Distribution. — Arkansas:  Stone  Co.:  upland  oak-hickory  woods, 
6  miles  east  of  Mountain  View,  holotype  205896  and  paratypes 
205895  U.M.M.Z.,  other  paratypes  15917,  collection  of  the  author: 
4  miles  northwest  of  Allison;  2  miles  northeast  of  Rushing. 

Polygyra  lithica  is  related  to  P.  dorfeuilliana  Lea,  differing  in 
the  teeth.  The  parietal  tooth  is  lower  and  is  rounded  rather  than 
squarish.  The  lip  teeth  are  smaller  and  are  not  deeply  immersed. 
It  is  apparently  a  species  of  quite  limited  range,  being  known  only 
from  a  small  area  in  north-central  Arkansas. 
Stenotrema  calvescens,  new  species.  PI.  4,  figs.  P  &:  Q. 

Shell  buffy  brown  to  bister,  globose-conic,  imperforate,  of  about 


Height 

Diameter 

6.5  mm. 

9.7  mm. 

5.6  mm. 

8.1  mm. 

6.0  mm. 

9.3  mm. 

6.7  mm. 

10.2  mm. 

July,   I9()l  NAUTILUS  29 

5.5  whorls;  periphery  above  the  middle,  rounded  or  slightly  sub- 
angular  in  [rent  of  the  aperture.  Embryonic  whorls  sculptured 
with  radially  elongated  granules,  later  whorls  with  fine  radial 
lines,  last  three  whorls  with  numerous  soft  deciduous  liairs. 
Aperture  naiTow,  very  pale  brown,  outer  edge  of  the  basal  lip 
wholly  adnate,  inner  edge  with  a  distinct  U-shaped  central  notch; 
interdenticular  sinus  rounded,  moderately  deep;  there  is  no  tooth 
within  the  outer  arc  of  the  lip.  Parietal  tooth  rather  low  and 
slender,  curved,  leaning  towards  the  basal  lip.  There  is  no  buttress 
connecting  the  parietal  tooth  to  the  end  of  the  outer  lip.  Edge  of 
the  parietal  callus  extending  well  beyond  the  parietal  tooth. 
Fulcrum  thin  with  slightly  convex  edge. 

Whorls     H./D. 
5.7  0.67  Holotype. 

5.2  0.69  Paratype. 

5.6  0.65  Paratype. 

5.5  0.63  Hixon. 

Distribution. — Tetinessee:  Marion  Co.:  Cumberland  Mtn.,  1 
mile  east  of  Monteagle,  holotype  205898  and  paratype  205897 
U.M.M.Z.,  other  paratypes  A9690,  collection  of  the  author;  Cimi- 
berland  Mtn.,  2.4  miles  southeast  of  Monteagle;  West  Fork  of 
Pr)'or  Cove,  2  miles  northeast  of  Jasper;  under  pine  logs,  summit 
of  Suck  Creek  Mtn.,  15  miles  west  of  Chattanooga  (W.  F.  Shay, 
coll.)  Hamilton  Co.:  vacant  lot,  Thrasher  Pike,  Hixon  (James 
R.  Hood,  coll.)  ;  near  Silver  Creek,  300  Signal  Mtn.  Road,  North 
Chattanooga. 

In  the  West  Fork  of  Pryor  Cove,  Stenotrema  calvescens  was 
found  associated  with  5.  stenotrema  (Pfr.)  from  which  it  could 
be  readily  separated  by  its  smaller  size,  less  reddish  color,  finer, 
more  numerous  hairs,  and  by  the  absence  of  a  buttress.  The 
hairs  are  found  only  in  young  and  recently  matured  shells.  They 
are  lost  in  old  shells.  Scrubbing  of  the  shells  in  cleaning  will 
remove  most  of  the  hairs.  Stenotrema  calvescens  is  apparently 
most  closely  related  to  S.  florida  Pilsbry,  from  which  it  differs  in 
its  smaller  size,  larger  lip  notch,  and  finer,  more  numerous  hairs. 
Praticolella  berlandieriana  campi  Clapp  &  Ferriss. 

Praticolella  campi  Clapp  &  Ferriss,  1919,  Naut.  32:  78,  PL  6, 
figs.  1-4. 

Praticolella  berlandieriana  taeniata  Pilsbry,  1940,  Land  Moll. 
N.  Amer.  /,  p.  696,  figs.  427g,  h. 

Praticolella  campi  is  a  winter  resting  stage  of  the  form  later 
described  as  P.  berlandieriana  taeniata  Pilsbry.  It  is  found  only 
in  the  winter  and  is  always  found  associated  with  adult  shells.  It 
always  agrees  in  color  pattern  with  the  adult  shells  found  with 
it.  Adult  shells  sometimes  show  an  iiTegularity  in  the  spire  corres- 


30  NAUTILUS  Vol.    75     (1) 

ponding  to  this  resting  stage.  Animals  dissected  by  both  Dr. 
Pilsbry  and  myself  were  found  to  be  sexually  undeveloped.  It  is 
a  stage  in  which  there  is  a  high  mortality,  as  they  are  common  in 
river  drift  and  fossil  deposits  while  other  immature  stages  are 
rare.  The  name  campi  will  have  to  replace  taeniata  for  the 
Praticolella  berlandieriana  X  P.  griseola  hybrid  populations  of 
southern  Texas  and  northeastern  Mexico. 

Similar  resting  stages  are  also  found  in  Praticolella  griseola 
(Pfeiffer)   and  Praticolella  pachyloma   ('Menke'  Pfeiffer). 
SucciNEA  soLASTRA,  new  Species.  PI.  4,  figs.  A-C;  text-fig.  1,  B. 

Shell  pale  buff,  translucent,  shining,  with  about  3.5  whorls, 
thin  but  firm,  elongate-ovate  in  shape,  sculpture  of  unevenly 
spaced  growth  wrinkles.  Spire  acute,  moderately  long,  sutures 
well  marked,  periphery  somewhat  flattened.  Aperture  ovate, 
occupying  about  sixty  percent  of  the  length  of  the  shell,  outer 
and  basal  margins  well  rounded.  Columella  nearly  straight. 

Animal  pale,  mantle  pale  with  scattered  brownish  spots.  Herma- 
phrodite duct  well  pigmented.  Talon  moderately  pigmented,  club- 
shaped  with  two  terminal  lobes  and  a  longitudinal  groove  down 
the  center,  giving  it  the  appearance  of  being  bifurcate.  Prostrate 
gland  large,  irregularly  oval,  compressed  by  surrounding  organs, 
unpigmented.  Vas  deferens  not  very  long,  entering  the  penis  a 
little  below  the  apex.  Vagina  and  atrium  very  short.  Vaginal 
retractor  muscle  long  and  stout.  Penis  rather  stout,  with  a  con- 
striction about  two-thirds  the  way  up,  above  this  it  is  somewhat 
inflated  and  free  from  the  sheath.  Penial  retractor  muscle  thick 
and  broad.  Spermatheca  globose,  the  duct  slender  throughout 
its  length. 


Height 

Diam. 

Ap.  H. 

Ap.  W. 

Whorls 

15.3  mm. 

7.3  mm. 

9.7  mm. 

5.7  mm. 

3.5  Holotype. 

16,0  mm. 

7.3  mm. 

9.3  mm. 

5.7  mm. 

3.5.  Paratype. 

16,0  mm. 

8.7  mm. 

9.7  mm. 

6.3  mm. 

3.5  Paratype. 

1 1 .7  mm. 

6.0  mm. 

7.0  mm. 

4.0  mm. 

3.5  Combes,  Texas. 

9.5  mm. 

4.7  mm. 

5.7  mm. 

3.2  mm. 

3.5  Combes,  Texas. 

Distribution. — Texas:  Terrell  Co.:  Sanderson;  5  miles  north  of 
Sanderson;  side  of  mesa,  5  miles  southeast  of  Sanderson.  Val- 
Verde  Co.;  Shumla.  Uvalde  Co.;  Uvalde,  Dimmit  Co.;  just 
north  of  Caterina;  near  culvert,  6.5  miles  south  of  Caterina; 
near  culvert,  2.5  miles  northwest  of  Canrizo  Springs.  Frio  Co.: 
culvert,  5.5  miles  southwest  of  Moore,  Bexar  Co.:  culvert,  on 
US-281,  9.5  miles  north  of  San  Antonio.  Live  Oak  Co.:  near 
Ramirena  Creek,  13  miles  south  of  George  West.  Bee  Co.:  culvert, 
2  miles  south  of  Pettus  Jim  Wells  Co.:  Premont.  Starr  Co.:  cul- 
vert, 6  miles  northwest  of  Roma;  1,5  miles  southeast  of  Rio 
Grande  City.  Hidalgo  Co.:  near  Bentzen  Rio  Grande  Valley  State 
Park,  holotype  205900  and  paratype  205899  U.M.M.Z.,  other  para- 


July,   1961 


NAUTILUS 


31 


types  14322,  collection  ot  the  author.  Willacy  Co.:  roadside,  0.7 
miles  north  of  Santa  Monica,  Cameron  Co.:  6.3  miles  southwest  of 
Boca  Chica;  0.5  mile  northwest  of  Combes;  dump,  3.4  miles  east 
of  La  Paloma.  Mexico:  Tamaulipas:  1  mile  southeast  of  Ciudad 
Mier;  San  Fernando.  Nuevo  Leon:  near  arroyo,  4  miles  north- 
east of  Pesqueria  Chica;  32.8  miles  east-northeast  of  General 
Bravo. 


Figure  1,  genitalia:  A  &  C,  Succinea  urbana  Hubricht.  B,  S.  solastra  Hu- 
bricht.  D  &  E,  Catinella  pugilator  Hubricht.  F-H,  C.  texana  Hubricht.  (ap., 
appendix;  at,  atrium;  h.d.,  hermaphrodite  duct;  p.,  penis;  p.g.  prostate;  v., 
vagina;  s.,  spermatheca;  v.d.,  vas  deferens;  t.,  talon.) 

Succinea  solastra  is  frequently  found  associated  with  Succinea 
luteola  Gould,  from  which  it  may  be  readily  separated  by  the 
anatomical  differences,  its  smaller  size,  more  acute  spire,  and 
translucent  shell.  The  shell  is  usually  covered  with  dirt  while 
that  of  S.  luteola  is  usually  clean.  It  is  most  closely  related  to 


32  NAUTILUS  Vol.   75    (1) 

Succinea  grosvenori  which  differs  in  its  more  globose,  more 
opaque  shell;  and  dark  mantle.  The  genitalia  are  not  distinguish- 
able from  those  of  S.  grosvenori  Lea. 

An  examination  of  many  lots  of  anatomical  material  of  suc- 
cineids,  collected  over  the  past  several  years  by  the  author,  has 
shown  that  the  answer  to  the  species  problem  in  Succinea  will 
not  be  found  in  the  study  of  the  genitalia.  Species  with  readily 
distinguishable  shells  have  very  similar  genitalia.  Thus,  Succinea 
pronophobus  Pilsbry,  S.  wilsoni  Lea,  and  S.  urbana,  new  species, 
are  not  separable  by  the  genitalia.  Succinea  campestris  Say,  S. 
luteola  Gould,  and  5.  floridana  Pilsbry,  form  another  similar 
group,  (The  difference  in  the  thickness  of  the  penial  sheath  re- 
ported by  Pilsbry  does  not  exist  in  my  material.  Possibly  the 
thickness  may  vary  with  the  age  of  the  animal.)  The  genitalia 
will  be  of  value  in  the  recognition  of  species  groups,  but  shell 
characters  will  have  to  be  used  in  many  cases  for  species  identifi- 
cation. On  the  other  hand,  in  the  genus  Catinella  very  good  spe- 
cific differences  are  to  be  found  in  the  anatomy,  especially  in  the 
shape  of  the  penis.  In  this  genus  this  will  probably  prove  to  be 
a  more  reliable  character  than  the  shape  of  the  shell. 
Succinea  urbana,  new  species.  PL  4,  figs.  D  &  E;  Text-fig.  1,  A  &  C. 

Shell  pale  golden-brown  in  color,  subtranslucent,  dull,  with 
about  3.5  whorls,  elongate-ovate,  rather  thin,  sculpture  of  un- 
evenly spaced  growth  wrinkles.  Spire  acute,  long,  sutures  only 
moderately  impressed;  whorls  moderately  convex.  Aperture  ovate, 
rather  small,  occupying  about  55%  of  the  length  of  the  shell, 
outer  and  basal  margins  well  rounded.  Columella  nearly  straight. 

Color  pattern  variable,  some  with  mantle  grayish,  others  with 
a  bold  pattern  of  black  and  pale  grayish-brown,  margin  of  foot 
gray.  Hermaphrodite  duct  moderately  pigmented.  Talon  mod- 
erately pigmented,  club-shaped,  two  lobed  and  with  a  median 
groove.  Prostate  gland  large,  almost  square  due  to  pressure  from 
the  other  organs,  moderately  pigmented.  Vas  deferens  not  very 
long,  entering  the  penis  below  the  terminal  loop,  somewhat 
expanded  below  where  it  joins  the  penis.  Penis  moderately 
slender,  with  an  inflated  loop  at  the  upper  end.  Penial  retractor 
connected  to  the  penis  below  the  terminal  loop,  Atrium  and 
vagina  very  short.  Spermatheca  large,  globose,  duct  slender. 

Length        Diam.        Apt.  L.      Ap.  W 

11.1mm.     6.0  mm.     6.0  mm.     4.0  mm.         Holotype. 

10.9  mm.     5.7  mm.     6.0  mm.     4.0  mm.         Paratype. 

11.7  mm.     6.3  mm.     6.6  mm.     4.2  mm.         Paratype. 

Type  Locality.  Alabama:  Montgomery  Co.:  vacant  lot,  Dudley 
St.,  and  Fairview  Ave.,  Montgomery,  holotype  205901  and  para- 


July,  1961  NAUTILUS  33 

types  205902  U.M.M.Z.,  other  paratypes  23392,  collection  oi  the 
author. 

Succinea  urbana  appears  to  be  most  closely  related  to  S.  wilsoni 
Lea,  which  it  resembles  in  its  genitalia.  It  differs  in  its  thicker 
shell  with  smaller  aperture  and  more  obese  spire. 
(To  be  concluded) 

PUERTO  RICAN  PUPILLIDS  AND  CLAUSILIOIDS 

By  H.  BURRINGTON  BAKER 

The  symbols  used  for  Puerto  Rican  localities  were  explained 
recently  (1961)  .^  Very  few  of  the  smallest  species,  especially  the 
leaf  arboreal  ones,  which  may  mature  in  a  few  months,  were 
obtained,  probably  because  these  may  be  much  reduced  in  num- 
bers or  represented  only  by  eggs  and/or  young  during  the  dry 
periods,  because  cultivated  areas  were  avoided,  and  since  no 
humus  was  sifted  or  saved.  All  the  pupillids  are  distributed 
widely,  probably  by  human  agencies. 

Pupoidcs  (s.s.)  nitidulus  (Pfeiffer) .  Enl;  lowland  species, 
mapped  by  van  der  Schalie,  1948:38,  from  Ps,  Wn  &:  Ws,  and 
reported  by  Dall  &  Simpson,  1901:372,  from  Fajardo   (Ee) . 

Gastrocopta  (s.s.)  pellucida   (Pfeiffer) .  Lowlands,  Es2,  Ps2. 

G,  (s.s.)  servilis   (Gould).  Lowlands,  Es2,  Ps2,  Ws. 

Pupisoma  (Ptychopatula)  dioscoricola  (C.  B.  Adams).  Pnl 
(young) .  Two  other  leaf  arboreal  Nesopupinae,  Bothriopupa 
tenuidens  (C.  B.  Adams)  and  P.  minns  Pilsbry,  reported  by  van 
der  Schalie,  1948:36,  39. 

Vertigo  (s.s.)  ovata  Say,  subspecies?  So  reported  by  Pilsbry, 
1919  (3a):  87,  pi.  13,  fig.  16,  from  Humacao  (Es)  ;  since  he  also 
stated  "there  is  nothing  to  differentiate  it,"  why  not  drop  Pupa 
hexodon  Adams,  from  Jamaica,  as  obsolete? 

Cerion  (Strophiops)  striatella  (Guerin)  +  C.  crassilabris  (Sow- 
erby);  see  Pilsbry,  1943  (5)  :34,  but  striatella  retains  its  original 
ending.  Psl,  under  bunch  grass;  Ps2,  roosting  on  tree  trunks; 
apparently  limited  to  southwest  limestone  rim.  Shells  from  19.0 
mm.  long  with  814  whorls  (Ps2)  to  31.1  with  11  (Psl),  so  C. 
monaense  Clench,  1951:274,  not  always  smaller,  but  does  have 
stronger  and  more  widely  spaced  costae  than  in  any  P.  R.  shells 
seen.  Color  varying  from  whitish,  through  those  with  brown 
flammules,  to  almost  uniformly  brownish. 

Pallial  complex  of  C.  striatella  (Ps2)  and  of  C.  uva  (L)  from 
just  south  of  Willemstad,  Curacao,  fundamentally  similar  to  that 
of  C.  incanum  [Pilsbry,  1946(11)  :160,  fig.  76,  G]  but  much  more 

1  In  the  remarks  about  Cepolis  boriquenae,  p.  147,  change  Ps4  to  Es4 
(Humacao) . 


34  NAUTILUS  Vol.   75    (1) 

elongate  (less  contracted?)  ;  lung  wall  5  times  as  long  as  kidney  or 
7  times  its  apical  width;  sometimes  with  vague,  whitish  strip 
along  hindgut  (Cf.  Nenin)  especially  towards  pneumostome; 
kidney  about  1 14  length  of  pericardium,  with  broad  base  (much 
as  in  Nenia) ,  without  ureter  but  with  similar  subapical  opening. 
Mantle  collar  with  weak  lappets   (similar  to  those  in  Nenia) . 

Genitalia  of  C.  strintella  very  similar  to  those  of  C.  inconum 
(Pilsbry,  figs.  H  to  K);  only  differences,  most  of  which  might  be 
due  to  contraction  and/or  sexual  stage,  will  be  noted.  Ovotestis 
of  4  lobes  of  alveoli,  imbedded  in  3rd  whorl  of  liver.  Carrefour 
also  without  definite  talon.  Spermatheca  similar  but  without 
diverticulum  in  5  examples  dissected;  stalk  more  swollen  near 
base;  in  2  animals,  containing  spermatophores,  each  of  which  is 
a  fragile,  thin  walled,  very  slender,  horny  tube  (about  as  long  as 
spermatheca  and  its  stalk)  and  with,  for  at  least  part  of  its  length, 
2  thickenings,  that  are  separated  by  about  \\  its  circumference. 
Vagina  similarly  receiving  branches  from  right  ocular  retractor, 
which  does  not  pass  through  penis-vaginal  angle.  Vas  epiphallus 
evidently  involving  most  of  free  loop  and  thus  about  equalling 
spermatophore  in  length;  beginning  as  gradual  enlargement  of 
vas  lumen  near  penis-vaginal  angle;  mainly  with  thin  wall  which 
internally  has  irregularly  transverse,  lobulate  thickenings;  attain- 
ing its  maximum  diameter  in  descending  limb;  tapering  and  with 
wall  becoming  much  thicker  and  more  muscular  (at  expense  of 
lumen)  towards  entrance  into  penis  near  base  of  last.  Penis  and 
largely  atrial  "stimulator"  very  similar;  penial  retractor  arising 
low  on  diaphragm  and  inserting  on  apex  and  down  along  one 
side  for  about  i/l  length  of  caecum. 

Genitalia  of  C.  nva  (pi.  3,  figs.  3,  4)  and  C.  uva  bonaircnsis  also 
similar  to  preceding,  but  with  salient  differences  as  noted.  Sperma- 
theca (S)  extending  to  about  2/^  length  of  spermoviduct;  di- 
verticulum (SD)  present  in  those  examined,  recurved  apically  but 
not  extending  above  aorta;  common  stalk  (SS)  longer  and  more 
slender  but  no  spermatophores  seen  (resting?) ,  about  1 1/^  length 
of  free  oviduct  (UV) .  Long  vas  epiphallus  (E)  swollen  (but  with 
smaller  lumen)  near  opening  (EP)  at  bifurcation  of  large  penial 
pilaster.  Penis  (P)  relatively  longer,  with  only  I/3  its  length  above 
vas  entrance.  Penial  apex  internally  (fig.  3)  with  heavy  longi- 
tudinal folds  but  basal  2/5  with  smooth  wall  except  for  pilaster 
and  strictly  penial  "stimulator"  (PP)  which  is  similar  in  outline 
but  is  simply  a  discontinuous,  elliptic  thickening  (i.e.,  without 
free  tip).  Atrium  relatively  shorter. 

The  differences  in  at  least  the  penis  of  C.  xiva  may  represent 
subgeneric  characteristics  of  Cerion  s.s.,  of  which  it  is  the  only 
species.  On  the  other  hand,  the  presence  or  absence  of  a  sper- 
mathecal  diverticuhmi  may  be  of  little  significance;  Pilsbry  stated 


July,  1961  NAUTILUS  35 

it  was  variable  in  C.  incanum  and  it  evidently  is  a  vestige  of  the 
long,  more  primitive  kind  of  geophile  spermatheca,  in  which  the 
sac  is  imbedded  above  the  aorta.  Also,  the  swelling  (Pilsbry:  fig.  I, 
epi)  of  the  long  vas  epiphallus  near  its  entrance  into  the  penis 
apparently  is  caused  by  the  contraction  of  this  region  with  more 
muscular  wall  but  smaller  lumen. 

Nenia  (s.s.)  tridens  (Schweigger) ,  Usually  roosting  on  tree 
trunks  or  vines  up  to  8  ft.,  but  rarely  on  fallen  leaves  or  on  rocks; 
Er-1,  2,  3,  Es3,  Jn I,  100  to  2400  ft.,  eastern  Puerto  Rico.  Foot  quite 
short  but  broad;  sole  with  2  or  3  coarse  waves  on  middle  zone; 
animal  moves  along  some  distance  and  then  jerks  shell  up  after 
it.  Embryonic  whorls  of  shell  about  2i/^,  teat-shaped,  relatively 
smooth  and  without  major  costulae  but  assuming  at  about  14 
whorl  close,  microscopic,  growth  wrinkles,  which  become  fine, 
punctulate  threadlets  by  end;  2nd  whorl  much  higher  than  first; 
later  whorls  assuming  major  costulae  and  crenulate  suture  very 
quickly;  young  shells  without  apertural  teeth. 

Pallial  complex  (pi.  3,  fig.  1)  fundamentally  similar  to  that  of 
Cerion;  lung  wall  about  5  times  as  long  as  its  apical  width  or  3 
times  length  of  kidney;  similarly  without  much  secondary  vena- 
tion, although  minute,  pigmented  tributaries  are  visible  along 
sides  of  principal  vein  (HV)  ;  kidney  (K)  about  twice  as  long  as 
its  base  or  length  of  pericardium  (H) ,  with  similarly  subapical, 
external  opening  (KX)  and  without  ureter.  [Although,  in  some 
examples,  discontinuous,  whitish  zones  (mucus  or  thickening?) 
are  visible  along  hindgut  (HG)  and  kidney,  these  certainly  are 
nothing  like  the  definite  thickening  along  the  hindgut  in  those 
sigmurethrous  geophiles  (e.g.,  Oleacinidae)  in  which  the  external 
ureteric  opening  is  in  the  apical  corner  of  the  lung.  Possibly  these 
vague  zones  were  why  Wiegmann,  1893:226,  described  Phaedusa 
as  sigmurethrous;  this  apparently  was  copied  by  Thiele  in  his 
"Handbuch."]  Mantle  collar  (MC)  with  moderate  parietal  (left) 
and  small,  separate  palatal   (right)  lappets. 

Genitalia  as  represented  by  P.  Hesse,  1925;  only  additions  and 
differences  will  be  noted.  Ovotestis  (not  unlike  that  of  Clausilia 
s.  s.;  Cf.  Steenberg,  1914)  consisting  of  3-4  conical  groups  of  simple 
and  bifurcate  alveoli;  hermaphroditic  duct  much  swollen  and 
convoluted  for  y^  its  length.  Carrefour  (Steenberg: pf)  with  very 
short  talon  (Steenberg:  vs) .  Sac  of  spermatheca  extending  almost 
to  tip  of  diverticulum,  which  reaches  to  near  apical  end  of  sper- 
moviduct;  spermathecal  stalk  relatively  longer  than  in  Hesse's  fig. 
(apparent  differences  may  be  due  to  preservation  and/or  sexual 
stage) .  Penis  (P.  fig.  2)  thin  walled,  with  several  internal  pilas- 
ters, passing  gradually  (EP)  into  vas  "epiphallus"  (Steenberg: 
canal  deferent)  which  has  thick,  muscular  walls  and  small  lumen 
(Cf.  Steenberg:  fig.  20)  and  looks  more  like  an  ejaculatory  organ 


36  NAUTILUS  Vol.   75    (1) 

than  an  epiohallus;  penial  retractor  (PR)  arising  low  on  diifram 
and  inserting  little  below  middle  of  "epiphallus"  (DE  to  EP)  ; 
base  of  penis  and  terminal  end  of  vas  proper  (up  to  DE)  bound 
by  thin  sheath.  Left  ocular  retractor  in  penis-vaginal  angle  (as 
stated  by  Hesse) . 

Undoubtedly,  the  Clausiliidae  (at  least  Nenia)  ^  and  the  Cerii- 
dae  (Cerionidae  Pilsbry)  are  closely  "related;"  although  the 
traditional  family  distinction  should  be  retained,  they  belong  in 
the  same  superfamily;  its  name  (-oidea  or  -acea)  would  come  from 
the  oldest  family,  Clausiliidae  Moerch,  1864.  Both  belong  in  the 
Mesurethra  (1955)  near  the  pupilloids  (Orthurethra) ,  in  which 
the  pallial  complex  mainly  differs  by  the  proximity  of  the  external 
renal  opening  to  the  pneumostome;  even  in  this  respect,  the  Par- 
tulidae  somewhat  approach  the  Mesurethra. 

Incidentally  Thiele  in  his  "Handbuch"  retained  this  position 
for  the  Clausiliidae  and  it  approximates  the  traditional  location 
of  the  Ceriidae  as  well;  Cerion  uva  (L.)  is  the  type  species  of  Pupa 
Lamarck,  in  which  genus  most  of  the  older  pupillid  species  were 
described,  and  the  obsolete  Pupidae  Fleming,  1818,  is  the  oldest 
name  for  the  family.  In  this  case,  the  resemblances  in  shell  forms 
perhaps  are  not  secondary  convergences.  As  Pilsbry,  1904  (0)  :  176, 
suggested,  the  Megaspiridae  probably  also  belong  near  the  Clau- 
siliidae (and  the  Ceriidae) . 

Added  references   (See  1961) 
1955,  Naut.  ^5:109.  1961,  Naut.  74:  142-149. 
Clench,  William  J.  1951,  J.  de  Conch.  PO: 269-276  &  pi. 
Hesse,  P.  1925,  Proc.  Malac.  Soc,  London  7^:154-155. 
Steenberg,  C.  M.  1914,  Mindeskrift,  f.  J.  Steenstrup,  no.  29:46 

pp.  &  pi. 
Wiegmann,  F.  1893,  ZooL  Erg.  Niederl.  Ost-Indien  2:226. 


JEANNE  SANDERSON  SCHWENGEL,  Sc.D. 
1889  to  1961 

Dr.  Schwengel  was  born  Jeanne  Sanderson,  in  Stockton,  Cali- 
fornia, July  12,  1889.  She  attended  public  schools  at  her  birth- 
place, and  also  St.  Xavier's  Academy,  Chicago,  Illinois.  To  help 


2  Since  the  above  was  written,  through  the  generosity  of  Dr.  F.  E.  Loosjes, 
of  Wageningen-Hoog,  The  Netherlands,  animals  of  Clausilia  bidentata 
(Stroem)  and  Lacinaria  plicata  (Draparnaiid)  have  been  examined.  In  both 
of  these  species,  the  pallial  complex  is  quite  similar  to  that  of  Nenia. 


July,   1961  NAUTILUS  37 

her  with  her  conchological  studies,  she  also  studied  biology  and 
Latin  at  Fordham  University,  New  York  City.  On  January  24, 
1918,  she  was  married  to  Frank  R.  Schwengel,  who  subsequently 
retired  from  the  U.  S.  Army  as  Brigadier  General,  and  became 
Chairman  of  Joseph  E.  Seagram  and  Sons,  Inc.  On  February  17, 
1961,  while  attending  the  annual  meeting  of  the  St.  Petersburg 
Shell  Club,  at  which  she  was  scheduled  to  deliver  an  address,  her 
over-willing  heart  failed  her.  She  died  suddenly,  after  her  return 
to  the  hotel,  where  she  was  staying  temporarily.  She  is  buried  in 
Scarsdale,  where  she  resided  for  many  years.  Her  husband  survives 
her. 

She  did  many  of  her  studies  on  mollusks  at  the  Academy  of 
Natural  Sciences  of  Philadelphia,  where  she  worked  with  Dr. 
Pilsbry,  and  was  appointed  Research  Associate  in  1935.  In  1943, 
the  University  of  Dayton  conferred  on  her  the  honorary  degree 
of  Doctor  of  Science,  in  recognition  of  her  contributions  to  the 
science  of  conchology.  In  1950,  the  Board  of  Overseers  of  Harvard 
College  appointed  her  a  member  of  the  visiting  committee  of  the 
Museum  of  Comparative  Zoology. 

In  addition  to  her  laboratory  studies,  she  was  an  indefatiguable 
collector  in  the  field,  and  helped  in  much  dredging,  especially  off 
the  coasts  of  Florida.  She  assisted  Dr.  Louise  M.  Perry  in  the 
compilation  of  molluscan  records  for  the  latter's  "Marine  shells 
of  southwest  Florida,"  1940,  helped  revise  it,  and  became  co- 
author of  "Marine  shells  of  the  western  coast  of  Florida." 

Dr.  Schwengel  was  elected  Vice-President  of  the  American 
Malacological  Union  in  1950;  became  its  President  in  1951; 
graciously  presided  over  the  eighteenth  annual  meeting  at  Boston 
in  August,  1952;  and  helped  its  progress  as  a  member  of  the 
council  for  the  rest  of  her  life.  As  many  of  the  members  will 
remember,  she  gave  delightful  evening  parties  at  many  of  the 
meetings,  before  and  after. 

She  was  a  member  of  the  following  scientific  associations: 
Academy  of  Natural  Sciences  of  Philadelphia  (since  1935),  Ameri- 
can Malacological  Union  (1936) ,  American  Association  for  the 
Advancement  of  Science  (1940),  Malacological  Society  of  London 
(1947) ,  Paleontological  Research  Institute,  Ithaca,  N.  Y.  (1953) , 
Natural  Science  Foundation,  Philadelphia,  Pa.  (Director  from 
1955  to  1958),  Philadelphia  Shell  Club    (1955),  and  Hawaiian 


38  NAUTILUS  Vol.   75    (1) 

Malacological  Society   (1955) . 

She  also  shared  in  many  other  activities.  She  was  a  member 
of  the  American  Red  Cross  in  Houston,  Texas  (1918) ,  in  Chicago, 
Illinois  (1920-1932)  and  in  Scarsdale,  N.  Y.  (1935-1945)  ;  and  was 
Chairman  of  Production  from  1943  to  1953.  Since  1943,  she  also 
belonged  to  the  Scarsdale  Woman's  Club  and  was  Chairman  of 
the  Garden  Section  from  1954  to  1956. 

Jeanne,  as  she  was  known  to  her  many  friends,  will  be  long 
remembered  for  her  ready  smile  and  happy  presence.  The  name  of 
Dr.  Schwengel  will  be  carried  on  by  her  publications  about  con- 
chology  and  by  her  large  shell  collection,  from  which  she  gave 
many  lots  to  various  institutions,  and  left  the  main  part  to  the 
Museum  of  Comparative  Zoology  at  Harvard  College.  —  Editors. 

Contributions  to  conchology 
1938    (1)  May  13,  Zoological  results  of  the  George  Vanderbilt 

South  Pacific  Expedition,   1937.  Part  I,  —  Galapagos 

Mollusca.  Proc.  Acad.  Nat.  Sci.  Philadelphia  90:]-S,  3  figs. 

[Daphnella  thalia,  Marginella  rosa,  Tralia  x)anderbilti]. 
1938    (2)    July  22.  Note  on  unreported  marine  molluscs  from 

Sanibel,  Florida,  By  Louise  M.  Perry,  J.  S.  S.  &  Ted  Dranga. 

Naut.  52(1) -.27-28. 
1940    (0)    Jan.  28.  [Cerithium  auricoma,  Latirus  cymatiiis;.  See 

1940  (0)  ].  Naut.  55  (3)  :pl.  12,  figs.  6-8a. 
1940    (1)    April  29.  Two  new  Floridan  marine  shells.  Naut.  53 

(4):  109-1 10.  [See  1940(0).] 

1940  (2)  Nov.  2.  New  Mollosca  from  Florida.  Naut.  54  (2): 49-52, 
pi.  3,  figs.  3,  3a,  6-9,  12.  [Marginella  jaspidea,  Crassispira 
phasma,  Fenimorea  halidorema,  Glyphostoma  pilsbryi,  Bella- 
spira  (?)  pentapleura.J 

1941  (1)  Oct.  24.  A  genus  and  family  of  marine  mollusks  new  to 
the  United  States.  Naut.  55  (2)  :  37-40,  pi.  3,  figs.  1-5.  [Lobiger 
pilsbryi.] 

1941  (2)  Oct.  24.  Marginella  hartleyana.  Naut.  55  (2)  :65,  pi.  3, 
figs,  6,  7. 

1942  (1)  May  7.  Living  Miira  florida.  Naut.  55  (4)  :144. 

1942  (2)  July  23.  Some  new  and  interesting  marine  shells  from 
northwest  Florida.  By  J.  S.  S.  k  Thomas  L.  McGinty.  Naut. 
56  (1)  :  13-18,  pi.  3,  figs.  2,  3,  5;  pi.  4,  f.  d.  [Calliostoma  (Eutro- 
chus)  faustum,  C.  fascinans,  Douglassia  benliana,  Trivia 
maltbiana,  Aclis  hypergonia,  Cyclostrema  (Aorotrema)  pon- 
togenes  (n.  subg.)  ] 

1942  (2a)  July  23.  [Pteria  xnnthia,  Phos  adelus,  Tritiaria  vir- 
giniae,  Lamellaria  leucosphaera.]  Naut.  5<5  (l):pl.  3,  figs.  1, 
la,  4,  6-9.  See  1942(3). 


NAUTILUS  75    (1) 


PIAIK  5 


JEANNE  SANDERSON  SCHWENGEL 


July,  1961  NAUTILUS  39 

1942  (3)  Oct.  14.  New  Floridan  marine  mollusks.  Naut.  56(2)  : 
62-66,  pi.  6,  figs.  1-3.  [Terebra  glossema;  see  also  1942  (2a).] 

1943  (1)  Feb.  15.  New  marine  shells  from  Florida.  Naut.  5(5  (3)  : 
75-78,  pi.  7,  figs,  1-7.  [Marginella  denticulata  destina,  M. 
idiochila,  Eubela  mcgintyi,  Drupa  didyma,  Epitonium 
(Cirsostrema)  Unteatum.] 

1943  (2)    July  23.  Diadora  jaumei.  Naut.  57  (1)  :32. 

1944  (1)    Feb.  9.  Smaragdia  viridis  viridemaris.  Naut.  37  (3)  :106. 
1944    (2)    Aug.   17.  A  new  Modiolaria  from  Florida.  By  R.  A. 

McLean  and  J.  S.  S.  Naut.  58  (1)  :  16-17,  pi.  1,  fig.  10.  [Modio- 
laria skomma.] 

1944  (3)  Aug.  17.  A  new  Floridian  Lamellaria.  Naut.  55(1):17- 
18,  pi.  1,  figs.  3-6,  6a.  [Lamellaria  koto.] 

1949  (1)  Mar.  18.  A  new  Japanese  limpet.  Naut.  62  (1)  :97-98,  pi. 
6,  figs.  1-lc.  [Acmaca  langfordi.] 

1949  (2)  Mar.  18.  Fusinus  spectrum,  a  new  record  for  the  Gulf 
of  Mexico,  Naut.  62  (1):  101-102. 

1949  (3)    Nov.  1.  Liguus  enlarges  its  menu.  Naut.  63  (2)  :72. 

1950  (1)  Feb.  13.  Two  Pacific  species  of  Phos.  Naut.  63  (3)  :80-82, 
pi.  5,  figs,  3,  4.  [Phos  lannumi,  P.  amoenus.] 

1951  (1)  Mar.  7.  New  marine  mollusks  from  British  West  Indies 
and  Florida  keys.  Naut.  64  (4)  :  1 16-1 19,  pi.  8,  figs.  1-8.  [Crassi- 
spira  drangai,  Oliva  drangai,  Pitaria  cordata,  Calliostoma 
jujubinum  adelae.] 

1955  (1)  Marine  shells  of  the  western  coast  of  Florida.  By  Louise 
M.  Perry  and  J.  S.  S.  Paleont.  Research  Inst.,  198  pp.,  55  pis. 

1955  (2)  Aug.  1.  New  Conns  from  Costa  Rica.  Naut.  69  (\)  :13- 
15,  pi.  2,  figs.  1-15.  [Conus  drangai,  C.  andrangae,  C.  gradatus 
thaanumi,  C.  recurvus  helenae.] 

1957  (1)  April  29.  Theodore  Thomas  Dranga,  1901-1956.  Naut. 
7(?  (4):  138- 140, 

1958  (1)  Mar.  4.  Dr.  H.  A.  Pilsbry  in  marine  malacology.  Naut. 
71  (3) :  87-89, 

NOTES  AND  NEWS 
Dates  of  the  Nautilus.  —  Vol.  74,  no.  I,  pp.  1-40,  pis.  1-4,  was 
mailed  July  1,  1960,  No.  2,  pp.  41-84,  Oct.  5,  1960.  No.  3,  pp.  85- 
124,  pis.  5-8,  Jan  11,  1961.  No.  4,  pp.  125-166,  pis.  9  &  10,  April  6, 
1961.  — H.  B.  B. 

First  European  Malacological  Congress.  —  Under  the 
auspices  of  a  committee  appointed  by  the  Conchological  Society 
of  Great  Britain  and  the  Malacological  Society  of  London,  scien- 
tific and  field  meetings  are  planned  in  London,  September  17  to 
21,  1962.  All  aspects  of  European  malacology  and  the  formation 


40  NAUTILUS  Vol.    75     (1) 

of  a  European  Malacological  Union  will  be  discussed.  However, 
the  Congress  will  be  open  to  all  interested  in  malacology  from 
any  part  of  the  world.  Inquiries  should  be  addressed  to  the  Honor- 
able Secretary: 

Rev.  H.  E.  J.  Briggs,  19  Si  ward  Road,  Bromley,  Kent,  England. 

Directory  of  conchologists,  1962.  —  I  wish  to  include  all  per- 
sons and  institutions  interested  in  the  study  or  collection  of 
mollusks  in  this  new  directory,  which  will  be  published  next 
January.  A  copy  need  not  be  purchased,  but  those  who  remit  $2.50 
now  will  receive  the  first  copies  printed.  When  known,  the  names 
will  be  followed  by  numbers,  indicating  interests  from  following 
key  (or  other  special  interests  will  be  listed  when  mentioned): 
1,  world  wide  shells,  2,  land  shells.  3,  fresh  water  shells.  4,  fossil 
shells.  5,  exchange  shells.  Secretaries  of  shell  clubs  or  institutions 
will  please  send  me  a  list  of  their  members  and  their  specialties. 
— John  Q.  Burch,  4206  Halldale  Ave.,  Los  Angeles  62,  California. 

On  the  delayed  description  of  Xylophaga  atlantica. — The 
wood-boring  clam  Xylophaga  atlantica  was  described  as  a  new 
species  by  Richards  in  1942  (Nautilus  5(5:58) .  Turner  subse- 
quently remarked  in  her  monograph  on  the  Pholadidae  that, 
"It  is  surprising  that  this  western  Atlantic  species  remained 
undescribed  until  1942"    (Johnsonia  5:153.  1955). 

Recently  the  writer  discovered  a  letter  in  the  file  of  E.  S. 
Morse  papers  deposited  at  the  Peabody  Museum  of  Salem  which 
indicates  that  this  bivalve  was  recognized  as  a  new  species  at  a 
much  earlier  date.  Following  is  the  letter,  published  here  with 
the  permission  of  its  writer.  Dr.  Henry  Jackson,  Jr.,  and  the 
Director  of  the  Peabody  Museum,  Ernest  S.  Dodge: 

"My  dear  Mr.  Morse, 

I  suppose  you  are  very  busy,  but  could  you  lend  me  the 
drawings  you  made  of  the  Xylophaga  I  found  at  North  Haven. 
It  is,  without  much  doubt,  a  new  sp.  This  I  will  be  able  to 
ascertain  very  soon  as  Mr.  Henshaw  has  sent  to  England  for  all 
the  species  of  the  genus.  When  I  describe  it  I  shall  like  to  have 
all  the  drawings  I  can  of  it.  I  have  prepared  a  very  excellent  one 
of  Xylophaga  dorsalis  Turton  and  also  one  of  my  [sic]  species 
which  with  your  permission  I  will  name  Xylophaga  [of  Morse] 


July,   1961  NAUTILUS  41 

nov.  sp.  I  will  be  very  careful  of  your  drawings  if  you  condescend 
to  have  them  published  by  me  in  the  Proceedings  of  the  Boston 
Society  of  Natural  History." 

The  letter  was  undated,  but  in  the  judgment  of  Dr.  Jackson, 
it  was  written  in  1907  or  1908.  In  Dr.  Turner's  monograph 
(ibid-p.l54) ,  specimens  of  X.  atlantica  are  listed  from  "off 
North  Haven  (Maine)  in  ten  fathoms"  which  are  deposited 
in  the  U.S.  National  Museum.  These  specimens  were  the  ones 
collected  by  Dr.  Jackson  in  1907.  He  does  not  now  recall  why 
his  study  was  never  published.  Possibly  because  of  his  age  (about 
15  years  at  that  time),  his  paper  was  not  accepted  for  publica- 
tion, although  he  did  publish  two  short  papers  in  the  Nautilus 
in  1907  and  1908.  Or,  it  might  have  been  that  interest  in  the 
matter  was  lost  while  preparing  for  a  trip  to  Europe  that  year. 
Possibly  he  was  discouraged  by  Dr.  Morse  from  publishing  a 
description  at  that  time.  Thirty-four  years  later  the  situation 
was  corrected  by  Dr.  Richards. — Ralph  W.  Dexter,  Kent  State 
University,  Kent,  Ohio. 

PUBLICATIONS  RECEIVED 

Pages  in  italics  include  new  taxons. 

1959 

Adam,  William.  Les  cephalopodes  de  la  mer  Rouge.  Miss.  R.  P. 
Dollfus  en  Egypte.  Res.  Sci.  3   (28)  :  125-193,  pis.  1-9,  25  figs. 

Basch,  Paul  F.  Two  new  molluscan  intermediate  hosts  for  Para- 
gonimus  kellicotti.  J.  Parasit.  45:212).  Studies  on  the  develop- 
ment and  reproduction  of  the  fresh-water  limpet,  Ferrissia 
shimekii  (Pilsbry) .  Trans.  Amer.  Microsc.  Soc.  75:260-276. 

Burch,  John  Bayard.  Chromosomes  of  aquatic  pulmonate  snails 
(Basommatophora) .  Dissertation  abstracts  20  (4)  :  2  pp. 

Emerson,  William  K,  The  gastropod  genus  Pterorytis.  Amer.  Mus. 
Novitat.  no.  1974,  8  pp.,  4  figs. 

Ghose,  Krishna  Chandra.  Observations  on  the  mating  and  ovi- 
position  of  two  land  pulmonates,  Achatina  fulica  Bowdich 
and  Macrochlamys  indica  Godwin-Austen.  J.  Bombay  Nat. 
Hist.  Soc.  55:183-187. 

Schalie,  Henry  van  der  ^  Dee  S.  Dundee.  Transect  distribution  of 
eggs  of  Pomatiopsis  lapidaria  Say,  an  amphibious  proso- 
branch  snail.  Trans.  Microsc.  Soc.  75:409-420,  5  figs. 

1960 

Adam,  William.  Cephalopoda  from  the  Gulf  of  Aqaba.  Israel 
Sea  Fish.  Res.  Sta.,  Bui.  26:26  pp.,  pi.  1,  10  figs.  Contribution 


42  NAUTILUS  Vol.   75    (1) 

a  la  connaissance  de  I'hectocotyle  chez  les  Ommastrephidae. 
Bui.  Inst.  roy.  Sci.  nat.  Belgique  36  (\9)  :10  pp.  A  propos  de 
Chlamys  (F.Acc.  Hinnites)  abscondita  (P.  Fischer,  1898)  de 
la  cote  occidentale  de  I'Afrique.  Ibid.  (20)  :10  pp.,  2  pis.  Les 
mollusques  teiTestres  et  dulcicoles  de  la  Belgique.  Quelques 
additions  et  rectifications.  Ibid.  (22):  10  pp.,  7  figs.  Les 
c^phalopodes  de  I'lnstitut  Francais  d'Afrique  Noire,  2.  Bui. 
ibid,  22:465-511,  4  figs. 

Barbosa,  Frederico  S.,  Elizabeth  Carneiro,  Ivete  Barbosa  &  Jose  F. 
Magalhaes.  Manual  de  malacologia  medica  (Trabalhos 
praticos) .  Pp.  182,  110  figs.  Fundacao  Goncalo  Moniz,  Salva- 
dor, Bahia,  Brasil. 

Branson,  Bradley  A.  Gastropoda  of  the  Rob  and  Bessie  Welder 
Wildlife  Foundation  Refuge,  San  Patricio  County,  Texas. 
Southwest.  Nat,  5:143-159. 

Burch,  John  B.  Chromosomes  of  Gyraulus  circumstriatus,  a  fresh- 
water snail.  Nature  75<5:497-498,  1  fig. 

Cooke,  C.  Montague,  Jr.  &  Yoshio  Kondo.  Revision  of  Tornatelli- 
nidae  and  Achatinellidae  (Gastropoda,  Pulmonata).  Bui. 
Bishop  Mus.  221:1-303,  123  figs,  ($6.50). 

Emerson,  William  K.  Remarks  on  some  eastern  Pacific  muricid 
gastropods.  Amer.  Mus.  Novitat.  no.  2009:15  pp.,  7  figs. 

Hubricht,  JLeslie.  The  cave  snail,  Carycliium  stys^ium  CalL  Trans. 
Ky.  Acad.  Sci.  27:35-38,  2  figs. 

Kuroda,  Tokubei.  A  catalogue  of  molluscan  fauna  of  the  Okinawa 
Islands.  Pp.  106,  3  pis. 

Leonard,  A.  Byron  &  John  C,  Frye.  Wisconsinan  molluscan 
faunas  of  the  Illinois  valley  region.  111.  State  Geol.  Surv., 
Circular  304:  32  pp.,  4  pis.,  3  figs. 

Morton,  J.  E.  Molluscs:  an  introduction  to  their  form  and  func- 
tions. Harper  Torchbooks,  The  Science  Library;  232  pp.,  23 
figs.  (11.40). 

Paraense,  W.  Lobato  &  Newton  Deslandes.  " Drepanotrema  sur- 
inamense,"  with  an  addendum  on  "D.  petricola"  (Plan- 
orbidae) .  Rev.  Brasil.  Biol.  20:257-263,  7  figs. 

Schalie,  Henry  van  der.  Egypt's  new  high  dam — asiset  or  liability. 
Biologist  ^2:63-70,  3  maps. 

Scott,  M.  I.  Hylton.  Nueva  familia  de  pulmonado  Basomatoforo, 
Neotropico  6:65-69,  fig.  1,  Sobre  la  presencia  del  genero 
Pupisoma  en  la  Argentina.  Ibid.: 25-29,  figs.  1-9. 

Smith,  Allyn  G.  A  new  species  of  Megomphix  from  California. 
Occ.  Papers  Calif.  Acad.  Sci.  no.  28:5  pp.,  3  figs. 

Soot-Ryen,  T.  Pelecypods  from  Tristan  da  Cunha.  Results 
Norweg.  Sci.  Exp.  to  Tristan  da  Cunha,  no.  49:'^7  pp.  (3 
pis.) ,  9  figs. 

Voss,  Gilbert  L,  Bermudan  cephalopods.  Fieldiana,  Zoology 
39:419-446,  figs.  73-75. 


THE  NAUTILUS 

Vol.   75  October,    1961  No.  2 

TWO  NEW  STROBILOPIDS  FROM  THE  PLEISTOCENE 

OF  THE  HIGH  PLAINS 

By  TONG-YUN  HO  and  A.  B.  LEONARD 

In  the  summers  of  1958  and  1959,  studies  on  the  Pleistocene 
geology  and  molluscan  faunas  of  the  White  (Blanco)  River  area 
in  northwestern  Texas  were  carried  out  by  us.  In  the  course  of 
these  studies,  numerous  fossil  shells  of  Strobilops  of  problematical 
taxonomic  status  were  found  associated  with  other  pulmonate 
mollusks  in  fluviatile  deposits  ranging  in  age  from  Kansan  to 
Wisconsinan.  About  200  specimens  of  Strobilops  have  been  ob- 
tained from  the  White  River  area.  We  were  unable  to  assign 
these  shells  to  any  known  species  owing  to  their  consistent  char- 
acteristics intermediate  between  extinct  S.  sparsicostata  F.  C. 
Baker  (1938:127)  and  living  S.  texasiana  Pilsbry  and  F'erriss 
(Pilsbry,  1948:856) .  For  these  reasons  and  because  of  their  wide 
geographic  distribution  and  stratigraphic  range,  it  seems  best  to 
propose  a  new  name  for  these  fossil  shells.  Series  of  several 
Pleistocene  Strobilops  from  the  post-Nebraskan  of  Central  High 
Plains  were  restudied.  Those  series  of  Strobilops,  which  had 
been  confused  with  S.  sparsicostata,  appear  to  belong  to  the  new 
species  described  here. 
Strobilops  loxsdalei,  new  species.      Plate  6,  figs.  1-2;  text  figs.  1-2 

Diagnosis:  Shell  large  for  Strobilops,  having  trochiform  out- 
line; decidedly  angulate  last  whorl;  SI/q  to  6  convex  whorls  sculp- 
tured by  40  to  50  moderately  spaced,  moderately  thickened, 
oblique  ribs  passing  over  base  without  diminishing;  weak  and 
nodose  infraparietal  lamella. 

Holotype:  Catalogue  number  12180,  University  of  Kansas 
Museum  of  Natural  History,  obtained  by  A.  B.  Leonard  and 
Tong-yun  Ho,  June  9,  1959.  Original  number  TYH  34. 

Dcscriptioji  of  Jwlotype:  Shell  large,  trochiform;  whorls  6  in 
number,  moderately  convex,  slowly  increasing  in  size;  last  whorl 
decidedly  angulate;  base  broadly  rounded;  first  W/o  whorls  finely 
granulose,  remaining  whorls  covered  by  moderately  spaced, 
moderately  thickened  ribs  passing  over  base  Avithout  reduction  in 
size,  except  for  those  immediately  in  front  of  aperture;  diameter 

43 


44  NAUTILUS  Vol.  75  (2) 

of  umbilicus  equivalent  to  i/g  greater  diameter  of  shell;  aperture 
lunate,  expanded,  having  moderately  thick  parietal  callus  and 
peristome;  parietal  lamella  nodose,  high,  emerging  to  edge  of 
parietal  callus  and  penetrating  about  %  of  a  whorl  inward;  inter- 
parietal lamella  nodose,  weak,  shortly  emerged  and  penetrating 
as  deep  as  parietal  lamella;  infraparietal  lamella  nodose,  weak, 
shortly  emergent  and  penetrating  as  far  as  parietal  lamella;  col- 
umellar  lamella  weak;  first  basal  fold  moderate,  triangular;  sec- 
ond basal  fold  high,  broadly  triangular,  nearly  as  long  as  first 
basal  fold;  third  basal  fold  low,  about  i/^  length  of  second  basal 
fold;  fourth  and  fifth  basal  folds  low,  each  approximately  twice 
length  of  third  basal  fold;  palatal  fold  low,  longer  than  fifth  basal 
fold   (internal  anatomy  from  paratypical  shell)  . 

Paratypes:  Many  variations  occur  among  more  than  200  para- 
types  in  the  length,  thickness,  and  number  of  baso-palatal  folds, 
and  in  the  development  of  ribs.  The  baso-palatal  folds  in  most 
shells  are  5  in  number  but  in  a  few  specimens  a  small  fold  is  pres- 
ent between  the  third  and  fourth  basal  folds.  The  inter-  and 
infra-parietal  lamellae  are  extremely  weak  and  may  become  dis- 
continuous in  some  examples.  The  number  of  ribs  ranges  from 
40  to  50.  In  some  examples,  a  faint  riblet  is  between  major  ribs. 
The  ribs  on  the  base  may  become  weak  on  the  last  half  of  the 
base  or  near  the  umbilicus  in  some  shells;  as  a  rule  they  are 
conspicuous  on  last  half  of  base. 

Comparisons:  S.  lonsdalei  is  clearly  distinguished  from  S. 
sparsicostata  (Plate  6,  figs.  5-6;  text  figs.  4-5)  by  having  ribbed 
base,  weak  nodose  infraparietal  lamella,  and  fine  and  narrowly 
spaced  oblique  ribs,  always  more  than  forty  in  number.  The 
denticles  as  a  group  are  always  weaker  than  are  those  of  S. 
sparsicostata.  The  parietal  lamellae  are  nodose  in  lonsdalei,  but 
most  of  them  are  always  smooth  in  sparsicostata.  The  palatal 
fold  is  generally  present  in  lonsdalei,  but  it  is  absent  in  some 
specimens  of  sparsicostata. 

The  shell  of  S.  lonsdalei  differs  from  that  of  S.  texasiana  (see 
Pilsbry,  1948,  fig.  464:5-11)  in  being  large  and  trochiform  instead 
of  small  and  dome-shaped.  The  last  whorl  is  subangulate  or 
rounded  in  texasiana,  whereas  it  is  decidedly  angulate  in  lons- 
dalei. The  ribs,  which  are  always  more  than  50  in  number  in 
texasiana,  are  always  less  than  50  in  lonsdalei.  The  ribs  are  also 
more  widely  spaced  and  coarser  in  lonsdalei  than  in  texasiana. 
The  internal  structure  of  the  shell  of  lonsdalei  is  also  different 


October,   1961  nautilus  45 

from  that  of  texasiana.  The  latter  has  a  tongue-shaped  (blunt- 
topped)  first  basal  fold,  and  high,  rather  flat-topped  second 
basal  fold,  while  in  the  former  both  the  first  and  second  basal 
folds  are  triangular  in  shape  with  subacuate  tips.  Unlike  tex- 
asiana, lonsdalei  has  a  low  second  basal  fold  and  intraparietal 
lamella. 

The  comparisons  of  dimensions  in  millimeters  and  numbers  of 
whorls  are  summarized  in  the  following  table.  In  the  table,  speci- 
men HI  is  the  holotype,  and  specimens  PI  and  P3  are  paratypes. 

Species  S.  sparslcostata         S.  lonsdalei       S.  texasiana 

Specimen     12    3    4  HI   PI   P2   P5     12   3   4 

Diameter     2.7   2.7   2.6   2.5  2.8   2.8   2.7   2.7    2.4  2.3  2.2  2.0 

Height       1.9   2.0   2.1   1.9  2.4   2.5   2.3   2.1    1.9  1.9  1.6  1.6 

No.  of  ribs  34   36   36   34  41   44   48   45  52   50  52  51 

No.  of       5.5   5.5   5.5   5.5        6    6    5 .9   5 .9    5.5  5.5  5.3  5.5 

whorls 

Type  locality:  Kansan  terrace  deposits  in  left  bank  of  White 
River,  near  bridge  on  Texas  Highway  261,  6.5  miles  east  of 
Kalgary,  Crosby  County,   Texas. 

Areal  and  stratigraphic  distributions:  At  present  S.  lonsdalei  is 
known  from  the  Pleistocene  deposits  of  Kansan  to  Wisconsinan 
age  at  13  localities  distributed  from  western  Oklahoma  to 
northern  Texas.  Representative  localities,  age  of  deposits,  and 
approximate  number  of  specimens  obtained  at  each  locality  are 
as  follows: 

Kansan  deposits,  NW  14,  sec.  23,  T.  23N.,  R.  18W.,  2.5  miles 
west  and  %  mile  north  of  Ouinlan,  Woodward  County,  Okla- 
homa. No.  4337,  KU,  70  specimens. 

Kansan  deposits  in  road  cut  on  Texas  Highway  70,  1  mile 
north-northeast  of  Turkey,  Hall  County,  Texas.  No.  1  1818,  KU, 
10  specimens. 

Kansan  deposits  in  left  bank  of  White  River,  near  bridge  on 
Texas  Highway  261,  6.5  miles  east  of  Kalgary,  Crosby  County, 
Texas.  No.  11650,  KU,  60  specimens.    (Type  locality.) 

Kansan  deposits  exposed  in  cut  bank,  2.2  miles  west  of  Post, 
Garza  County,  Texas.  No.  10854,  KU,  40  specimens. 

Wisconsinan  terrace  deposits  on  north  side  of  U.S.  Highway  83, 
4.1  miles  east  of  Crosbyton,  Crosby  County,  Texas.  No.  10237, 
KU,    150  specimens. 

Wisconsinan  terrace  deposits,  5  miles  northeast  of  bridge  on 


46  NAUTILUS  Vol.  75  (2) 

U.S.  Highway  77  over  Red  River,  northern  central  Cook  County, 
Texas.  No.  12123,  KU,  10  specimens. 

Strobilops  lonsdalei  cansasiana,  new  subspecies.  Plate  6,   figs. 
3-4;   text  fig.   3. 

Diagnosis:  Shell  broadly  conic,  moderately  elevated;  whorls 
5i/2  in  number  sculptured  by  43  to  52  moderately  spaced  ribs; 
base  almost  smooth  or  finely  striate;  parietal  lamellae  penetrating 
from  1/2  to  Ys  of  last  whorl. 

Holotype:  Catalogue  number  12181,  KU,  obtained  by  Dr.  C. 
W.  Hibbard,  August  9,  1943. 

Description  of  Holotype:  Shell  moderate,  broadly  convex; 
whorls  51/2  in  number,  convex;  last  whorl  decidedly  angulate; 
base  more  or  less  narrowly  rounded;  first  I14  whorls  finely 
granulose,  remaining  whorls  sculptured  by  moderately  spaced, 
moderately  thickened  ribs  passing  over  base  in  extremely  fine 
striae;  diameter  of  umbilicus  contained  about  8  times  in  diameter 
of  shell;  aperture  lunate,  expanded,  having  moderate  peristome 
and  parietal  callus;  parietal  lamella  nodose,  penetrating  one- 
half  a  whorl  inward;  interparietal  lamella  nodose,  weak,  shortly 
emerged  and  penetrating  as  deep  as  parietal  lamella;  infraparietal 
lamella  shortly  emerged,  longer  than  interparietal  lamella,  pen- 
etrating as  far  as  parietal  lamella;  baso-palatal  folds  similar  to 
those  of  S.  lonsdalei  lonsdalei  (internal  structure  of  shell  based 
on  paratype) . 

Paratypes:  Few  variations  are  noted  in  more  than  2000  para- 
types  from  8  localities.  The  parietal  lamellae  pentrating  1/9  last 
whorl  inward  in  most  specimens,  but  about  5/^  last  whorl  in  the 
specimens  from  Reno  County,  Kansas,  and  Greer  County,  Okla- 
homa. The  base  is  smooth  or  finely  striate  in  most  examples, 
but  a  few  ribs  are  present  immediately  behind  the  peristome 
of  a  few  specimens.  The  baso-parietal  folds  are  as  variable  as 
those  of  S.  lonsdalei  lonsdalei.  Variations  in  size  of  shell,  and 
number  of  ribs  are  exemplified  in  the  following  measurements: 

Holotype  Paratypes 

Diameter    (mm.)  3.1  2.6         2.7         3.1 

Height    (mm.)  2.2  2.1         2.0         2.1 

No.  of  ribs  45  52         44  43 

No.  of  whorls  5.5  5.5        5.5        5.5 

Comparisons:  This  subspecies  differs  from  Strobilops  lonsdalei 
lonsdalei  in  having  smaller  shell  with  broad  and  smooth  base, 
less  elevated  spire,  more  ribs,  and  shorter  parietal  lamellae.  All 
other  characteristics  appear  to  be  similar  to  S.  lonsdalei  lonsdalei. 
S.  lonsdalei  cansasiana  resembles  Strobilops  texasiana  in  having 
the  same  number  of  ribs,  but  the  former  differs  from  the  latter 


October,    19(51 
NAU  riLUS  75  (2) 


NAUTILUS 


47 

PLAIK  (i 


Kigs.  1,  2.  Strobilops  lonsdalei  lonsdalei  Ho  and  Leonard.  Lateral  and  basal 
\  iews  of  holot\  pica]  shell.  Figs.  3,  4.  S.  lonsdalei  cansasiana  Ho  and  Leonard. 
Lateral  and  basal  views  of  holotypical  shell.  Figs.  5,  6.  S.  sparsicostata  Baker, 
Lateral  and  basal  views  of  topotypical  shell  from  Rexroad  Ranch,  9  miles 
south  and  7  miles  west  of  Meade,  Meade  C'oiintv,  Kansas.  All  rigurcs  enlarged 
approximately    15    times. 


48 


NAUTILUS 


Vol.  75  (2) 


A^ 


-(fc 


ii 


k. 


vv 


Sections  of  last  whorl  of  Sirobilops  showing  internal  lamellae  antl  toicls  ol 
ihe  shell.  All  fie;iires  enlarged  approximately  14  times.  Figs.  1,  2.  Strobilops 
loiisdah'i  lousdalci  Ho  and  Leonard,  paratvpes.  Fig.  3.  .S.  lousdalei  cansasiana 
Ho  and  Leonard,  paratype.  Figs.  4,  .5.  S.  sparsicoslata  Raker,  lopotvpes. 

in  bearing  rather  coarse  and  widely  sj^ared  ribs,  smooth  base, 
and  a  decidedly  angulate  last  whorl. 

Type  locality:  Kansan  deposits,  SW  14,  sec.  2,  T.  .81,S,  R.  28W., 
6  miles  north  of  Meade,  Meade  County,  Kansas. 

Area!  and  strntigraphic  distributions:  S.  lousdalei  ca)isasiaua 
is  now  known  only  Irom  Kansan  deposits  in  Iowa,  Kansas  and 
Okhdioma.  The  southern  range  of  S.  lousdalei  causasiaua  over- 
laps witli  the  noiithern  range  of  .S.  lousdalei  lousdalei  in  western 
Oklahoma.  Representative  localities  and  apj^roximate  number 
of  specimens  in  otir  collections  are: 

Kansan  deposits,  I  mile  east  and  4  miles  north  ol  Little  Sioux, 
Harrison  County,  Iowa.  No.  4517,  KU,  1 1  specimens. 

Kansan  deposits,  SW  14,  sec.  2,  T.  .8 IS.,  R.  28W.,  6  miles  north 
of  Meade,  Meade  County,  Kansas.  No.  ,8702,  KV ,  500  sjiecimens. 
(Tyj)e    locality.) 

Kansan  deposits,  SW  i/, ,  sec.  20,  T.  I  IS.,  R.  2i:.,  I  miles  west 
of  Navarre,  Dickenson  County,  Kansas.  No.  l.^^iOl,  Kl',  10  speci- 
mens. 


October,   19(5 1  nautilus  49 

Kansan  deposits,  sec.  36,  T.  24S.,  R.  7W.,  7  miles  east  and 
1.1  miles  north  of  Arlington,  Reno  County,  Kansas.  No.  9414, 
KU,   15  specimens. 

Kansan  deposits,  sec.  8,  T.  5N.,  R.  28E,  neaj-  north  border  of 

Gate,  Beaver  County,  Oklahoma.  No.  4743,  KU,  150  specimens. 

Kansan  deposits  in  right  bank  of  North  Fork  of  Red  River, 

north  of  Rock  Island  Railroad,  2  miles  east  of  Granite,  Greer 

County,  Oklahoma.  No.   10751,  KU,  70  specimens. 

Ecology  and  relationships:  The  ecological  requirements  of  this 
extinct  species  are  not  certainly  known.  The  association  of  it 
with  Retinella  electrina,  Pupilla  muscorum,  Stenotrema  leai,  and 
its  occurrence  in  the  alluvium  of  fine  and  well-sorted  sands  and 
silts  seems  to  imply  that  the  species  thrived  in  rather  humid  and 
cool  situations  on  the  flood  plains  where  vegetation  was  available. 
The  exact  cause  of  extinction  is  unknown,  but  the  initiation  of 
semiarid  and  otherwise  severe  climate  in  the  High  Plains  at  the 
time  of  the  Bradyan  interglacial  interval  may  have  been  respon- 
sible. This  view  is  strengthened  by  the  fact  that  no  single  species 
of  Strobilops  and  no  other  northern  gastropod  that  lived  in  the 
area  during  pre-Wisconsinan  time,  is  found  alive  in  the  White 
River  region  today. 

S.  lonsdalei  is  morphologically  and  stratigraphically  inter- 
mediate between  5.  sparsicostata  of  the  Nebraskan  and  existing 
S.  texasiana,  which  extends  to  the  Wisconsinan,  but  is  more 
closely  allied  to  5.  sparsicostata  than  to  5.  texasiana  on  the  basis 
of  size  and  shape  of  shell.  In  the  light  of  our  present  knowledge, 
5.  lonsdalei  seems  to  be  on  a  side  branch  of  the  line  leading  from 
S.  sparsicostata  to  5.  texasiana.  S.  sparsicostata  may  have  given 
rise  to  S.  lonsdalei  by  development  of  ribs  on  the  base  of  the 
shell,  by  increase  in  the  number  of  ribs,  and  by  reduction  in  size 
of  denticles. 

S.  lonsdalei  is  named  in  honor  of  the  late  Dr.  John  T.  Lonsdale, 
formerly  Director,  The  University  of  Texas  Bureau  of  Economic 
Geology,  in  recognition  of  his  continued  interest  and  support  of 
our  studies  in  Texas. 

Literature  cited 
Baker,  Frank  C.  1938.  New  land  and  freshwater  mollusca  from 

the  upper  Pliocene  of  Kansas  and  new  species  of  Gyraulus 

from  early  Pleistocene  strata.  Naut.  5/  (4)  :  126-131. 
Pilsbry,  Henry  A.  1948.  Land  Mollusca  of  North  America  (north 

of  Mexico)  .  Acad.  Nat.  Sci.  Philadelphia  Monographs  no.  3, 

2(2):856-858,  fig.  464:5-11. 


50  NAUTILUS  Vol.   75   (2) 

THE  STATUS  OF  THYASIRA  INSIGNIS,  T.  PLANA. 

AND  T.  INAEQUALIS,  ALL  VERRILL  AND  BUSH 

By  K.  W.  OCKELMANN 

Marine  Biological  Laboratory,  Helsingor,  Denmark 

While  working  up  the  marine  lamelHbranchs  of  East  Green- 
land (Ockelmann,  1959)  a  list  was  compiled  of  all  the  species 
known  from  the  Atlantic  sector  of  the  Arctic  and  adjacent  re- 
gions and,  in  many  cases,  I  had  to  consult  the  original  descrip- 
tions. It  thereby  appeared  that  the  validity  of  some  of  the  species 
of  Thyasira,  described  by  Verill  &  Bush  (1898)  from  the  east 
coast  of  North  America,  was  rather  doubtful  and  that  they  prob- 
ably were  synonymous  with  certain  earlier  described  species.  In 
particular,  this  seemed  to  apply  to  Thyasira  insignis,  T.  plana, 
and  T.  inaequalis. 

During  a  stay  in  the  United  States  in  1959,  I  therefore  took 
the  opportunity  to  examine  a  major  part  of  the  collection  of 
Thyasira  kept  in  the  U.S.  Nat.  Museum.  This  material  includes 
the  types  as  well  as  most  of  the  specimens  of  Thyasira  identified 
by  Verrill  &  Bush.  To  facilitate  a  later  thorough  revision  of  the 
whole  group,  which  is  highly  needed  in  spite  of  the  useful  papers 
by  Dall  (1901) ,  and  Lamy  (1920) ,  the  results  of  my  examination 
will  be  given  here. 

Thyasira  insignis:  This  form  was  described  as  Cryptodon 
insignis  by  Verrill  &  Bush  (1898,  p.  785,  pi.  91,  figs.  1,  2).  Two 
specimens  were  figured  and,  apparently,  the  authors  did  not 
designate  a  holotype.  However,  one  of  these  specimens  is  now 
in  the  type  collection  of  the  U.S.  Nat.  Museum,  probably  selected 
by  Dall  when  he  prepared  his  'Synopsis  of  the  Lucinacea  and  of 
the  American  species'  (1901).  It  is  an  empty  shell  (U.S.N.M. 
52596)  with  the  left  valve  drilled.  This  is  depicted  on  fig.  2  in 
the  paper  by  Verrill  Sc  Bush,  while  fig.  1,  same  plate,  is  drawn 
from  a  single  left,  drilled  valve  (U.S.N.M.  52733) .  Both  speci- 
mens are  from  U.S.F.C.  station  2499,  N.44°46'30''  W.59°55'45", 
depth  130  fath.  ('Albatross',  1885).  Further,  28  valves  (U.S.N.M. 
52557  and  52733)  from  stations  of  the  U.S.F.C.  2498  and  2499, 
were  present.  Of  the  whole  material,  consisting  of  1  shell  and  29 
valves,  11  still  had  remains  of  the  ligament,  but  no  trace  of  the 
soft  parts  whatever.  Thus,  tliey  may  vei7  well  have  rested  in  the 
bottom  deposits   for  a  considerable  time  after  death.   A  single 


October,   1961  nautilus  51 

valve  from  off  Cape  Cod  which  at  first  was  identified  as  Crypto- 
don  sarsii  by  Verrill  (1880,  p.  399),  but  later  (Verrill  k  Bush, 
1898)  referred  to  C.  insignis,  seems  not  to  be  present  in  the 
material.  According  to  the  latest  lists  of  the  marine  mollusks  of 
the  east  coast  of  North  America,  T.  insignis  occurs  from  New- 
foundland to  Cape  Cod,  Mass.,  65-471  fms.  (Johnson,  1934;  La 
Rocque,  1953) .  This  distribution,  however,  is  simply  based  on 
the  data  of  Verrill  &  Bush  (1898)  whose  material  solely  consists 
of  empty  shells  and  valves.  From  this,  apparently  T.  insignis  has 
never  been'  taken  alive. 

My  examination  of  the  type  as  well  as  the  other  material 
strongly  suggests  that  T.  insignis  is  not  specifically  distinct  from 
T.  sarsi  (Philippi)  1845,  b,  p. 91,  as  Axinus  sarsii).  The  outline 
of  the  shell,  its  proportions,  the  appearance  of  its  surface,  the 
hinge  margin,  the  size  and  the  position  of  the  ligament,  as  well 
as  the  scars  and  marks  on  the  inner  side  do  not  differ  in  the  two 
forms.  The  only  observable  differences  are  that,  in  some  of  the 
valves,  the  lower,  posterior  undulation  from  the  umbo  to  the 
posterior-basal  maigin  ends  a  little  more  ventrally  than  in  most 
of  the  specimens  of  T.  sarsi,  and  that  T.  insignis  may  become  32 
mm.  long,  while  the  largest  specimens  of  T.  sai'si  (valves  from 
W.  Norway)  seen  by  me  have  a  length  of  23.5  mm.  However, 
T.  sarsi  varies  rather  much  in  shape,  as  do  several  other  species 
of  Thyasira,  and  it  also  seems  to  attain  fairly  different  maximum 
sizes  in  different  localities.  G.  O.  Sars,  who  gives  good  figures  of 
the  typical  form  (1878,  pi.  19,  fig.  5  a-b) ,  shortly  describes  a 
'monstr.  oblonga'  collected  along  with  typical  specimens  at 
Lofoten.  This  form  especially  is  virtually  indistinguishable  from 
T.   insignis. 

According  to  my  own  studies,  T.  sarsi  now  occurs  from  the 
Sound  (Denmark)  through  the  Kattegat,  along  the  Swedish 
west  coast  and  the  Norwegian  coast  northward  to  at  least  Lofoten, 
and  it  is  also  found  in  some  fjords  of  northwest  Icelantl.  G.  O. 
Sars  (1878),  however,  states  that  it  occurs  as  far  northward  as 
Vadso  (E.  Finmark) ,  and  this  may  be  correct,  although  I  have 
not  seen  animals  from  that  region.  It  has  even  been  recorded 
farther  north  and  east  by  other  authors.  Lamy  (1920,  p.  297), 
for  instance,  mentions  it  from  the  Kara  Sea.  However,  since 
specimens  of   T.  gouldi    (Philippi)   often   have   been   identified 


52  NAUTILUS  Vol.  75  (2) 

and  listed  as  T.  sarsi,  the  northern  limit  of  distribution  is  not 
known  with  certainty.  T.  sarsi  has  also  been  recorded  from  W. 
Greenland  (Posselt  &  Jensen,  1898,  p.  80,  as  Axinus  flexuosus  var. 
sarsii).  The  Zool.  Museum,  Copenhagen,  however,  only  possesses 
a  single,  empty  shell  from  W,  Greenland,  viz.  from  Godthaab. 
The  ligament  of  this  shell  is  fairly  well  preserved,  but  soft  parts 
are  completely  absent  and  the  shell  surface  does  not  look 
'fresh'.  There  are  thus  three  possibilities:  1)  This  specimen  is  a 
quaternary  fossil,  and  T.  sarsi  is  now  extinct  at  W.  Greenland, 
2)  The  specimen  is  wrongly  labelled,  and  not  collected  at  W. 
Greenland  at  all,  or  3)  T.  sarsi  occurs,  although  sparsely,  in 
west  Greenland  water.  At  present,  none  of  these  interpretations 
can  be  excluded. 

From  the  preceding  data,  I  conclude  that  T.  insignis  merely 
represents  a  large  form  of  T.  sarsi,  and  that  the  former  should 
be  regarded  as  a  subspecies  of  the  latter  and  referred  to  as 
Thyasira  sarsi  insignis  (Verrill  &:  Bush) .  Further,  no  proof 
exists  that  this  is  an  extant  member  of  the  recent  fauna  of  the 
east  coast  of  N.  America,  since  all  the  specimens  referred  to  it 
may  very  well  be  Quaternary  fossils,  thus  indicating  a  wider  dis- 
tribution of  T.  sarsi  in  former  times,  viz.  from  the  Scandinavian 
coasts  to  the  region  around  Newfoundland  and  New  England. 
This  supposition  also  agrees  with  the  record  of  an  empty,  pos- 
sibly fossil,  shell  of  T.  sarsi  from  W.  Greenland,  and  with  the 
existing  isolated  population  of  N.W.  Iceland. 

Thyasira  plana:  This  was  described,  with  good  figures,  as 
Cryptodon  planus  by  Verrill  &  Bush  (1898,  p.  788-9,  pi.  LXXX- 
VIII,  figs.  3,  4).  The  type  (U.  S.  N.  M.  159893)  is  a  specimen  from 
U.S.F.C.  sta.  254,  Cape  Cod  Bay,  Tishing  ledge'.  Wood  End  Light 
N.  50°E.  7  miles,  21  fath.  ('Speedwell',  1879).  A  close  examin- 
ation of  it  left  no  doubt  that  it  is  an  about  half-grown  specimen  if 
Thyasira  gouldi  (Philippi)  (1845  a,  p.  74-75).  It  has  a  well- 
rounded  outline  which  is  often  exhibited  by  smaller  specimens 
of  T.  gouldi.  Its  hinge-margin  is  comparatively  strong  and 
fairly  well-rounded  below  the  umbo,  a  feature  also  found  in  T. 
gouldi.  Furthermore,  Verrill  &  Bush's  statement  as  to  the  ab- 
sence of  a  distinct  tubercle  or  tooth-like  projection  is  not  quite 
correct.  There  is,  just  as  in  T.  gouldi,  a  rather  stout,  though 
small,  tooth  in  the  right  hinge  close  to,  and  below,  the  umbo, 


October,    19bl  nautilus  53 

and — as  also  shown  by  Venill  &:  Bush,  fig.  3 — a  distinct  pit  or 
groove  in  the  left  hinge  corresponding  to  it.  A  highly  distinctive 
character  of  T.  goiildi  is  the  large,  conspicuous  prodissoconch 
wliich  most  often  measures  210-250  microns  across.  This  feature 
is  also  present  in  the  type  of  T.  plana,  the  prodissoconch  of 
which  measures  about  215  microns  across.  Finally,  all  the  other 
material  investigated  and  originally  labelled  as  Cryptodon 
planus  also  turned  out  to  belong  to  T.  gouldi,  mostly  being 
smaller  specimens  of  it.  This  material  originates  from  the  follow- 
ing localities:  Casco  Bay,  1873.  Seal  Cove,  8-10  fms.,  Y.  M.,  1872, 
Chelsea  Beach,  Mass.,  Dall.  Friars  Head,  Maine,  Henderson  Coll. 
U.S.F.C.  stations  72+73,  42B+43B  +  58B  +  60B-f  66B-f  67B,  134, 
136,  140,  IGO,  165,  181,  191,  199,  210,  283+285,  296,  and  340 
(serial  numbers  U.S.N.M.  74281,  74283-74286,  74289,  108868, 
159848,  159849,  159876-159884,  and  445797).  There  is  also  one 
sample  from  Maine,  collected  by  Stearns,  but  without  a  serial 
nmnber.  Living  specimens,  identified  as  Cryptodon  planus,  weie 
found  at  depth  from  15-19m.  to  188  m.  and  at  temperatures 
between  about  4.0°  and  9.4°C.  (These  data  are  based  upon 
Smith,  1887,  according  to  the  station  numbers  of  the  samples). 
The  distribution  agrees  with  that  given  by  Johnson  (1934) ,  and 
La  Rocque  (1953) ,  viz.  from  Halifax,  N.S.,  to  Cape  Cod,  Mass., 
and,  according  to  these  authors,  the  range  of  T.  gouldi  along  the 
east  coast  of  N.  America  is  from  Greenland  to  Connecticut,  5-400 
fath.  Abbott  (1954,  p.  384) ,  however,  gives  the  range  as  "Labra- 
dor to  North  Carolina,"  but  without  any  references  or  details. 
T.  plana  thus  occurs  well  within  the  range  of  T.  gouldi,  regard- 
less of  the  depth  record  of  400  fath.  for  T.  gouldi  which  seems 
doubtful  and  may  not  apply  to  living  specimens.  Actually,  T. 
gouldi  is  uncommon  at  depth  exceeding  100  m.  and  even  at 
depths  of  about  50  m.  often  only  smallei"  specimens  are  found. 
These  are  usually  more  rotund  than  are  the  larger  ones;  and 
apparently  such  specimens  led  Verrill  &:  Bush  to  describe  Crypto- 
don planus. 

Thyasira  inaequalis:  This  was  described  as  Cryptodon  (Ax- 
inulus)  inequalis  by  Verrill  &  Bush  (1898,  p.  791,  pi.  90,  figs.  1, 
2).  Dall  (1901,  p.  786)  is  correct  in  regarding  it  as  a  Thyasira 
S.S.,  not  an  Axinulus.  The  type  (U.S.N.M.  159850)  is  an  animal 
from  U.S.F.C.  sta.  98,  99,  midway  between  Sandwich  Point  and 


54  NAUTILUS  Vol.  75  (2) 

McNab's  Island  Light,  Halifax  Harbor,  depths  of  1 8  fms.  (Speed- 
well, 1877).  On  a  close  inspection,  it  turned  out  to  be  nothing 
but  an  aberrant  specimen  of  T.  gouldi,  aberrant  in  so  far  as  the 
outline  of  the  shell  is  concerned.  Otherwise  it  shows  all  the 
features  typical  of  T.  gouldi;  the  prodissoconch,  for  instance, 
measures  about  215  microns  across.  Also  the  other  material  in- 
vestigated (U.S.F.C.  stations  72-f  73,  293,  296,  and  340  (U.S.N. M. 
74287,  159847,  202836,  and  202837)  comprises  only  specimens  of 
T.  gouldi,  but  of  a  more  normal  outline  and  shape.  This  again 
points  towards  the  true  nature  of  the  type:  An  individual  variant 
of  T.  gouldi.  Just  as  is  the  case  with  T.  plana,  T.  inaequalis  is 
sympatric  with  T.  gouldi,  as  it  occurs  from  Halifax,  N.S.,  to 
Cape  Cod,  Mass.,  14  to  49  fath.  (Johnson,  1934;  La  Rocque, 
1953). 

It  is  concluded,  then,  that  both  Thyasira  plana  (Verrill  & 
Bush) ,  and  T.  inaequalis  (Verrill  &  Bush)  must  be  regarded  as 
junior  synonyms  of  T.  gouldi  (Philippi).  The  available  data  do 
not  justify  a  claim  for  even  their  subspecific  status. 

Most  figures  of  T.  gouldi  in  the  literature  are  on  a  small  scale 
and  do  not  show  the  distinctive  features  satisfactorily.  For  con- 
venience, therefore,  references  to  usable  figures  may  be  given 
here:  G.  O.  Sars,  1878,  pi.  19,  fig.  6  a-b  (as  Axinus  gouldii), 
and  Ockelmann,  1959,  pi.  2,  figs.  4-5. 

My  cordial  thanks  are  due  to  Dr.  Harold  A.  Rehder  for 
kindly  placing  the  collections  and  facilities  of  the  U.S.  National 
Museum  at  my  disposal. 

A  few  living  adult  T.  sarsi  have  been  found  by  me  since,  in  the  Gotthab 
Fjord,  W.  Greenland,  July,  1961. 

Literature  cited 
Abbott,  R.  T.  1954.  American  seashells.  Van  Nostrand,  New  York. 
Ball,  W.  H.  1901.  Proc.  U.S.  nat.  Mus.  25(1237)  :779-833. 
Johnson,  C.  W.  1934  List  of  Marine  Mollusca  of  the  Atlantic 

Coast  from  Labrador  to  Texas.  Proc.  Boston  Soc.  Nat.  Hist. 

-/O(l):  1-204. 
Lamy,  E.  1920.  J.  de  Conchyliol.  ^5:233-318    (3e  Part.). 
La  Rocque,  A.  1953.  Catalogue  of  the  recent  Mollusca  of  Canada. 

Bull.  nat.  Mus.  Ottawa  129   (Biol.  ser.  no.  44)  :  1-406. 
Ockelmann,  K.  W.   1959.  Marine  Lamellibranchiata.    (In:   The 

Zoology  of  East  Greenland.)    Medd.  om  Gronland  122    (4)    : 

1-256. 
Philippi,  R.  A.  1845a.  Zeitschr.  f.  Malakozool.  2:68-79. 
Philippi,  R.  A.  1845b.  Zeitschr.  L  Malakozool.  2:87-91. 


October,   19()1  nautilus  55 

Possclt,  H.  J.  S:  Jensen,  A.  S,  1898.  Gronlands  Brachiopoder  og 

Bloddyr.    (In:  Conspectus  Faunae  Groenlandicae)  .  Medd.  cm 

Gronland  25;  I-XIX,  1-298. 
Sars,  G.   ().    1878.   Bidrag  til   Kundskaben  om   Norges  Arktiske 

Fauna.  I.  Mollusca  regionis  arcticae  Norvegiae.  Christiania. 
Smith,  S,   1889.   Rep.  U.S.  Comm.  Fish  and' Fisheries  for   1886 

(part  XIV):  871-1017. 
Verrill,  A.  E.  1880.  Proc.  U.  S.  nat.  Mus.  5:356-405. 
Verrill,  A.  E.  R:  Bush,  K.  J.  1898.  Proc.  U.S.  nat.  Mus.  20   (1 139)  : 

775-901. 


MARINE  SHELLS  OF  WATER  ISLAND,  VIRGIN  IS. 

Bv  J.  A.  WEBER 

Watei"  Island  is  located  about  y^  of  a  mile  east  of  St.  Thomas. 
It  consists  mainly  of  volcanic  rocks  which  rise  to  a  height  of  294 
feet  at  the  highest  point,  leaving  a  very  rough  topography.  The 
water  aurrounding  the  island  varies  from  shallow  to  a  depth  of 
50  to  85  feet  in  the  channels.  The  bays,  sand  beaches,  rock  ledges 
and  coral  reefs  furnish  a  great  variety  of  ecological  situations  for 
mollusks.  Collecting  is  very  convenient,  as  the  proprietor  of  the 
island,  Mr.  Walther  H.  Phillips,  maintains  tourist  cottages, 
where  wet  clothes  and  specimens  may  be  handled.  The  writer 
made  two  trips  to  the  island,  one  in  August,  1956,  and  the  other 
in  July,  1958,  which  included  shallow-water  collecting  and  some 
dredging.  The  species  collected  follow: 

Gnstropods.  Arene  cruentata 

Acmaea  antillarum  A.  miniata 

A.  cubensis  A.  riisei 

A.  leucopleura  Aspella  elizabethae 

A.  jamaicensis  A.  paupercula 

A.  pustvdata  Astraea  americana  cubana 

A.  pustulata  pulcherrima  A,  caelata 

Acteon  punctostriata  A.  longispina 

Alaba  incerta  A.  tuber 

Alabina  adamsi  Atys  caribaea 

Alvania  auberi  A.  sharpi 

Anachis  albella  Batillaria  minima 

A.  catenata  Bittium  cerithioides 

A.  nitens  B.  varium 

A.  obesa  Bulla  occidentalis 

A.  subcostulata  Bullata  ovuliformis 

Antillophos  candei  Bursa  thomae 

Architectonica  nobilis  Caecum  cooperi 


56 


NAUTILUS 


Vol.  75  (2) 


C.  decussatum 

C.  regulare 

C.  floridanvim 

C.  pulchellum 

Calliostoma  jujubinum 

C.  pulcher 

C.  zonamestum 

C.  sarcodum 

Cantharus  auritulus 

Cassis  flammea 

C.  madagascariensis 

C.  tuberosa 

Cerithides  costata 

Cerithiopsis  iota 

C.  flavum 

C.  fusiforme 

C.  greeni 

C.  rugulosum 

Cerithium  algicola 

C.  eburneum 

C.  literatum 

C.  muscarum 

C.  variabile 

Cerodrillia  thea 

Charonia  variegata 

Coliimbraria  lanceolata 

Columbella  mercatoria 

C.  ovulata 

C.  ovuloides 

Conus  spurius 

C.  jaspideus 

C.  mus 

C.  regius 

C.  cardinalis 

C.  dominicanus 

Coralliophila  abbreviata 

C.  bracteata 

C.  caribaea 

Crassispira  ebenia 

C.  nigrescens 

C.  ostrearum 

Crepidula  aculeata 

C.  maculosa 

C.  plana 

Cyliiidrobulla  beaui 

Cymatium  gemmatum 


C.  muricium 
C.  nicobairium 
C.  parthenopium 
C.  pileare 

Cyphoma  gibbosum 
C.  macgintyi 
Cypraea  cinerea 
C.  spurca  acicularis 

C.  zebra 

Cypraecassis  testiculus 
Cythara  trilineata 
Daphnella  lymneiformis 
Decipifus  pulchellus 
Diodora  arcuata 

D.  cayenensis 
D.  dysoni 

D.  minuta 
Drillia  elatoir 
Drupa  nodulosus 
Echininus  nodulosus 
Emarginula  phrixodes 

E.  pumila 
Engina  turbinellus 
Engoniophos  unicinctus 
Epitonium  albidum 

E.  candianum 
E.  echinaticostum 
E,  lamellosum 
E.  occidentale 

E.  unifasciatum 
Erato  maugeiiae 
Euchelus  guttarosea 
Fasciolaria  tulipa 
Fenimorea  fucata 

F.  phasma 
Fissurella  augusta 
F.  barbadensis 

F.  barbouri 
F.  nimbosa 
F.  fascicularis 
F.  nodosa 
F.  rosea 

Fossarus  orbignyi 
Fusilatirus  cayohuesonicus 
Glyphotunis  diminuta 
Haliotinella  patinaria 


October,   1961 


NAUTILUS 


57 


Haminoea  antillarum 
H.  elegans 

Hemitonia  emarginata 
H.  octo-radiata 
Hipponix  antiquatiis 
H,  submfus 
Hyalina  albolineata 
H.  avena 
H.  avencia 
H.  tenuilabra 
H.  parkeri 

Ithycythara  lanceolata 
Jaspidella  jaspidea 
Kinziella  (juadrilineata 
Latirus  brevicaudatus 
Leucozonia  nassa 
L.  leucozonalis 
Litiopa  melanostoma 
Littorina  angulifera 
L.  meleagiis 
L.  mespillum 
L.  ziczac 
Livona  pica 
Lucapina  aegis 
L.  philippiana 
L.  sowerbyi 
L.  sufftisa 

Lucapinella  limatula 
Mangelia  biconica 
M.  fusca 
M.  melantica 
M.  quadrilineata 
Marginella  sulcata 
Meioceras  nitidum 
Melampus  coffeus 
Melanella  jamaicensis 
Micromelo  undata 
Mitra  albicostata 
M.  albocincta 
M.  barbadensis 
M.  cubana 
M.  floridana 
M.  hanleyi 
M.  hanleyi  gemmata 
M.  moisei 
M.  nodulosa 


M.  straininea 

Mitrella   duclosiana 

M,  fusilormis 

Modulus  carchedonius 

M.  modulus 

Monilispira  albocincta 

M.  jayana 

M.  leucocyma 

Morum  oniscus 

Murex  cabritti 

M.  cailleti  kugleri 

M.  micromeris 

M.  pomum 

Muricopsis  oxytatus 

Nassarina  glypta 

Nassarius  albus 

N.  vibex 

Natica  canrena 

N.  livida 

Neodrillia  jamaicensis 

Nerita  peloronta 

N.  tessellata 

N.  versicolor 

Neritina  virginea 

Nitidella  dichroa 

N.  idalina 

N.  laevigata 

N.  nitidula 

N.  ocellata 

Nodilittorina  tuberculata 

Odostomia  laevigata 

Oliva  reticularis 

Olivella  adelae 

O.  dealbata 

O.  nivea 

O.  perplexa 

Opalia  pumilo 

Parvi turbo  weberi 

Pedipes  mirabilis 

Persicula  miniata 

Petaloconchus  erectus 

Phalium  cicatricosum 

Phenacolepas  hamillei 

Pira  monile 

Pisania  pusio 

Planaxis  lineatus 


58 


NAUTILUS 


Vol.  75  (2) 


P.  nucleus 

Polinices  lacteus 

P.  uberinus 

Prunum  virginianum 

Psarostula  monilifera 

Puperita  pupa 

P.  tristis 

Pyramidella  crenulata 

P.  dolabrata 

Pyrunculus  caelata 

Pyrgocythara  coxi 

P.  emeryi 

Retusa  caniculata 

Rissoa  gradata 

Rissoina  bryeria 

R.  cancellata 

R.  chesneli 

R.  decussata 

Rhizorus  acutus 

Sigatica  semisulcata 

Sinum  perspectivum 

Siphonaria  alternata 

S.  pectinata 

Smaragdia  viridis 

Spartophos  floridanus 

Stigmaulax  sulcata 

Synaptocochlea  picta 

S.  coccina 

Tectarius  muricatus 

Tegula  excavata 

T.  fasciata 

T.  hotessieriana 

T.  lividomaculata 

T.  substriata 

T.  viridula 

Terebra  dislocata 

T.  glossima 

T.  hastata 

Thais  deltoidea 

T.  haemastoma  floridana 

T.  rustica 

Tonna  maculosa 

Torinia  infundibulum 

T.  cyclostoma 

Trailia  ovula 

Tiicola  adamsi 


T.  bella 

T.  tessellata 

Trigonostoma  rugosum 

Trimusculus  carinata 

Triplioia  decorata 

T.  mirabilis 

T.  nigrocincta 

T.  pulchella 

Tritonalia  intermedia 

Trivia  pediculus 

T.  quadripunctata 

Truncatella  caribaeensis 

Turbo  canaliculatus 

T.  castanea 

Turbonilla  curta 

T.  interrupta 

Vasum  muricatum 

Vermicularia  spirata 

Xenophora  conchyliophora 

Pelecypods. 

Abra  aequalis 

Aequipecten  acanthodes 

A.  muscosus 

A.  gibbus 

Americardia  guppyi 

A.  medium 

Anadara  notabilis 

Anadontia  alba 

Anomalocardia  brasiliana 

Antigona  listeri 

A.  rugatina 

Area  umbonata 

A.  zebra 

Arcopagia  fausta 
Arcopsis  adamsi 
Asaphis  deflorata 
Barbatia  cancellaria 

B.  Candida 

B.  domingensis 
B.  tenera 

Basterotia  newtoniana 
B.  quadrata 
Botula  fusra 
Brachidontes  citrinus 
B.  exustus 
Cuspidaria  costellata 


October,   1961 


NAUTILLIS 


59 


Chama  congregata 

C.  florida 

C.  macerophylla 

C.  sinuosa 

Chione  cancellata 

C.  grus 

C.  mazycki 

C.  pygmaea 

Chlamys  benedicti 

C.  imbricata 

C.  multisguamata 

C.  ornata 

Codakia  costata 

C.  orbicularis 

C.  orbiculata 

C.  pectinella 

C.  portoricana 

Congeria  leucophaeta 

Coralloiphaga  coralloiphaga 

Corbula  operculata 

Crassinella  guadeloupensis 

C.  lunulata 
Cumingia  coarctata 
Cyathodonta  cruziana 
Diplodonta  punctata 
Divaricella  dentata 

D.  quadrisulcata 
Donax  denticulata 
Echinochama  archinella 
Ervilia  nitens 
Glycymeris  decussata 
G,  pectinata 

G.  undata 

Gouldia  cerina 

G.  insularis 

Isognomon  alatus 

I.  bicolor 

I.  radiatus 

Laevicardium  laevigatum 

L.  1.  sybariticum 

L.  mortoni 

Lima  pellucida 

L.  lima 

L.  scabra 

L.  tenera 

Limea  bronniana 


Lioberus  castaneus 

Lithophaga  antillarum 

L.  bisulcata 

L.  nigra 

Lucina  multilineata 

L.  pensylvanica 

L.  radians 

L.  trisidcata  blandus 

Lysonsia  beana 

Lyropecten  antillarum 

L.  nodosus 

Macoma  brevifrons 

M.  constricta 

M.  tenta 

Macrocallista  maculata 

Mactra  fragilis 

Modiolus  americanus 

Mulina  lateralis 

Muscukis  lateralis 

Ostraea  frons 

Papyridea  semisulcata 

P.  soleniformis 

Pecten  ziczac 

Petricola  lapicida 

Phacoides  pectinatus 

Pholadomya  Candida 

Pinctata  radiata 

Pinna  rudis 

Pi  tar  albida 

P.  aresta 

P.  circinnata 

P.  fulminata 

Plicatula  gibbosa 

P.  sp? 

Pododesmus  rudis 

Pseudochama  radians  variegata 

Pteiia  colymbus 

Quadrans  lintea 

Rocellaria  cuneiformis 

R,  ovata 

Semele  bellastriata 

S.  nuculoides 

S.  proficua 

S.  proficua  radiata 

Solemya  occidentalis 

Solecurtus  cummingia 


60 


NAUTILUS 


Vol.  75  (2) 


S.  sancta-marthae 
Spengleria  rostrata 
Spondylus  aniericanus 
Strigilla  carnaria 
S.  mirabilis 
S.  piciformis 
Tagelus  plebeius 
T.  divisus 
Tellina  alternata 
T.  candeana 
T.  crystallina 
T.  cuneata 
T.  interrupta 
T.  guildingii 


T.  laevigata 
T.  promera 
T.  similis 
T.  sybaritica 
T.  versicolor 
Thyasira  trisinuata 
Tivela  abaconis 
Trachycardium  isocardia 
T.  magnum 
Transenella  cubaniana 
T.  culebrana 
Trigonicardia  antillarum 
Venericardia  tridentata 


EIGHT  NEW  SPECIES  OF  LAND  SNAILS  FROM 

THE  SOUTHERN  UNITED  STATES 

By  LESLIE  HUBRICHT 

(Concluded  from  July  no.) 

Succinea  Indiana  lives  in  places  which  are  so  dry  that  no  other 
species  of  native  land  snail  can  survive  there.  Not  only  can  it 
withstand  sun  and  drought,  but  the  ravages  of  roadgraders  and 
bulldozers  as  well.  Below  the  Clark  Hill  Dam,  it  is  found  abun- 
dant on  land  from  which  some  twenty  feet  of  earth  had  been 
removed  during  the  construction  of  the  dam.  Land  that  even 
the  weeds  did  not  find  entirely  hospitable.  At  this  locality  the 
snails  were  heavily  infested  with  leucochloridium.  Every  speci- 
men opened  contained  larvae  or  sporocysts. 
Succinea  campestris  Say 

Succinea  campestris  Say,  1817  Jour.  Acad.  Nat.  Sci.  Phila  /.•  281. 

Succinea  campestris  vagans  Pilsbry,  1900,  Naut.  14:  74. 

Qiiickella  vagans  (Pilsbry) ,  Pilsbry,  1948;  Land  Moll.  N.  Amer. 
//,  p.  848,  figs.  443d. 

An  examination  of  the  shells  of  the  type  lot  of  Succinea  cam- 
pestris vagans  Pilsbry,  disclosed  that  it  was  a  mixture  of  two 
species  of  shells.  One  species,  of  which  there  were  5  or  6  spe- 
cimens, had  a  very  short  spire,  large  aperture  and  almost  hyaline 
shell,  and  were  not  distinguishable  from  shells  of  Succinea 
campestris  Say  of  the  same  size.  The  remaining  shells,  of  which 
there  were  25  to  30,  were  smaller,  with  longer  spire  and  smaller 
aperture.  These  were  not  distinguishable  from  Catinella  vermeta 
(Say)  of  the  same  size.  Pilsbry's  original  description  was  clearly 


October,   1961  nautilus  61 

based  on  the  shells  of  S.  campestris.  Later,  when  he  described  the 
animal,  he  used  the  animals  of  C.  vermeta.  Since  the  original 
description  was  based  on  shells  of  C.  campestris,  the  name  x/agans 
must  be  placed  in  the  synonomy  of  that  species. 

Succineids  become  sexually  mature  when  very  small.  After 
they  have  mated  they  will  add  at  least  a  full  whorl  to  the  shell. 
Dr.  Pilsbry,  apparently,  was  not  aware  of  this,  as  vagens  was  dis- 
tinguished from  campestris  only  by  its  smaller  size  and  fewer 
whorls. 

Catinella  texana,  new  species.  PI.  4,  figs.  F  to  H;  text-fig.  1,  F 
to  H. 

Shell  pale  golden-brown  in  color,  translucent,  rather  dull,  with 
about  four  whorls,  elongate-ovate,  sculpture  of  unevenly  spaced 
growth  wrinkles.  Spire  acute,  long,  sutures  well  marked,  whorls 
well  rounded.  Aperture  ovate,  occupying  about  58%  of  the  length 
of  the  shell,  outer  and  basal  margins  well  rounded.  Columella 
nearly  straight. 

Mantle  and  sides  of  foot  covered  with  a  dark  gray  flecking. 
Hermaphrodite  duct  well  pigmented.  Talon  strongly  pigmented, 
almost  black;  club-shaped  and  undivided  but  with  a  terminal 
notch.  Prostate  gland  large,  oval  and  unpigmented.  Vas  deferens 
not  very  long,  entering  the  penis  at  the  apex.  Penis  rather  broad 
unpigmented,  with  a  large  pouch,  connected  near  the  base  on 
the  anterior  side.  There  is  no  trace  of  a  sheath.  Penial  retractor 
connected  to  the  side  of  the  penis  near  the  middle.  Spermatheca 
large,  globose,  duct  slender. 

Height  Diameter        Ap.  H.  Ap.  W. 

13.3  mm.         6.7  mm.         7.7  mm.         5.0  mm.         Holotype. 

13.3  mm.         7.3  mm.         8.3  mm.         5.7  mm.         Paratype. 

10.8  mm.         6.0  mm.         6.3  mm.         4.0  mm.         Paratype. 

Type  Locality. — Texas:  Milan  Co.:  near  small  spring,  near 
Brazos  River,  4,7  miles  northeast  of  Cause,  holotype  205892  and 
para  types  205891  U.M.M.Z.,  other  paratypes  15427,  collection  of 
the   author. 

Catinella  texana  is  most  closely  related  to  Catinella  vermeta 
(Say) ,  differing  in  having  a  more  slender  shell,  and  in  having 
the  appendix  basal  rather  than  in  the  middle  of  the  penis. 
Catinella  pugilator,  new  species.  PI.  4,  figs.  I  to  K;  text-figs.  1, 
D  &  E. 

Shell  straw-yellow,  translucent,  shining,  oblong-ovate,  with 
about  three  whorls,  sculpture  of  fine  growth  wrinkles.  Spire 
rather  short,  sutures  moderately  impressed,  whorls  convex.  Aper- 
ture ovate,  occupying  about  68%  of  the  length  of  the  shell,  outer 


62  NAUTILUS  Vol.  75  (2) 

and  basal  margins  well  rounded.  Columella  nearly  straight. 

Mantle  gray,  overlayed  with  black  blotches.  Hermaphrodite 
duct  well  pigmented.  Talon  moderately  pigmented,  club-shaped, 
with  a  terminal  notch.  Prostate  gland  rather  large,  somewhat 
rectangular  in  shape,  lightly  pigmented.  Vas  deferens  rather 
long,  slender,  abruptly  enlarging  before  it  enters  the  penis, 
entering  the  penis  at  the  apex.  Penis  broad,  unpigmented,  with 
a  small  thumb-like  appendix  near  the  center  of  the  posterior 
side.  There  is  no  trace  of  a  sheath.  Penial  retractor  connected  to 
the  anterior  side  of  the  penis  near  the  middle  and  to  the  thick- 
ened portion  of  the  vas  deferens.  Spermatheca  rather  large, 
globose,  duct  rather  short,  thick.  Atrium  and  vagina  very  short. 
Height  Diameter        Ap.  H.  Ap.  W. 

13.2  mm.         7.1mm.         9  3  mm.         5.1mm.         Holotype. 
11.1mm.         6.0  mm.         7.8  mm.         4.7  mm.         Paratype. 

10.3  mm.         5.3  mm.         7.0  mm.         4.0  mm.         Paratype. 
Type  Locality. — North  Carolina:  Currituck  Co.:  salt  marsh,  0.5 

mile  south  of   Maple,   holotype   205889   and   paratypes   20589D 
U.M.M.Z.,  other  paratypes  20980,  collection  of  the  author. 

Catinella  pugilator  resembles  C.  hubrichti  Grimm  in  its  Oxy- 
Zoma-like  shell,  but  differs  in  its  boxing-glove  shaped  penis.  The 
very  small  penial  appendix  plus  the  thick  spermatheca  duct  will 
readily  separate  this  species  from  all  other  described  species. 

Vertigo  teskeyae,  new  species.  Text-fig.  2. 

Shell  ovate,  dark  chestnut,  with  about  five  whorls.  Spire  con- 
vexly  conic,  with  obtuse  summit;  whorls  increasing  rapidly,  the 
last  much  the  largest,  its  height  equal  to  almost  two-thirds  of  the 
shell,  sculpture  of  fine  irregular  growth  lines.  Lip  thin  and 
expanded,  with  a  distinct  crest  behind  it.  Aperture  with  a  dis- 
tinct sinulus.  Parietal  lamella  rather  strong,  about  twice  as  long 
as  high;  angular  lamella  small;  an  infraparietal  tubercle  is  never 
present.  Columellar  lamella  strong,  horizontal,  located  in  about 
the  center  of  the  columella.  Basal  fold  variable,  rarely  very 
large,  often  nearly  obsolete,  located  on  the  end  of  a  callous  ridge 
which  extends  along  the  basal  and  lower  outer  walls  just  inside 
of  the  lip.  Infrapalatal  tubercle  often  present,  although  usually 
very  small.  Lower  palatal  fold  moderately  well  developed,  upper 
palatal  fold  somewhat  smaller.  Suprapalatal  fold  never  present. 


Height 

Diameter 

Whorls 

2.6  mm. 

1.6  mm. 

5         Holotype. 

2.1  mm. 

1.5  mm. 

4         Chocowinity.   (smallest) 

2.9  mm. 

1.8  mm. 

5         Lightwoodknot  Creek,   (largest) 

Distribution — Maryland:  Dorchester  Co.:  subfossil,  near  beach, 
0.5  mile  south  of  Elliott.  North  Carolina:  Camden  Co.:  swamp,  1 


October,   1961 


NAUTILUS 


63 


Figure  2.  Vertigo  teskeyae:  A,  holotype.  B,  paratype.  C,  from  Light woodkiiol 
Creek. 


mile  south  of  South  Mills.  Gates  Co.:  near  Bennetts  Creek,  Gates- 
ville.  Chowan  Co.:  swamp,  1.2  miles  south  of  St.  Johns.  Beaufort 
Co.:  swamp,  4.8  miles  north  of  Pantego;  near  Pamlico  River, 
opposite,  VVashington;  swamp,  1.4  miles  northeast  of  Chocowinity. 
Craven  Co.:  near  Little  Creek,  1  mile  north  of  Askin;  swamp, 
1.8  miles  south  of  Vanceboro.  Columbus  Co.:  bank  of  canal,  west 
side  of  Lake  Waccamaw,  holotype  205903  and  paratypes  205904 
U.M.M.Z.,  other  paratypes  20870,  collection  of  the  author.  South 
Carolina:  Williamsburg  Co.:  near  Black  Mingo  Creek,  north  of 
Rhems.  Georgia:  Screven  Co,:  near  Brier  Creek,  10  miles  east- 
northeast  of  Sylvania.  Toombs  Co.:  swamp,  6.5  miles  south  of 
Parkers.  Florida:  Flagler  Co.:  near  Pellicer  Creek,  13.5  miles 
north  of  Brunnell.  Holmes  Co.:  low  woods,  2.5  miles  west-north- 
west of  Sweet  Gum  Head.  Alabama:  Montgomery  Co.:  low  woods, 
0.5  mile  north  of  Mt.  Meigs.  Covington  Co.:  near  Lightwoodknot 
Creek,  3.7  miles  west  of  Opp.  Mississippi:  Wanen  Co.:  low  woods, 
near  Big  Black  River,  6.2  miles  east-northeast  of  Bovina. 

Vertigo  teskeyae  is  most  closely  related  to  V.  ovata  Say,  from 
which  it  may  be  readily  distinguished  by  its  distinctly  darker 
color  (when  fresh) ,  larger  size,  and  fewer  teeth.  The  infraparietal 
and  suprapalatal  tubercles  are  never  present.  The  columellar 
lamella  is  horizontal,  not  sloping  upward,  and  is  lower  in  posi- 
tion on  the  columella.  The  basal  fold  is  smaller  and  is  lower 
down.  The  aperture  is  much  more  open  than  is  usual  in  V.  ovata. 
V.  teskeyae  is  the  largest  Vertigo  known  from  the  southeastern 
United  States.  It  is  a  species  of  the  southern  Atlantic  and  Gulf 
Coastal  Plains.  In  North  Carolina,  it  is  found  cra^vling  on  the 


64  NAUTILUS  Vol.  75  (2) 

wet  mud  and  debris  of  swamps.  Farther  south  it  may  be  found 
on  the  undersides  of  palmetto  leaves  in  company  with  Pupisoma. 
It  is  frequently  found  associated  with  V.  ovata. 

Vertigo  teskeyae  is  named  in  honor  of  Mrs.  Margaret  C. 
Teskey,  in  recognition  of  her  services  in  behalf  of  the  American 
Malacological  Union. 

PUERTO  RICAN  CAMAENIDAE 

By  H.  BURRINGTON  BAKER 

The  symbols  used  for  Puerto  Rican  localities  were  explained 
recently    (I96I) . 

Caracolus  carocoUa  (Linne).  Under  leaves  on  ground  when 
dry,  but  fair  climber  on  tree  trunks,  up  to  10  ft.  around  El 
Yunque;  En,  Er,  En3,4,  Jn,  Pn,  Pr,  Wn,  Wr,  Ws  (dead)  ;  every- 
where except  in  driest  places;  not  even  empty  shells  recorded 
from  Ps  or  Ww;  0-4000  ft.  Animal  (added  to  Wurtz,  1955:112) 
sluggish;  foot  large,  with  sides  varying  from  slate  color  to  dark 
chestnut  with  buff  spots;  all  tentacles  dark;  sole  trizonal,  with 
coarse  waves. 

The  obsolete  name  Helix  arecibensis  Pfeiffer,  1856,  evidently 
was  based  on  a  juvenile  shell  of  Caracolus,  probably  this  species. 

Caracohis  marginella  (Gmelin) .  Very  good  climber  on  tree 
trunks,  nearly  subarboreal,  often  seen  15-20  ft.  above  ground; 
Ee,  En,  Es,  Jn,  Pn,  Wn;  mainly  on  lowlands,  especially  near 
cultivation;  not  seen  above  1500  ft.  in  Luquillo  Mts.  or  in  the 
eastern  Cordillera  Central  (Pr) .  Animal  dorsally  almost  black, 
but  shading  into  whitish  near  sole;  ommatophores  dark  but 
eyes  darker.  Uniformly  light  greenish  shells  with  whitish  animals 
(Jnl,  near  which  "albinos"  were  found  in  1864,  v.  Bland, 
1875:80). 
Caracolus  marginella  mayaguezi,  new  subspecies. 

Pleurodonte  bornii  (Pfr.)  Pilsbry,  1889  (6b)  :  127,  in  part,  pi. 
56,  fig.  6  type  (ANSP.  20861)  from  Swift  Collection,  labeled 
"Mayaguez."  Type  locality  (the  older  collectors  often  gave  the 
nearest  port):  top  of  Pico  Montoso,  near  Maricao,  elevation  2300 
ft.  (Wr2) ,  where  shells  match  Pilsbry's  figs,  very  closely.  Shell 
differs  from  typical  C.  marginella  by  its  broader  dark  zones, 
which  on  the  apical  side  restrict  paler  to  narrow,  sutural  and 
carinal  zones.  Also  found:  Maricao  Forest  (Wr3) ,  near  Cabo  Rojo 
(Ws)  and  near  Guanica  Bay  (Psl) ,  0-3000  ft.  Also  noted  by 
Bland,  1875:80. 

All  shells  collected  further  east,  including  those  around  mouth 
of   Rio  Guajataca    (Wn),   which   slightly   approach    mayaguezi, 


October,   1961  nautilus  65 

are  quite  typical  marginella.  Probably  the  2  forms  intergrade, 
since  the  typical  one  thrives  in  lowland,  cultivated  places;  in 
fact,  from  field  notes,  I  saw  living  "marginella"  (which  form?) 
on  trees  in  the  garden  of  the  Agricultural  Station  in  Mayaguez. 

Polydontes  (Luquillia)  luquillensis  (Shuttleworth)  .  Adults  on 
trunks  of  trees,  5-12  ft.  up,  but  young  (looking  like  those  of  P. 
lima)  usually  on  ground;  Luquillo  Mts.,  above  2000  ft.,  where  it 
replaces  the  "closely  allied"  P.  lima.  Animal  bright  ochraceous 
(but  one  almost  white);  eye  and  inferior  tentacles  almost  black; 
sole  blackish  slate. 

Although  these  lots  show  considerable  variation  in  banding 
and  height  (Cf.  Pilsbry,  1889  (2)  :pl.  7,  figs.  63,  64)  none  matches 
that  of  van  der  Schalie,  1948: pi.  6,  fig.  9,  which  seems  immature 
and  weather  worn.  Despite  considerable  differences  in  its  later 
whorls,  Granodomiis  Pilsbry,  1931,  seems  only  a  section  of  the 
subgenus  Luquillia  Crosse,  1892   (Cf.  Wurtz,  1955:132). 

Polydontes  (Granodomus)  lima  (Ferussac,  1821)  Fair  climber 
on  tree  trunks  and  brush,  but  less  so  than  C.  marginella;  espe- 
cially common  near  cultivation;  Ee2,  Es,  Jn,  Js,  Ps2  (dead) ,  Wn, 
Ws  (1  shell  form  castrensis) ,  Ww;  lowlands.  Secretes  quantities 
of  disagreeable  mucus;  seen  in  reciprocal  copulation.  ANSP. 
1125  (from  Bland,  without  exact  locality)  contains  2  shells  of 
form  castrensis  and  one  of  subspecies  maricao;  see  Pilsbry, 
1889  (6b)  :pl.  1,  figs.  3  (castrensis)  and  4    (maricao). 

P.  (G.)  lima  maricao  Clench.  Apparently  climbs  less  than 
typical  lima;  Prl-3,  Wr2,3;  Cordillera  Central,  2500-4000  ft. 

Dr.  van  der  Schalie's  (1948:79)  nice  "curves"  show  long  and 
careful  study,  but  is  a  "random  sample"  from  a  large  area 
actually  possible?  Might  not  the  "normal  distribution"  of  his 
composite  one  be  due  to  his  larger  series,  as  shown  by  his  maps, 
of  shells  from  lower  altitudes?  If  based  on  enough  animals,  that 
from  over  1500  ft.  may  be  significantly  bimodal,  and  the  habitat 
of  typical  maricao  seems  considerably  higher. 

My  lots  of  P.  lima  maricao  often  show  less  prominently  the 
peripheral  brown  band  and  sometimes  additional,  finer  ones 
like  the  larger  shells  from  the  Virgin  and  Leeward  Islands.  For 
this  reason,  in  part,  the  very  involved  nomenclature  of  the  latter 
is  outlined.  Perhaps,  rnaricao  and  asperula  may  be  "microspecies," 
and  "typical"  lima,  which  thrives  in  cultivated  areas,  might 
consist  of  very  unstable  hybrids  between  them? 
P.   (G.)  LIMA,  VAR.   (?)  iNCERTA   ("Fer."  Bcck)   or  (Ferussac) . 


66  NAUTILUS  Vol.  75  (2) 

Helix  incerta  Ferussac,  1823,  livT.  18: pi.  105,  figs.  2  (named  on 
cover?) .  (Helix)  Otala  incerta  Beck,  1837:36,  from  Lesser  Antilles 
(sic) ,  founded  on  figs,  cited.  Not  Helix  incerta  Draparnaud,  1805. 
(?)  H.  incerta  Pfeiffer,  1841,  Symb.  Hist.  Helic.:37,  from  St. 
Thomas. 

Dir.  Pilsbry  and  I  always  disagreed  on  the  delicate  question  as 
to  whether  Beck's  name  should  be  preoccupied  in  his  genus 
Helix,  or  only  in  his  subgenus  Otala,  but  Beck  himself,  1837a, 
when  he  added  description  to  a  few  of  his  nude  names,  regularly 
used  his  genus  or  its  initial;  e.g.,  Nanina  (Microcystis)  ornatella 
(p.  2) .  In  any  case,  "incerta"  should  be  returned  to  the  synonymic 
oblivion,  from  which  Pfr.  snatched  it.  The  Philadelphia  Acad. 
Nat.  Sci.  has  3  lots  of  the  fossils  from  St.  Thomas  (ANSP.  1033 
from  Bland)  and  I  agree  with  R.  Swift  (Mss.  in  ANSP.  30634 
from  St.  John  I.,  and  in  30636  from  St.  Thomas)  that  Ferussac's 
figures  look  most  like  one,  worn,  deformed  shell  of  P.  lima.  The 
apparently  dark  band  of  his  left  fig.  is  absent  from  his  right  one 
and  Deshayes  made  no  mention  of  it. 

P.      (G.)    LIMA    ASPERULA     (Beck)  . 

Helix  lima,  var.  notab.  (sic)  Ferussac,  1832,  livr.  22,  Explic. 
pis.  livr.  22-27:  p.  ij;  livr.  24,  pi.  46a,  figs.  4  &  5  [copied  by  Pils- 
bry, 1889  (2)  :pl.  4,  figs.  36  &  37];  certainly  not  a  valid  name. 
(Helix)  Otala  asperula  Beck,  1837:36,  founded  solely  on  figs, 
cited  above,  from  Curacao  (sic) ,  but  with  H.  Hgen.  lima  var. 
notabilior  Fer.,  in  synonymy.  Helix  notabilis  Shuttleworth, 
1854:40,  partly  founded  on  figs,  cited  above,  from  Is.  San  Juan 
(St.  John)  and  Anegada,  and  with  Helix  lima  var.  notabilis 
Fer.  cited.  Thelidomus  notabilis  (Sh.)  Pilsbry,  1889  (6b)  &  1890 
(5b)  :  pi.  1,  fig.  1  (ANSP.  1047,  Tortola,  from  Thos.  Bland)  and 
var.,  pi.  1,  fig.  1  [ANSP.  30634,  labeled  as  from  San  Jan  (St. 
John  I.)  by  Krebs  (from  Swift  Collection)  ]. 

Since  they  all  are  founded  on  exactly  the  same  figures,  I  now 
select  Ferussac's  fig.  4  (the  adult,  cited  above)  as  representing 
the  type  shell,  and  Tortola,  Virgin  Islands,  as  the  type  locality 
of  notab.  Fer.,  asperula  Beck  and  notabilior  Beck.  The  ANSP. 
lots  from  St.  John  Island  (now  designated  as  type  locality  of 
H.  notabilis  Sh.)  seem  usually  a  little  thinner  and  more  com- 
monly have  additional  bands  above  and  below  periphery,  but 
intergrade. 

Incidentally,  all  the  records  of  "incerta"    (actually  asperula) 
from  Puerto  Rico  are  due  to  Kreb's  label  of  "San  Jan,"  which 


October,   1961  nautilus  67 

I  certainly  (and  probably  Pilsbry)  took  to  be  a  misspelling 
of  San  Juan,  Puerto  Rico,  where  asperula  does  not  occur.  To 
add  to  the  confusion,  the  principal  town  of  Antigua  apparently 
was  named  after  the  same  saint,  and  shells  similar  to  the 
Tortola  form  in  banding,  but  thinner  (like  those  from  St. 
John  I.)  lived  on  St.  Bartholomew  (ANSP.  1051  from  Bland), 
which  also  is  in  the  Leeward  Islands.  Anegada  (ANSP.  1049 
from  Bland)  is  another  Virgin  I.  Finally,  one  shell  (ANSP. 
30628  from  W.  G.  Binney)  is  labeled  Martinique,  and  looks 
like   the  Tortola   form. 

Polydontes  (Parthena)  acutangula  (Burrows) .  Adults  truly 
arboreal,  but  young  animals  often  found  under  leaves  on 
ground;  Enl,  Er,  Es4,  Janl,  Pn2  (long  dead),  Prl;  0-3400  ft. 
east  of  San  Juan,  but  westward  only  found  living  at  4000  ft. 
Green  mantle  of  adults  showing  through  shell,  but  young  ones 
ochraceous  brown.  Animal  (added  to  Wurtz,  1955:112)  in- 
capable of  withdrawal  into  shell,  extending  out  to  8  inches 
(20  cm.)  long,  including  about  I1/2  inches  of  tentacles,  but 
mantle  only  extending  slightly  over  columella;  tail  long,  angu- 
late  dorsally  with  median  groove;  sides  of  foot  becoming  brown- 
ish near  sole:  ommatophores  darkish  slate;  inferior  tentacles 
lighter;  mantle  collar  orange-brown;  mucus  yellow;  seen  in 
reciprocal  copulation  on  trees. 

Van  der  Schalie,  1948:82,  obtained  another  camaenid,  Zachry- 
sia  auricoma  havanensis  Pilsbry,  a  Cuban  species. 
Additional  references    (See  1961) 
Baker,  H.  Burrington.  1961    (1),  Naut.  7^:142.  1961    (3)  „  Naut. 

75:33. 
Beck,  H.  1837,  Index  molluscorum,  etc.:  1-124. 

1837a,  Specierum  novarum,  etc.:  1-8. 

Wurtz,    Charles    B.    1955,    Proc.    Acad.    Nat.    Sci.    Philadelphia 
707:99-143,  pis.   1-19. 

ADULT  LIVING  SPECIMEN  OF  PAPUINA  FERUSSACI 

Bv  J.  B.  HENRARD 

Oegstgeest,  The  Netherlands 

To  identify  Papuina  ferussaci  (Lesson) ,  I  looked  up  the  history 
of  the  species  and  its  whereabouts.  The  description,  published 
in  1831,  is  rather  short  but  sufficient;  moreover  a  figure  was 
given  by  him.  Unfortunately  his  type  specimen  is  probably 
lost,  Tapparone  Canefii  tried  in  vain  to  locate  it  in  the  museum 
at  Paris.    Pfeiffer,   who  had  in    1848   to  deal   with   the   species, 


68  NAUTILUS  Vol.  75  (2) 

could  only  copy  the  description  in  his  Mon.Hel.Viv.,  p.231,  but 
in  the  year  1868,  in  Vol.  5.  of  his  Monograph,  he  identified 
material  from  Cumniing's  collection  as  being  Lesson's  species 
and  gave  a  new  description  of  his  own.  This  new  description, 
however,  was  in  many  important  characters,  very  different  from 
the  original  one.  Tapparone  recognized  the  error  and  demon- 
strated convincingly  in  the  year  1883,  that  Pfeiffer's  so-called 
Helix  ferussaci,  both  the  description  and  the  colored  plate  in 
the  Novitates  Conch,  tab. 56  fig.  7-9,  was  a  characteristic  new 
species  and  named  it  Helix  exsultans.  In  the  well-known  Manual 
of  Conch.  Vol.  7,  p.30,  f.51  the  true  species  of  Lesson  was  placed 
by  Pilsbry  in  the  genus  Papiiina;  he  gave  a  good  description, 
taken  from  the  original  one  and  from  the  plate.  This  description 
runs  as  follows:  "Shell  imperforate,  trochiform,  deep-chestnut, 
with  a  golden-yellow  line  at  the  carina,  and  ascending  the  spire 
to  the  apex,  whorls  6,  completely  flat,  the  apex  is  enlarged, 
convex;  the  last  whorl  has  a  salient  acute  carina.  Aperture 
much  depressed,  triangular,  wider  than  high,  very  acute  at  the 
junction  of  the  appressed  basal  lips;  peristome  reflexed,  sinuous 
at  the  angle." 

In  the  year  1933,  Lesson's  species  was  mentioned  by  Madame 
V.  Benthem  Jutting  on  the  list  of  all  the  species  of  the  genus 
Papuina,  known  at  that  time.  This  important  and  useful  list 
was  published  in  the  periodical  Nova  Guinea,  Vol.  17  (Zoology) 
Livr.  1,  p.  411.  She  placed  in  it  var.  exsultans  (Tapp.  Canefri) , 
the  color  being  much  diluted  to  cinnamon.  Because  this  color 
is  the  typical  one  of  Tapparone's  species,  the  identification,  in 
my  opinion  is  probably  correct. 

The  most  recent  mention  of  Papuina  ferussaci  is  given  by  Adam 
and  Leloup  in  Vol.  2,  p.29  (Mus.  Roy.  Sc.  Hist.  Nat.)  in  the 
year  1939,  together  with  fig.  16  on  Plate  2.  The  only  specimen 
they  had,  was  found  in  1929  near  Manokwari,  which  is  the 
modern  name  of  the  type  locality.  Port  Dorey  of  Lesson.  Their 
specimen  was  in  bad  condition,  worn  and  nearly  totally  bleached, 
but  with  the  yellow  keel  still  present  at  the  base  of  the  shell 
and  with  the  typical  brown  color.  I  agree  with  both  authors 
that  their  specimen  belongs  to  the  true  P.  ferussaci  of  Lesson, 
after  comparison  with  their  figure  16.  Since  that  time,  no  further 
data  are  known  to  me. 

In  the  year  1949,  Father  Rombouts,  a  missionary  at  Sorong, 


October,  1961  nautilus  69 

told  me  that  he  would  penetrate  what  he  called  the  Aifat  Anoek 
Tifa  Region,  in  the  northeast  part  of  "The  Vogelkop,"  to  visit 
some  papua-settlements.  Because  he  was  much  interested  in 
natural  history  and  the  region  was  never  visited  by  collectors, 
I  asked  him  if  he  could  give  some  attention  to  living  land-mol- 
lusks.  He  promised  to  do  that.  Although  he  was  obliged  to 
return  earlier,  on  account  of  the  unkind  and  hostile  natives, 
he  collected  a  very  interesting  lot.  Upon  his  arrival  home,  he 
handed  the  specimens  to  me  and  we  examined  them  together. 
Among  them  there  were  species  I  had  already  found  myself, 
but  others  were  quite  new  to  me;  among  them  was  a  very  beau- 
tiful species  of  Papuina,  the  subject  of  this  paper. 

Pointing  to  this  Papuina  Father  Rombouts  said:  "This  one 
must  be  very  rare  and  I  found  it  so  beautiful,  that  I  tried  to 
find  some  more,  but  could  not  obtain  another  one." 

All  the  characters,  given  by  Lesson  and  from  the  translation 
by  Pilsbry,  given  above,  agree  perfectly  with  this  specimen. 
Lesson  mentioned,  however,  only  the  general  and  easily  rec- 
ognizable ones,  so  that  the  more  subtle  ones  (mostly  visible 
under  a  strong  lens)  are  wanting  in  his  description.  I  give  here 
an  additional  description  of  my  specimen. 

High  trochiform  shell,  somewhat  higher  in  proportion  to 
the  width,  dark  castaneous,  the  faintly  convex  base  shining, 
^vhereas  the  whorls  above  the  very  sharp  keel  are  of  a  dull 
color.  Whorls  nearly  totally  flat,  only  very  slightly  concave 
to  their  lower  sutures,  the  profile  very  different  from  those  of 
the  allied  species  P.  exsultans  (Tapp.)  and  P.  pythonissa  (Tapp.). 
Keel  with  a  golden-yellow,  1/2  mm  broad  band,  visible  also  along 
the  sutures  and  below  the  keel.  The  common  oblique  growth- 
lines  very  distinct,  under  a  strong  lens;  the  whorls,  especially 
the  lower  one,  with  about  5  regular  spiral-lines,  base  radiately 
striated,  towards  the  umbilical  region  somewhat  deeper  and 
wider  grooved.  Whorls  just  below  the  suture  slightly  darker 
colored,  forming  on  the  ultimate  one  a  faint,  not  sharply  de- 
fined zone.  Apex  somewhat  darker  colored.  Umbilicus  a  faint 
perforation,  obtected  by  the  columellar  margin.  Peristome 
darker,  more  blackish-brown  and  the  aperture  provided  within 
with  a  grayish-white,  rather  sharply  defined  margin,  the  inner 
part  of  the  aperture  brown.  Columellar  margin  rather  broadly 
reflexed  and  provided  at  its  base  with  a  distinct,  although 
small  tooth-like  pale  projection,  the  columella  passing  rather 
bluntly  to  the  nearly  straight  lower  margin  of  the  peristome; 
outer  margin  of  the  peristome  beaked,  with  a  sharp  keel. 


70  NAUTILUS  Vol.   75   (2) 

The  small  columellar  tooth,  mentioned  in  my  description,  was 
not  indicated  by  Lesson  and,  in  the  lithographic  picture  by 
the  artist,  this  character  did  not  show  fully.  Moreover,  the 
dimensions  are  not  indicated  on  Lesson's  plate.  In  the  descrip- 
tion the  diameter  is  given  as  11  lines;  accepted  as  French  lines, 
this  becomes  24.8  mm,  and  my  specimen  measures  28.2  mm. 
My  shell  is  therefore  somewhat  larger.  Because,  however,  in 
many  species  of  the  genus  Papuina,  there  occur  fluctuations  in 
the  dimensions,  there  is,  for  the  time  being,  no  reason  to  accept 
a  special  variety  for  my  larger  one. 

Pilsbry  placed  P.  ferussaci  in  his  6th  group,  the  "Group  of 
the  P.  pileolus."  He  characterized  this  group  as  "subperforate 
or  imperforate,  pyramidal,  sharply  keeled  and  flat  at  the 
base."  Unfortunately,  many  species,  mentioned  by  him,  were 
not  sufficiently  known  to  him,  especially  if  there  were  no 
figures,  and  this  group  is  very  heterogeneous.  P.  leonardi  (Tapp.) 
and  P.  turris  (Adams)  are  not  sharply  keeled  at  the  base;  they 
do  not  belong  to  this  group.  The  former  certainly  belongs  to 
the  5th  group  of  Pilsbry,  called:  "the  group  of  the  P.  antiqua." 
This  also  was  recognized  by  Madame  v.  Benthem  Jutting,  who 
accepted  P.  leoyiardi  simply  as  a  synonym  of  P.  antiqiin  (Adams 
&  Reeves) ,  in  the  list  of  the  species  of  Papuinn  in  Nova  Guinea 
Vol.  17,  cited  above  p.  121.  Papina  turris  (H.  Adams)  has  only 
a  compressed  last  whorl  with  a  very  blunt  keel  and  must  be 
eliminated  also.  The  remaining  species  of  the  group  are,  as 
to  form  and  outline,  very  different  and  it  is  thus  very  heterogen- 
eous. We  can,  however,  divide  the  group  easily  into  two  homo- 
geneous ones,  if  we  accept  for  the  true  "group  of  the 
P.  pileolus"  only  the  low-spired  species  without  a  subcolumellar 
tooth,  viz.  P.  pileolus  (Ferussac),  P.  rhynchostoma  (Pfeiffer)  and 
P.  fergusoni  (H.  Adams).  The  other  group,  containing  high-spired 
species  with  a  very  distinct  subcolumeller  tooth  is  to  accept  as  the 
"group  of  the  P.  ferussaci." 

The  new  group  is  not  only  very  homogeneous  but,  moreover, 
zoogeographically  sharply  limited,  because  all  the  species,  be- 
longing to  this  group,  inhabit  the  northwest  part  of  Dutch  New 
Guinea.  All  its  members  are  represented  in  my  collection.  I 
shall  summarize  them  afterwards  more  in  detail  in  another 
paper. 


October,   1961  nautilus  71 

AN  EXCURSION  TO  LAKE  MIRAGOANE,  HAITI 
By  WALTER  JACOB  EYERDAM 

Lake  Miragoane  in  the  southwest  peninsula  of  Haiti  is  a 
small  muddy  lake  which  is  about  three  square  miles  in  area 
and  lies  near  the  coast  about  half-way  between  the  towns  of 
Petit  Goave  and  Miragoane,  approximately  60  miles  on  the 
road  west  of  Port-au-Prince.  It  is  the  largest  fresh- water  lake 
in  Haiti  and  its  vicinty  is  sparsely  populated.  There  are  two 
other  large  lakes  on  Hispaniola  which  are  several  hundred 
square  miles  in  area  which  are  saline  and  were  originally  inlets 
of  the  Gulf  of  Haiti  in  geological  times.  These  are  the  Etang 
Saumatre  and  the  Lac  Enriquilla  (150  ft.  below  sea  level) ,  both 
of  which  lie  due  east  of  Port-au-Prince  about  50  miles.  The 
latter  lake  is  all  within  the  Dominican  border. 

In  1927  while  collecting  plants  and  land  snails  for  the 
Smithsonian  institution  and  reptiles  for  Dr.  Thomas  Barbour 
in  the  southwestern  peninsula  of  Haiti  and  the  islands  of  Gonave 
and  Grande  Cayemite,  I  also  made  a  special  effort  to  collect 
all  the  freshwater  shells  that  could  be  found.  More  than  1000 
lots  of  land  shells  were  collected  and  sent  to  Dr.  Bartsch.  Most 
of  them  were  taken  alive  or  in  a  good  condition.  For  one  thing 
I  found  that  Haiti  compares  in  no  manner  with  Cuba  in  rich- 
ness of  its  land  shell  fauna.  There  is  only  one  species  of  Liguiis, 
and  only  a  very  few  species  of  Cerion  and  Urocoptis,  of  which 
there  are  many  hundreds  of  species  in  Cuba.  The  only  family 
of  land  snails  that  seems  fairly  well  represented  is  the 
Annulariidae. 

There  are  many  areas,  some  of  which  are  of  considerable 
extent  where  snails  are  exceedingly  scarce.  The  topography 
of  Haiti  is  very  rough  and  mountainous  with  more  than 
3,000,000  people  of  which  over  99  percent  are  negro.  Most  of 
the  popvilation  are  peasants. 

It  was  my  good  fortune  to  team  up  with  the  great  Swedish 
botanist,  and  naturalist.  Dr.  Erik  Ekman,  the  foremost  authority 
on  Caribbean  plants  who  knew  intimately  the  flora  of  Cuba 
and  Haiti.  Before  he  died  in  the  Dominican  Republic  in  1930, 
he  had  discovered  over  2,000  new  species  of  plants  in  Cuba 
and  Haiti  during  his  20  years  of  botanical  rambles. 

I  spent  nearly  5  months  in   hard  marches   and   almost  con- 


72  NAUTILUS  Vol.  75  (2) 

tinuous  biological  collecting  with  Ekman,  who  would  quickly 
drop  a  companion  if  he  couldn't  keep  up  with  his  pace  and 
live  the  primitive  life  to  which  he  had  adapted  himself.  We 
never  hired  natives  or  pack  animals  and  we  carried  all  our 
supplies  on  our  own  backs;  we  lived  on  native  food,  drank  water 
wherever  it  could  be  found,  often  from  stinking  pools;  we 
shot  birds,  stuffed  the  skins,  and  ate  the  meat.  Guinea  fowl  are 
abundant  in  some  areas  and  the  natives  seldom  molest  them. 
Haitian  peasants  are  usually  very  poor,  but  invariably  hospitable, 
so  generally  we  slept  in  native  huts  while  collecting  in  the 
mountains  and  islands. 

At  that  time  the  American  Marines  still  occupied  Haiti  be- 
cause of  the  Caco  rebellion  a  few  years  before.  Ekman,  a  very 
eccentric  man  who  cared  nothing  about  formalities  and  hated 
shams  or  snobbishness  was,  nevertheless,  a  very  learned  man 
and  a  delightful  companion.  He  was  well  posted  on  a  wide 
variety  of  subjects,  was  conversant  in  all  the  native  dialects  and 
could  speak  at  least  8  European  languages.  He  was  known  per- 
sonally throughout  Haiti  by  much  of  the  peasantry,  the  military 
officers  and  by  the  president.  The  American  Marine  officers 
called  him  the  "Iron  Man  of  Haiti"  or  the  "Crazy  Swede,"  be- 
cause very  few  men  had  the  stamina  and  drive  to  keep  up  a 
continual  pace  of  hard  marches  into  the  rough  jungles  and 
live  on  native  food  and  sleep  wherever  he  happened  to  be 
when  darkness  fell. 

Our  regular  routine  was  to  be  up  at  4  A.M.,  start  off  imme- 
diately, but  without  a  breakfast,  eat  a  few  mangoes  or  wild 
fruits  if  we  found  them,  or  purchase  a  couple  of  pennies'  worth 
from  peasants  during  the  day,  and  at  sundown  we  usually  had 
a  dish  of  rice  and  a  boiled  fowl  and  Haitian  hard  tack  with 
plenty  of  tea  and  sugar,  around  a  small  campfire. 

Lake  Miragoane  is  quite  round  and  has  a  very  muddy  bottom 
of  unknown  depth.  It  is  completely  surrounded  by  reeds  and 
dangerously  deep  mud.  The  flora  is  particularly  interesting 
around  the  outlet  of  the  lake.  A  large  waterlily  grows  here 
with  stems  12  to  15  feet  long.  Ekman  always  knew  right  away 
when  he  found  a  new  species  of  plant  and  was  almost  always 
correct.  Here  I  found  a  new  fern  and  a  duckweed.  Numerous 
species  of  ferns  a  nd  several  beautiful  species  of  Canna  grew 


I 


October,   19(il  nautilus  73 

in  the  swamp.  While  Ekman  and  I  gazed  longingly  at  the  lake 
and  contemplated  its  floristic  possibilities  for  our  botanical  col- 
lection, wondering  how  to  find  an  area  where  we  would  not 
disappear  into  the  treacherous  mud,  we  finally  found  a  small 
boy,  who  had  a  hollowed  out  log  boat,  and  who  agreed  to  take 
us  wheie  we  wanted  to  go.  He  paddled  and  poled  the  boat 
along  the  margin  of  the  lake  until  an  ample  series  of  desired 
plants  was  made.  The  marsh  around  the  lake,  and  the  surface 
of  tiie  water,  was  a  regular  aviary  of  interesting  birds.  Glossy 
ibis,  two  kinds  of  egrets,  several  species  of  ducks,  four  kinds 
of  herons,  the  blue,  the  green,  the  Louisiana  and  night 
herons,  were  there.  Curlew,  yellowlegs,  grebes,  coots  and  sand- 
pipers and  the  curious  jacanas,  skipping  and  running  about 
on  the  lily  pads  after  insects.  To  a  real  ornithologist  or  a 
naturalist  with  proper  equipment  and  plenty  of  time,  this  lake 
has  a  great  deal  of  interest. 

On  this  special  excursion  we  asked  the  little  boy  to  put  us 
ashore  on  the  margin  of  the  lake  of  the  opposite  side.  There 
the  mud  appeared  firm  enough  to  walk  on  through  the  marsh 
to  the  land.  The  objective  was  some  rare  species  of  ferns  and 
grasses  that  could  be  seen  from  the  lake.  The  boy  left  us,  and 
we  went  our  way  to  the  shore.  Soon  we  started  to  break  through 
the  mud  crust,  and  were  floundering  around  in  the  deep  ooze 
for  some  time,  often  falling  through  the  upper  crust  only  to 
become  more  deeply  mired.  We  began  to  grow  quite  anxious 
as  to  how  we  should  get  back  to  safer  ground,  and  out  of  the 
swamp.  Suddenly  we  came  upon  a  black  woman  intently  engaged 
in  setting  snares  for  wild  fowl.  She  hadn't  noticed  us  at  all 
until  we  were  almost  upon  her  when  she  became  terribly 
frightened  at  our  appearance  out  of  the  mud.  She  screamed 
and  tried  to  run  away,  but,  after  a  small  gift  of  a  few  cigarettes, 
she  calmed  down  and  showed  us  a  sort  of  path  out  of  the  big 
swamp.  It  was  really  a  lifesaver  for  us. 

Upon  gaining  safer  ground  on  the  lake  shore,  I  found  a  few 
spots  in  shallow  water  inhabited  by  moUusks.  During  about 
one  half  hour,  spent  searching  for  freshwater  shells,  I  collected 
the  following  species,  from  which  Dr.  Clench  (1936)  and 
Clench  Sc  Aguayo  (1937)  named  a  subgenus  and  4  forms. 
Physa  (Haitia)  elegans  Dr.  Clench.  Made  type  of  a  subgenus. 


74  NAUTILUS  Vol.  75  (2) 

because  not  closely   related   to   the  other  forms  in   the  West 

Indies. 
Helisoma  eyerdnmi  Clench  and  Agiiayo. 
Helisoma  caribaea    (Orbigny) 
Tropicorbis  pallida    (C.  B.  Adams) 
Tropicorbis  albicans    (Pfeiffer) 
Drepanotrema   cimex    (Moricand) 
Drcpanolrcrna    anatiniim     (Orbigny) 
Potamopyrgus    coronatus   ariomus    Clench    &    Aguayo. 
Ferrissia    (Laevapex)    haitiana  Clench  &:  Aguayo. 

Up  to  that  time,  less  than  50  species  of  freshwater  shells  were 
known  from  the  West  Indies  and  Bahama  islands. 

Later  in  the  afternoon,  we  found  a  beautiful  waterfall  where 
we  had  a  delightful  bath,  collected  some  Tare  plants  and  snails, 
and  I  also  collected  a  rare  frog,  Eleotherodactylus  inoptatus 
(Barbour)  and  a  rare  snake  Uromacer  oxyrhyricliys  (Dumeril 
and  Bibron) . 

Several  times  before  I  had  come  alone  to  the  south  side  of  the 
lake  to  collect  specimens.  There  in  the  jungles  many  fine  land 
snails  were  taken,  including  Biilimulus,  Cepolis  and  Caracoliis, 
and  a  colony  of  Liguiis  xiirgirieiis  on  the  lignum  vitae  and 
logwood  trees  yielded  several  hundred  fine  specimens.  On  the 
ground,  amongst  these  trees,  I  also  collected  scorpions,  taran- 
tulas, centipedes  and  several  specimens  of  the  giant  Haitian 
armoured  millepede,  over  8  inches  long  and  about  one  inch 
thick.  When  disturbed  it  sprays  acid  at  its  enemies  a  distance 
of  more  than  a  foot.  The  substance  is  exuded  from  pores  along 
the  sides.  It  smells  and  burns  like  sulphuric  and  prussic  acid.  I 
did  not  realize  how  dangerous  these  creatures  could  be  with 
careless  handling  until  I  met  Dr.  Cook,  American  specialist  of 
palm  trees,  centipedes  and  millipedes  a  few  days  afterward  in 
Port-au-Prince.  Most  of  the  skin  of  one  side  of  his  face  was 
burned  off,  and  his  helper  had  one  of  his  hands  badly  burned 
as  a  result  of  heedless  collecting  of  these  large  millipedes. 

No  serious  work  has  ever  been  done  in  collecting  the  fauna 
of  lake  Miragoane  except  a  casual  excursion  made  by  Dr.  Wm. 
Beebe  in  1927  a  lew  weeks  before  I  came  to  Haiti  when  he 
discovered  two  new  endemic  fishes.  One  of  them,  a  tiny  guppy 
about  14  inch  long,  fully  grown,  was  rather  common  in  a 
ditch  near  the  lake.  With  its  high  degree  of  endemism,  a  careful 
collecting    survey    of    its    limnological    fauna    by    a    couple    of 


October,  1961  nautilus  75 

competent  biologists  should  bring  to  light  a  surprising  number 
of  new  and  interesting  species  of  animals. 

LiTF.RATURF.    iMV    > 

Clench.  William  J.  1936.  Mem.  Soc.  Cuba  Hist.  Nat.  /0:335-342, 
pi.  25. 

&  C.  C.  Aguayo.  1937.  Mem.  cit.  //:61-76,  pi.  7. 

Cochran,  Doris   M.   1928.   Biol.   Soc.   Washington  41:53-60. 

APPARENT  GEOTACTIC  BEHAVIOR  IN  PHYSA 

By  ANDREW  McCLARY 

University  of  Wisconsin-Milwaukee 

This  note  describes  an  experiment  in  which  snails  of  the  genus 
Pliysa,  identified  as  P.  Integra  Haldeman^  were  tested  for  geotac- 
tic  behavior  in  a  mud  medium. 

Method.  Six  populations  of  P.  Integra,  collected  from  the 
locations  given  in  Table  1,  were  used  in  the  study.  Experiments 
were  conducted  between  September  25  and  Octobei-  8,  1960.  Each 
population  was  collected  early  on  a  given  day,  and  tested  in 
the  afternoon  or  evening  of  the  same  day.  The  medium  in  which 
the  snails  were  tested  was  a  silty  mud  obtained  from  the  banks 
of  the  Milwaukee  River  at  Estabrook  Park,  on  the  north  side 
of  the  city  of  Milwaukee.  The  populations  designated  as  "2", 
"4"  and  "5"  in  Table  1  were  collected  in  the  same  area  as  the 
mud.  The  mud  was  washed  in  tap  water,  sieved  through  a  fine 
mesh  screen,  and  allowed  to  dry  to  a  tacky  consistency.  Some 
oligochaetes  of  the  family  Tubificidae  were  present  in  the  mud 
after  sieving;  no  other  forms  of  life  were  obvious.  Before 
testing  a  given  population,  tap  water  was  added  to  a  portion  of 
the  mud,  which  was  then  used  in  the  test  as  described  below. 
In  some  cases,  the  mud  utilized  had  been  employed  in  previous 
tests,  in  other  cases,  fresh  mud  was  used. 

Twenty  snails  were  selected  at  random  from  the  population 
to  be  tested,  each  snail  being  measured  at  the  time  of  selec- 
tion. The  average  shell  length  of  the  animals  used  was  about 
7mm.  Each  snail  was  placed  alone  in  a  150  ml.  glass  beaker 
filled  to  a  level  of  4mm.  with  mud.  Although  the  orientation 
of  the  snails  was  not  controlled,  most  animals  appeared  to  land 
in  an  upright  postion.  Care  was  taken  to  place  the  snails  in  the 


1  The  writer  wishes  to  thank  Dr.  H.  van  der  Schalie,  Museum  of  Zoology, 
University  of  Michigan,  for  kindly  identifying  the  specimens. 


76  NAUTILUS  Vol.  75  (2) 

middle  of  the  mud  surface.  Mud  was  then  added  over  the 
snail  in  each  beaker  to  form  a  mud  column  totaling  50mm.  in 
height.  The  mud  column  was  typically  of  a  consistency  such  that 
after  a  half  hour  approximately  5mm.  of  water  had  collected 
above  its  surafce.  At  the  start  of  a  test,  each  snail  was  therefore 
immersed  in  a  mud  medium  at  an  approximate  distance  of 
4mm.  from  the  beaker  floor,  25mm.  from  the  beaker  wall,  and 
45mm.  from  the  mud  surface.  The  20  beakers  were  then  placed 
on  a  table  4  feet  below  a  100  watt  lamp.  Room  temperature 
during  the  tests  ranged  from  19-22°C.  Beakers  were  inspected 
at  5  minute  intervals,  mud  surface,  beaker  wall,  and  beaker  floor 
being  watched  for  the  appearance  of  snails.  As  the  mud  medium 
was  homogeneous  in  texture,  snails  appearing  could  readily  be 
identified.  Each  beaker  was  observed  for  a  period  of  two  hours. 
If  a  snail  had  not  appeared  by  the  end  of  this  period,  its 
position  in   the  mud  was  ascertained  by   probing. 

Results.  Of  the  total  of  120  snails  used  in  the  experiment, 
61  surfaced  on  the  mud.  Surfacing  usually  occurred  at  a  point 
roughly  between  the  center  of  the  beaker  and  the  beaker  wall 
so  that,  assuming  travel  in  a  straight  line,  the  61  snails  moved 
towards  the  surface  along  a  path  that  was  at  an  average  angle 
of  70°  from  the  horizontal.  The  average  time  lapse  between 
start  of  a  run  and  surface  appearance  was  42  minutes.  Eight  of 
the  120  snails  were  recorded  as  appearing  at  beaker  walls  at 
points  ranging  from  half  way  up  the  beaker  wall  to  just  below 
the  mud  surface.  Again  assuming  travel  along  a  straight  line, 
snails  moving  towards  a  beaker  wall  did  so  along  a  path 
that  was  at  an  average  angle  of  50°  from  the  horizontal.  Average 
time  lapse  between  start  of  a  run  and  appearance  at  a  beaker 
wall  was  6.^  minutes.  Fifty-one  of  the  102  test  snails  did  not 
appear  at  either  the  surface  or  the  wall  of  a  beaker.  Probing 
indicated  that  15  of  these  had  moved  up  through  the  mud 
medium,  having  traveled  from  I/3  to  14  the  distance  to  the  sur- 
face. The  remaining  36  snails  were  found  to  be  in  approximately 
the  location  where  they  had  been  placed  at  the  start  of  the 
two-hour  period. 

Snail  size  did  not  appear  to  be  a  significant  factor  in  the 
behavior  shown.  The  consistency  of  the  mud  medium  differed 
slightly  between  tests,  and  this  may  have  been  a  partial  cause 


October,   1961 


NAUTILUS 


77 


of  the  behavior  variations  found  between  populations.  Possibly 
also  more  exact  tests  would  reveal  actual  behavior  differences 
between  snails  of  different  populations. 
A  summary  of  the  results  is  given  in  Table  1. 

Table  1 


Behavior  < 

3f  Physa 

in  a  Mud  Medium 

Date 
Tested 

No. 

of  Snails  Recovered  in  Each 
Four  Locations 

of 

Population 

Number 

Mud       Part  Way 
Surface    to  Surface 

Beaker 
Wall 

Original 
Location 

Total 

1 

9-25-60 

8 

2 

0 

10 

20 

2 

9-30-60 

Hi 

3 

1 

2 

20 

3 

10-2-60 

11 

2 

1 

6 

20 

U 

10-7-60 

12 

2 

2 

I 

20 

? 

10-8-60 

9 

1 

2 

8 

20 

6 

10-8-60 

7 

? 

2 

6 

20 

Total 

61 

15 

8 

36 

120 

Collection  sites  of  popiilations  were  as  follows.  1:  Aquarluw  in  Botany  Department 
at  University  of  Wisconsin — Milwaukee.  2,  h,  $:   Milwaukee  River  bank  at  Estabrook 
Park,  Milwaukee  County.  3:  Stone  fountain  at  Whltnall  Park,  Milwaukee  Coonty. 
6?  Stream  at  Kletzsch  Park,  Milwaukee  County. 

Discussion.  Possibly  vertical  gradients  involving  physical- 
chemical  properties  exist  in  a  mud  medium  and  serve  to  guide 
an  immersed  snail  to  the  surface  of  the  mud.  Gradients  of  this 
type  might  conceivably  involve  differences  in  temperature,  light, 
oxygen,  or  in  the  concentration  of  various  organic  or  inorganic 
materials  in   the  mud. 

An  alternate,  and  perhaps  more  likely,  explanation  of  the 
upward  movement  of  the  snails  in  the  present  experiment  is 
that  the  mud  itself  provided  no  cue  as  to  direction,  but  that 
the  movement  was  a  true  geotactic  behavior,  or  response  to 
gravity. 

Snails  are  usually  considered  to  orient  to  gravity  either  by 
statocysts,  organs  designed  to  ascertain  position  in  space;  or  by 
proprioceptors,  sensitive  to  the  differential  stresses  in  muscle 
which  occur  in  an  organism  subjected  to  the  pull  of  gravity 
(Carthy,    1958). 

While  statocysts  are  present  in  snails  (Bouvier,  1887;  Lacaze- 
Duthiers,   1872) ,  these  organs  have  not  been  clearly  shown  to 


78  NAUTILUS  Vol.  75  (2) 

I'unction  in  gcotaciic  behavior,  for  the  experimental  work  which 
has  been  done  in  this  respect  is  open  to  criticism,  having  proved 
difficult  to  repeat  (Pieron,  1928;  Crozier  and  Navez,  1930).  On 
the  other  hand,  a  strong  case  has  been  made  for  the  existence 
of  a  proprioceptive  control  of  snail  geotactic  behavior.  Experi- 
ments have  shown  that  snails  which  normally  show  negative 
geotactic  movement  will  change  their  direction  of  travel  if 
muscle  stress  becomes  greater  on  one  side  of  the  body  than 
the  other.  This  latter  condition  can  be  brought  about  by 
attaching  a  thread  to  the  shell  apex  and  gently  twisting  the 
shell  out  of  line  with  the  snail's  body.  Snails  so  manipulated 
turn  so  that  their  body  lies  along  the  same  axis  as  the  shell, 
and  travel  in  the  new  direction  determined  by  this  axis,  even 
if  the  new  direction  is  one  that  is  positively  geotactic.  It  has 
been  concluded  from  experiments  of  this  sort  that  snails  show- 
ing negative  geotactic  behavior  have  oriented  themselves  so 
that  there  is  an  equal  muscle  tension  on  both  sides  of  the  body, 
as  would  presumably  be  true  when  the  shell  hangs  ventrally. 
A  snail  traveling  in  any  direction  but  upward  woidd  presumably 
experience  unequal  muscle  tension  due  to  the  weight  of  the 
shell   (Crozier  and  Navez,  op.  cit.) . 

While  possibly  snails  may  orient  to  gravity  by  this  method 
in  air  or  water,  orientation  of  this  kind  would  seem  less  likely 
to  occur  in  a  medium  such  as  mud.  In  a  mud  medium,  there  is 
probably  very  little  tendency  for  a  snail's  shell  to  sink  faster 
than  its  body  and  so  create  muscle  tension,  forcing  the  body  to 
orient  upward.  This  is  almost  certainly  true  when  a  snail  is  at 
rest  in  a  mud  medium,  and  is  probably  true  to  a  large  extent 
when  a  snail  is  moving  through  the  medium,  although  this 
remains  to  be  verified.  In  any  case,  the  results  of  the  present 
experiment  indicate  that  a  re-examination  of  the  possible  role 
of  statocysts  in  snail  orientation  seems  desirable. 

The  utility  of  negative  geotactic  behavior  in  some  species 
of  snails,  such  as  the  tree  forms,  may  be  that  of  enabling  an  ani- 
mal to  regain  its  normal  habitat  after  accidental  dislodgement 
(Crozier  and  Navez,  op.  cit.) . 

In  aquatic  pulmonates,  negative  geotactic  behavior  may  be  a 
mechanism  which  sends  an  animal  to  the  surface  to  gain 
oxygen    (Cheatum,    1934) ,  although  other  factors  may  also  be 


October,   1901  nautilus  79 

involved,  as  many  pulmonates  appear  never  to  surface  (Russell- 
Hunter,  1953).  The  negative  geotactic  behavior  (if  such  it 
should  prove  to  be)  described  here  may  have  been  a  response 
to  oxygen  loss,  the  latter  perhaps  resulting  from  sudden  immer- 
sion in  the  mud  medium.  Possibly  also  a  medium  such  as 
mud  will  stimulate  receptors  in  all  or  a  part  of  the  snail's  body 
and  that  negative  geotactic  movement  results  from  the  stimula- 
tion. This  latter  possibility  is  strengthened  by  the  fact  that, 
wliile  no  quantitative  records  were  kept,  snails  often  ceased 
their  upward  movement  as  soon  as  they  had  surfaced  on  the 
mud,  although  they  still  had  no  direct  access  to  air,  being 
below^  the  surface  of  the  water  which  was  above  the  mud. 

References 
Bouvier,  E.  L.   1887.  Systems  nerveux,  morphologie  generale  et 

classification    des   gastropodes    prosobranches.    Ann.    Sc.    Nat. 

Zool.  (7)  iii:  1-570. 
Carthy,  J.  D.  An  introduction  to  the  behavior  of  invertebrates. 

New  York.  1958. 
Cheatum,  E.  P.  1934.  Trans.  Amer.  Microsc.  Soc.  55:318-407. 
Crozier,  W.  J.  and  Navez,  A.E.  1930.  Jour.  Gen.  Psychol.  5:3-37. 
Lacaze-Duthiers,   H.    1872,  Arch.   Zool.   Exp.   Gen.   7:97-166. 
Pieron,  H.  1928.  Jour.  Genet.  Psychol.  55:3-17. 
Russell-Hunter,  W.    1953.   Proc.   Roy.   Soc,  Edinburgh.   ^^5  (ii)  :- 

143-165. 

LOUISIANA  LAND  SNAILS  WITH  NEW  RECORDS 

By  dee  SAUNDERS  DUNDEE  and  PATTI  WATTi 
Louisiana  State  University  in  New  Orleans 

Collections  made  during  the  past  year  have  revealed  three 
mollusks  not  previously  reported  in  Louisiana. 

Gulella  bicolor  (Hutton) ,  known  previously  from  this  area 
by  Dr.  Harold  Harry  but  not  reported  by  him  (personal  com- 
munication, 1959),  has  been  taken  from  20  localities  in  and 
around  New  Orleans.  Nowhere  is  it  abundant;  colonies  appear 
to  consist  of  few  individuals  which  are  found  only  after  rains 
or  in  very  damp  habitats. 

A  small  colony  of  Praticolella  griseola  (Pfeiffer)  was  dis- 
covered in  the  yard  of  the  Laclede  Steel  Corporation  on  France 


1  This  paper  is  an  outgrowth  of  a  current  investigation  supported  by  a 
research  grant,  RG  7194,  from  the  National  Institutes  of  Health,  Public 
Health   Service. 


80  NAUTILUS  Vol.   75   (2) 

Road  in  New  Orleans.  We  have  found  it  nowhere  else  even 
though  extensive  searches  have  been  made.  At  this  locality  it  is 
living  in  and  around  a  pile  of  steel  girders.  The  pile  shows  signs 
of  having  been  there  undisturbed  for  some  time.  It  is  part  of 
a  barge  which  used  to  make  trips  to  Texas.  Possibly  this  is  how 
this  colony  came  to  be  here.  It  is  an  interesting  find  in  view  of 
Pilsbry's  statement  (1940,  p.  689)  concerning  Praticolella  dis- 
tribution: "The  herd  of  eastern  Mexico  and  Texas  is  separated 
from  that  of  the  southeastern  states  by  the  Mississippi  River  and 
State  of  Louisiana,  300  miles  more  or  less." 

Vaginula  sp.  (?)  first  came  to  our  attention  while  collecting 
in  Mobile,  Alabama.  There  it  is  found  in  large  localized  colonies. 
Up  to  that  time  we  had  not  discovered  it  in  Louisiana  (probably 
because  of  a  dry  period),  but  shortly  thereafter  we  began  finding 
it  in  New  Orleans.  To  date  it  is  known  from  4  localities  in  the 
city. 

Since  little  is  known  of  the  mollusks  of  Louisiana,  we  consider 
it  useful  at  this  time  to  publish  a  list  of  the  terrestrial  mollusks 
which  have  been  reported. 

In  this  list  letters  following  localities  indicate  the  source  of 
the  information  as  follows:  (G)  =  Goodrich,  (Ha)  =  Haas, 
(HH)  =  Harry,  (H)  =  Hubricht,  (?)  =  Pilsbry,  (T)  =  Taylor, 
(V)  =  Vanatta,  (Vi)  =  Viosca.  Localities  are  parishes  unless 
otherwise  noted. 

Helix  (ispefsa  (Miiller) — Orleans,  East  Baton  Rouge  (HH,  P, 
Vi).2 

H.  aperta  Born— Orleans    (HH,  Vi).^ 

Otala  vermiculata   (Miiller)  — Orleans.    (P)  ^ 

O.  lactea  Miiller— Orleans    (HH)  .2 

Polygyra  septemvola  febigeri  (Bland) — Orleans,  Vermillion  (P)  ; 
Plaquemines  (HH). 

P.  auriformis  (Bland)  — Orleans,  Calcasieu   (P) . 

P.  leporina   (Gould)— DeSoto   (P)  ;  Grant   (H)  ;  Catahoula   (V) . 

P.  triodontoides  (Bland)  — Calcasieu  (P) . 

P.  texasiana  (Moricand) — DeSoto  (P,  HH)  ;  Grant,  Rapides, 
Caddo,  Orleans,  Calcasieu,  Nachitoches,  Tensas    (HH)  .-^ 

P.  dorjeuilliana  Lea — Nachitoches,  DeSoto,  Bienville  (P)  ;  Clai- 
borne (H) . 

2  These  were  no  longer  found  in  1948  (Harry)  and  we  have  found  none 
of  them  in  Orleans  Parish. 

3  Specimens  may  be  P.  triodontoides   (Harry,  1951) . 


Odobci.    HXil  NAUTILUS  81 

Stcnoticnut  lahio.stnii    (lihmcl) — -Rapides,  I'nioii    (P). 

S.    sIcnotreiiKi     (Plcillcr)  — Madison,    Rapides    (P)  . 

S.     nionodoii    (illciac     (Pilsbry) — DeSoio,     Bienville,     (Calcasieu 

(lype  loealiiy    (P):   Evangeline,  Allen,  Vernon,  Rapides    (H)  . 
S.   iiionodon   loiiu  f)  icrsoni  Pilsbry — -Catahoula     (V)  . 
Mcsodon     tliyroidus     (Say) — Morehouse,     I'laukiin.     Calahoida 

(\'):   Caleasieu,  Grant,  Ouaehita    (H)  . 
M.  nijlfitus  (S'ly) — Morehouse,  Franklin   (V)  ;  (Caleasieu   (H,  P)  ; 

Lxangeline,  Vernon,  Grant,  LaSalle    (H)  . 
Triodopsis  vulluosa    (Goidd) — C^alcasieu    (P)  . 
T.  oagini  Call — Naehitoches    (P)  ;  Grant,  Claiborne    (H)  . 
T.  carolifiioisis    (Lea) — -DeSoto,  Caddo    (P)  . 
T.  jostcri   (F.  C^.  Baker) — Lake  Pontchaitraiu    (no  parish  given) 

/.  (ilhohihris  (Say) — included  in  the  range  given  by  Pilsbry — 
no  dehnite  localities  listed    (P):  Oua(  iiiia    (H)  . 

T.  dn'csta    (Gould) —DeSoto    (P)  . 

I Idplotrciiui  concavum    (Say) — LaSalle    (H) . 

Hiilniiiiliis  dealbatiis    (Say) — DeSoto    (P)  . 

liiDiniiti  decollata  (Linnaeus) — Orleans  (Ci,  P,  Vi)  ;  East  Baton 
Rouge   (HH). 

I junellaxis  gracilis  (Button) — Orleans  (HH,  P)  ;  East  Baton 
Rouge   (HH). 

I:iigl(indinti  rosea  (Feussac) — Orleans,  Franklin,  Iberia,  Iber- 
ville. \V.  Carroll    (P)  :  Plaquemines    (HH)  . 

J.,  rosea  biillata  (Gould) — Orleans,  St.  Landry,  Lake  Pontchar- 
train  (no  parish  given)  ,  Iberia  (P);  Franklin  (P,  V)  . 

liuconulus  chersinus  (Say) — State  of  Louisiana  (P)  ;  Catahoula 
(V);   Plaquemines    (HH)  . 

/:'.   cliersi)ius   trocJndus    (Reinhardt) — Caddo    (P)  . 

E.   chersinus   dentatus    (Sterki) — Catahcjula     (\');    Rapides    (P)  . 

Guppya  sterkii    (Dall) —Bienville    (?) . 

Retinella  indentata  (Say) — LaSalle  (H);  Morehouse,  Catahoula 
(V)  ;   Plaquemines    (HH). 

''.  indentata  paucilirata  (Morelet) — State  of  Louisiana  (P)  ; 
C:iaiborne    (P,  H)  ;   Morehouse    (V). 

Mesonipliix  friabilis    (W.   G.   Binney) — Morehouse    (P,   V). 

M.  inilgatus  H.  B.  Baker — Calcasieu,  Iberia,  Morehouse,  Franklin 
(P)  ;    Vernon,    LaSalle    (FI);    Morehouse,    Franklin     (\')  . 

Haivaiia  rniniscula  (Binney) — included  in  the  range  given  by 
Pilsbry — no  definite  localities  given  (P)  ;  Plaquemines  (HH); 
.Morehouse,  Catahoula    (V) . 

I'entridens   demissus    (Binney) — Vernon,    Grant,    LaSalle    (H)  ■ 

/'.  demissus  brittsi    (Pilsbry) — DeSoto,  Bienville,  Calcasieu    (P). 

/'.  intertextus  (Binney) — Calcasieu,  Claiborne.  Evangeline,  La- 
Salle   (H)  ;  Bienville    (P). 


82  NAUTILUS  Vol.  75  (2) 

Zonitoidcs  arboreus  (Say)  — State  of  Louisiana  (P)  ;  Morehouse, 
Richland,  Ouachita,  Franklin,  Catahoula  (V)  :  Plaquemines 
(HH) . 

Limax  fiaviis  Linnaeus — Lincoln    (T). 

Limax  marginatiis  Miiller — Caddo    (HH). 

Deroceras  sp. — Plaquemines    (HH) . 

Deroceras  Uicvc  Miiller — East  Baton  Rouge.  Ciaddo,  DeSoto 
(HH). 

Angiaspira  (iltern/iln  (Say) — Catahoula  (P,  V):  Morehouse, 
Franklin  (V). 

A.  alternata  crassa  Walker — Caddo,  Franklin,  Morehouse,  De- 
Soto (P)  ;  Claiborne,  Ouachita,  Grant,  Calcasieu,  Evangeline, 
Vernon    (H)  . 

HeUcodiscus  singlcyanus  (Pilsbiy) — Morehouse  (P,  \^)  ;  Oua- 
chita  (V). 

H.  singleyaniis  inermis  H.  B.  Baker — Morehouse,  Ouachita    (P). 

Pliilomycus  carolinianus  (Bosc) — Calcasieu,  Evangeline,  Rich- 
land, Ouachita   (H) . 

P.   carolinianus  flexuolaris  Rafinescjue — Grant    (P)  . 

Fallifera   marmorea    (Pilsbry)  — Vernon,  Grant    (H)  . 

Oxylonia  sallaena     (Pteifter)  — Orleans,   DeSoto    (P). 

Succinca   nnicolor  Tryon — Orleans    (type  locality)     (P). 

.S'.  grusx'etwri  Lea — Rapides    (P) . 

S'.  lutcula  Gould — Vermillion,  Plaquemines    (HH) . 

S.  concordialis  Gould — Concordia    (type  locality)     (P)  . 

Strobilops  tcxasiana  (Pilsbry  and  Ferriss) — Morehouse,  Rapides 
(P.  V). 

S.  labyrinthica  (Say) — Morehouse,  Catahoula  (V). 

S.  aenea  Pilsbry — State  of  Louisiana  (P)  ;  Morehouse,  Richland, 
Catahoula,  Franklin;  form  micrompliala:  Rapides,  Morehouse, 
DeSoto,  Richland,  Franklin,  Catahoula    (P)  . 

Gastrocopta  contracta  clnneana  (Vanatta) — Orleans,  Franklin 
(V)  ;  Morehouse,    (P,  V)  ;  Plaquemines    (HH)  . 

G.  corticaria   (Say) — Rapides   (P). 

G.  pentodon  Say — Plaquemines   (HH)  . 

G.    tappaniana     (C.    B.    Adams)  — INIorehouse,    Franklin     (V)  . 

G.  rupicola  (Say)  — included  in  the  range  given  by  Pilsbry — no 
definite    localities   given     (P)  :    Plaquemines     (HH)  . 

G.  pelliicida  hnrdaccUa    (Pilsbry) —Gulf  States    (P). 

Pupoides  albilabris   (C.  B.  Adams)— Gulf  States    (P)  . 

P.  modicns  Gould — Placpiemines    (HH)  . 

Vertigo   inilium   Gould — Plaquemines    (HH)  . 

V.  oscariana  Sterki — Madison    (P)  . 

V.  rugosula  Sterki — State  of  Louisiana    (P)  ;   Morehouse    (V)  . 

IJcUcina   orbiculata    (Say) — DeSoto,  Jefferson,   Orleans    (P)  . 

Bradybaena  similaris — (Ferussac) — Orleans    (Ha,   HH,  G). 


October,  1961  nautilus  83 

Literature  cited 
Goodrich,  C.  1940.  Naut.  53  (3)  :  105. 
Haas,  F.  1945.  Fieldiana.  Zoology  31:2,  3-14. 
Harry,  H.  VV.  1942.  Occ.  Papers  of  La.  State  Univ.  Marine  Lab., 

No.   1. 

1948.  Naut.  62  {\)  :  20-24. 

1951.  Naut.  64{?,):  96-99. 

Hubricht,  L.  1956.  Naut.  69(4)  :  124-26. 

Pilsbry,  H.  A.  1939-41.  Land  Mollusca  of  North  America   (North 

of  Mexico)  Acad.  Nat.  Sci.  Monographs  3,  vols.  1  &  2. 
Taylor,  W.  E.  1899.  Gulf  Fauna  and  Flora  Bull.,  1  (3):69-73. 
Vanatta,  E.  G.  191 1.  Proc.  Acad.  Nat.  Sci.  Phil.:  525-531. 
Viosca,  P.  1928.  Naut  41  (4):  139-40. 

THE  AMERICAN  MALACOLOGICAL  UNION 

Twenty-eighth  Annual  Meeting,  June  20-23,  1961 

For  the  third  time  in  its  30  year  history,  the  American  Mala- 
cological  Union  convened  at  the  U.  S.  National  Museum  in 
VV^ashington,  D.  C.  Attendance  broke  all  records  (150)  while  the 
four-day  program  rolled  smoothly  along  thanks  to  careful  plan- 
ning and  the  dedicated  services  of  volunteer  members  of  the 
National  City  Shell  Club.  Behind  the  scenes  was  Dr.  Harald 
A.  Rehder  together  with  his  henchmen  Drs.  Morrison  and  Rose- 
water,  all  veterans  of  past  A.M.U.  meetings,  therefore  anticipating 
the  requirements  of  their  guests  well  in  advance. 

President  Thomas  E.  Pulley  presided  over  the  academic  ses- 
sions; it  was  due  to  his  vigilance  in  enforcing  time  limits  that 
every  paper  on  an  unusually  full  program  was  heard. 

An  evening  garden  party  at  the  beautiful  home  of  Mr.  and 
Mrs.  A.  Lothrop  Lutrell  in  Rockville,  Maryland  was  the  high- 
light of  opening  day,  and  on  Thursday  the  annual  dinner  was 
held  at  the  Cumberland  Country  Club  where  food,  service  and 
sumptuous  surroundings  left  nothing  to  be  desired. 

The  Executive  Council  met  on  Wednesday  evening  and  at  the 
annual  business  meeting  on  the  following  afternoon  it  was  an- 
nounced that  it  had  been  found  necessary  to  raise  the  annual 
dues  one  dollar  per  year  with  joint  and  life  memberships  to  be 
advanced  accordingly.  (This  advance  will  not  become  effective 
until  1962.) 

No  site  had  been  selected  for  the  1962  meeting,  but  a  com- 
mittee has  been  appointed  to  consider  the  matter  and  it  is  ex- 


84  NAUTILUS  Vol.  75  (2) 

pected  that  the  time  and  place  will  be  announced  in  the  1961 
report  bulletin. 

The  following  slate  of  officers  for  the  coming  year  was  pre- 
sented by  the  nominating  committee  and  elected  by  unanimous 
vote  of  the  members  present: 

President,  William  K.  Emerson. 

Vice-president,  Albert  R.  Mead. 

2nd  Vice-president,  Robert  W.  Talmadge. 

Secretary-treasurer,  Margaret  C.  Teskey. 

Publications  Editor,  Morris  K.   Jacobson. 

Councillors-at-Large,   John  B.  Burch,  H.  B. 

Herrington,  Edward  H.  Michelson,  Virginia  Orr. 

The  field  trip  on  Friday  was  a  50  mile  excursion  to  Scientists 
Cliffs  on  Chesapeake  Bay  where  the  clay  banks  of  the  Calvert 
formation  yield  a  seemingly  endless  series  of  Miocene  fossils. 
The  trek  back  to  the  city  was  the  final  featine  of  the  1961  meet- 
ing, another  to  be  added  to  the  long  list  of  memorable  reunions 
of  the  AMU.  —  Margaret  C.  Teskey 


NOTES  AND  NEWS 


Beckianum — New  genus  (or  subgenus  of  Leptinaria  Beck, 
1837?)  in  Achatinidae  (Subulininae) .  Type  species  is  Synopeas 
beckianum  (Pfeiffer)  H.  B.  Baker,  1945,  Naut.  58:9\;  1947,  Naut. 
61  (1)  :  plate  1,  from  Cordoba,  Mexico.  As  stated  by  Zilch,  1959, 
Handb.  Palazool.  <5(2):351,  Synopeas  Jousseaume,  1889,  is  a 
homonym  of  Foerster,  1856. 

As  pointed  out  in  1945:88,  this  unique  species  is  very  distinct 
from  either  Opeas  (Ferussaciinae)  or  Lamellaxis  and  Leptinaria, 
sensu  stricto  (Subulininae)  but,  from  its  shell  sculpture,  it  may 
belong  near  Pelatrinia  Pilsbry,  1907  (1):324,  which  was  proposed 
as  a  subgenus  of  Leptinaria.  Pelatrinia  has  a  larger,  more  conoid 
and  imperforate  shell,  which  carries  its  columellar  cord  into  the 
aperture. — H.  Burrington  Baker. 

Maxwell  Smith — Dr.  Joshua  L.  Baily,  Jr.,  writes  from  Ashe- 
ville,  N.  C,  that  Maxwell  died  there  Sept.  12,  and  that  a 
biographic  sketch  will  be  prepared  for  a  future  number. — H.B.B. 

Southern  California  slugs,  additional  locality  records. — The 
following  records  are  here  listed  because  of  their  importance  in 
establishing  knowledge  of  distribution. 

Lehrnannia   poiricri     (Mabille)  .    Santa    Cruz    Island,    Stanton 


October,  1961  nautilus  85 

Ranch  headquarters  (introduced) ,  23-1-1960.  Dr.  Carey  Stanton! 
Prol)abIy  all  records  of  Limax  marginatvs  iMiill.  from  North 
America,  and  certainly  all  California  records  of  same  should  be 
referred  to  this  species^. 

Anadenulus  cockerelli  (Hemphill).  Kern  County,  soutli  of 
sawmill  at  head  of  Tejon  Canyon,  Tehachapi  Mts.,  altitude 
about  5,300  feet,  3-V-1958.  W.  O.  Gregg,  W.  B.  Miller! 

Ariolimax  coliimbianiis  stromineus  Hemphill.  Ventura  County, 
Santa  Paula  Canyon,  altitude  about  2,700  feet,  29-V-1955.  Ed 
Peterson! 

Hesperarion  niger  (J.  G.  Cooper) .  Kern  County,  west  of  Poso 
Creek  and  about  one  half  mile  north  of  Glenville — Woody  road, 
8-VI-1945.  Under  oak  log  in  damp  meadow.  W.  O.  Gregg,  M.  L. 
Walton!  This  is  the  record  which  Dr.  Pilsbry-  refers  merely  to 
"northern  Kern  Co."  Tulare  County,  along  Arrastre  Creek,  about 
3  miles  east  of  White  River,  altitude  about  2,200  feet,  26-V-1957. 
W.  O.  Gregg,  W.  B.  Miller,  W.  B.  Miller  Jr.! 

Hesperarion  hemphilli  (W.  G.  Binney) .  Santa  Cruz  Island, 
Canada  del  Puerto,  1.3  mile  from  Prisoners  Harbor,  17-1-1960. 
Found  under  old  log.  W.  B.  M.,  W.  O.  G.!  Santa  Barbara  County, 
along  Salsipuedes  Creek  near  Calif.  Hwy.  1,  about  3.4  miles 
south  of  junction  with  Hwy,  150,  10-III-1951.  W.  O.  G.,  W.  B.  M., 
W.  B.  M.  Jr.!  San  Luis  Obispo  County,  near  tributary  of  San 
Luis  Obispo  Creek  at  old  ranch  road  .8  mile  north  of  U.  S.  Hwy. 
101,  south  side  Cuesta  Pass,  Santa  Lucia  Mts.,  13-n-1960.  W.  O. 
G.,  W.  B.  M.! 

Binneya  notnbilis  J.  G.  Cooper.  Santa  Barbara  Island,  El 
Primero  Canyon,  19-IV-1959.  W.  O.  G.,  W.  B.  M.!  Shells  only, 
but  with  epidermis  remaining  on  some  shells.  Very  dry  at  time 
of  collecting.  —  Wendell  O.  Gregg. 

Mass  movement  of  a  colony  of  the  mud  snail  Ilyanassa. — In 
a  survey  of  the  marine  life  at  Cape  Ann,  Massachusetts,  a  dense 
colony  of  Ilyanassa  obsoleta  was  discovered  on  a  mud  flat  border- 
ing a  salt  marsh  about  midway  up  Little  River  in  1933.  Little 
River  is  a  side  channel  to  the  Annisquam  Tidal  River,  a  marine 
inlet  which  divides  the  Cape  Ann  Promontory.  The  habitat  was 
very  soft,  black,  sticky  mud.  The  colony  in  question  occupied 
an  area  about  85  feet  wide  and  extended  outward  from  the  edge 
of  the  salt  marsh  for  some  80  feet  downshore.  In  1935  a  series  of 
quadrat  counts  gave  a  range  of  25-61  with  an  average  of  43  snails 


1  Quick,  H.  E.,  Proc.  Mai.  Soc.  London,  vol.  29,  pt.  5,  pp.  181-189;  British 
Slugs'.  Bull.  British  Museum,  Zool.,  vol.  6,  no.  3,  p.   197-200. 

-  I'ilsbi7,  H.  A.,  Land  MoUusca  of  North  .America,  vol.  2,  pt.  2,  p.  724. 


86  NAUTILUS  Vol.  75  (2) 

per  quarter  square  meter.  Measurements  the  next  year  indi- 
cated the  same  abundance.  In  1956,  quadrat  counts  ranged  from 
7-36  with  an  average  of  21  snails.  Counts  made  the  next  two  years 
gave  a  range  of  9-53  and  an  average  of  26  in  1957;  a  range  of 
8-64  with  an  average  of  26  in  1958.  I  observed,  however,  that  a 
band  of  snails,  usually  not  far  from  the  marsh,  had  a  density  of 
about  3  snails  per  square  inch.  This  band  of  maximum  density 
seemed  to  shift  with  fluctuations  of  tidal  levels  between  spring 
tides  and  neap  tides.  The  5  quadrat  levels  at  which  the  counts 
were  made  and  are  quoted  above,  never  fell  on  this  narrow  band 
of  maximum  density.  In  1960  this  colony  was  first  visited  on 
August  6.  The  water  level  at  the  time  of  observation  did  not 
permit  an  extensive  sample,  but  abundance  seemed  to  be  about 
the  same  as  in  the  past  4  years.  On  September  6,  however,  a 
single  snail  was  found  in  the  area  formerly  occupied  by  the 
colony  described  above.  A  search  disclosed  that  the  entire  colony 
had  moved  to  a  new  area  some  150  feet  northeastward,  to  the 
entrance  of  a  ditch  which  cuts  back  into  the  marsh.  No  explan- 
ation seems  apparent  for  this  mass  movement.  —  Ralph  W. 
Dexter,  Department  of  Biology,  Kent  State  University,  Kent, 
Ohio. 


PUBLICATIONS  RECEIVED,  1960 

Pages  in  italics  include  new  taxons 

Benthem  Jutting,  W.  S.  S.  van.  Non-marine  Mollusca  of  the  lime- 
stone hills  in  Malaya.  Some  zoogeographical  remarks  on  the 
non-marine  Mollusca  of  the  Moluccas.  Proc.  Cent.  &  Bicent. 
Congress  of  Biology,  Singapore,  1958:  63-68,  4  figs.;  164-169, 
1   fig. 

Chernin,  Eli,  Edward  H.  Michelson  &  Donald  L.  Augustine. 
Daubaylia  potomaca,  a  nematode  parasite  of  Helisoma  tri- 
volvis,  transmissible  to  Australorbis  glabratus.  J.  Parasitology 
-^^.•599-607    (2  pis.). 

Michelson,  Edward  H.  Chemoreception  in  the  snail  Australorbis 
glabratus.  Amer.  J.  Tropic.  Med.  &  Hygiene  9:480-487,  3  figs. 

1961 

Abbott,  R.  Tucker.  The  genus  Lnmbis  in  the  Indo-Pacific.  Indo- 
Pacific  Mollusca  1  (3)  '.28  pp.,  18  pis.   (1  colored) ,  maps. 

Abbott,  R.  Tucker.  How  to  know  the  American  marine  shells. 
Pp.  222,  12  col.  pis.,  many  text  figs.  Signet  Key  Book,  P.  O.  Box 
2310,  Grand  Central  Station,  New  York  17,  N.  Y.  {Ibi  +5«s 
postage) . 


THE  NAUTILUS 

Vol.   75  January,   1962  No.   3 

THE  DISTINCTION  BETWEEN  ACTEOCINA  CANDEI 
AND  RETUSA  CANALICULATA 

Bv   HARRY  W.  WELLS  and  MARY  JANE  WELLS 
Deparimeni  of  Biological  Sciences,  Florida  State  University,  Tallahassee 

Abbott  (1954)  has  suggested  that  the  opisthobranch  gastropods 
Acteocina  candei  (Orbigny)  and  Retusa  canaliculata  (Say)  may 
belong  to  the  same  species  and  may  be  merely  geographic  races  or 
subspecies.  The  recovery  of  a  large  number  of  Acteocina  candei 
from  offshore  waters  in  the  vicinity  of  Cape  Hatteras  has  per- 
mitted us  to  compare  this  form  with  an  equally  large  number  of 
superficially  similar  specimens  of  Retusa  canaliculata  collected  in 
Pamlico  Sound.  Our  observations  and  conclusions  are  presented 
in  this  report.  Our  comparisons  led  to  the  discovery  of  certain  as- 
pects of  the  biology  of  Retusa  canaliculata  not  previously  re- 
corded, including  the  presence  of  a  radula  and  the  occurrence  of 
non-pelagic  reproduction  in  this  species. 

Material.  More  than  300  specimens  of  Acteocina  candei  were 
recovered  from  the  stomachs  of  seastars  (Astropecten  articulatus) 
collected  from  a  sand  bottom  off  Ocracoke  Inlet,  North  Carolina, 
at  a  depth  of  4  to  7  fathoms  (Wells  et  al  1961)  .  Most  of  these 
were  fresh  specimens,  from  the  soft  parts  of  which  radulae  could 
be  recovered.  A  similar  number  of  specimens  of  Retusa  canalicu- 
lata were  collected  from  sand  flats  2  to  3  inches  below  mean  low 
tide  along  the  eastern  shore  of  Pamlico  Sound  1  mile  north  of 
Avon,  N.  C.  These  two  collecting  stations  are  separated  by  a 
distance  of  less  than  30  nautical  miles.  While  one  habitat  is 
oceanic,  the  other  is  estuarine,  with  a  greater  range  in  tempera- 
tures, more  rapid  temperature  changes,  and  reduced  salinities. 

In  addition  to  these  specimens  from  North  Carolina,  shells  of 
both  species  from  different  parts  of  their  ranges  have  been  ex- 
amined at  the  U.  S.  National  Museum. 

Observations.  These  "forms"  differ  in  several  consistent  charac- 
ters: Oxierall  shell  shape.  Variations  in  the  shell  shape  of  Acteo- 
cina candei  and  Retusa  canaliculata  are  shown  in  Figures  8-10 
and  1-3,  respectively.  Specimens  from  the  oceanic  collection    {A. 

87 


NAUTILUS 


Vol.  75    (3) 


Figs.  1-3.  Retusa  canaliculata,  showing  variation  in  shell  shape.  Fig.  3.  R. 
canaliculata  with  extreme  development  of  the  spire  (protoconch  eroded)  . 
Fig.  4.  Acteocina  candei,  protoconch.  Fig.  5.  R.  canaliculata,  protoconch. 
Fig.  6.  R.  canaliculata,  apical  view.  Fig.  7.  A.  candei,  apical  view.  Figs.  8-10. 
A.  candei,  showing  variation  in  shell  shape.  (Fig.  10  shows  juvenile  condition.) 
(Figures  were  prepared  with  aid  of  a  camera  lucida.) 


January,   1962 


NAUTILUS 


89 


Fig.  11.  Egg  mass  of  Retusa  ca?ialiciilala.  Fig.  12-  Radiilar  teeth  of 
Acleoci)ia  candei.  Fig.  13.  Radiiiar  teeth  of  R.  canaliculata.  Fig.  14.  R.  canali- 
culata,  egg  membrane  containing  veliger.  Figs.  15-16.  Shell  of  newly  hatched 
R.  canaliculata.    (Figures  were  prepared  with  the  aid  of  a  camera  lucida.) 

candei)  are  generally  spindle-shaped,  with  an  elevated  spire  and 
a  tapered  base,  so  that  the  shell  is  widest  in  the  middle  (Figs.  8 
and  9) .  However,  juvenile  specimens  of  A.  candei  may  appear 
more  or  less  truncate  (Fig.  10) ,  because  of  the  lesser  development 
of  the  spire.  The  aperture  tapers  gradually  at  its  apical  end. 
Specimens  from  the  estuarine  collection  [R.  canaliculata)  are 
more  cylindrical  in  shape,  usually  with  lower  spire,  with  a  more 
rounded  base,  and  with  nearly  parallel  sides,  so  that  the  shell  is 
shouldered  and  nearly  as  wide  near  its  apical  end  as  it  is  in  the 
middle  (Figs.  1  and  2)  .  However,  there  is  some  variation  in  the 
elevation  of  the  spire  in  R.  canaliculata,  occasionally  approaching 
(as  in  Fig.  3)  the  development  characteristic  of  A.  candei.  The 
existence  of  individuals  with  such  "intermediate"  shell  characters 
has  no  doubt  led  to  confusion  in  identification.  Specimens  of  both 
species  that  show  a  broad  shallow  depression  in  the  middle  of  the 


90  NAUTILUS  Vol.   75    (3) 

last  whorl  (as  in  Fig.  2) ,  producing  a  sinuous  outer  lip  such  as 
that  figured  by  Marcus  (1958,  fig.  26)  for  Acteocina  {:=Torna- 
tina)  candei  have  been  examined.  A  similar  depression  is  shown 
in  the  shells  of  Retusa  obtusa  figured  by  Lemche  (1948,  figs.  43- 
53) ,  but  in  that  species  also  this  character  is  variable  and  not 
always  present.  Because  of  its  occurrence  in  several  similar  species 
and  the  variability  of  its  expression,  such  a  shallow  depression 
cannot  be  used  as  a  reliable  character  for  the  separation  of  species. 

Protoconch.  Whereas  in  A.  candei  the  protoconch  is  obvious 
and  projecting,  in  R.  canaliculata  only  a  small  portion  is  visible 
as  a  small  protuberance,  and  this  is  often  eroded.  The  hetero- 
strophic  protoconch  of  A.  candei  is  composed  of  about  one  and 
one-half  whorls  (Fig.  4)  set  at  an  angle  of  80°  from  the  axis  of 
the  adult  shell.  It  is  about  i/4  submerged  in  the  first  adult  whorl 
(Fig.  7) .  In  contrast,  the  heterostrophic  protoconch  of  R.  canali- 
culata contains  only  about  3/^  of  a  whorl  (Fig.  5)  set  at  an  angle 
of  80°  from  the  adult  shell  axis.  It  is  about  %  hidden  by  the  first 
adult  whorl  (Fig.  6) .  This  difference  in  protoconchs  would  indi- 
cate that  the  planktonic  stage  is  of  moderate  length  in  the  larva  of 
A.  candei,  while  the  planktonic  stage  is  suppressed  in  the  larva  of 
R.  canaliculata. 

Radular  teeth.  Using  the  technique  described  by  Turner 
(1960) ,  the  radulae  were  mounted  for  examination.  The  radular 
teeth  of  a  specimen  of  A.  candei  2.5  mm.  in  length  are  shown  in 
Figure  12.  The  median  tooth  is  18  microns  wide,  10  microns 
high,  and  bears  4  or  5  forward  projecting  denticles  on  each  of  two 
arches.  The  lateral  tooth,  with,  an  overall  length  of  45  microns, 
has  a  base  33  microns  long  and  a  moderately  curved  cusp  30 
microns  long;  it  bears  5  to  7  sharp  denticles  on  an  expansion  at 
the  angle  formed  at  the  base  of  the  cusp.  By  the  same  technique, 
a  radula  was  discovered  in  R.  canaliculata,  the  teeth  of  which  are 
shown  in  Figure  13.  For  a  specimen  3  mm.  in  length,  the  median 
tooth  is  30  microns  wide,  19  microns  high,  and  is  comprised  of 
two  arches,  each  bearing  ten  or  eleven  denticles.  The  lateral  tooth 
is  approximately  65  microns  long,  with  a  base  46  microns  long 
and  a  strongly  curved  cusp  41  microns  long  that  bears  16  to  20 
small  denticles  on  a  curved  ridge.  For  each  species,  the  radulae 
of  at  least  10  specimens  were  examined.  The  shape  of  the  teeth 
and  the  distribution  and  relative  size  of  the  denticles  clearly  dis- 


January,   1962  nautilus  91 

tinguish  the  radulae  of  the  two  species. 

Because  of  the  minute  size  of  the  radulae  in  these  species,  ade- 
quate radular  preparations  are  difficult.  Evidently,  the  radulae 
of  both  species  escaped  detection  by  early  workers.  Pilsbry  (1893) 
treated  both  species  as  having  no  radula;  however,  later  workers 
recognized  the  presence  of  a  radula  in  Acteocina  (Thiele  1931; 
Marcus  1958,  as  Tornatina) .  Thiele  (1925)  could  find  no  radula 
in  Retusa  canaliculata,  and  many  workers  (Marcus  1958,  Zilch 
1959,  and  others)  have  utilized  the  absence  of  a  radula  to  charac- 
teiize  the  Retusidae,  and  to  place  R.  canaliculata  in  this  family. 
The  possession  of  a  radula  is  used  for  the  placement  of  Acteocina 
candei  in  the  Scaphandridae.  Since  R.  obtusa  (Montagu)  is  the 
type  species  of  Retusa,  the  discovery  of  a  radula  in  canaliculata 
makes  a  re-examination  of  R.  obtusa  for  the  presence  of  a  radula 
necessary'  in  order  to  clarify  the  position  of  canaliculata  in 
opisthobranch  classification.  Until  the  presence  or  absence  of  a 
radula  in  R.  obtusa  is  ascertained,  we  prefer  to  retain  canaliculata 
in  the  genus  Retusa. 

Although  the  possession  of  a  slightly  modified  shell  may  reflect 
environmental  differences  between  two  habitats  occupied  by  the 
same  species,  the  differences  considered  above  can  scarcely  be 
attributed  to  the  environment.  On  the  basis  of  this  comparison, 
these  forms  should  be  regarded  as  biologically  distinct  species. 
This  group  of  differentiating  characters,  particularly  the  differ- 
ences in  protoconchs  and  radular  teeth,  provide  unquestionable 
grounds  for  the  recognition  of  two  species. 

Acteocina  candei  occurs  from  North  Carolina  south  to  Argen- 
tina (Carcelles  &  Parodiz  1938),  principally  in  offshore  or  oceanic 
habitats.  In  contrast,  Retusa  canaliculata  occurs  from  Cape  Cod 
to  the  West  Indies,  primarily  in  estuarine  habitats.  As  Parker 
(1959)  noted  for  the  Texas  coast,  R.  canaliculata  is  a  character- 
istic inhabitant  of  enclosed  bays  of  variable,  low  to  intermediate 
salinities. 

Reproduction  in  Retusa  canaliculata.  Egg  masses  produced  by 
Retusa  canaliculata  (Fig.  11)  were  collected  on  sand  flats  in  Pam- 
lico Sound,  where  they  were  found  to  be  relatively  common  from 
May  to  October.  The  egg  masses  of  this  species  bear  a  resem- 
blance to  one  figured  by  Thorson  (1946,  fig.  151)  as  that  of 
Philine  scabra.  They  are  composed  of  a  spherical  jelly  mass  about 


92  NAUTILUS  Vol.  75    (3) 

2.0  to  2:5  mm  in  diameter,  attached  by  a  strand  about  5  mm  long 
to  stems  and  blades  of  marine  grasses,  worm  tubes,  shell  frag- 
ments, and  other  objects.  Each  mass  contained  a  number  of  large 
eggs,  each  within  its  own  egg  membrane  (Fig.  14) .  The  envelop- 
ing egg  membranes  are  ovoid  and  0.30  to  0.35  mm  in  length. 
Although  eggs  were  usually  found  only  in  the  enlarged  spherical 
part  of  the  mass,  in  some  cases  they  were  also  contained  in  the 
attachment  strand,  as  figured.  A  number  of  egg  masses  were  re- 
moved to  the  laboratory  for  observation.  Development  progressed 
within  the  egg  capsule,  through  a  shelled  veliger  stage,  to  a  min- 
iature crawling  juvenile.  These  young  snails  were  observed  crawl- 
ing within  their  individual  egg  capsules,  from  which  they  event- 
ually emerged  to  crawl  about  the  aquarium.  These  observations 
constitute  the  first  record  of  non-pelagic  development  in  Retusa 
canaliculata. 

The  retention  of  the  veliger  stage  within  the  egg  capsule  un- 
doubtedly serves  to  maintain  this  species  in  estuarine  areas  wdiere 
the  net  flow  of  water  seaward  could  carry  planktonic  larvae  away 
from  an  otherwise  favorable  environment.  The  evolution  of 
such  a  mechanism  would  have  considerable  survival  value  for 
the  larvae  of  species  that  inhabit  estuaries.  Indeed,  non-pelagic 
development  insures  a  more  stable  population  size  (Thorson, 
1950) .  Such  a  non-pelagic  development  has  been  demonstrated 
for  some  species  of  echinoderms,  polychaete  annelids,  prosobranch 
gatropods,  and  pelecypods  (Thorson,  1950) .  Generally,  tecti- 
branchs  have  been  considered  to  produce  planktonic  larvae. 

Lemche  (1948)  indicated  that  studies  of  the  apical  whorls  of 
tectibranch  gastropods  may  give  reliable  evidence  about  their 
larval  development,  and  Thorson  (1950)  successfully  applied 
similar  correlations  of  apical  shell  form  to  the  development  of 
prosobranch  gastropods.  According  to  Lemche's  standards,  the 
blunt,  relatively  coarse  protuberance  formed  by  the  protoconch 
of  Retusa  canaliculata  fits  the  general  protoconch  morphology 
that  is  correlated  with  a  reduced  or  suppressed  planktonic  devel- 
opment. Significantly,  Lemche  found  the  protoconch  of  Retusa 
obtusa  to  be  wanting,  and  suggested  that  it  probably  exhibits  a 
non-pelagic  development.  In  his  svnvey  of  Danish  planktonic 
larvae,  Thorson  (1946)  had  not  found  larvae  that  he  could  assign 
to  R.  obtusa,  although  adults  of  that  species  had  been  known 


January,   1962  nautilus  93 

from  the  area  studied.  Evidently,  R.  obtusa  shares  with  R.  canali- 
culdta  a  non-pelagic  type  of  reproduction. 

The  relatively  large  size  of  the  eggs  and  the  prolonged  repro- 
ductive period  of  R.  canaliculata  are  features  that  have  been 
correlated  with  non-pelagic  development  in  other  marine  inverte- 
brates (Thorson  1950) .  By  increasing  the  chances  of  successful 
reproduction,  these  characters  contribute  to  the  effectiveness  of 
non-pelagic  development  as  an  effective  mechanism  for  the  pro- 
duction of  a  new  generation  without  the  waste  usually  associated 
with  plauktonic  reproduction. 

Acknowledgments.  These  specimens  were  obtained  and  studied 
during  the  course  of  research  supported  by  a  grant  (G-5838)  from 
the  National  Science  Foundation  to  Dr.  I.  E.  Gray  of  Duke  Uni- 
versity and  aided  by  the  Cape  Hatteras  National  Seashore  of  the 
National  Park  Service.  The  authors  wish  to  express  their  appreci- 
ation to  Dr.  Harald  A.  Rehder  and  Dr.  J.  P.  E.  Morrison  of  the 
Division  of  Mollusks,  United  States  National  Museum,  for  making 
available  specimens  and  literature. 

References 

Abbott,  R.  T.  1954.  American  Seashells.  D.  Van  Nostrand  Co., 
New  York.  541  pp. 

Carcelles,  A.,  and  J.  J.  Parodiz.  1938.  Physis,  12:  251-266. 

Lemche,  H.  1948.  K.  Danske  Vidensk.  Selsk.  Skr.  (Biol.) ,  5:  1-136. 

Marcus,  E.  1958.  Bol.  Oceanogi-.  S.  Paulo,  7  (1-2)    (1956)  :  31-80. 

Parker,  R.  H.  1959.  Bull.  Amer.  Assoc.  Petrol.  Geol.,  43:  2100- 
2166. 

Pilsbry,  H.  A.  1893.  George  W.  Tryon,  Jr.,  Manual  of  Conchology; 
structural  and  systematic.  Order  Opisthobranchia,  vol.  15:  134- 
436.  Philadelphia. 

Ihielc,  J.  1925.  Gastropoden  der  Deutschen  Tiefsee-Expedition, 
Teil  2.  Wiss.  Ergebn.  Deutschen  Tiefsee-Exped.,  17 {2)  Opistho- 
branchia: 257-288,  348-352.  Jena. 

1 93 1.  Handbuch  der  systematischen  Weichtierkunde,  vol.  /. 

G.  Fischer,  Jena.  778  pp. 

Thorson,  G.  1946.  Meddel.  Komm.  Danmarks  Fiskeri-  og  Havun- 
dersogelser,  Scr.  Plankton  •/(!):  1-523. 

1950.  Biol.  Rev.,  25:  1-45. 

Turner,  Ruth.  1960.  Nautilus,  75(4):   135-137. 

Wells,   H.  W.,  M.  J.  Wells,  and  I.  E.  Gray.   1961.   Biol.  Bull., 

120  (2) .  In  press. 
Zilch,   A.    1959.   Gastropoda   von   Wilhelm   Wenz;    Euthyneura. 

Handbuch  der  Palaozoologie,  6(2):   1-200.  Berlin. 


94  NAUTILUS  Vol.  75    (3) 

RANGE  EXTENSION  FOR  CYMATIUM  CARIBBAEUM 
WITH  A  NOTE  ON  ADVENTITIOUS  DISPERSAL 

By  ARTHUR  S.  MERRILL 

U.  S.  Department  of  the  Interior,  Fish  and  Wildlife  Service 

Bureau  of  Commercial  Fisheries  Biological  Laboratory 

Woods  Hole,  Massachusetts 

A  specimen  of  Cymatium  caribbaeum  CI.  Sc  T.  was  recently  sent 
to  me  by  Mr.  Richard  Spencer  of  Charleston,  South  Carolina.  He 
had  collected  the  example  alive  from  a  navigation  buoy  after  it 
was  brought  in  from  its  station  off  Port  Royal,  South  Carolina,  to 
the  U.  S.  Coast  Guard  Base  at  Charleston  for  cleaning  and  servic- 
ing. Clench  and  Turner  (Monograph  of  the  family  Cymatiidae 
in  the  western  Atlantic.  Johnsonia:  3  (36)  :  206,  1957.)  record 
the  northernmost  range  for  this  species  as  Lake  Worth,  Florida. 
The  buoy  record  thus  extends  the  range  northward  along  the 
Atlantic  coast  about  350  miles. 

The  specimen  is  a  thin-lipped  juvenile,  height  31.7  mm.,  width 
16.1  mm.  This  is  the  second  species  of  this  genus  whose  range  has 
been  considerably  extended  northward  as  a  result  of  buoy  sam- 
pling. The  first  was  a  juvenile  Cymatium  labiosum  (Wood) 
which  I  collected  in  1948  from  a  buoy  brought  in  from  off  Cape 
Romain,  South  Carolina.  This  record  appears  in  the  monograph 
by  Clench  and  Turner  (1957)   on  page  202. 

The  Cymatiidae  are  stenothermic  and  range  widely  in  all  tropi- 
cal waters.  A  probable  long  larval  stage  aids  in  their  wide  dispersal 
according  to  Clench  and  Turner.  The  larvae  of  the  Cymatium 
species  found  on  the  South  Carolina  buoys  probably  chanced  to 
be  dispersed  northward  from  Florida  by  way  of  rapid  transport 
in  the  Gulf  Stream.  Caught  in  the  coastal  waters  below  Cape 
Hatteras,  they  finally  settled  on  the  convenient  buoy  surfaces. 

The  normal  longitudinal  distribution  of  a  species  with  pelagic 
larvae  is  controlled  for  the  most  part  by  the  temperature  ex- 
tremes that  it  can  tolerate  as  an  adult.  The  chance  dispersal 
beyond  the  consistent  range  of  such  a  species  is  probably  common. 
When  this  occurs  and  when  the  larvae  happen  to  settle  in  a  favor- 
able habitat  during  a  season  in  which  the  temperatures  are  also 
favorable,  the  organisms  may  survive  and  grow  until  such  time  as 


*  Wells,  W.  Harry  and  \.  E.  Gray.  The  seasonal  occurrence  of  Mytilus 
ediilis  on  the  Carolina  coast  as  a  result  of  transport  around  Cape  Hatteras. 
Biol.  Bull.:  119  {3):  550-559,  1960. 


January,   1962  nautilus  95 

the  temperature  becomes  lethal.  The  tact  that  an  adult  of  the 
genus  Cymatium  has  never  been  recorded  from  the  bottom  in  off- 
shore Carolinian  waters  suggests  that  juveniles  are  winter-killed. 

Wells  and  Gray*  recently  reported  on  a  cold  water  species, 
Mytilus  ediilis,  which  commonly  spreads  beyond  its  normal  south- 
ern limit  at  Cape  Hatteras.  The  species  is  unable  to  survive  sum- 
mer temperatures  in  the  northern  part  of  the  Carolinian  sub- 
province.  However,  the  larvae  of  the  fall  spawning  colonize  this 
area  after  the  water  temperatures  fall  below  lethal  values. 

There  is  no  need  to  stress  the  importance  of  accurate  range 
records  for  ecological  purposes.  In  order  to  increase  the  accuracy 
of  range  records,  one  must  distinguish  between  that  part  of  the 
range  within  which  a  species  is  able  to  maintain  itself  and  propa- 
gate, and  that  portion,  usually  at  the  extremes,  where  it  is  unable 
to  complete  its  life  history.  It  is  also  important  to  record  whether 
the  specimen  was  alive  or  dead  when  collected;  especially  near  the 
borders  of  its  geogiaphic  or  bathymetric  range.  Knowledge  of 
extreme  range  is  particularly  useful  to  indicate  possible  modifica- 
tions in  the  usual  biota  of  an  aiea  should  long-range  environ- 
mental changes  alter,  even  slightly,  in  any  direction. 

ARCIDENS  CONFRAGOSUS  AND  PROPTERA  CAPAX 
IN  KANSAS 

By  HAROLD  D.  MURRAY 

Department  of  Zoology,  The  University  of  Kansas 

The  extended  ranges  of  the  two  species  of  mussels  herein  re- 
ported are  the  result  of  studies  of  the  unionid  fauna  of  Kansas 
from  the  years  1956  to  1959.  Call  (1885,  1886  and  1887)  listed 
numerous  species  of  unionids  in  Kansas  without  descriptions  or 
illustrations,  and  Scammon  (1906)  described  and  illustrated  the 
species  of  fresh-water  mussels  occuiTing  in  Kansas.  Neither  Call 
nor  Scammon  reported  these  two  species  as  occurring  in  Kansas. 
A  search  of  the  literature  indicates  that  neither  has  been  recorded 
for  Kansas.  Catalogue  numbers  refer  to  the  mollusk  collection  of 
the  Museum  of  Natural  History,  The  University  of  Kansas, 
Lawrence,  Kansas. 

Arcidens  confragosiis  (Say) ,  KUMNH.  11083.  This  species  was 
not  collected  by  the  author  but  was  uncovered,  quite  by  accident, 
in  the  museum  collection.  R.  W.  Reese  obtained  a  single  gravid 


96  NAUTILUS  Vol.  75    (3) 

specimen  of  Arcidens  confragosus  from  the  Marais  des  Cygnes 
River,  3  miles  east  of  Ottawa,  Franklin  County,  Kansas,  on  Sep- 
tember 25,  1949.  Unfortunately,  only  a  half  shell  of  this  specimen 
remains  in  the  museum  collection.  An  empty  shell  of  A.  con- 
fragosus was  found  at  a  later  date  at  the  same  locality,  but  has 
subsequently  been  lost.  The  half  shell  remaining  in  our  collection 
measures  60  mm.  in  length  and  42  mm.  in  height.  A.  confragosus 
was  taken  from  a  gravel  shoal  where  the  water  ranged  in  depth 
from  one  to  five  feet.  Subsequent  attempts  to  collect  additional 
examples  have  been  unsuccessful. 

Because  Utterback  (1916:103)  reported  A.  confragosus  as  oc- 
curring in  northern  and  central  Missouri  and  because  the  Osage 
River  (Marais  des  Cygnes  River  in  Kansas)  flows  through  central 
Missouri,  the  presence  of  A.  confragosus  in  Kansas  is  not  surpris- 
ing. Possibly  A.  confragosus  may  occur  in  isolated  areas  in  eastern 
Kansas  not  yet  adequately  sampled. 

Proptera  capax  (Green) ,  KUMNH.  10486.  A  gravid  specimen 
oi  Proptera  capax  was  recovered  by  the  author  on  August  27,  1956 
from  the  Neosho  River,  7i/2  miles  east  and  1  mile  south  of  Em- 
poria, Lyon  County,  Kansas.  Simpson  (1914:47)  reported  the 
westernmost  range  of  P.  capax  as  the  St.  Frances  River  in  eastern 
Arkansas  and  two  almost  certainly  invalid  records  from  the  Elk- 
horn  and  Blue  Rivers,  Nebraska. 

Inasmuch  as  the  St.  Frances  River  in  Arkansas  empties  into  the 
Mississippi  River  a  short  distance  from  the  Arkansas  River  and 
inasmuch  as  the  Neosho  River  is  a  tributary  of  the  Arkansas 
River,  it  is  not  surprising  that  P.  capax  should  occur  in  the 
Neosho  River  in  Kansas.  Although  Utterback  (1916:163)  reported 
P.  capax  in  Missouri,  his  records,  as  best  as  can  be  determined,  are 
from  the  extreme  eastern  portion  of  Missouri.  P.  capax  has  not 
been  reported  to  this  date  in  those  streams  flowing  from  Kansas 
through  Missouri  to  the  Mississippi  River,  which  is  somewhat 
surprising  considering  the  appearance  of  P.  capax  in  the  Neosho 
River. 

The  single  example  of  P.  capax  thus  far  known  for  Kansas  is 
typical  in  most  respects  for  the  species.  The  nacre  is  a  somewhat 
darker  purple,  and  the  shell  is  somewhat  more  elongate  than 
specimens  from  the  Mississippi  River. 

The  Kansas  specimen  measures  155  mm.  in  length  and  101  mm. 


January,   19(32  nautilus  97 

in  height.  In  length,  this  specimen  exceeds  the  previously  re- 
ported maximum  length  of  P.  capax  by  12  mm.  (Haas,  1941 : 261) . 
P.  capax  ^vas  recovered  from  slowly  moving  water  3i/^  feet  deep 
in  a  substrate  of  small  rocks  and  sand  having  very  little  silt. 

Because  A.  confragosus  is  a  species  with  unusual  morphological 
features  and  not  likely  to  be  confused  with  other  species,  and 
because  Call  (1885,  1886  and  1887)  and  Scammon  (1906)  did  not 
report  this  species  in  Kansas,  it  seems  most  likely  that  A.  con- 
fragosus has  made  its  appearance  in  Kansas  after  1906.  It  is  my 
opinion  that  P.  capax  may  have  occurred  sparsely  in  Kansas  for 
many  years  and  may  possibly  have  been  confused  by  Call  and  by 
Scammon  with  Lampsilis  ovata  ventricosa   (Barnes) . 

Literature  Cited 
Call,  R.  E.  1885.  Bull.  Washburn  College  Lab.  Nat.  Hist.,  i.-48- 
123. 

1886.  Bull.  Washburn  College  Lab.  Nat.  Hist.,  7.177-183. 

1887.  Bull.  Washburn  College  Lab.  Nat.  Hist.,  2.11-25. 

Haas,  F.  1941.  Zool.  Series,  Field  Mus.  Nat.  Hist.  Chicago,  2^.-259- 

270. 
Scammon,  R.  E.    1906.   Univ.  Kansas  Sci.   Bull.,  5.-279-373,   pis. 

52-86. 
Simpson,  C.  T.   1914.  A  descriptive  catalogue  of  the  naiades  of 

pearly  fresh-water  mussels.  Detroit,  1540  pp. 
Utterback,  W.  L  1916.  Amer.  Midi.  Nat.  4:  (1-10)  :  200  pp.,  29  pis. 

RADULAE  OF  NORTH  AMERICAN  ANCYLID  SNAILS 
I.  SUBFAMILY  RHODACMEINAE  ^ 

Bv  PAUL  F.  BASCH 
Department  of  Biology,  Kansas  State  Teachers  College,  Emporia 

freshwater  limpet  snails,  constituting  a  distinct  subfamily  of  the 
family  Ancylidae,  and  limited  in  distribution  to  the  southeastern 
United  States.  Walker  (1917)  recognized  two  "sections"  or  sub- 
genera of  Rhodacmea  based  partly  upon  conchological  characters, 
but  principally  upon  features  of  the  radula,  to  be  discussed  below. 
In  connection  with  a  continuing  study  of  North  American  fresh- 
water limpets,  I  have  reported  briefly  on  the  anatomy  of  one 
species  of  Rhodacmea  (Basch,  1960) ,  and  more  recently  have  ex- 
amined the  radulae  of  other  species  in  an  attempt  to  understand 
better  this  unusual  genus  of  mollusks.  I  am  indebted  to  Dr.  Henry 


1  Supported  bv  grant  G-14125  from  the  Xaiional  Science  Fouiidaiion. 
The  genus  Rhodacmea  consists  of  about  half  a  dozen  species  of 


98  NAUTILUS  Vol.  75    (3) 

van  der  Schalie  of  the  University  of  Michigan  Museum  of  Zoology 
(UMMZ)  for  kindly  making  available  to  me  the  material  used 
in  this  study. 

The  radulae  upon  which  Walker  based  his  systematic  con- 
clusions were  prepared  for  him  by  Rev.  H.  M.  Gwatkin  of  Cam- 
bridge, England.  The  specimens  from  which  those  slides  were 
made  were  collected  in  the  Tennessee  and  Coosa  River  drainages 
in  Alabama,  presumably  by  A.  A.  Hinkley  or  H.  H.  Smith,  in  the 
first  decade  of  this  century.  In  Walker's  1917  paper,  and  again 
the  following  year  (Walker,  1918) ,  two  figures  illustrate  the  radu- 
lae of  Rhodacmea  filosa  (Conrad)  and  R.  rhodacme  Walker. 
These  drawings  were  the  work  of  Mrs.  Lydia  M.  H.  Green.  Al- 
though probably  Mr.  Walker  examined  the  preparations  himself, 
there  is  to  my  knowledge  no  direct  evidence  that  he  did  so. 

I  have  examined  7  slides  of  radulae  prepared  by  Rev.  Gwatkin, 
including  those  from  which  Mrs.  Green  made  her  original  illus- 
trations. The  fact  that  his  preparations  have  remained  in  ex- 
cellent condition  for  over  half  a  century  is  a  tribute  to  the  skill  of 
Rev.  Gwatkin,  who  died  in  November,  1916.  In  addition,  I  have 
prepared  24  slides  of  radulae  extracted  from  dried  animals  found 
within  their  shells  in  the  UMMZ  collection.  The  wide  open  aper- 
ture of  the  limpets  and  the  lack  of  spiral  coiling  allowed  easy  re- 
moval of  the  dry  animals  without  damage  to  the  shells.  After 
removal,  the  animals  were  soaked  for  several  hours  in  a  0.5% 
solution  of  trisodium  phosphate  to  soften  the  tissues,  and  the 
radulae  then  dissected  out.  Although  large  portions  of  these  frail 
ribbons  could  be  removed  intact,  I  could  not  obtain  an  unbroken 
radula  in  this  manner.  The  number  of  transverse  rows  of  teeth 
per  ribbon  could  therefore  not  be  determined.  In  some  cases, 
where  bits  of  tissue  adhered  to  the  radula,  the  entire  piece  of 
ribbon  was  immersed  briefly  in  a  dilute  solution  of  sodium  hypo- 
chlorite (clorox)  to  clean  it.  Such  treatment  does  not  harm  the 
teeth  if  performed  judiciously,  and  is  far  more  convenient  than 
the  traditional  sodium  hydroxide  bath.  All  ribbons  were  rinsed 
in  70%  ethyl  alcohol  and  mounted  in  polyvinyl  lactophenol,  a 
mounting  medium  which  dries  fairly  rapidly  and  does  not  require 
previous  dehydration  and  clearing  of  the  specimen. 

The  following  specimens  were  examined,  all  UMMZ  numbers: 
A.  Prepared  radulae  from  the  Gwatkin  collection 


January,   1962  nautilus  99 

#946,  947,  948  — 7^.  rhodacme  Walker,  Coosa  River. 

•^9A9  —  R.  gwatkiniana  Walker,  Coosa  River. 

:jijt950,  951  — R.  clatior  (Anthony),  Tennessee  drainage,  Flor- 
ence, Alabama. 

#976  —  R.  filosa    (Conrad) ,  Coosa  River 
B,  Radulae  extracted  and  mounted  from  dry  specimens 

#65998  —  R.  cahawbensis  Walker,  Little  Cahaba  River,  Bibb 
County,  Alabama.  2  specimens. 

#69213  —  R.  filosa,  Coosa  River,  Talladega  County,  Alabama. 
8  specimens. 

#69221 — R.  giuatkiniana,  Coosa  River,  Coosa  County,  Ala- 
bama. 3  specimens. 

#69223  —  R.  giuatkiniana,  Coosa  River,  Chilton  County,  Ala- 
bama. 4  specimens. 

#69237  —  R.  rhodacme,  Coosa  River,  St.  Clair  County,  Ala- 
bama. 7  specimens. 

All  the  identifications  of  species  are  those  of  Walker.  A  sixth 
species,  Rhodacmea  hinkleyi  (Walker)    was  unavailable. 

The  specific  features  utilized  in  establishing  the  two  subgenera 
of  Rfiodacmea  are  as  follows   (Walker,  1917)  : 

"Section  Rhodacmea,  s.s.  Shell  elevated.  Radula  with  a  uni- 
cuspid  central,  which  has  the  base  triangularly  expanded;  laterals 
with  the  cusp  of  the  mcsocone  extending  bvit  little  beyond  the 
base  and  not  overlapping  the  base  of  the  central  tooth."  Included 
in  this  section  were  R.  filosa,  R.  cahawbensis,  R.  elatior,  and  R. 
hinkleyi. 

"Section  Rhodocephala,  n.  sect.  Shell  depressed.  Radula  with  a 
faintly  bicuspid  central  which  has  the  sides  of  the  base  straight 
and  not  expanded;  laterals  with  the  cusp  of  the  mesocone  extend- 
ing far  beyond  the  base  and  overlapping  the  base  of  the  central 
tooth."  In  this  section  were  placed  R.  rhodacme  and  R.  gxvat- 
kiniana. 

From  my  study  of  these  species  I  have  come  to  the  conclusion 
that  there  is  no  constant  character  by  which  it  is  possible  to  dis- 
tinguish species  or  species  groupings  within  Rhodacmea.  I  submit 
that  the  characters  utilized  by  Walker  are  largely  artifacts,  which 
are  based  upon  two  conditions  —  the  amount  of  wear  on  the  teeth 
examined,  and  variations  in  techniques  of  mounting  the  ribbon 
for  study.  In  figs.  A  to  C  are  shown  3  rows  of  teeth  from  a  single 
radula  of  R.  cahawbensis.  Changes  in  the  conformation  of  indi- 
vidual teeth  as  a  consequence  of  the  amount  of  wear  may  be  easily 
seen.  The  central  tooth,  originally  bicuspid,  becomes  worn  down 
to  a  single  cusp;  the  length  of  the  mesocone  may  vary  by  30% 


100 


NAUTILUS 


Vol.  75    (3) 


^^.'f 


3  4  5  6  7         8 


mi^ij 


r-  ,.'     "t 


Representative  teeth  from  one  radula  of  Rhodacmea  cahaxubensis,  UMMZ. 
#65998,  prepared  and  mounted  January,  1961,  from  a  dried  specimen.  A,  teeth 
near  the  front  of  ribbon,  showing  severe  wear;  B,  teeth  from  an  intermediate 
area,  showing  some  wear;  C,  unworn  teeth  from  a  region  near  end  of  radular 
sac.  Only  central  and  8  of  the  11  teeth  on  one  side  are  shown.  Features  marked 
by  small  letters  in  C  ilhistrate  variable  characters:  a,  central  tooth  bicuspid 
or  unicuspid;  b,  degree  of  overlapping  between  rows;  c,  amoimt  of  "shoulder- 
ing" on  medial  surface  of  first  tooth;  d,  configuration  of  finer  denticles  on  first 


January,   1962  nautilus  101 

within  the  same  ribbon,  and  the  amount  to  wliich  it  overlaps  the 
base  and  the  base  of  the  central  tooth  are  both  dependent  partly 
upon  wear  and  partly  upon  the  amount  of  pressure  applied  to 
the  coverslip  in  making  the  slide.  Depending  upon  the  manipu- 
lation of  the  preparation,  teeth  may  be  rotated  into  various  posi- 
tions, and  may  be  separated  from  each  other  to  a  greater  or  lesser 
degree.  The  specific  areas  of  greatest  apparent  variation  in  the 
sample  studied  are  pointed  out  on  figure  C. 

Previous  studies  on  the  reliability  of  the  radula  as  a  taxonomic 
tool  (Howe,  1930;  Van  Cleave  and  Richey,  1936)  suggest  that  this 
structure  alone  is  not  a  dependable  criterion  of  specific  differences 
within  a  genus  (in  those  cases,  Vivipariis) .  In  the  present  genus, 
although  the  radula  is  of  the  greatest  importance  in  separating 
Rlwdacmca,  sensu  lato  from  other  patelliform  fresh-water  mol- 
lusks,  I  feel  that  no  reliance  can  be  placed  upon  it  in  making 
critical  taxonomic  decisions  within  the  genus. 

With  regard  to  the  elevated  versus  depressed  shell,  this  may  be 
a  character  useful  for  species  differentiation,  but  I  do  not  believe 
it  to  be  of  sufficient  importance  to  separate  subgenera,  particularly 
when  the  radulae  are  indistinguishable  in  all  species  studied.  I 
must  conclude  that  the  subgenera  (or  sections)  Rhodacmea,  s.s. 
and  Rhodocephala  are  based  largely  upon  artifact  rather  than 
biological  differences,  and  therefore  invalid.  In  a  later  paper  I 
hope  to  discuss  the  relationships  of  Rhodacmea  within  the  Ancy- 
lidae,  and  of  the  various  species  to  one  another. 

Literature  cited 

Basch,  Paul  F.  1960.  Naut.  73  (3)  :  89-95. 

Howe,  Sam  W.  1930.  Naut.  •/-/  (2)  :  53-63. 

Van  Cleave,  Harley  J.,  and  Emily  M.  Richey.  1936.  Trans.  Amer. 

Micros.  Soc.  55(2):  223-229. 
Walker,  Bryant.  1917.  Naut.  31  (1)  :  1-10. 
1918.  A  synopsis  of  the  classification  of  the  fresh-water  Mol- 

lusca  of  North  America,  north  of  Mexico,  and  a  catalogue  of 

the  more  recently  described  species,  with  notes.  Misc.  Publ. 

Mus.  Zool.  Univ.  Mich.  6:  1-213. 


row  of  teeth;  e,  lateral  distance  between  teeth,  and  basal  characters;  f,  pro- 
nounced notch  present  or  absent  on  4th  tooth;  g,  ninnber  of  teeth  per  row 
(10,  11,  or  12,  with  1  ribbon  of  13).  All  figures  drawn  with  the  aid  of  a 
camera  lucida. 


102  NAUTILUS  Vol.   75    (3) 

NEW  SPECIES  OF  HELICODISCUS 
FROM  THE  EASTERN  UNITED  STATES 

By  LESLIE  HUBRICHT 

HELICODISCUS  MULTiDENS,  new  species.  Plate  7,  D-F;  Text  fig.  1 
Shell  discoidal,  spire  flat  or  nearly  so;  whorls  4i/2  to  5;  pale 
greenish-yellow,  dull,  opaque.  Umbilicus  wide  and  shallow,  show- 
ing all  the  whorls,  occupying  from  45  to  50%  of  the  diameter  of 
the  shell.  Whorls  well  roundfed,  slowly  increasing,  the  last  slowly 
descending;  sculptured  with  numeious,  fine,  spiral  threads.  Aper- 
ture lunate,  the  peristome  somewhat  thickened  within.  Within 
the  last  quarter  whorl  there  are  3  pairs  of  teeth  on  the  outer  and 
basal  walls.  These  teeth  are  radially  elongate,  raised  on  a  heavy 
callous  ridge,  and  separated  by  a  rounded  sinus.  Alternating  with 
these  are  3  teeth  on  the  parietal  wall.  These  teeth  extend  out  to 
about  the  center  of  the  whorl,  are  about  twice  as  broad  as  high, 
the  ends  are  turned  forward,  the  upper  end  more  so  than  the 
lower.  Of  the  3  sets  of  teeth  the  center  set  is  usually  more  fully 
developed  than  the  others.  As  the  shell  grows,  the  teeth  farthest 
within  are  absorbed  and  a  new  set  added  near  the  aperture. 

Height,  1.88  mm.  Diameter,  4.75  mm.  Umbilicus  diameter,  2.22 
mm.  Aperture  height,  1.55  mm.  4.5  whorls.  Holotype. 

Distribution:  —  Tennessee:  Putnam  Co.:  in  Jared  Hollow  Cave, 
3  miles  northeast  of  Chestnut  Mound  (Thomas  C.  Ban",  Jr., 
coll.)  .  DeKalb  Co.;  in  Jim  Cave,  1.5  miles  southeast  of  Dowell- 
town,  holotype  207798  U.M.M.Z.,  paratypes  17063,  collection  of 
the  author;  in  Avant  Cave,  1  mile  east  of  Dowelltown. 


Helicodiscus  multidens  Hubricht;  A,  view  of  central  pair  of  teeth.  B, 
parietal  tooth  from  above.  C,  diagram  of  tooth  arrangement. 

Helicodiscus  multidens  is  most  closely  related  to  H.  triodus 
Hubricht.  In  H.  triodus  the  teeth  are  smaller,  and  the  sets  of  3 
teeth  are  placed  at  irregular  intervals  in  the  last  whorl,  not 
crowded  near  the  aperture.  H.  multidens  is  a  rare  snail,  found 


NAUTILUS 


Map  1.  Distribution  of  Helicodiscus  shimeki  Hubricht  as  represented  by 
specimens  in  the  collection  of  the  author  and  in  the  collection  of  the  Uni- 
versity of  Michigan. 

only  in  caves.  Of  the  nine  specimens  so  far  collected  only  one  was 
found  alive. 

Helicodiscus  shimeki,  new  species.  Plate  7,  A-C;  Map  1. 

Shell  discoidal,  pale  yellow,  somewhat  shining,  translucent, 
spire  flat  or  slightly  convex.  Umbilicus  wide,  shallow,  showing  all 
the  whorls,  occupying  about  50%  of  the  diameter  of  the  shell. 
Whorls  5  to  6,  well  rounded,  very  narrow  and  slowly  increasing; 
nuclear  whorls  with  faint  spiral  striae;  later  whorls  with  numer- 
ous spiral  threads.  Aperture  lunate,  peristome  thin.  Within  the 
last  whorl  there  are  usually  three  pairs  of  small  conical  teeth;  on 
the  outer  and  basal  walls,  the  earlier  teeth  are  absorbed. 

Height,  1.7  mm.  Diameter,  4.2  mm.  Umbilicus  diameter,  2.1 
mm.  Aperture  height,  1.7  mm.  5.2  whorls.  Holotype. 

Type  locality:  loioa:  Delaware  Co.:  Backbone  State  Park,  holo- 
type  207796  and  paratypes  207797  U.M.M.Z.,  other  paratypes 
13807,  collection  of  the  author. 

Helicodiscus  shimeki  is  a  species  of  the  northern  United  States 
and  probably  southern  Canada,  although  the  author  has  seen  no 
specimens  from  there.  It  ranges  from  Iowa  eastward  to  northern 
New  York.  Map  no.  1. 

Helicodiscus  shimeki  may  be  readily  distinguished  from  H. 
parallelus  (Say)  by  its  more  slender  whorls  and  its  broader,  shal- 
lower umbilicus.  It  resembles  H.  salmonaceus  W.  G.  Binney  in  its 
proportions,  but  the  thread  striae  are  coarser,  and  the  umbilicus 
is  not  quite  so  broad.  It  stands  somewhat  intermediate  between 
these  two  species.  It  is  named  in  honor  of  the  late  Bohumel 
Shimek  of  Iowa  City,  Iowa. 


104 


NAUTILUS 


Vol.  75    (3) 


Map  2.  Distribution   of   Helicodiscus   notius   Hubrifcht   as   represented   by 
specimens  in  the  collection  of  the  author. 


Helicodiscus  notius,  new  species.  Plate  9,  N-P;  Map  2. 

Shell  discoidal,  the  spire  flat  or  nearly  so;  whorls  5  to  5i/^;  pale 
yellowish,  dull,  translucent  when  young,  becoming  opaque  with 
age.  Umbilicus  wide  and  shallow,  showing  all  the  whorls,  occupy- 
ing from  40  to  45%  of  the  diameter  of  the  shell.  Whorls  well 
rounded,  slowly  increasing,  sutures  well  impressed;  sculptured 
with  numerous  spiral  threads.  Aperture  lunate,  peristome  thin. 
Within  the  last  whorl  there  are  usually  2  or  3  pairs  of  conical 
teeth,  one  on  the  basal  wall  and  one  above  it  on  the  outer  wall. 

Height,  1.66  mm.  Diameter,  3.66  mm.  Umbilicus  diameter,  1.62 
mm.  Aperture  height,  1.04  mm.  5  whorls.  Holotype. 

Type  locality:  Alabama:  Jackson  Co.:  side  of  Keel  Mtn.,  Paint 
Rock,  holotype  207792  and  paratypes  207793  U.M.M.Z.,  other 
paratypes  17588,  collection  of  the  author. 

Helicodiscus  notius  is  found  over  most  of  the  southeastern 
United  States,  ranging  from  the  Gulf  north  to  Virginia  and  west 
to  Missouri  and  Oklahoma.  See  map  no.  2. 

Helicodiscus  notius  differs  from  H.  parallelus  (Say)  in  being  a 
little  larger  due  to  the  additional  whorl;  in  having  a  broader,  shal- 
lower umbilicus;  and  in  having  the  thread  striae  on  the  embry- 


I 


NAUTILLfS  75    (.H) 


PLATE 


j^ 


^' 


HoldlNlK's.    \-(  .  Hclirodiscus  shhnrhi  Huhiidii.  HI'.  H.  innllidrii.s  Huhrichl. 


NAUTILUS  75    (3) 


PLATE  8 


I 


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M 


II()h)l\|)LS.  (,-1.  flrli((>ilis(iis   noliiis  spcciis   Hiihiitlil.    |   .<:   R.  //.  UadciuH'ciis 
Ihihiiilu.  L  c<:  M,  //.  jucksoiii  Hubricht. 


January,   1962  nautilus  105 

onic  whorls  more  strongly  developed.  It  is  most  closely  related  to 
H.  eigenmanni  Pilsbry,  from  which  it  differs  in  being  smaller  and 
in  having  a  larger  umbilicus. 
Helicodi,sc;u.s  notius  specus,  new  subspecies.  Plate  8,  G-1. 

Differing  from  the  typical  form  in  being  without  the  internal 
teeth  in  shells  laiger  than  two  and  one-half  whorls.  The  last  half 
whorl  slowly  descends  to  the  aperture,  and  the  peristome  is  often 
much  thickened. 

Type  locality:  Kentucky:  Barren  Co.:  in  Burnette  Cave,  0.6 
mile  west  of  Park  City,  holotype  207794  and  paratypes  207795 
U.M.M.Z.,  other  paratypes  17027,  collection  of  the  author. 

Helicodiscus  notius  specus  is  known  only  from  the  type  locality. 
In  this  cave,  it  was  feeding  on  the  guano  of  the  cave-cricket, 
Hadenoecus  subterraneus  (Scudder) ,  in  company  with  Carychium 
stygium  Call.  Typical  H.  notius  is  occasionally  found  in  caves 
also,  but  only  where  leaves  have  been  washed  or  blown  in.  It 
never  has  become  a  guano  feeder. 

Helicodiscus  saludensis    (Morrison) 

Gastrodonta  (Clappiella)  saludensis  Morrison,  1937,  Proc.  Biol. 
Soc.  Wash.  50-58,  PI.  4,  figs.  1-4. 

Clappiella  saludensis  (Morrison) ,  Pilsbry,  Land  Moll.  N. 
Amer.  II,  p.  433,  fig.  233. 

I  believe  that  this  species  should  be  placed  in  the  genus 
Helicodiscus.  It  is  so  like  H.  parallelus  (Say)  in  its  general  ap- 
pearance that  one  needs  a  lens  to  distinguish  it.  H.  fimbriatus 
Wetherby  has  a  similar  pattern  of  internal  teeth,  although  they 
are  not  as  regularly  alternating. 

Troglodiscus,  new  subgenus. 

Shell  with  numerous,  fine,  revolving,  epidermal  fringes;  but 
without  the  revolving  ridges  of  Helicodiscus  s.s.  There  are  no 
internal  teeth  at  any  stage  of  growth. 

Type  species:  Helicodiscus  barri. 
Helicodiscus  barri,  new  species.  Plate  9,  R-T. 

Shell  small,  pale  gieenish-yellow,  subtranslucent,  thin,  de- 
pressed; spire  low,  convex;  whorls  4  to  41/2,  well  rounded,  sutures 
well  impressed.  Umbilicus  moderately  large  and  deep,  contained 
about  3  times  in  the  diameter  of  the  shell.  Sculpture  of  numerous, 
fine,  revolving  epidermal  fringes.  There  being  from  40  to  50  of 
these  fringes  on  the  last  whorl.  Aperture  lunate,  peristome  thin. 
There  are  no  internal  teeth  at  any  stage  of  growth. 

Height,  1.8  mm.  Diameter,  3.9  mm.  Umbilicus  diameter,  1.4 
mm.  Aperture  height,  1.3  mm.  4.5  whorls.  Holotype. 

Distribution:  Tennessee:  Dickson  Co.:  in  Columbia  Caverns, 
2  miles  southwest  of  Van  Leer,  holotype  207799  and  paratypes 


106  NAUTILUS  Vol.   75    (3) 

207800  U.M.M.Z.,  other  paratypes  17446,  collection  of  the 
author.  Davidson  Co.:  in  Bull  Run  Cave,  2.5  miles  northwest 
of  Scottsboro. 

Helicodiscus  barri  is  known  only  from  the  total  darkness  of 
caves.  It  was  found  feeding  on  raccoon  dung.  It  is  named  for  its 
discoverer,  Thomas  C.  Barr,  Jr. 

Helicodiscus  hadenoecus,  new  species.  Plate  8,  J-K;  PI.  9,  U. 

Shell  very  small,  yellowish,  opaque,  subdiscoidal;  whorls  4, 
well  rounded,  sutures  well  impressed.  Umbilicus  moderately 
large  and  deep,  contained  from  21/2  to  3  times  in  the  diameter  of 
the  shell.  Embryonic  whorl  smooth,  later  whorls,  with  fine,  re- 
volving, epidermal  fringes,  which  are  found  on  both  the  upper 
and  lower  surfaces  and  in  the  umbilicus.  There  are  from  30  to  40 
of  these  fringes  on  the  last  whorl.  Aperture  lunate,  peristome 
sinuous,  not  thickened.  There  are  no  internal  teeth  at  any  stage 
of  growth. 

Height,  1.2  mm.  Diameter,  2.7  mm.  Umbilicus  diameter,  1  mm. 
Aperture  height,  1  mm.  4  whorls.  Holotype. 

Distribution:  Kentucky:  Barren  Co.:  in  Beckton  Cave,  0.5  mile 
northwest  of  Beckton.  Tennessee:  DeKalb  Co.:  in  Avant  Cave, 
2  miles  east  of  Dowelltown.  Van  Buren  Co.:  in  McElroy  Cave, 
1.5  miles  northeast  of  Bone  Cave  P.  O.,  holotype  207801  and 
paratypes  207802  U.M.M.Z.,  other  paratypes  17444,  collection 
of  the  author.  White  Co.:  in  Indian  Cave,  2.5  miles  southeast 
of  Quebeck.  Jackson  Co.:  in  Hargis  Cave,  1  mile  north  of  Gran- 
ville (Thomas  C.  Barr,  Jr.,  coll.)  Alabama:  Madison  Co.:  in 
Aladdin  Cave,  7  miles  northeast  of  Maysville. 

Helicodiscus  hadenoecus  is  known  only  from  the  total  dark- 
ness of  caves.  It  feeds  on  the  guano  of  the  cave-cricket,  Hade- 
noecus subterraneus  (Scudder) .  It  is  related  to  Helicodiscus 
barri,  differing  in  its  smaller  size.  Dead  shells  from  which  the 
epidermis  has  been  removed  show  no  sculpture  other  than  weak 
growth  lines.  They  show  no  ridges  like  those  in  H.  parallelus. 
Dead  shells  resemble  those  of  Hawaiia  minuscula  (Binney) . 
Helicodiscus  punctatellus  Morrison. 

The  type  locality  for  this  species  is  in,  not  near.  Whites  Cave, 
near  Mammoth  Cave.  The  author  found  dead  shells  fairly  com- 
mon in  Whites  Cave,  but  no  living  specimens.  The  author  visited 
nearly  every  accessible  cave  in  Mammoth  Cave  National  Park 
without  finding  this  species  in  any  other  cave.  It  apparently 
occurred  only  in  Whites  Cave  and  is  now  extinct. 
Helicodiscus  jacksoni,  new  species.  Plate  8,  L-M;  PI.  9,  Q. 

Shell  very  small,  depressed,  thin;  whorls  4  to  414,  well  rounded, 


January,   1962  nautilus  107 

sutures  well  impressed,  spire  low  but  convex.  Umbilicus  wide, 
occupying  about  40%  of  the  diameter  of  the  shell.  Embryonic 
whorl  smooth,  later  whorls  with  irregularly  spaced  growth  wrin- 
kles, lower  surface  of  last  whorl  nearly  smooth.  Aperture  lunate, 
peristome  simple. 

Height,  1.11  mm.  Diameter,  2.44  mm.  Umbilicus  diameter, 
0.96  mm.  Aperture  height,  0.77  mm.  4.5  whorls.  Holotype. 

Type  locality:  Maryland:  Dorchester  Co.:  subfossil,  in  shell 
mound  near  beach,  0.5  mile  west  of  Elliott,  holotype  207803  and 
paratypes  207804  U.M.M.Z.,  other  paratypes  22476,  collection 
of  the  author. 

Helicodiscus  jacksoni  is  most  closely  related  to  Helicodiscus 
singleyanus  inermis  H,  B.  Baker,  with  which  it  was  found.  It 
differs  in  its  distinctly  larger  umbilicus  and  in  the  more  prom- 
inent growth  wrinkles  on  the  upper  surface  of  the  last  whorl.  The 
whorls  are  a  little  more  slender  and  not  so  tightly  coiled.  The 
aperture  is  smaller.  Only  dead  shells  were  found,  and  most  of 
these  were  chalk  white.  Some  were  fresh,  these  were  translucent, 
with  a  very  pale  yellow  shell. 

It  is  named  in  honor  of  Ralph  W.  Jackson,  who  discovered  this 
interesting  shell  locality. 
Paravitrea  roundyi  Morrison. 

PiLSBRYNA  TRIDENS   MorrisOU. 

These  two  species  seem  out  of  place  in  the  genera  to  which 
they  have  been  assigned.  Possibly  when  they  are  better  known, 
they  will  prove  to  belong  to  the  genus  Helicodiscus. 


TYPE  OF  THE  GENUS  ENGINA  (BUCCINIDAE) 

By  VIRGINIA  ORR 

The  type  species  of  the  genus  Engina  Gray,  1839,  is  Engina 
zonata  Gray,  1839,  by  subsequent  designation  (Gray,  1847) .  This 
species  has  not  been  correctly  identified  by  authors  because  it 
was  not  figured;  the  color  pattern  is  uncommon  for  the  species 
but  principally  because,  in  error.  Gray  described  it  as  II/2  inches 
long   (misprint?) . 

Mr.  S.  P.  Dance,  of  the  British  Museum,  recently  showed  me 
a  lot  of  two  shells,  BV.  113,  marked  Enzina  (sic)  zonata  Gray  in 
Gray's  handwriting.  They  came  from  Mr.  Guilding  and  the 
locality  was  "West  Indies".  Neither  the  adult  nor  the  juvenile 
shell  shows  white  knobs  around  the  base,  the  common  color-form 


108 


NAUTILUS 


Vol.  75    (3) 


Radula  of  Engina  turbinella  (Kiener)  ,  ANSP.  195841,  Buccoo  Reef,  Tobago 
Island,  B.  W.  I.  app.  970  X- 

of  the  species,  but  both  are  easily  identified  as  the  common  west- 
ern Atlantic  species,  Engina  turbinella  (Kiener,  1836) .  The  adult 
shell  measures  11.5  mm.  in  length  (approximately  14  inch)  and 
is  here  designated  lectotype  of  Engina  zonata  Gray.  I  figure  it 
(Plate  10,  fig.  A)  with  a  similarly  colored  specimen  from  St. 
Thomas,  Virgin  Islands,  ANSP  34755    (fig.  B) . 

The  lot  from  which  the  lectotype  is  taken  is  the  one  referred 
to  by  Tomlin  (1928)  as  "a  possible  type  lot"  and  misidentified 
as  Engina  leucozona  (Philippi,  1844).  My  figure  A  is  from  the 
photograph  published  by  Bartsch  (1931) .  However,  I  have  in- 
cluded a  scale  showing  in  the  original  photograph  but  not  previ- 
ously published  because  it  explains  why  Bartsch  did  not  recognize 
the  species. 

Bartsch  saw  only  the  photograph  of  the  specimen  and  had  no 
notes  on  the  size  of  the  shell  except  the  scale  photographed  with 
it.  Reading,  "Axis:  II/2  inches"  in  Gray's  description  he  pre- 
sumed the  scale  was  an  inch  rule  and  translated  the  size  to  27  mm. 
He  made  no  attempt  to  identify  this  outsized  Engina. 

Although,  from  the  misspelling  of  the  generic  name  noted 
earlier,  this  lot  of  Engina  zonata  evidently  was  not  labeled  by 
Gray  in  1839  (he  misspelled  Enzina  in  1842  and  1847),  it  prob- 
ably was  in  his  hands  when  he  designated  the  type  of  the  genus. 
It  is  also  the  only  Gray  material  of  that  name  which  has  come  to 
light.  Except  for  the  discrepancy  in  size,  the  specimens  agree  well 
with  Gray's  description. 

Fortunately,  the  better-known  name  for  the  species,  Engina 


NAUTILUS  75    (3) 


PLATE  9 


■s 


Holotypcs.  N!'.  Ilclicodiscus  noliiis  Ilubviclit.  ().  H.  jacksoiii  Hubrichi. 
R-  1  ,  H.  harri  Hubrichi.  U,  H.  hadrnocrtis  Hiibiichl.  I'holographs  bv  John  V,. 
iV.iuh.  I'liiversity  of  Michigan. 


NAUTILUS  75    (3) 


PLATE  10 


4    i:  n 


:-  'f^y. 


/      . '» 


3  4 

Suci'lla  lupillii.s:  Figs.  I -L'.  spec  inuii  lioni  liaiut-  (I'.S.X.M.  (at.  No.  12{i,'i(j)  . 
Figs.  3-4,  original  figures  in  .\Iaiiini.  ITSO.  pi.  124.  ligs.  I  IPS,  111).  (All  ligines 
approximateh   natural  size.) 


I'ig.  .V.  LctlotNpc  oi  luio/iui  zoitdla  (.ia\.  H\  .  Ii;f.  liiitisli  Miisciiin.  West 
Indies.  .Scale  in  nun.  Fig.  M.  l-:>ioi)in  linb'niclla  (Kiencr),  AN.SP,  'H7."i,").  St. 
Ihomas,  Virgin  Islands. 


January,  1962  nautilus  109 

turbinella  ( Kiener)  precedes  E.  zonata  by  3  years.  Therefore,  the 
type  species  of  the  genus  Engina  Gray  is  Engina  zonata  Gray, 
1839  (non  Reeve  1846)   =  Purpura  turbinella  Kiener,  1836. 

References  cited 
Bartsch,  Paul.  1931.  Proc.  U.  S.  Nat.  Mus.,  79  (15)  :   1-10,  pi.  1, 

fig.  6. 
Gray,  J.  E.  1839.  Zoology  of  the  Blossom,  pp.  112-113. 

1847.  Proc.  Zool.  Soc.  London,  p.  133. 

Tomlin,  J.  R.  leB.  1928.  Nautilus,  -/2  (2)  :  40. 

THE  STATUS  OF  NUCELLA  ROEDING 

By  HARALD  a.  REHDER 

U.  S.  National  Museum,  Smithsonian  Institution 

The  genus  Nucella  was  instituted  by  Roding  (Musuem  Bol- 
tenianum,  1798,  p.  130-131)  for  the  5  species  listed  below.  Fol- 
lowing each  name  I  have  given  what  I  believe  to  be  the  present 
correct  or  proper  assignment  of  the  species  in  question: 

reticulata Cancellaria  reticulata  (Linnaeus) . 

moschatellina Species  dubia. 

macina Nomen  nudum. 

lapillus Paralagena  smaragdula   (Linnaeus) 

theobroma Nucella  lapillus  (Linnaeus) . 

It  will  be  noted  that  I  have  been  unable  to  identify  one  of  the 
species:  N.  moschatellina.  Roding  rechristened  with  this  name 
Gmelin's  Buccinum  laeve,  repeating  the  latter's  reference  to  Mar- 
tini's figures  in  volume  4  of  the  Conchylien-Cabinet  (1780,  pi. 
124,  fig.  1150).  Martini's  description  of  this  species  in  the  ac- 
companying text  (Conchylien-Cabinet,  vol.  4,  1780,  pp.  36-37) 
does  not  quite  agree  with  the  figures,  and  later  authors  have 
been  unable  satisfactorily  to  identify  the  species.  Gray  (Zoology 
Beechey's  Voyage,  1839,  p.  126)  placed  it  in  his  new  genus 
Bullia  and  gave  as  a  synonym  sqiialida  King.  However,  the  finely 
striated  sculpture  and  grooved  interior  of  the  aperture  mentioned 
by  Martini  speak  against  this  assignment.  The  same  criticism  can 
be  leveled  against  Pfeiffer's  (Krit.  Register  Martini  &  Chemnitz 
Konch. — Kat.,  1840,  p.  33)  suggestion  that  the  species  may  repre- 
sent the  juvenile  stage  of  Buccinum  undatum  Linnaeus. 

Clench,  in  his  treatment  of  the  western  Atlantic  species  of 
Purpura  and  Thais  (Johnsonia,  vol.  2,  no.  23,  1947),  has  dis- 
cussed on  pages  85-86  the  problem  of  the  type  of  Nucella,  but 


110  NAUTILUS  Vol.    75     (3) 

because  1  have  come  to  a  conclusion  differing  from  his  I  am 
placing  my  views  on  record,  as  I  feel  that  the  nomenclature  of 
this  common  boreal  group  should  be  settled. 

Nucella  was  first  actively  revived  by  Dall  in  1909  (Prof.  Paper 
U.  S.  Geol.  Survey,  no.  59,  1909,  pp.  46-48,  50)  as  a  subgenus  of 
Thais  Roding.  Clench  (I.e.)  is  coiTect  in  stating  that  Dall  did 
not  specifically  designate  a  type,  although  he  undoubtedly  in- 
tended to  do  so  in  his  outline  of  the  classification  on  page  50, 
where  he  cites  T.  [hais]  lapillus  after  the  name  Nucella  s.s.  Later 
on,  in  his  paper  on  the  members  of  this  group  from  northwestern 
America  (Proc.  U.  S.  Nat.  Mus.  49,  no.  2124,  1915,  p.  558)  he 
did  definitely  designate  N.  lapillus  (Linnaeus)  as  type.  As  Clench 
has  pointed  out,  this  is  an  invalid  designation  since  Linnaeus' 
species  was  not  included  by  Roding  in  his  genus.  Therefore, 
Suter  (Manual  of  New  Zealand  Mollusks,  1913,  p.  425),  Iredale 
(Trans.  Proc.  New  Zealand  Inst.  47,  1915,  p.  472),  and  Grant 
and  Gale  (Mem.  San  Diego  Soc.  Nat.  Hist.  1,  1931,  p.  716)  were 
wrong  in  accepting  Dall's  type  designation.  Wenz  (Handbuch 
der  Palaozool.,  vol.  6,  Pt.  1,  Lief,  6,  1941,  p.  1123)  credits  Dall 
with  citing  filosa  Gmelin  (lapillus  L.)  as  type,  while  Winkworth 
(Proc.  Malac.  Soc.  London  26,  1945,  p.  141),  credits  him  with 
giving  N.  theohroma  as  the  type  species.  Both  these  statements 
are  also  erroneous,  since  Dall,  on  the  one  hand,  merely  listed 
filosa  Menke  (not  filosa  Gmelin)  as  an  example  ( 1909:  46) ,  and, 
on  the  other  hand,  as  Clench  states,  makes  no  mention  at  all  of 
Roding's  theobroma. 

Clench,  however,  has  overlooked  Winkworth's  earlier  type 
designation  (Journ.  of  Conch.  19,  1932,  p.  229) ,  where  he  un- 
equivocally gave  N.  theobroma  Roding  as  type,  without  crediting 
it  to  Dall.  This  is  the  earliest  valid  type  designation  for  Nucella 
that  I  have  been  able  to  discover. 

The  principal  problem  seems  to  be  the  identity  of  Nucella 
theohroma  which  Clench  concludes  must  be  considered  a  species 
dubia.  Roding  refers  his  species  to  Bucciniim  filosum  Gmelin 
(Syst.  Naturae,  Ed.  13,  vol.  1,  Pt.  6,  p.  3486),  and  follows  the 
latter  author  in  citing  two  figures  in  Martini  (Conch.-Cabinet, 
vol.  3,  1777,  p.  433,  pi.  121,  fig.  1113-1114)  as  illustrating  the 
species. 

Clench  feels  that  these  figures  are  unidentifiable,  but  to  me  it 


January,   1962  nautilus  111 

seems  to  be  evident  on  a  number  of  counts  that  they  represent  a 
form  of  the  common  Nucella  lapillus  (Linnaeus) .  First  of  all. 
Martini  himself,  in  the  accompanying  text,  considers  the  shell  a 
variety  of  lapillus,  stating  that  it  differs  only  in  the  color  pattern; 
the  locality  he  doubtfully  gives  as  East  Indies.  Secondly,  Roding 
has  renamed  the  same  shell  again  on  page  132  of  the  "Museum 
Boltenianum"  as  Nossa  ligata,  again  referring  to  Buccinum 
filosum  Gmelin,  and  to  the  same  figures  in  Martini.  Here  it  is 
placed  immediately  next  to  Nassa  rudis  Roding,  which  as  Clench 
points  out,  is  merely  a  new  name  for  lapillus.  Thus  it  seems  cer- 
tain that  Roding  regarded  Buccinum  filosum  as  being  close  to 
lapillus  Linnaeus.  Finally,  an  examination  of  the  U.S.N.M.  col- 
lection of  Nucella  lapillus  revealed  a  specimen,  collected  in 
France,  that  very  closely  resembles  the  figure  given  by  Martini. 
On  plate  10  I  give  a  photograph  of  this  specimen  together  with 
a  reproduction  of  the  figures  of  Martini.  Considering  the  varia- 
bility of  the  species,  I  am  sure  that  specimens  could  be  found 
that  match  the  figure  of  Martini  even  more  closely. 

Polytropa  Swianson,  1840  (Treatise  Malacology,  pp.  80,  305) 
is  the  next  available  name,  and  the  one  that  Clench  (op.  cit.) 
used  instead  of  Nucella.  Swainson  included  in  this  genus  Purpura 
lapillus,  which  Gray  in  1847  (Proc.  Zool.  Soc.  London,  pt.  15, 
p.  138)  designated  as  type.  Thus  Polytropa,  it  is  true,  is  the  first 
taxon  that  was  fixed  unequivocally  on  lapillus,  but  is  a  name 
that  has  been  little  used,  and  Nucella,  as  explained  above,  is 
clearly  available  for  this  group,  and  is  earlier  by  42  years. 

A  survey  of  the  literature  over  the  past  15  years,  using  the 
Zoological  Record  as  a  source,  reveals  that  18  authors  used  the 
name  Thais,  without  subgeneric  assignment,  for  the  species 
lapillus,  4  (all  American) ,  used  Polytropa  (as  a  subgenus  of 
Thais) ,  and  14  authors  used  Nucella,  all  but  two  as  a  distinct 
genus,  a  procedure  that  Clench  (op.  cit.)  suggested  probably 
should  be  followed  for  this  group. 

AGGREGATIONS  OF  THE  TERRESTRIAL  PULMONATE 
CIONELLA  LUBRICA 

By  ERNEST  J.  ROSCOE 
Chicago  Natural  History  Museum 

Widely  distributed  in  Europe,  Asia,  and  north  Africa,  Cionella 
luhrica    (Miiller)    is  generally  spread  over  the  North  American 


112  NAUTILUS  Vol.   75    (3) 

continent  from  northern  Alaska  to  northern  Mexico  (Pilsbry, 
1948) .  The  species  is  quite  ubiquitous  within  its  range.  Although 
there  has  been  little  work  done  on  population  density,  collectors 
generally  report  that  C.  Inbrica  is  difficult  to  obtain  in  t|uantity. 
AVhenever  large  aggregations  have  been  noticed,  the  observation 
has  been  thought  of  sufficient  importance  to  place  on  record. 
Such  recorded  observations  are  summarized  in  the  accompanying 
table,  to  which  the  following  may  now  be  added. 

On  October  27,  1960  Miss  Maidi  Wiebe,  Staff  Artist,  Geology 
Department,  Chicago  Natural  History  Museum,  observed  a  large 
aggregation  of  C.  lubrica,  probably  totaling  several  hundred  in- 
dividuals, on  the  sidewalk  in  front  of  her  apartment  in  Oak 
Park,  Illinois,  at  about  7:30  A.M.  The  pavement  was  moist  from 
a  brief  shower  the  preceding  evening,  although  the  adjacent 
ground  was  not  particularly  wet,  the  moisture  having  permeated 
into  the  soil.  Air  temperature  was  about  47°  F.  No  other  organ- 
isms were  observed  in  association  with  the  snails.  At  5:00  P.M. 
Miss  Wiebe  noticed  that  about  a  fourth  of  the  number  of  snails 
observed  that  morning  were  still  present  on  the  pavement.  At 
7:30  A.M.  the  following  morning  the  snails  had  almost  vanished. 
The  temperature  was  somewhat  higher,  about  54°  F.,  and  the 
pavement  was  dry.  Heavy  rains  fell  on  each  of  the  following  two 
days.  A  few  snails  were  observed  on  the  walk  for  a  period  of 
about  14  days,  none  being  noticed  thereafter. 

The  available  data  are  insufficient  to  warrant  drawing  final 
conclusions.  Some  tentative  suggestions  do  seem  in  order,  and 
the  summation  will  at  least  serve  to  call  attention  to  the  need 
for  additional  observations  and  to  indicate  the  type  of  informa- 
tion required. 

Aggregations  of  C.  lubrica  have  been  observed  in  widely  scat- 
tered localities  in  North  America,  from  Massachusetts  and  Penn- 
sylvania to  British  Cohnnbia.  I  have  been  unable  to  find  refer- 
ences to  any  observed  aggregations  in  the  Old  World.  Quick 
(1954)  states  that  C.  lubrica  is  much  more  abundant  that  its 
close  relative  C.  lubricella  Porro  in  England. 

The  aggregations  have  occurred  from  late  May  or  early  June 
to  late  October.  The  majority  of  observations  have  been  made  in 
late  summer  or  fall.  Krull  and  Mapes  (1952)  have  noted  that 
these  snails  became  increasingly  numerous  after  September  1  in 


January,   1962  nautilus  113 

central  New  York.  They  ascribed  the  greater  numbers  of  snails 
observed  in  association  with  rocks  at  this  time  to  the  increased 
warmth  and  protection  afforded  by  the  stones. 

1  he  observed  aggregations  have  generally  occurred  either  dur- 
ing or  preceding  a  storm.  This  would  tend  to  support  Hender- 
son's (1905)  suggestion  that  they  are  a  response  to  physical  fac- 
tors. However,  Mapes  and  KruU  (1951)  found  collecting  difficult 
even  after  a  rain.  Additional  data  on  response  to  various  environ- 
mental conditions  is  given  in  Krull  and  Mapes  (1952) .  That  such 
aggregations  are  for  the  purpose  of  reproduction  has  been  sug- 
gested (Anon.,  1946;  Pilsbry,  1948) .  Possibly  both  physical  and 
biological  factors  are  involved,  perhaps  depending  upon  the 
season. 

A  hint  of  a  cyclic  phenomenon  is  contained  in  Clapp's  (1914) 
statement  that  "in  25  years  collecting  in  the  Sewickley  Valley 
[Pennsylvania]  I  have  found  this  species  decidedly  rare."  Long- 
time observations  on  populations  of  C.  lubrica  are  necessary.  I 
plan  to  keep  tab  on  the  Oak  Park  population  during  succeeding 
years. 

Stephens  (1918)  found  large  numbers  of  these  snails  fastened 
by  their  secretions  to  the  walls  of  a  building,  all  with  the  apex 
of  the  shell  directed  downwards.  This  suggests  some  sort  of 
oriented  reaction  (oriented  behavior  is  discussed  in  Allee  et  al., 
1949  and  Warden  et  al.,  1940) .  Possibly  this  behavior  was  a  re- 
sponse to  excessive  ground  moisture,  Mapes  and  Krull  (1951) 
noting  that  C.  lubrica  "is  quickly  activated  and  stimulated  to 
climb  up  when  submerged."  About  100  of  the  Oak  Park  snails 
were  kept  under  my  observation  for  several  days  in  a  small  petri 
dish,  the  bottom  of  which  was  covered  by  moistened  newspaper. 
There  was  a  noticeable  tendency  for  the  snails  to  concentrate  in 
the  center  of  the  dish,  in  fact  to  form  a  pile  of  snails,  when  the 
paper  was  quite  moist.  As  the  paper  dried  out  the  snails  tended 
to  move  out  and  distribute  themselves  more  uniformly  over  the 
bottom.  No  mating  activity  was  noticed  during  the  relatively 
short  observation  period.  Oriented  behavior  is  discussed  in  Allee 
et  al.  ( 1949)  and  in  Warden  et  al.  (1940) . 

These  aggregations  of  C.  lubrica  are  in  contrast  to  the  difficulty 
experienced  by  Mapes  and  Krull  in  collecting  this  snail  in  central 
New  York  for  experimental  studies.  Some  of  their  collecting  was 


114 


NAUTILUS 


Vol.  75    (3) 


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done  near  Cazenovia,  the  general  locality  of  Henderson's  (1905) 
observations.  Mapes  and  KruU  found  it  necessary  to  resort  to  the 
use  of  traps  consisting  of  wet  gunny  sacks  spread  over  the  ground 
in  suitable  areas  in  order  to  obtain  sufficient  specimens.  These 
traps,  examined  two  or  four  times  weekly,  resulted  in  only  26 
specimens  per  trap  even  in  favorable  sites  and  under  favorable 
conditions.  These  authors  are  of  the  opinion  that  the  requisite 
factors  for  large  populations  of  C.  lubrica  are  moist  soil,  almost 
constant  shade,  and  a  large  quantity  of  suitable  vegetation. 

The  materials  required  for  the  culture  of  C.  lubrica  are  inex- 
pensive, but  such  cultures  do  require  a  considerable  amount  of 
attention.  Detailed  instructions  may  be  found  in  Mapes  and 
Krull  (1951) .  There  is  here  an  excellent  opportunity  for  some 
interested  amateur  with  the  time  at  his  disposal  to  make  some 
very  worthwhile  observations  on  the  biology  of  this  snail. 

References 
Allee,  W.  C.   1931.  Animal  Aggregations.  Univ.  Chicago  Press, 

Chicago,  111. 
Allee,  W.  C,  et  al.   1949.  Principles  of  Animal  Ecology.  W.  B. 
Saunders  Co.,  New  York.  See  especially  Chapter  23,  pp.  393- 
419. 
Anon.,   1946.  Naut.  60:72. 

Binney,  Amos.  1851.  The  Terrestrial  Air-breathing  Mollusks  of 
the  United  States  and  the  Adjacent  Territories  of  North  Amer- 
ica, vol.  1.  Charles  C.  Little  Sc  James  Brown,  Boston.  Edited 
by  A.  A.  Gould. 
Binney,  W.  C,  and  T.  Bland.  1869.  Land  and  Freshwater  Shells 
of  North  America,  Part  1,  Pulmonata  Geophila.  Smith.  Misc. 
Coll.,  194. 
Clapp,  Geo.  H.  1914.  Naut.  28:96. 
Fromming,  Ewald,  1954.  Biologic  der  mitteleuropaischen  Land- 

gastropoden.   Duncker  &  Humblot,   Berlin. 
Gould,  A.  A.  1870.  Report  on  the  Invertebrata  of  Massachusetts. 

2nd  ed.  Wright  8c  Potter,  Boston    (1st  ed.,  1841,  not  seen). 
Henderson,  J.  B.  1905.  Naut.  /c9;109-l  10. 
Mapes,  C.  R.  1951.  Cornell  Veterinarian  ^/. -382-432. 
Mapes,  C.  R.,  and  W.  H.  Krull.   1951.  Ibid,  II.  Cornell  Veter- 
inarian ■/;. -433-444. 
Pilsbry,  Henry  A.  1948.  Land  Mollusca  of  North  America  (north 
of  Mexico)  .  Vol.  2,  part  2.  Phila.  Acad.  Nat.  Sci.,   Monogr. 
No.  3. 
Quick,  H.  E.  1954.  Proc.  Malacol.  Soc.  London  50.-204-214. 
Stephens,  T.  C.  1918  Science,  n.s.  43:21]. 

Warden,  C.  J.,  et  al.  1940.  Comparative  Psychology,  vol.  2,  Plants 
and  Invertebrates.  Ronald  Press  &  Co.,  New  York. 


IHi  NAUTILUS  Vol.   75    (3) 

PUERTO  RICAN  HOLOPODOPES 
By  H.  BURRINGTON  BAKER 

The  name  Holopodopes  (plural  ot  holopod-ops)  is  proposed 
here  lor  an  infraorder  ot  the  suborder  Sigmurethra,  to  include 
the  achatinoids  (Achatinidae  and  Spiraxidae) ,  Streptaxidae, 
rhytidoids  (Acavidae  proper  -{-  Caryodinae,  Haplotrematidae, 
Rhytididae  and  Chlamydephoridae)  and  orthalicoids  (Urocop- 
tidae  and  Orthalicidae  or  Bulimulidae) .  In  a  systematic  ar- 
rangement, they  should  precede  the  Aulacopoda  (arionoids, 
limacoids  and  testacelloids)  and  the  restricted  Holopoda  (poly- 
gyroids,  oleacinoids  and  helicoids) .  Some  of  the  reasons  for  the 
separation  of  the  holopodopes  have  been  outlined  in  an  earlier 
paper  (1955) . 

The  symbols  used  for  Puerto  Rican  localities  were  explained 
recently    (1961). 

Opeas  piimilum   (Pfeiffer) .  Es2   (coconut  plantation) . 

The  older  name,  Helix  hannensis  Rang,  1831,  from  West 
Africa,  which  may  have  been  this  species,  now  can  be  dropped  as 
obsolete.  The  other  widely  distributed  Opeas,  O.  pyrgula 
Schmacker  &  Boettger,  which  has  less  arcuate  growth  striae, 
probably  also  occurs  in  Puerto  Rico,  although  it  has  not  been 
reported,  unless  the  older,  but  obsolete  name,  Stenogyra  alabas- 
trina  Shuttleworth,  from  near  San  Juan,  represented  it.  The  more 
typical  Ferussaciinae,  Cecilioides  (Geostilbia)  aperta  (Swainson) 
-|-  gundlaclii  (Pfeiffer)  and  C.  (Karolus)  consobrinus  (Orbigny) , 
found  by  van  der  Schalie,  1948:50,  51,  also  have  been  distributed 
widely  by  commerce. 

Lamellaxis  (Leptopeas?)  micra  (Orbigny)  and  var.  rnargari- 
taceus  (Shuttleworth).  Es2  (coconuts),  Pnl  (yams),  Wr2  (cof- 
fee) ,  Wr3.  All  my  dry  shells  (more  in  alcohol)  nearer  the 
smoother  form,  which  is  not  limited  to  Puerto  Rico;  the  obsolete 
name,  Stenogyra  gompharium  Shuttleworth,  from  near  San  Juan, 
may  have  been  based  on  the  typical  one. 

Martens,  1877:345,  reported  the  Cuban  L.  (Leptopeas)  pahi- 
dinoides  (Orbigny)  from  Aguadilla  (Ww),  but  I  got  neither 
it  nor  the  ubiquitous  L.  (Allopeas)  gracilis  (Hutton) ,  which  may 
mean  they  are  more  limited  to  cidtivated  areas. 

Lamellaxis  (s.  s.)  rnonodon   (C.  B.  Adams)   and  var.  opalescens 
(Shuttleworth).  Pnl,  Pr3,  Wr2,  Wn;  my  only  dry  shell    (Wr2) 
the  imperforate  form,  which  also  occurs  in  Jamaica. 


January,   19G2  nautilus  117 

Obeliscus  (Stenogyra)  terebraster  (Lamarck) .  Deep  in  humus, 
Er3,  5,  Prl-4,  6,  Wr2,  3,  1800-4000  ft.;  typical  larger  form. 
O.  (S.)  TERKBRASTER  RARisiNiSTER,  new  subspecies.  Enl,  Es3,  4,  Jnl 
(type  locality),  Pnl;  lowland  form  smaller  throughout,  not  sim- 
ply with  less  whorls.  The  type,  Pilsbry,  1906  (I6a)  :pl.  32,  fig.  31, 
a  sinistral  shell  (ANSP.  59320  from  R.  Swift)  of  this  variety.  It 
measures:  14.4  mm.  by  28  (4.1  mm.)  with  lOi/^  whorls. 

O.  (S.)  swiftianus  (Pfeiffer) .  Ps2  (attains  length  of  10.5  mm., 
with  8.2  whorls)  &  Ws  (smaller)  ;  this  species  has  gone  around 
the  world. 

O.  (Pseudobalea)  hasta  (Pfeiffer) .  Under  leaves  on  ground  and 
in  moss  on  tree  trunks;  Er2,  Jnl,  Pn,  Pr3,  6,  Wr,  100-3400  ft. 

Since  Balea  dominicensis  Pfr.,  1853,  from  "I.  Haiti,"  is  identi- 
fiable only  from  its  inclusion  by  Pfeiffer  himself  in  the  synonymy 
of  O.  hasta,  the  slightly  older  name  is  clearly  obsolete,  even  if 
dated  from  Pilsbry,  1906  (16a)  :272. 

Subulina  octona  (Bruguiere) .  On  ground  under  dead  leaves; 
Ee,  En,  Es2-4,  Jn,  Js,  Pn,  Pr2,  3,  Ps,  Wn,  Wrl,  Ws,  Ww;  0-3400  ft. 

The  obsolete  Stenogyra  (Subulina)  acicularis  Shuttleworth, 
based  on  one  shell  from  near  Fajardo,  seems  to  be  represented  in 
some  eastern  lots  (Ee,  Es2)  from  near  the  seashore  by  occasional 
dwarfed  shells  (with  fewer  whorls,  but  sexually  mature)  which 
have  more  deeply  etched,  growth  striae,  that  do  crenulate  the 
sutures. 

Leptinaria  unilamellata  ( Orbigny) .  Widely  reported  by  van 
der  Schalie,  1948:56,  probably  from  cultivated  places. 

Very  unfortunately,  I,  1945:91,  adopted  Orbigny's  prior  name 
before  the  more  widely  used  L.  lamellata  (Potiez  &  Michaud) 
would  have  been  saved  by  the  50  year  "rule."  Recently,  Aguayo, 
1961:94,  added  the  other,  equally  ubiquitous,  but  more  terrestrial 
L?  (Beckianum)  beckianum  (Pfeiffer) ,  which  already  was  known 
from  both  the  Virgins  and  Haiti. 

Austroselenites  (Zophos)  alticola  H.  B.  Baker.  Deep  in  leaf 
humus,  Er2-4,  and  probably  Erl  and  5  (too  young  for  exact 
identification)  ;  certainly  above  2500  ft,  on  El  Yunque  and  proba- 
bly Luquillo  Mts.  above  2000. 

A.  (Z.)  co77color  (Ferussac) .  Also  terrestrial.  En  3,  4,  Jn,  Pn, 
Prl,  3,  4,  Wr3,  and  probably  Pr  2,  6,  Wn,  Wr2  (too  young)  ;  cer- 
tainly on  lowlands,  and  apparently  represented  by  a  smaller, 
darker  race  up  to  4000  ft.  in  the  Cordillera  Central,  but  my 
material  is  too  scanty  for  certainty. 

The    widely    transported    streptaxid,    Diaphera    (Huttonella) 


118  NAUTILUS  Vol.   75    (3) 

bicolor  (Hutton)  was  found  by  van  der  Schalie,  1948:68.  Dia- 
phera  Albers,  1850,  of  which  Diaphora  Albers-Martens,  1860,  is 
a  homonymic  emendation,  is  prior  to  Gulella  Pfr.,  1856,  and 
D.  bicolor  is  the  type  (Nevill,  1878)  of  Huttonella  Pfr.,  also 
1856,  of  which  Indoennea  Kobelt,  1904,  is  a  subjective  synonym. 

Brachypodelln  (s.  s.)  riisei  (Pfeiffer).  Lowlands,  En,  Jn,  Pn, 
Wn,  Ws. 

B.  (s.  s.)  pallida  ("Pfeiffer"  Philippi) .  Lowlands,  En,  Jn;  Ps2 
(beattyiy,  VVs    (approaching  beattyi)  . 

B.  (Brevipedella)  portoncnnn  (Pfeiffer) .  En,  Jn,  Pn,  \Vn,  Wr2, 
0-2300  ft. 

These  3  species  of  Brachypodella  may  occur  within  a  few  feet 
of  each  other  on  limestone  rocks,  but  B.  portoricana,  much  the 
poorest  climber,  was  the  only  one  found  (Wr2)  far  from  the 
limestone  rim,  and  B.  pallida,  the  best  climber,  was  the  only 
species  collected  at  the  driest  station  (Ps2) ,  where  it  was  buried 
quite  deeply  under  rocks.  Besides  their  difference  in  habits,  these 
shells  from  Ps2  do  average  smaller  and  commonly  their  last 
whorls  are  less  widely  solute  (some  almost  adnate)  but  the  "dif- 
fers" in  riblets  seem  individual  variation.  Some  are  about  the 
size  of  typical  beattyi  Clench,  1951:251,  fig.  3,  from  Mona  Island; 
one  with  all  17  whorls  measures  10.3  mm.  (over  all)  by  2.1  (not 
including  aperture) .  Many  more  were  found  intact  but  even  a 
little  shaking  rendered  them  decollate. 

Pseudopineria  viequensis  (Pfeiffer) .  In  holes  and  under  over- 
hang of  cliffs,  on  or  beneath  the  limestone  outcrops  of  the  north- 
ern rim,  commonly  in  shaded  places  where  Brachypodella  was 
absent;  Jnl,  2.  Foot  whitish;  ommatophores  translucent,  cylin- 
dric,  with  black  eyes;  inferior  tentacles  small  but  certainly  pres- 
ent; sole  unizonal,  with  1  or  2  waves  that  involve  its  full  width. 

Macroceramiis  rnicrodon  (Pfeiffer).  Pilsbry,  1903  (25a)  :  1 15- 
1 16,  pi.  24,  figs.  71-72  (ANSP.  251 17  &  25115,  from  R.  Swift)  ;  van 
der  Schalie,  1948:map  62,  pi.  7,  figs.  2a,  2b.  En,  Jn;  growth 
threads  narrower  than  their  interspaces  (much  more  so  on  sub- 
apical  whorls)    and  irregularly  spaced  on  later  whorls. 

The  obsolete  M.  johannis  Pfeiffer  appears  indistinguishable, 
and  Aguadilla  falls  within  the  range  of  the  typical  subspecies. 

M.  rnicrodon,  var.  sliultleiuorthi  (ALirtens) ,  1877:352  (?) , 
without  exact  locality.  Pilsbry,  fig.  74  (ANSP.  2572,  from  Bland)  ; 
van  der  Schalie:  fig.  2c.  Ws:  largest  18.7  mm.  long  with  12  whorls. 
Ps2:  1  adult  18.8  mm.  long  (decollate)  and  2  immature,  one  of 
which  approaches  loeryi  in  growth  threads. 

M.  rnicrodon  loeryi  Jacobson,  1955.  Ps2    (near  type  locality)  ; 


January,  1962  nautilus  119 

one  adult  and  1  immature,  empty  shells;  look  as  i£  started  life  like 
var.  a,  but  assumed  later  whorls  ot  var.  b,  which  see.  Adult  14.1 
mm.  by  55    (7.7  mm.)    with  7  whorls  remaining. 

M.  rnicrodun  loeryi,  var.  a.  Ps2,  3.  Shells  about  size  and  form  of 
typical  microdon;  one  (Ps2)  measures:  14.1  mm.  by  36  (5.0  mm.) 
with  11.2  whorls;  ranging  to  17.2  mm.  long  with  12.6  whorls; 
without  bluish  tinge  when  alive;  1  shell  unicolor,  without  white 
patches. 

AI.  microdoji  loeryi,  var.  b.  (?)  M.  shuttleworthi  Martens,  1891: 
132,  Penuelas.  Under  bunch  grass,  but  climbing  4  to  6  ft.  during 
rain;  near  Tallaboa  (Psl).  Shell  with  growth  threads,  especially 
on  later  whorls,  broader  than  their  interspaces  and  quite  evenly 
and  closely  spaced;  and  those  on  subapical  whorls  less  widely 
spaced  than  in  typical  microdon.  Predominant  color  of  shell  light 
sky-blue  on  living  examples  but  now  (1960)  almost  completely 
faded  to  whitish  (opaque)  with  narrow,  light  brownish  (trans- 
lucent) growth  (axial)  bands,  as  were  empty  shells  found  nearby 
(one  of  last  unicolor,  without  whitish  patches)  ;  with  almost  no 
trace  of  basal  angulation  in  many;  largest  21.1  by  37  (7.8  mm.), 
minor  diam.  33  (6.9  mm.)  with  13.2  whorls;  smallest  19.8  mm. 
long  wuth  1 1  whorls  remaining.  Living  animal  light  slate  color, 
darker  near  sole  and  on  ommatophores;  mantle  collar  dark 
brownish  gray,  Avith  minute  light  spots;  sole  unizonal,  with  1  or  2 
locomotor  waves  across  its  full  width. 

In  1939,  var.  b.  was  considered  a  subspecies  of  M.  miaodon 
(Cf.  van  der  Schalie:96)  but  I  hesitated  to  name  it  because  of  the 
doubt  about  the  true  M.  shuttleworthi,  which  Martens  later 
located  at  the  nearby  Penuelas.  The  above  presents  the  old  notes, 
with  a  few  changes;  typical  loeryi  and  var.  a  were  included,  as 
now. 

Microceramus  (s.  s.)  guanicus  H.  B.  Baker.  On  rocks  near 
ground;  only  known  from  type  locality  (Ps2)  and  Mona  Island 
(Clench,  1951). 

Bulimulus  (s.  s.)  guadalupensis  (Bruguiere) .  On  ground  and 
tree  trunks  up  to  5  ft.,  Ee,  Es2  (coconut  plantation),  Es4,  Jnl, 
Ps4    (garden) ,  Wwl    (garden) . 

For  this  widely  disseminated  pest,  van  der  Schalie,  1948:87,  re- 
turned to  B.  exilis  (Gmelin). 

B.  (s.  s.)  diaphanus  (PfeiflPer) .  More  terrestrial,  Pnl,  Pr2,  4, 
Ps2,  3,  Wn,  Ws;  0-3100  ft.  Shell  epidermis  rufous  (typical)  to 
whitish. 

This  may  be  a  native  species,  although  described  originally  from 
St.  Thomas.  Pilsbry's,  1897  (9b)  : 46-47,  transference  to  this  species 
of  the  Puerto  Rican  records  of  the  now  obsolete  B.  fraterculus 


120  NAUTILUS  Vol.   75    (3) 

("Fer."  Potiez  &  Michaud,  1838)  is  accepted;  certainly  nothing 
like  the  original  (P.  &  M.)  figures  has  been  found  since.  The 
last  name  was  nude  in  Ferussac,  1821,  livr.  11:54,  and  in  Beck, 
1837:67,  although  Beck  questioningly  (?)  referred  it  to  Helix 
tenuissirna  Ferussac,  1832    (-|-  Orbigny,  1835?). 

Drymaeus  (Mesembrinus)  virgulatus  (Ferussac,  1821)  and  color 
form  (?)  liliaceus  (Fer.,  1832).  Arboreal,  edges  of  Psl  (30% 
Uliaceus)  and  Ps2  (45%  liliaceus)  ;  empty  shells  seen  elsewhere 
but  only  near  cultivated  places. 

The  separation  of  these  two  lots  into  liliaceus  (complete  ab- 
sence of  brownish  color)  and  virgulatus  (with  even  a  few  streaks, 
up  to  axial  and/or  spiral  bands  of  variable  continuity)  seems 
highly  arbitrary.  Confessedly,  most  of  the  "lilies"  have  a  less 
elongate  (Roding)  form  and  a  slightly  swollen  last  whorl,  but 
they  include  the  most  elongate^  shell  in  the  series,  and  typical 
virgulatus  covers  almost  the  entire  range  of  shell  form.  These 
may  be  hybrids  (of  an  introduced  with  a  native  form?)  but  they 
cast  doubt  on  any  specific  separation,  even  if  it  has  continued 
for  130  years. 

D.  (M.)  multilineatus  (Say)  ;  Cf.  form  osmenti  Clench  and/or 
eboreus  Grimshawe.  Along  road  to  Wn.  One  shell  from  living 
adult  resembling  in  form,  texture  and  coloration  those  of  Pilsbry, 
1946(11)  :27,  figs.  15b,  with  similar  columellar  blotch  (absent  in 
thicker  and  chalkier  D.  virgulatus)  and  slightly  wider,  sutural 
stripe,  but  lacking  all  other  dark  bands  except  4  (plus  traces) 
axial  streaks  near  aperture,  and  showing  no  signs  of  characteristic 
bluish  color  near  apex.  Another  shell  too  broken  and  bleached 
to  be  sure  of  its  form,  with  similar  columellar  patch  and  axial 
streaks,  but  without  sutural  stripe. 

Neither  fits  the  obsolete  Drymaeus  hjalmarsoni  (Pfr.) ,  from 
"near  Manati"  (between  Jn  and  Pn) ,  which,  as  Pilsbry  (1899) 
suggested,  seems  to  have  been  close  to  his  (1946)  Leptodrymaeits, 
and  even  might  be  the  subsequently  named  D.  dormani  (W.  G. 
Binney) . 

Simpulopsis  (Eudioptusf)  psidii  (Martens,  1877).  Not  ob- 
tained. 

From  field  notes,  the  young  shells  of  Platysuccinea  from  the 
Cordillera  Central,  which  approach  it  in  height  of  spire,  were 
confused  with  this  arboreal  species,  and  thus  no  special  search 


1  More  so  than  beattyi  Clench,  1951:  fig.  6,  but  less  so  than  his  type,  figs. 
4  &  5. 


Januai7,  1962  nautilus  121 

was  made  tor  it  west  of  Utuado  (between  Wr6  and  Pnl) .  How- 
ever, a  shell  ot  the  sagdid  at  the  size  of  S.  psidii  (5  by  5  mm.) 
has  one  less  whorl  (2  vs.  3)  ;  being  small,  mature  5.  psidii  might 
be  found  only  near  the  end  of  a  long  rainy  period.  Pilsbry, 
1902  (31)  :lxvii,  included  both  Eudioptus  Albers-Martens  (not 
a  homonym)  and  Platysiiccinea  in  the  bulimulid  Simpulopsis, 
but  no  geophile  systematist  in  this  century  has  guessed  either  to 
be  a  "Bulimulus."  The  type  species  of  Eudioptus,  in  which 
Martens  placed  his  "Bulimus"  psidii,  is  Brazilian,  but  a  dweller 
on  guava  leaves  might  be  carried  by  commerce. 

Gaeotis  nigrolineata  Shuttleworth.  Mainly  on  leaves  of  palms, 
but  also  on  those  of  Cecropia  and  other  trees,  Er3,  4,  5;  in  axils 
of  palm  pinnae,  Wr3;  from  Luquillo  Mts.  to  western  end  of  the 
Cordillera  Central;  2000-3000  ft.  Living  animal  (Er)  3  by  li/g 
inches  (76  by  38  mm.)  ;  yellowish  gieen,  but  variable;  dorsum 
of  foot  slaty  with  yellow  middorsal  stripe  (flavolineata)  which 
often  becomes  double  anteriad;  and  often  with  frosty,  whitish 
patches  (like  albopunctulata.})  ;  sides  and  inferior  tentacles 
lighter,  but  sole  often  with  orange  margin  on  anterior  I/3 
(flavolineata);  ommatophores  blue  green,  but  eyes  lighter;  mantle 

(over  shell)  pea  green;  often  with  (internal)  black  streaks 
visible  (nigrolineata). 

Very  flimsy  shell  "sigaretiform"  but  thin  lower  surface  of 
apical  whorls  present  and  containing  those  of  animal,  so  that 
separation  in  preserved  examples  means  damage  to  one  or  the 
other  (shells  still  on  most  of  mine) .  Lower  surface  of  apical 
whorls  broken  away  from  all  6  shells  studied  by  Pilsbry,   1899 

(35):227-231,  as  follows:  ANSP.  4613,  one  labeled  "albopunctu- 
lata" and  one  (4614)  "nigrolineata"  from  Humacao  (Bland) . 
ANSP.  26052,  type  lot  of  malleata;  2  shells  from  "near  San  Juan," 
from  Swift,  collected  in  1855.  ANSP.  4959,  one  juvenile  labeled 
"flavolineata"  by  Bland  (not  by  Pilsbry)  from  "Luquillo." 
ANSP.  26051,  labeled  "nigrolineata"  from  "Pto.  Rico"  (Swift). 

G.  nigrolineata  is  the  type  species  of  Gaeotis,  by  subsequent 
designation  of  Kobelt,  1880,  111.  Conch.:264.  Although  only  pre- 
liminary dissections  have  been  made  as  yet,  all  my  material  looks 
like  one  variable  species,  but  none  of  it  comes  from  the  lowlands 

(secretive  during  dry  weather?) .  On  El  Yunque,  this  bulimulid 
slug  was  fairly  common,  but  its  coloration  blends  with  the  leaves. 

Additional  references    (See   1961) 
1961  (I),  Naut.  74:142.  1961  (5),  Naut.  75:64. 
Aguayo,  Carlos  G.  1961,  Caribb.  J.  Sci.  /.•89-106. 
Jacobson,  Morris.  1955,  Archiv  f.  Molluskenk.  84:91-99. 


122  NAUTILUS  Vol.  75    (3) 

JULIA  ANNA  GARDNER 
1882-1960 

Julia  Anna  Gardner,  known  around  the  world  for  her  work  in 
stratigraphy  and  molluscan  paleontology,  died  after  a  long  ill- 
ness on  November  15,  1960.  She  was  a  geologist  with  the  United 
States  Geological  Survey  for  32  years. 

Julia  Gardner  was  born  in  Chamberlain,  South  Dakota,  on 
January  26,  1882.  Her  undergraduate  studies  were  done  at  Bryn 
Mawr  College  where  she  received  the  Bachelor  of  Arts  degree  in 
1905  and  the  Masters  degree  in  1907.  Miss  Gardner  entered  The 
Johns  Hopkins  University  in  the  fall  of  1907,  the  first  woman 
regularly  admitted  to  the  Department  of  Geology.  After  receiving 
her  doctoral  degree  in  1911,  she  continued  there  as  Assistant  in 
Paleontology  until  1915.  Her  studies  of  the  Late  Cretaceous 
Mollusca  of  Maryland  and  other  smaller  groups  were  published 
by  the  Maryland  Geological  Survey  in  1916.  In  1915  she  began 
work  on  the  Miocene  Alum  Bluff  Group  of  Florida  under  con- 
tract with  the  United  States  Geological  Survey. 

During  World  War  I,  the  broad  and  specific  humanitarianism 
which  characterized  her  life  impelled  her  to  go  to  France  with 
the  Red  Cross  in  1917.  She  saw  strenuous  service  as  an  Auxiliary 
Nurse  and  after  the  war  with  the  American  Friends  in  the 
devastated  areas  of  France. 

Upon  her  return  to  the  United  States  in  1920,  Miss  Gardner 
joined  the  Geological  Survey  and  took  up  again  her  professional 
career,  which  was  to  become  long  and  distinguished. 

She  was   the  author  of  over  40  reports.   Pre-eminent  among 
these  are: 
The  molluscan  fauna  of  the  Alum  Bluff  group  of  Florida:  LJ.  S. 

Geol.  Survey  ProL  Paper  142,  1926-1947. 
The   Midway   group   of  Texas:    Texas   LTniv.    Bull.    3301,    1933 

[1935]. 
The  Mollusca  of  the  Tertiary  form;ttions  of  northeast  Mexico: 

Geol.  Soc.  America  Mem.  11,  1947. 

Their  scientific  value  was  equaled  by  their  usefulness  in  the 
field  of  petroleum  geology,  and  they  are  standards  of  reference 
in  the  economic  investigations  of  Tertiary  strata  of  both  North 
and  South  America.  Her  published  works  are  foundation  stones 
and  bench  marks  in  Coastal  Plain  stratigraphy  and  paleontology 


January,   1962  nautilus  123 

that  insure  Julia  Gardner  a  high  place  with  the  pioneers  in  the 
geology  ot  the  region. 

Among  the  societies  of  which  Miss  Gardner  was  a  member  are 
Phi  Beta  Kappa;  Sigma  Xi;  the  Geological  Society  of  America, 
of  which  she  was  a  fellow  and  vice  president  in  1953;  the  Ameri- 
can Association  of  Petroleum  Geologists;  and  the  Paleontological 
Society,  which  she  served  as  president  in  1952.  She  was  a  charter 
member  of  the  American  Malacological  Union  and  was  widely 
known  and  loved  by  collectors  everywhere. 

She  was  an  official  delegate  of  the  United  States  to  the  Inter- 
national Geological  Congress  in  Madrid  in  1926,  and  to  the  In- 
ternational Congress  in  Moscow  in  1937. 

Julia  Gardner  made  a  great  contribution  in  her  chosen  field  of 
geology;  an  equally  great  contribution  in  the  field  of  human  rela- 
tionship may  be  exemplified  by  her  encouragement  of  the  Japan- 
ese scientists  while  on  a  tour  of  duty  in  Japan  after  World  War  II. 
They  confirm  that  her  kindness  and  generosity  heartened  them 
to  carry  on  their  researches.  In  her  wide  circle  of  friends,  few  do 
not  have  cause  to  remember  some  kindness,  some  encouragement, 
some  tangible  assistance,  timely,  but  given  unexpectedly.  Julia 
Anna  Gardner  will  live  long  in  the  hearts  of  her  friends.  —  Druid 
Wilson. 


NOTES  AND  NEWS 


PoMACEA  PALUDOSA  IN  ALABAMA  —  In  1943,  John  Richardson 
II  collected  Pomacea  paludosa  (Say)  at  Gainesville,  Florida,  and 
introduced  it  into  ponds  on  his  farm  7  miles  north  of  Jackson- 
ville, Florida.  From  here,  in  1953,  Frank  Lyman  introduced  it 
into  a  small  pond  on  the  shore  of  Gantt  Lake,  5  miles  northeast 
of  Gantt,  Covington  Co.,  Alabama.  It  soon  escaped  into  Gantt 
Lake,  and  is  now  to  be  found  along  the  shore  for  a  mile  in  either 
direction  from  Lyman's  pond.  —  Leslie  Hubricht. 

S\ic<:iriea  Indiana  Pilsbry. 

Succinea  Indiana  Pilsbry,  1905,  Nautilus,  19:  28. 

Siiccinea  vaginocontorta  Lee,  1951,  Occas.  Papers  Mus.  Zool. 
Univ.  Mich.  no.  533,  pp.  1-7,  PI.  1-2. 

A  careful  examination  of  the  anatomy  and  shell  of  topotypes 


124  NAUTILUS  Vol.  75    (3) 

Succinea  indiana  showed  no  character  which  could  be  used  to 
distinguish  it  from  S.  vaginacontorta.  The  same  twisted  vagina 
is  found  in  S.  indiana.  S.  vaginacontorta  Lee  must  be  placed  in 
the  synonomy  of  S.  indiana. 

Additional  records  for  Succinea  indiana  are  as  follows:  Xorth 
Carolina:  Craven  Co.:  sandy  roadside,  4.4  miles  west  of  New 
Bern.  South  Carolina:  McCormick  Co.:  below  Clark  Hill  Dam. 
Georgia:  Richmond  Co.:  roadside,  1600  Gordon  Highway,  Au- 
gusta. Alabama:  Montgomery  Co.:  waste  ground,  Atlanta  High- 
way and  Forest  Hills  Ave.,  Montgomery.  Dallas  Co.:  near  Ala- 
bama River,  opposite  Selma.  —  Leslie  Hubricht.' 

ViviPARUs  suBPURPUREUs  IN  OKLAHOMA.  —  A  single  Specimen, 
33.0  mm.  in  height  and  32.0  mm.  in  greater  diameter,  of  this 
species  was  discovered  in  a  small,  mud-bottomed,  heavily-vege- 
tated stream  at  the  south  boundary  of  Tom,  McCurtain  County, 
Oklahoma  on  16  July,  1961.  The  spire  was  greatly  eroded.  Al- 
though this  specimen  represents  a  new  record  for  Oklahoma,  it 
is  not  a  particularly  surprising  one.  The  Austroriparian  Biotic 
Province  extends  into  Oklahoma  at  this  point  but  it  is  separated 
from  the  main  body  of  that  ecological  division  by  the  Red  River. 
Viviparus  subpurpureus  (Say)  is  a  fairly  common  species  in  ad- 
jacent Texas  and  Louisiana  but  apparently  has  met  some  diffi- 
culty in  traversing  the  river  barrier  into  Oklahoma,  as  I  have 
collected  rather  extensively  in  this  part  of  the  state  and  have  not 
found  the  species  before.  —  Branley  A.  Branson,  Dept.  of  Biol- 
ogy, Kansas  State  College,  Pittsburg. 

Helisoma  anceps  transported  by  a  giant  water  bug.  —  On 
5  May,  1961,  I  collected  a  giant  water  bug,  Lethocerus  americanus 
(Leidy),  which  had  been  attracted  (at  night)  to  a  lOO-watt 
mercury  lamp  on  a  lawn  at  the  University  of  Michigan's  Erwin  S. 
George  Reserve,  Livingston  County,  Michigan.  The  bug  was 
carrying  on  its  scutellum  a  specimen  of  Helisoma  anceps 
(Menke) .  The  snail  was  alive  and  appeared  to  be  immature, 
measuring  5  mm.  high  and  7  mm.  long.  Giant  water  bugs  of  this 
species  and  a  similar  species,  Benacus  griseus  (Say) ,  frequently 
come  to  lights  in  early  spring  and  later  in  the  summer;  evidently 


1  Accidentally  omitted   from   Naut.   ?'':()(). —  H.   B.   B. 


I 


January,   1962 


NAUTILUS 


125 


there  is  much  movement  between  ponds,  presumably  associated 
with  mating  behavior.  Any  animal  that  happened  to  become 
attached  to  a  bug  would  be  similarly  transported.  During  1959- 
61,  I  examined  about  500  of  these  bugs,  but  this  is  the  only 
occasion  I  found  an  attached  snail.  But  among  several  hundred 
hydrophilid  beetles  similarly  attracted  to  light,  I  found  3  with 
immature  leeches  attached  to  their  elytra.  The  leeches  at  first  ap- 
peared dead,  but  quickly  became  active  when  placed  in  water.  In 
view  of  frequent  speculation  as  to  how  totally  aquatic  animals 
colonize  isolated  ponds,  these  records  may  be  of  interest.  —  D.  F. 
Owen,  Museum  of  Zoology,  University  of  Michigan,  Ann  Arbor. 


The  anatomy  of  glyphyalinia  junaluskana. — Glyphyalinia 
junaluskana  (Clench  &  Banks)  generally  has  been  treated  as 
a  subspecies  of  G.  sculptilis  (Bland) .  However,  a  study  of  the 
genitalia  shows  that  it  is  quite  distinct.  The  following  descrip- 
tion of  the  lower  genitalia  is  based  on  a  specimen  collected  at 


Glyphyalinia    junaluskana    (Clench    &    Banks).    Three    views   of    terminal 
genitalia,  drawn  with  aid  of  camera  lucida.  Scale  line  =  1  mm. 


126  NAUTILUS  Vol.  75    (3) 

4700  ft.,  0.6  mile  southeast  of  Beech  Gap,  GTaham  Co.,  North 
Carolina. 

Penis  (fig.)  rather  short  and  stout,  club-shaped,  basal  portion 
nearly  white;  upper  portion  pale  brownish,  globose.  Penial 
retractor  short  and  stout,  inserted  at  apex  of  penis.  Epiphallus 
parallel  to  penis,  long  and  stout,  as  large  as  the  penis;  upper 
end  deeply  cleft  into  two  equal  lobes;  lower  portion  pale  brown, 
becoming  darker  at  upper  end.  Spermatheca  narrowly  obovate, 
not  clearly  differentiated  from  the  thick  duct.  Vagina  and  free 
oviduct  short. 

It  differs  from  G.  sculptilis  in  being  without  the  penial  appen- 
dix of  that  species  and  in  having  the  epiphallus  divided  into  two 
distinct  lobes.  It  apparently  belongs  in  section  Glyphyalinia.  s.  s 
rather   than    in   section    Glyphognomon    H.    B.    Baker. — Leslie 

HUBRICHT. 

CoRBicuLA  FLUMiNEA  (Miiller)  from  the  Ohio  River — On  a 
field  trip  last  August  30,  1960,  to  the  Ohio  River  just  east  of 
Fort  Massac  and  Metropolis  in  Massac  County,  Illinois,  I  col- 
lected 1 1  recently  dead  specimens  of  Corbicula  (Corbicula)  flu- 
minea.  They  were  15-20  feet  from  the  north  bank  in  6''  of  water. 
The  bottom  was  more  gravel  than  sand  with  very  little  mud 
present.  The  areas  on  either  side  were  muddy  bottoms  and 
about  one  foot  deep,  yet  no  specimens  were  found. 

Corbicula  fiinninea,  a  native  of  eastern  Asia,  has  not  been 
reported  from  the  northeast  or  central  states  as  of  this  writing; 
therefore,  I  can  mention  those  collected  at  Metropolis,  Illinois, 
as  new  in  this  part  of  the  country.  Sinclair  and  Ingram  (1961, 
Naut.  7-/:l  14-1 18)  reported  the  species'  presence  in  the  Tennessee 
River  in  Hardin  County,  Tennessee.  Previously,  the  species  was 
known  only  from  the  south  to  north  western  states,  i.e.,  Arizona, 
California,  Idaho,  Nevada,  Oregon  and  Washington  (op.  cit.) . 
Corbicula  flumiyiea  seems  to  be  widely  distributed  over  the 
United  States.  Records  of  its  further  spread  are  to  be  expected 
with  additional  collecting. 

Since  I  am  not  an  expert  on  foreign  sliells,  the  specimens  were 
submitted  to  Dr.  Fritz  Haas  of  Chicago  Natural  History  Museum 
for  identification.  They  are  now  Chicago  Natural  History  Mu- 
seum No.  103678.  Sincere  appreciation  is  extended  to  Dr.  Haas 
for  his  assistance. — Frederick  R.  Fechtner. 


I 


THE  NAUTILUS 

Vol.   75  APRIL,   1962  No.  4 


A  NEGLECTED  WEST  ATLANTIC  STROMB 

By  a.  a.  OLSSON  and  D.  R.  MOORE 

In  1949,  Mr.  Leo  A.  Burry  described  and  figured  (Shell  Notes, 
December,  1949) ,  a  most  unusual  Strombus  under  the  name  of 
S.  canaliculntus,  reported  as  having  been  taken  in  a  fish-trap  set 
at  a  depth  of  175  fathoms  off  Hopetown,  Elbow  Key  in  the 
Bahamas.  The  whereabouts  of  the  type  is  unknown.  The  figure 
shows  an  immature  specimen  in  which  the  lip  is  not  yet  fully 
formed.  Subsequent  to  the  description  of  the  species,  the  junior 
author  saw  3  additional  specimens  which  Burry  had  obtained 
from  the  same  source.  Two  specimens  were  half  grown  and 
smaller  than  the  holotype,  the  third  was  fully  matured  with  a 
flared  and  thickened  outer  lip.  This  last  specimen,  a  topotype, 
is  in  the  reference  collection  of  the  Marine  Laboratory  of  the 
University  of  Miami  (Accession  no.  30:1839).  It  measures  242 
mm.  in  length.   (Plate  11.) 

The  full-grown  shell  has  a  high,  broad,  conic  spire  of  8  or 
more  whorls,  each  edged  with  a  sharp,  peripheral  keel  or  carina, 
the  main  surface  of  each  spire-whorl  smooth,  except  for  the  fine 
lines  of  growth.  There  are  no  axial  riblets  at  any  stage.  The 
peripheral  keel  becomes  more  strongly  emphasized  on  the  shoul- 
der of  the  body-whorl  and  at  its  termination  on  the  edge  of  the 
outer  lip  it  forms  into  a  deep  notch.  The  wall  of  the  shell  is 
quite  heavy,  interior  of  aperture  with  a  pink  coloration  and  the 
surface  is  covered  with  the  usual,  light-brown,  peeling  perio- 
stracum.  The  report  that  this  shell  was  taken  from  a  fish-trap 
seems  very  questionable.  There  seems  to  be  notliing  closely  re- 
lated to  S.  canaliculatus  in  the  living  fauna;  S.  leidyi  Heilprin  of 
the  Florida  Pliocene  is  perhaps  the  nearest,  but  even  in  this  case, 
the  resemblance  is  not  close. 

The  collection  of  at  least  4  typical  specimens  of  S.  canaliculatus 
in  a  relatively  short  time  and  from  the  same  general  area  seems 
to  preclude  a  chance  anomaly  or  a  deformed  or  diseased  shell. 
The  absence  of  axial  ribbing  on  any  part  of  the  whorls  of  the 

127 


128  NAUTILUS  Vol.  75  (4) 

spire  is  again  emphasized.  More  material  with  accurate  data  as 
to  station  and  environment  is  needed  and  we  are  calling  re- 
newed attention  to  this  interesting  species  so  that  collectors 
visiting  the  Bahamas  can  be  on  the  watch  for  it. 


THE  STATUS  OF  STROMBUS  CANALICULATUS 

By  ROBERT  ROBERTSON 

Assistant  Curator  of  Mollusks, 

Academy  of  Natural  Sciences  of  Philadelphia 

Olsson  &  Moore  (1962)  have  claimed  that  Strombus  canalicu- 
latus  Burry  (1949)  is  a  distinct,  "neglected"  species,  not  closely 
related  to  any  other  living  species.  They  have  discounted  the  idea 
that  the  4  known  specimens  are  abnormal. 

5.  canaliculatus  was  originally  described  on  the  basis  of  a  sin- 
gle, sub-adult  shell,  purportedly  from  a  fish  trap  in  175  fathoms 
off  Hope  Town,  Elbow  Cay,  Great  Abaco,  Bahama  Islands.  The 
trustworthiness  of  this  type  locality  has  bearing  on  the  validity 
of  this  species.  If,  as  Olsson  has  suggested  (m  litt.),  it  is  a  deep 
water,  "outer  shelf"  species,  it  could  have  been  overlooked. 

Unfortunately,  Burry  gave  no  information  as  to  how  he  ob- 
tained the  holotype.  According  to  D.  R.  Moore  {in  litt.) ,  Burry 
obtained  it  from  a  man  named  Ford  [not  Rev.  Paul  D.],  who  had 
a  shop  in  Riviera  Beach,  Florida,  and  who  imported  shells  from 
the  Bahamas.  Possibly,  Ford  bought  the  shell  from  a  fisherman. 
There  are  four  reasons  to  doubt  that  this  shell  (collected  alive) 
came  from  175  fathoms  off  Hope  Town:  (1) .  Although  detritus 
feeders  can  live  at  great  depths,  I  doubt  that  such  a  large, 
herbivorous  gastropod  (Robertson,  1961)  could  live  at  175 
fathoms,  well  below  the  photosynthetic  zone.^  (2) .  The  escarp- 
ment bounding  the  Little  Bahama  Bank  off  Hope  Town  is 
precipitous.  The  margin  of  the  Bank  is  at  a  depth  of  55-70 
fathoms  (personal  observations)  ;  soundings  of  about  1000 
fathoms  are  recorded  as  near  as  5  miles  to  the  Bank  (see  H.O. 
chart  0026e)  .  (3).  The  currents  off  Abaco  are  so  strong  (2-3 
knots)   that  retrieval  of  a  fish  trap  from  175  fathoms  would  be 


'  Various  Strombus  species,  all  smaller  than  5.  canaliculatus,  are  known 
definitely  to  live  in  depths  of  at  least  30-43  fathoms,  but  not  deeper.  A  tiny 
species  has  been  dredged  alive  in  the  Hawaiian  Islands  between  43  and  66 
fathoms  (see  Abbott,  1960,  pp.  83-84) . 


NAUTILI TS  75(1) 


PLATE   II 


ligs.  1:1  Siioinhu.s  canaliculalus  Bmry.  Length:  242 


NAUTILUS  75(4) 


PLATE   12 


^  i 


^'    L.^k 


Spires  of  iiialloniu'd,  tarinalc  Mxnnbus  (Ti  i<(>i  iii.s)  i^/iic/.s  Linn.  MalKima 
Islands  (?)  .  Fij^.  I.  Holotype  of  S.  caintlii  iihil us  lluiiy.  Drawing  l)v  (lilbcrt  L. 
Voss  in  Burrv  (1949).  Fig.  2.  Pui  ported  "topoiype"  of  ,S.  raiialirulatus.  Same 
slicll  as  tliat  figured  l)\  Olsson  &  Moore  (19(32).  Both  figs,  approx.  natural  size. 


April,   1962  nautilus  129 

impossible  without  equipment  not  available  to  local  fishermen, 
Avho  do  not  place  their  fish  traps  ("pots")  outside  the  reef. 
(4) .  During  3  summers  (1953-1955)  while  collecting  shells  at 
Hope  Town,  1  was  unable  to  get  any  information  from  the  local 
fishermen  about  the  finding  of  5.  canaliculatus.  These  fishermen 
remembered  details  about  shells  of  commercial  value  collected 
years  before. 

In  view  of  these  4  reasons  to  doubt  that  the  holotype  came 
from  175  fathoms  off  Hope  Town,  it  is  improbable  that  the 
other  3  specimens  of  S.  canaliculatus  (see  Olsson  &  Moore)  came 
precisely  "from  the  same  source  [locality]".  Olsson  &  Moore 
concede  that  they  doubt  that  the  "topotype"  figured  by  them 
was  taken  from  a  fish  trap.  Lyman  (1951)  reported  that  the 
adult  specimen  (presumably  the  "topotype")  "was  brought  in 
from  the  Bahamas  and  delivered  to  Mr.  Burry."  Thus,  there  is 
no  compelling  evidence  that  any  of  these  4  shells  came  from  deep 
water,  or  even  from  the  vicinity  of  Hope  Town. 

The  shell  figured  by  Olsson  &  Moore,  identified  by  them  as 
5.  canaliculatus,  differs  significantly  from  the  holotype.  Oriented 
conventionally,  the  holotype  (Plate  12,  fig.  1)  has  a  high  spiral 
ridge  projecting  vertically  from  each  whorl  of  the  spire,  forming 
a  deep  channel  near  the  suture.  A  similar  ridge  projects  laterally 
on  the  shell  figured  by  Olsson  &  Moore  (Plate  12,  fig.  2)  ;  there 
is  no  deep  channel.  In  addition,  the  spires  of  the  2  shells  are  of 
different  heights. 

In  my  opinion,  the  name  5.  canaliculatus  has  been  applied  to 
malformed  shells  of  S.  (Tricornis)  gigas  Linn.  The  holotype  of 
S.  canaliculatus  (255  mm.  long)  and  the  shell  figured  by  Olsson 
&  Moore  (242  mm.  long)  are  both  the  normal  size  of  adult  and 
sub-adult  S.  gigas.  A  congenital  defect  of  a  small  part  of  the 
mantle  laying  down  the  shell  material  of  the  posterior  part  of 
the  outer  lip,  or  an  injury  at  an  early  age,  would  account  for  all 
the  morphological  differences  (including  the  suppression  of 
spines  and  axial  sculpture) ,  and  for  the  differences  in  the  form 
of  the  spiral  ridge  if  the  position  of  the  defective  part  of  the 
mantle  varies.  There  are  other  striking  malformations  of  S.  gigas 
(see  M.  Smith,  1940,  and  M.C.Z.  no.  168409;  Salisbury,  1953), 
and  similar  carinate  malformations  due  to  mantle  defects  occur 
in  many  other  gastropods    (Pelseneer,  1920;  for  more  examples. 


130  NAUTILUS  Vol.   75(4) 

see  Crosse,  1881;  Sykes,  1903;  Dautzenberg,  1911). 

A  malformed  shell  of  5.  (Strombus)  pugilis  alatus  Gmelin 
(Plate  13)  corresponds  in  part  to  S.  canaliculatus.  On  the  last  II/2 
whorls  a  vertical  spiral  ridge  forms  a  deep  channel  near  the 
suture.  Growth  was  normal  until  the  outer  lip  was  twice  broken. 

La  Fontaine  (1875)  and  Abbott  (1960,  p.  112)  discuss  2  more 
malformed,  carinate  Strombus,  one  (S.  vittatus  Linn.)  canali- 
culate. 

A  simple  experiment  might  prove  beyond  question  that  S. 
canaliculatus  is  a  malformed  shell.  The  appropriate  portion  of 
the  mantle  of  juvenile  S.  gigas  could  be  artificially  damaged  in 
order  to  observe  the  effects  on  the  subsequently  formed  outer 
lip  of  the  shell. 

In  1951,  2  years  after  describing  5.  canaliculatus  and  about 
11/2  years  before  his  death  (in  Dec,  1952),  Leo  A.  Burry  ad- 
mitted to  William  J.  Clench  and  Ruth  D.  Turner  that  he  then 
believed  he  had  named  a  malformed  shell.  The  additional  speci- 
mens were  in  Burry's  possession  at  that  time. 

Acknoioledgment:  Axel  A.  Olsson  graciously  pro\'ided  the  ex- 
cellent photograph  of  the  spire  of  S.  canaliculatus,  and  courte- 
ously allowed  me  to  read  the  manuscript  of  his  paper  co-authored 
with  Donald  R.  Moore. 

References 

Abbott,  R.  T.  1960.  Indo-Pacific  Mollusca,  1  (2) . 

Burry,  L.  A.  1949.  Shell  Notes  [publ.  Frank  Lyman,  Lantana, 
Florida],  2:106-109. 

Crosse,  H.  1881.  J.  Conchyl.,  2P;341-342,  pi.  11,  fig.  3. 

Dautzenberg,  Ph.  1911.  J.  Conchyl.,  55:209,  pi.  10,  figs.  12-13. 

La  Fontaine,  J.  de.  1875.  Ann.  (Mem.)  Soc.  Malac.  Belgique, 
;0:21,  pi.  2.  figs.  1-2. 

Lyman,  F.  1951.  Shell  Notes,  2:195-196. 

Olsson,  A.  A.  &  D.  R.  Moore,  1962.  Naut.,  75:127-128. 

Pelseneer,  P.  1920.  Acad.  roy.  Belgique;  classe  sciences;  Mem. 
(8vo),   (2)5:22,  45. 

Robertson,  R.  196L  Notulae  Naturae  (Acad.  Nat.  Sci.  Philadel- 
phia) ,  no.  343. 

Salisbury,  A.  E.  1953.  Proc.  Malac.  Soc.  London,  30A7AS,  pi.  8 

Smith,  M,  1940.  World-wide  sea  shells,  p.  131. 

Sykes,  E.  R.  1903.  Proc.  Malac.  Soc.  London,  5:260. 


April,  1962  nautilus  131 

VARIATION  AND  CHANGE  IN  SURFACE 
SCULPTURE  IN  ANOMIA  ACULEATA 

By  ARTHUR  S.  MERRILL^ 

Surface  sculpturing  on  the  shell  of  Anomia  acule.ata  Gmelin 
varies  considerably  (Verrill  and  Smith,  187S;  Jackson,  1890; 
Whiteaves,  1901;  and  Foster,  1937)  ranging  from  smooth  to 
highly  spinose.  In  fact,  lacking  intermediates,  the  variants  at  the 
extremes  could  be  easily  mistaken  for  separate  species.  One,  ac- 
cordingly, might  expect  to  find  grave  nomenclatorial  difficulties 
in  the  literature  but  this  is  not  the  case.  The  smoother  of  the  in- 
dividuals are  still  occasionally  listed  with  "form"  or  "varietal" 
names  but  it  is  generally  well  understood  that  they  are  part  of  a 
single  species  complex. 

With  ample  material  available,  this  study  was  undertaken  sim- 
ply to  describe  the  degree  and  complexity  of  sculptural  variation 
normally  occurring  in  this  species.  As  the  study  progressed  I 
foimd  that  spinosity,  if  it  is  expressed,  develops  at  widely  differ- 
ing shell  sizes.  In  the  usual  lamellibranch,  a  character  which 
develops  in  the  postlarval  form  becomes  apparent  at  about  the 
same  age  in  all  specimens.  But  in  A.  aculeata,  this  laboratory  has 
individuals  whose  variability  for  the  characteristic  spinosity  may 
be  expressed  any  time  after  metamorphosis. 

This  paper,  then,  describes  variability  in  A.  aculeata  and  pre- 
sents data  to  show  the  change  in  ratio  of  smooth  to  spiny  indi- 
viduals with  reference  to  shell  size. 

Methods  of  Collecting  and  Measuring.  Most  of  the  material 
used  in  this  study  came  from  navigation  buoys  stationed  in  off- 
shore waters  south  of  Nantucket,  Massachusetts.  Since  buoys  are 
brought  in  periodically  for  cleaning  and  servicing,  the  many 
forms  of  life  found  attached  can  be  removed  for  study.  Anomia 
often  homesteads  the  convenient  buoy  surfaces,  after  the  initial 
pelagic  larval  stage,  instead  of  settling  in  its  usual  habitat  on  the 
bottom.  Because  smaller  Anomia  were  predominant  on  the  buoys 
sampled,  the  bottom  was  dredged  to  obtain  representatives  of 
the  larger  adult  sizes. 

A.   aculeata  usually   maintains   a   suborbicular   shape   during 


^  U.  S.  Department  of  the  Interior,  Fish  and   Wildlife  Service,   Bureau  of 
Commercial    Fisheries    Biological    Laboratory,    Woods    Hole,    Massachusetts. 


132  NAUTILUS  Vol.  75(4) 

growth.  Those  which,  by  some  accident  of  position,  meet  an  ob- 
stacle in  the  immediate  vicinity  assume  various  distorted  shapes 
in  an  effort  to  circumvent  the  impediments.  This  apparently  does 
not  affect  the  morphology  gieatly  other  than  to  shift  the  axis  of 
growth.  The  shape  may  become  transversely  elongated,  or,  more 
commonly,  irregularly  rounded.  Thus,  to  better  indicate  true 
shell  size,  measurements  of  both  the  height  and  length  were 
taken,  added  together,  and  the  result  divided  by  2.  Over  4,000 
specimens,  from  newly  settled  larvae  to  mature  adults,  were 
examined. 

Variation  in  surface  structure.  A.  aculeata  derives  its  specific 
name  from  the  spinous  and/or  foliaceous  character  of  its  surface. 
While  most  individuals  exhibit  some  degree  of  surface  ornamen- 
tation, many  individuals  are  entirely  smooth.  Foster  (1937), 
reporting  on  specimens  brought  up  attached  to  shells  from  fairly 
deep  water  on  Georges  Bank,  described  considerable  variation  in 
the  spines,  but,  lacking  smooth  individuals,  he  did  not  describe 
this  condition.  Verrill  (Verrill  and  Smith,  1873)  mentioned  vari- 
ations in  A.  aculeata  from  the  Bay  of  Fundy  and  the  Casco  Bay 
areas  and  described  a  variety  with  the  scales  "more  or  less 
absorptive  or  even  entirely  absent."  Jackson  (1890)  and  Whit- 
eaves    (1901)    briefly  discussed  the  smooth  form. 

Plate  14  illustrates  the  variation  found  in  the  surface  sculpture 
of  this  species.  On  individuals  with  well-developed  spines,  the 
spines  may  radiate  to  the  ventral  margin  at  regular,  well-spaced 
intervals.  On  others,  they  may  be  irregularly  or  more  closely 
spaced.  The  spines  may  be  seated  on  raised  ridges  or  may  simply 
arise   from   the  surface. 

Concentric  lines  form  at  intervals  as  the  shell  grows.  The 
spines  terminate  at  the  line  formed  at  the  end  of  each  of  these 
growth  intervals,  and  other  spines  commence  to  form  with 
renewed  shell  growth.  The  degree  of  fluted,  frilled,  or  other 
transverse  sculpture  depends  on  the  degree  of  development  of 
concentric  processes.  For  example,  on  those  with  closely  spaced 
and  conspicuous  growth  lines,  the  spines  are  blunted.  Often  those 
with  weaker  growth  lines  tend  to  have  spines  which  project  from 
radiating  ridges.  The  spines  may  also  lack  radial  symmetry  and 
simply  form  along  concentric  lines  apparently  at  random  over  the 
shell  surface. 


April,  1962  nautilus  133 

On  those  individuals  with  smooth  surfaces,  the  concentric 
growth  lines  are  slightly  undidate  causing  the  surface  to  appear 
craggy  or  scaly.  Very  few  have  the  smooth  and  glittering  surface 
of  the  allied  species,  A.  simplex  Orbigny;  usually  the  surface 
appears  dull  and  chalky. 

Practically  every  conceivable  combination  occurs  between 
plain  smooth  and  fully  scaly  individuals.  Those  not  quite  smooth 
may  have  faint  ridges,  perhaps  just  a  ray  or  two  in  some;  others 
may  have  a  full  fan  of  strong  ridges  with  no  sign  of  spines.  On 
some  with  ridges,  there  may  be  a  raised  spine  here  and  there,  or 
possibly  one  of  the  ridges  will  be  fidly  spined  with  none  of  the 
other  ridges  showing  the  slightest  tendency  to  spininess.  Some 
will  be  smooth  to  a  certain  stage  in  growth,  then  suddenly 
begin  producing  nicely  developed  spines.  Others  may  do  this  and 
then  just  as  suddenly  become  smooth  again.  There  is  seemingly 
no  end  to  the  possible  combinations.  For  the  purpose  of  this 
paper,  the  individuals  have  been  separated  into  two  recognizable 
categories:  those  lacking  surface  ornamentation,  and  those  show- 
ing surface  sculpture  to  various  degiees.  The  latter  include 
individuals  whose  sculpture  ranges  from  faintly  rayed  to  heavily 
spinose. 

Size-frequency  data  by  station:  As  mentioned  earlier,  popula- 
tion samples  of  A.  aculeata  were  taken  from  3  offshore  buoys  as 
well  as  from  the  bottom  by  dredging.  Table  1  lists  the  frequency 
distribution  from  all  stations  as  well  as  the  percentage  of  smooth 
Anomia  in  each  size  category.  The  station  locations  and  other 
data  relating  to  them  are  listed  below. 

Nantucket  Shoals  Lightship  Buoy  (NSLS).  The  NSLS  buoy, 
located  1  mile,  55°  true,  north  of  the  permanent  Nantucket 
Shoals  Lightship  at  40°  33'  N.  latitude  and  69°  28'  W.  longitude, 
was  placed  on  station  October  8,  1957.  When  removed  May  10, 
1958,  it  had  been  in  the  water  for  7  months.  An  estimated 
25,000-30,000  specimens  of  A.  aculeata  were  attached  to  the  buoy. 
The  sample  used  for  this  study  consisted  of  2,423  specimens. 

Davis  Shoals  Buoy  (4DS#1).  The  station  occupied  by  the 
}DS#I  buoy  is  about  25  miles  west-northwest  of  the  NSLS  buoy 
at  40°  57'  N.  latitude  and  69°  55'  W.  longitude.  This  buoy  was 
on  station  1  year  from  May  7,  1957,  to  May  15,  1958.  The  sample 
taken  from  this  buoy  included  537  specimens  of  A.  aculeata. 


134 


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April,  1962  nautilus  135 

Davis  Shoals  Buoy  {4DS#2).  The  4DvS#2  buoy  succeeded  the 
4DS#1  buoy.  It  remained  on  station  for  6  months  from  May  15, 
1958,  until  November  10,  1958.  The  sample  included  983  speci- 
mens of  A.  aculeata. 

Channel  Station.  At  a  station  about  44  miles  northeast  of 
Nantucket  Shoals  Lightship  (41^03'  N.  latitude;  68°45'  W. 
longitude)  several  bottom  tows  were  made  during  the  course  of 
sea  scallop  investigations  on  September  16,  1959.  From  the 
dredged  material  246  A.  aculeata  were  picked  from  shells,  stones, 
and  debris. 

Discussion:  The  larvae  of  A.  aculeata  pass  through  a  pelagic 
stage  in  the  upper  waters  during  which  time  they  are  subject  to 
variable  currents  and  winds.  Hence,  they  are  apt  to  settle  hither 
or  yon  in  due  time  on  the  ocean  floor.  It  is  not  surprising,  then, 
to  find  samples  made  up  of  individuals  that  vary  from  one 
another  widely.  Verrill  (Verrill  and  Smith,  1873)  observing 
variation  in  this  species  thought  it  might  possibly  be  a  variety 
of  the  true  A.  ephippium  which  has  a  smooth  surface.  He  was 
sure  it  was  not  A.  glabra  [=  A.  simplex]  of  our  coast.  Whiteaves 
(1901)  remarked  of  the  smooth  form  of  eastern  Canada  as  fol- 
lows: "The  little  smooth  form  of  this  region  that  has  been 
hitherto  referred  to  A.  ephippium,  is  regarded  by  Verrill  as  most 
probably  a  form  of  A.  aculeata."  Jackson  (1890)  was  able  to 
separate  the  two  species  of  Anomia  in  our  area  by  means  of  prc^- 
dissoconchal  characteristics.  He  found  that  larval  A.  aculeata 
had  a  byssal  notch  both  in  the  left  and  right  valve,  while  larval 
A.  glabra  had  a  notch  only  in  the  right  valve.  By  this  means,  we 
all  are  sure  of  which  species  he  was  referring  when  he  stated, 
".  .  .  associated  specimens  are  frequently  wanting  in  the  spinous 
character,  although  unquestionably  belonging  to  the  same 
species."  Bousfield  (1958,  1960)  followed  W^hiteaves'  lead  (per- 
sonal communication)  in  listing  the  smooth  form  as  A.  ephip- 
pium. 

Two  authors  (Jackson,  1890;  Foster,  1937)  have  remarked  on 
the  fact  that  the  surface  of  A.  aculeata  may  be  smooth  for  a 
period  of  time  before  spines  begin  to  form.  Jackson  was  referring 
to  the  young  growing  for  a  brief  period  of  time  before  any  spines 
were  produced;  Foster  was  commenting  on  occasional  large  speci- 
mens on  which  the  spines  were  represented  only  on  the  outer 


136 


NAUTILUS 


Vol.  75(4) 


100 


80 


-   ^ 


60 


40 


20 


I    i    r 


BUOY 


4DS 

4DS*2  BUOY   

NSLS     BUOY 

CHANNEL  


\ 

■:    \ 
\      \ 
\        \ 

\         \ 


\ 


I \ L_J I \ I     I      I 


6  8  10  12 

S/Z£    IN    MILLIMETERS 


14 


Fig.  1.  Data  from  Table  1  plotted  to  show  decrease  in  percentage  of  smooth- 
surfaced  Anomia  aculeata  with  increase  in  shell  size. 

edge  oi  the  shell.  These  miscellaneous  facts  can  now  be  con- 
nected and  a  clear  picture  of  variation  presented  by  means  of 
the  data. 

A.  aculeata,  in  truth,  is  spineless  for  a  brief  period.  The  first 
signs  of  spines  appear  on  occasional  specimens  at  about  0.7  mm. 
By  this  time  the  shell  has  possibly  tripled  in  height  since  setting. 
As  can  be  seen  on  the  graph  in  Figure  1,  the  percentage  fre- 
quency of  smooth  individuals  decreases  rapidly  at  first  and  levels 
off  at  about  25  percent  above  8  mm.  In  large  spiny  individuals 


NAUTILUS  75(4) 


PLATE  13 


.\[all()niic'(l.  caiinatc  Sliuuihus  < Sty(niil>u.\j  piiolli.s  aliiliis  (.nielin.  jiucnile. 
Shell  ct)ated  with  MgO  to  show  sculptuiL-  bin  not  color  pattern.  Saiiibel 
Island  (?),  Florida,  .\merican  Miiseiiin  ol  Xaiinal  History  (no.  46068).  Fig.  1. 
\]>ical  \iew.  x'-^"-  I'ig-  -■  Vix'rtural  \iew.  Fig.  .S.  ()l)li(iiic  \iew.  Figs.  2-3 
iiainral  size. 


NAUTILUS  75  (4) 


PLATE   14 


A)iomui    nnilctilti    C.mcliii.    Rcpicsciilativc    individuals    arranged    in    show 
variation  in  shell  stidpturc. 


April,   1962  nautilus  137 

one  can  easily  trace  back  along  the  shell  and  see  when  spines 
were  first  laid  down.  Although  this  usually  is  found  to  be  within 
the  first  few  millimeters  of  giowth,  as  reflected  by  the  graph,  it 
may  be  found  on  particular  specimens  at  any  size.  Even  a  speci- 
men of  almost  maximum  size  may  finally  express  potential  to 
produce  spines. 

It  should  be  pointed  out,  in  regard  to  using  material  mostly 
from  buoys  for  this  study,  that  the  environment  on  the  buoy  is 
probably  more  uniform  than  on  the  bottom.  However,  the  small 
amount  of  data  from  the  bottom  collections  fits  in  well  with  the 
data  from  buoy  samples.  This  tends  to  confirm  the  possibility 
that  variation  in  surface  sculpture  in  A.  aculeata  might  be  a 
genetic  rather  than  an  environmentally  induced  phenomenon. 
Conclusion  and  Summary 

1.  Surface  sculpture  in  the  shell  of  Ajiomia  aculeata  varies 
considerably  depending,  in  part,  on  size  (or  age)  .  The  sinface 
may  be  smooth  or,  to  a  highly  variable  degree,  spinose.  A  long 
series  of  similarly-sized  specimens  from  single  buoy  samples  can 
be  formed  in  which  graded  variability  in  surface  structure  is 
shown  easily.  Therefore,  in  keeping  with  the  usual  procedures  in 
taxonomy,  no  distinction  shoidd  be  shown  the  extreme  variants. 

2.  All  early  post-larval  A.  aculeata  have  smooth  siufaces  and, 
although  a  percentage  may  remain  smooth  throughout  life,  many 
start  early  to  put  on  spines,  the  percentage  of  spiny  individuals 
increasing  with  shell  size. 

3.  The  fact  that  smoothness  is  always  expressed  in  the  early 
dissoconch  shell  implies  that  spinosity  is  a  natural  later  develop- 
ment. The  early  smooth  shell  undoubtedly  represents  the  gen- 
eralized or  ancestral  form.  The  generalized  form  found  along  the 
western  Atlantic  coast  is  A.  simplex,  or  along  the  European 
coast,  A.  ephippium  (type  species  of  Anomia) .  Spinosity,  a 
divergence  from  the  generalized,  may  be  considered  a  derivative. 

Literature  cited 
Bousfield,  E.  L.  1958.  Proc.  Nova  Scotian  Inst.  Sci.  24  (3)  :  303-325. 
1960.   Canadian  Atlantic  sea  shells.   National   Museum   of 

Canada,  Ottawa,  72  pp. 
Foster,  Richard  W.  1937.  Naut.  50(3)  :  102-103. 
Jackson,  Robert  Tracy.  1890.  Phylogeny  of  the  pelecypoda,  the 

Aviculidae  and  their  allies.  Alem.  of  the  Boston  Soc.  of  Nat. 

Hist.  4  (8)  :277-400. 


138  NAUTILUS  Vol.  75(4) 

Whiteaves,  J.  F.  1901.  Catalogue  of  the  marine  invertebrates  of 
Eastern  Canada.  Geological  Survey  of  Canada,  Ottawa, 
271  pp. 

Verrill,  A.  E.  and  S.  I.  Smith.  1873.  Report  upon  the  invertebrate 
animals  of  Vineyard  Sound  and  adjacent  waters,  with  an 
account  of  the  physical  features  of  the  region.  Report  of 
the  U.  S.  Fish  Commission  for  1871-72,  448  pp. 

ENHANCEMENT  OF  LATENT  SHELL  FLUORESCENCE 
BY  GLYCERINE 

By  RICHARD  G.  WOODBRIDGE,  III 
c/o  Transspace  Laboratory,  Box  111,  Princeton  Junction,  New  Jersey 

At  the  close  of  a  recent  period  of  collecting  and  experimenting, 
an  interesting  observation  was  made  pertaining  to  the  fluores- 
cence of  shells  (1)  which  is  deemed  worthy  of  preliminary 
publication  at  this  time. 

All  specimens  were  obtained  at  and  below  (using  scuba)  the 
"low  tide  line",  Northwest  Harbor,  Deer  Isle,  Maine,  August 
28-29,  1961.  The  ultraviolet  light  used  was  one  originally  de- 
veloped for  underwater  work  and  littoral  exploration  (2)  emis- 
sion being  from  a  BLB  lamp  (approximately  .8500  to  4000 
angstroms) . 

While  attempting  to  ascertain  the  suitability  of  glycerine  as  a 
preservative  for  the  color  and  normal  fluorescence  of  "coralline 
algae"  (3) ,  a  small  "limpet"  adhering  to  one  specimen  was  inad- 
vertently coated  with  the  glycerine,  whereupon  (in  the  dark) 
under  the  ultraviolet  light  the  limpet  shell  burst  into  a  strong, 
glowing,  deep  red  fluorescence. 

Fifteen  living  specimens  of  Acmaen  testudinalis  testudinalis 
(Miiller)  (limpets)  were  subsequently  either  coated  with  or 
immersed  in  glycerine.  In  every  case  a  deep  red  fluorescence 
resulted  over  the  entire  shell  while  under  the  glycerine.  Prior  to 
treatment  these  specimens  had  no  apparent  visible  red  fluores- 
cence. 

Unfortunately  time  did  not  permit  an  examination  of  the 
effect  of  glycerine  on  the  fluorescent  areas  (feeding?,  homing?) 
surrounding  such  limpets  when  they  are  found  living  on  "bare" 
rocks  —  an  observation  made  by  the  writer  some  years  ago  (un- 
published) . 

Twenty-seven  living  specimens  of  Littorina  littorea    (Linn^) 


April,  1962  nautilus  139 

(the  common  periwinkle)  were  treated  with  glycerine.  In  all 
cases,  a  similar  deep  red  shell  fluorescence  of  considerable  inten- 
sity resulted,  particularly  strong  in  small  specimens.  Location  of 
the  area  of  maximum  shell  fluorescence  was  somewhat  variable, 
appearing  to  have  a  tendency  to  localize  in  areas  in  the  proximity 
of  the  apex.  Fifteen  specimens  kept  immersed  in  glycerine  ap- 
pear to  have  retained  their  red  fluorescence  unchanged  for  two 
months. 

In  the  case  of  the  periwinkles,  the  writer  has  felt  for  some 
years  that  occasionally  some  sort  of  dark  red  shell  fluorescence 
could  be  observed  but  it  was  so  illusive  and  ephemeral  that  such 
observations  could  be  considered  at  best  as  highly  questionable; 
however,  it  has  led  the  writer  to  refer  to  the  phenomena  as 
"enhancement"  of  fluorescence,  rather  than,  say,  "inducement." 

Vigorous  scraping  with  a  knife  blade  of  both  limpet  and  peri- 
winkle shells  subjected  to  the  above  glycerine  treatment  re- 
sulted in  no  apparent  diminution  of  shell  fluorescence  until  the 
shells  were  deeply  abraded  —  suggesting  that  the  phenomena  is 
not  due  to  a  superficial  shell  coating. 

Three  small,  living  specimens  of  Bucciniim  undatum  Linn^ 
(the  common  welk)  coated  or  immersed  in  glycerine  showed  no 
such  shell  fluorescence. 

A  small  specimen  of  Clanculus  puniceus  from  the  writer's  col- 
lection of  fluorescent  shells  showed  no  particular  enhancement 
of  its  normal  brilliant  rose  fluorescence. 

References 
Woodbridge,  R.  G.  Fluorescent  Shells,  a  Monograph,  etc.  Pub- 
lished by  Transspace  Laboratory,   1961. 
Ibid.  Underwater  Ultraviolet  Lights,  etc.  Published  by  Trans- 
space  Laboratory,  1960. 
Woodbridge,  R.  G.  and  Woodbridge,  R.  C.,  The  Application  of 
Ultra-Violet  Lights  to  Underwater  Research,  Nature  184:2b^ 
(July  25,  1959) . 

REPRODUCTION  OF  OLIVELLA  MUTICA^ 

By  ROBERT  T.  PAINE 

Scripps  Institution  of  Oceanography 

La  Jolla,  California 

Olivella   mutica    (Say)    occurs   commonly   in   warm,    shallow 


^Contribution    no.    171    from    the   Oceanographic   Institute,    Florida   State 
University,  Tallahassee,  Florida. 


140  NAUTILUS  Vol.   75(4) 

waters  from  North  Carolina  to  Texas,  and  in  the  West  Indies 
(Abbott,  1954) .  However,  little  appears  to  be  known  about  its 
general  biology.  The  following  observations  are  primarily  con- 
cerned with  the  external  form  of  the  egg  capsvde,  length  of  the 
developmental  period,  and  duration  of  the  spawning  season. 
Some  data  are  also  presented  on  the  nature  of  the  occupied 
habitat  and  population  density.  This  study  was  made  possible 
through  use  of  the  facilities  of  the  Alligator  Harbor  Marine 
Laboratory  of  Florida  State  University,  in  Franklin  County, 
Florida.  Menzel  (1956)  has  cited  O.  mutica  as  a  common  snail 
in  the  immediate  region.  Olivellas  were  collected  from  the  south 
end  of  a  sandbar  lying  across  the  mouth  of  Alligator  Harbor. 
Extensive  portions  of  this  bar  become  exposed  at  most  low  tides. 

Thirty- two  snail  egg  capsules  of  similar  appearance  but  of 
unknown  parentage  were  collected,  primarily  from  the  valves  of 
a  brachiopod  which  occurs  commonly  on  the  bar.  All  but  7 
of  these  embryos  were  reared  successfully  in  filtered  sea  water, 
and  many  of  them  were  then  maintained  until  they  were  suf- 
ficiently large  to  be  identified.  Most  were  identified  from  Abbott 
(1954)  and  Olsson's  monograph  (1956)  as  O.  mutica.  However, 
one  O.  pusilla  (Marrat)  which  is  also  present  at  Alligator  Har- 
bor, was  subsequently  discovered  included  with  the  specimens  of 
O.  mutica.  Because,  unfortunately,  laboratory-raised  and  field- 
collected  specimens  were  mixed,  I  do  not  know  whether  all  the 
eggs  were  O.  mutica,  or  whether  a  few  might  have  been  O.  pu- 
silla. This  confusion  of  two  very  similar  species  will  not  affect 
those  conclusions  based  on  egg  capsules  in  which  O.  mutica  is 
known  to  have  predominated.  However,  no  distinction  was 
drawn  between  these  two  olivellas  in  the  field  and  thus  the 
density  and  habitat  information  includes  both  species. 

Adult  snails  were  noted  from  September,  1959,  to  August, 
I960,  indicating  year-round  occupancy  of  the  intertidal  portions 
of  the  sandbar's  southern  end.  The  bottom  sediment  here  is 
predominately  a  mixture  of  fine-grained  quartz  sand  M'ith  an 
average  diameter  of  0.125  to  0.250  mm.,  and  a  few  shells  or  shell 
fragments.  The  silt  and  clay  fractions  of  the  sediment  are  mini- 
mal, and  little  decaying  organic  matter  has  accumulated.  The 
marine  grass  Diplanthera  wrightii  (Ascherson) ,  occurring  in 
patches,  has  consolidated  portions  of  the  bottom  sediment  but 
has  not  changed  the  physical  characteristics  of  the  substratum 


April,  1962  nautilus  141 

appreciably.  Olivellas  were  collected  from  both  consolidated  and 
unconsolidated  sandy  sediment,  with  the  maximum  density  of 
178  snails  per  square  meter  in  the  latter.  Snails  of  this  genus 
were  not  found  in  samples  from  soft  bottom  areas  overlain  by 
quantities  of  organic  material. 


670 


A 


1050/t 

Fig.   1.  Usual  egg  capsule  ot  Olivella  mutica. 

The  encapsulate  eggs  of  O.  mutica  (Figure  1)  are  laid  separ- 
ately, one  ovum  per  capsule,  with  no  trace  of  nurse  eggs,  and  are 
attached  to  any  solid  object  that  protrudes  above  the  surface  of 
the  sand.  The  geneial  cliaracteristics  of  the  egg  and  the  site  of 
oviposition  are  thus  similar  to  those  described  by  Marcus  and 
Marcus  (1959)  for  O.  verrcauxii  (Ducros) .  The  base  of  the  egg 
capsule  of  O.  mutica  is  roughly  circular  and  1.0  to  1.1  mm.  in 
diameter.  From  this  base  rises  a  smooth,  spherical,  transparent 
dome  about  670  microns  in  diameter.  Near  the  top  a  distinct 
ridge  indicates  the  line  along  which  the  capsule  lid  separates 
when  the  young  snail  escapes.  In  general  form  the  capsules  ap- 
pear indistinguishable  fiom  those  figured  by  Perry  and  Schwen- 
gel  (1955)  for  O.  pusilla.  In  contrast,  the  egg  capsule  of  O.  ver- 
reauxii  (Marcus  and  Marcus,  1959),  though  entirely  similar  in 
gross  shape,  can  readily  be  distinguished  by  the  surface  sculptur- 
ings  —  fine,  raised,  parallel  lines. 

The  majority  of  the  eggs  of  O.  mutica  were  found  attached  to 
the  anterior  end  of  the  valves  of  the  inarticulate  brachiopod, 
Glottidia  pyramidata  (Stimpson) ,  with  from  1  to  5  capsules  per 
brachiopod.  Freshly-deposited  eggs  or  developing  embryos  were 
obtained  on  25  March  and  8  April,  1960,  when  the  water  temper- 
ature ranged  from  16°  to  20°  C.  The  maximum  developmental 


142  NAUTILUS  Vol.  75(4) 

period,  based  on  laboratory-reared  eggs,  was  19  or  20  days  for 
both  collections.  The  cleavage  and  veliger  stages  are  spent  in  the 
capsule,  and  upon  hatching  the  capsule  lid  is  broken  off  and  a 
young  snail,  750  microns  in  length  crawls  away.  On  the  other 
hand,  Marcus  and  Marcus  (1959)  have  shown  that  O.  verreauxii 
hatches  as  a  veliger  after  8-9  days. 

Large  samples  of  the  brachiopod  were  also  closely  scrutinized 
on  26  February,  10  and  12  March,  22  and  23  April,  and  3,  8,  25, 
26,  and  28  May;  they  were  devoid  of  Olivella  egg  cases.  The  first 
egg  to  hatch  in  the  laboratory  did  so  on  6  April.  If  one  extrapo- 
lates back  20  days,  the  suggested  period  of  embryonic  develop- 
ment, the  initial  spawning  took  place  about  17  March.  Egg- 
laying  appears  to  have  ceased  sometime  between  8  and  22  April. 
These  data  suggest  that  O.  miitica  spawns  for  approximately  one 
month  during  the  spring,  and  that  egg-laying  is  closely  limited 
to  this  period.  No  information  is  available  on  spawning  in  the 
fall,  or  latitudinal  variations  in  the  duration  of  egg-laying. 

Literature  cited 
Abbott,  R.  T.  1954.  American  Seashells.  Princeton:  D.  vanNos- 

trand  Co.  541  pp. 
Marcus,  E.  and  E.  Marcus,  1959.  On  the  reproduction  of  Olivella. 

Univ,  Sao  Paulo,  Fac.  Filos.  Cienc.  e  Letras,  Bol.  Zool.  22: 

189-199. 
Menzel,  R.  W.  1956.  Annotated  check-list  of  the  marine  fauna 

and  flora  of  the  St.  George's  Sound  -  Apalachee  Bay  Region, 

Florida  Gulf  Coast.  Fla.  State  Univ.  Ocean.  Inst.,  Contrib. 

61. 
Olsson,  A.  A.  1956.  Studies  on  the  genus  Olivella.  Proc.  Acad. 

Nat.  Sci.,  Philad.  108:155-225. 
Perry,  L.  M.  and  J.  S.  Schwengel,   1955.  Marine  shells  of  the 

western  coast  of  Florida.  Ithaca:  Paleon.  Res.  Inst.  318  pp. 


PUERTO  RICAN  OLEACINOIDS 

By  H.  BURRINGTON  BAKER 

The  symbols  used  for  Puerto  Rican  localities  were  explained 
recently,  1961  (1) . 

Yunquea  denselirata  H.  B.  Baker.  Only  known  from  type 
locality  (Er3)  ;  on  last  whorl  of  type  (only  dry)  shell,  "very  fine" 
growtli  threads  average  1 1  microns  from  crest  to  crest,  which 
means  91  per  mm.,  and  that  the  "very  slightly  stronger"  ones  run 
13  to  15  per  mm.  Y.  monteplatonis  (Pilsbry)   has  similar  threads. 

Odontosagda  sp.  "Yunquea  denselirata"  van  der  Schalie,  1948: 


April,   1962  nautilus  143 

71,  pi.  6,  fig.  5.  Probably  an  unnamed  species,  from  about  3  km. 
south  of  Aguas  Biienas. 

The  anatomy  of  Odontosagda  is  unknown.  Although  its  shell 
has  internal  lamellae  (like  some  Sagdinae) ,  in  general  form  and 
size,  it  does  resemble  more  closely  that  in  Yunquea  (Yunqueinae). 

Hyalosagda  (Microsagda)  subaquila  (Shuttleworth)  .  Among 
dead  leaves  in  disturbed  places.  En,  Jn,  Pn,  Prl  (2500  ft.) ,  Wn, 
Wr2,  Ws,  Ww2;  mainly  lowlands.  Foot  light  near  sole  but  almost 
black  above,  with  lighter  middorsal  stripe  and  triangle  between 
ommatophores,  which  are  very  dark;  eyes  light  with  dark  centers. 

H.  (M.)  subaquila,  var.  a.  Psl-3;  shell  more  depressed,  with 
larger  umbilicus    (about  I/5  major  diameter)  . 

H.  (Lacteoluna)  selenina  (Gould) .  Probably  distributed  by 
commerce;  Es2,  Jn,  Pn,  Pr2-6,  Wn,  Wr,  Ws,  0-3400  ft.,  but  not 
found  in  Luquillo  forests. 

H.  (Aerotroc/ius?)  krugiana  (Martens) .  Size  (1877:346)  :  3.5  by 
171  (6  mm.)  with  5  whorls,  subangulate,  narrowly  umbilicate. 
Caguana,  west  of  Utuado. 

Martens  compared  the  last  species  to  the  Cuban  Helix  turbini- 
formis  Pfeifier,  which  at  the  time  was  confused  with  type  species 
of  Aerotrochiis,  the  Jamaican  Hyalosagda  subpyramidalis  (C.  B. 
Adams)  ;  the  last  is  a  higher,  but  angulate  shell,  which  lives  on 
trunks  of  trees.  These  3  Puerto  Rican  species  belong  in  the  sub- 
genus (or  genus?)  Lacteoluna;  Hyalosagda  (s.  s.)  and  Stauro- 
glypta  appear  to  be  mainly  Jamaican. 

Aquebana  (s.  s.)  velutina  (Lamarck) .  Living  animals  very  rare, 
under  rocks,  5-6  inches  below  surface  (Jnl)  and  under  dead 
leaves  between  limestone  ledges  (Enl)  during  and  at  end  of 
drouth,  but  dead  shells  numerous  (Jn2)  which  may  indicate 
that  it  has  periods  of  abundance.  Fresh  shells  with  greenish  tinge. 
To  1940c: 59,  is  added:  foot  pale,  with  chrome  spots,  obscured 
by  chalky  ones. 

Platysuccinea  portoricensis  (Shuttleworth) .  Terrestrial,  under 
upper  layer  of  dead  leaves,  especially  in  ruderal  places;  Er,  Pr, 
Wr,  1500-4000  ft.  Fresh  shells,  with  adherent  dirt  except  in 
columellar  region,  greenish  corneous  on  El  Yunque  and  often 
with  very  low  spires,  but  often  more  hyaline  with  higher  spires 
in  Cordillera  Central.  Animal  (added  to  1940c:61)  rather  slug- 
gish; side  of  foot  very  warty,  ochraccous  to  brown,  darker  in 
wrinkles,  dorsad  and  on  ommatophores;  mantle  collar  not  over- 
lapping peristome  but  with  umbilical  lobe  over  columellar  area; 
sole  dark  slate,  only  trizonal,  with  numerous  rapid  waves  in 
middle  zone. 

P.  portoricensis,  var.  a.  Under  rocks,  Enl;  also  empty  shell 
from  Pnl ;  less  than  200  ft.  Shells  much  smaller,  more  hyaline  and 


144  NAUTILUS  Vol.  75  (4) 

with  higher  spires,  but  not  distinguishable  from  young  ones  from 
Cordillera  Central;  also  quite  similar  in  form  to  "Simpulopsis" 
aenea  Pfeiffer,  1861,  as  figured  by  Fischer  &  Crosse,  1877,  Miss.. 
Mex.  Moll.  7;pl.  24,  figs.  12;  "S."  cumingi  Pfr.,  P.  Z.  S.,  1861:pl.  3, 
fig.  2,  from  Mexico,  looks  like  typical  P.  portoricensis  (some  El 
Yunque  shells  even  larger)  ;  and  ruderal  snails  do  get  around. 

Varicella  (Vagavarix)  portoricensis  (Pfeiffer) .  Terrestrial;  En, 
Es,  Jn,  Pn,  Pr3,6,  Wn,  Wr2,3,  Ws,  0-3000  ft.,  but  not  found  in 
Luquillo  Mts. 

V.  (Vagavarix)  calderoni  H.  B.  Baker.  On  ground;  Pr6,  Wr2, 
2000-3000  ft.,  Cordillera  Central. 

V.  (Vagavarix)  sulculosa  (Shuttleworth) .  Terrestrial;  En,  Es, 
Jn,  Pn,  Pr  1,5,6,  Ps2,3,  Wn,  Wr,  Ww,  0-4000  ft.,  but  not  found  in 
Luquillo  Mts.  Larger  lots  intergrade  completely  with  smoother 
form  terebraeformis  (Sh.) ,  which  has  more  widely  and  irregu- 
larly spaced  growth  striae;  mostly  size  of  sulculosa  but  one  (Ps2) 
12.1  mm.  long  with  8.2  whorls. 

V.  (Vagavarix)  sporadica  H.  B.  Baker.  On  ground;  Es3,4,  Er2, 
200-2500  ft.,  only  found  in  eastern  Puerto  Rico. 

Oleacina  (Laevaricella)  interrupta  (Shuttleworth) .  Usually 
subarboreal,  but  also  seen  alive  under  leaves  at  base  of  trees; 
Er2,  Pr2,3,  Wr2,3,  2000-3400  ft.,  ranging  from  Luquillo  Mts.  to 
western  Cordillera  Central.  Animal  (1941b:29)  seen  attacking 
Nenia;  lateral  zones  of  foot  narrow. 

Oleacina  (Boriquena)  glabra  (Pfeiffer,  1846) .  Terrestrial,  Er2- 
5,  above  2000  ft.  in  Luquillo  Mts. 

O.  (B.)  playa  (H.  B.  Baker) .  On  ground;  En,  Es3,  Jn,  Pn,  Wn, 
Ws,  lowlands,  0-800  ft.  This  "large,  elongate,  smooth  species  com- 
mon to  Puerto  Rico"  first  noticed  and  called  "gracilior"  by  Shut- 
tleworth, 1854:51;  all  I  did  (1940a)  was  to  prove  it  a  distinct 
species  and  name  his  "var.  beta"  in  Varicella;  if  his  descriptive 
term  be  considered  a  trinomial,  it  would  be  preoccupied  in 
neither  Gla7idina  nor  Oleacina. 

Although  the  animal  of  Oleacina  s.  s.  is  still  unknown,  Bori- 
quena (possibly  only  a  section)  and  Laevaricella  (a  very  distinct 
subgenus)  now  are  made  congeneric  with  it,  because  of  the 
close  resemblance  in  the  embryonic  shells  of  Oleacina,  Boriquena 
and  Varicella  s.s.,  which  was  pointed  out  by  Pilsbry,  1907  (12a): 
127,  and  because  the  differences  in  shell  form  between  O.  voluta 
(Gmelin)  and  O.  (voluta  var.?)  flexuosa  (Pfr.)  from  Haiti  so 
closely  parallel  those  between  the  Puerto  Rican  O.  glabra  and 
O.  playa.  In  1956:133,  135,  after  vacillating  in  I941d  and  1943c, 
the  almost  strictly  Antillean  Oleacinidae  (-f  Varicellarum) ,  with 
1  Cuban  species  in  Florida,  were  separated  from  the  Spiraxidae 
(including  Streptosylinae  and  Euglandininae) ,  which  are  most 


April,  1962  nautilus  145 

differentiated  on  the  mainland,  but  have  invaded  Jamaica  (Spir- 
axinae)  and  other  Antilles  [Streptostylinae:  Salasiella  (Laevolen- 
cina  &  Flavoleacina),  Streptostyla  (Rectoleacina),  etc.]. 

These  two  families  have  almost  nothing  in  common  except 
their  carnivorous  habits  and  the  correlated  convergences  in  the 
radulae  of  their  most  specialized  groups.  Even  in  their  radulae, 
the  Spiraxidae  almost  intergrade  with  the  Achatinidae  (sens. 
lat.) ,  which  also  have  little  epiphallic  differentiation  and  some- 
times (Ferussaciinae)  have  similar  "heterurethrous'  "  pallial  com- 
plexes. The  changes  in  the  radulae  within  Varicella  (especially 
Melaniella)  show  that  the  convergences  (uniformly  aculeate 
teeth)  have  come  from  different  sources,  and  the  Oleacinidae 
have  sigmurethrous  pallial  complexes  and  truly  penial  epiphalli, 
often  with  well  developed  verges,  much  more  like  those  in  the 
Sagdidae.  The  unroofing  of  the  adrectal  limb  of  the  ureter,  which 
is  paralleled  in  both  the  Oleacinidae  and  the  bigger  Spiraxidae 
(and  in  many  other  groups  of  geophiles)  is  found  also  in 
Yunquea,  which  has  the  most  generalized  radula  in  the  oleaci- 
noids.  Incidentally  as  far  as  known,  outside  the  small  species  of 
Hojeda,  and  the  riideral  Hyalosagda  selenina  and  (possibly) 
Platysuccinea,  the  Sagdidae  also  seem  to  be  restricted  to  the 
Antilles. 

Additional  references    (see   1962) 
1961  (1),  Naut.  /•/ .-142-149.  1962,  Naut.  75:116-122. 

RADULAE  OF  NORTH  AMERICAN  ANCYLID  SNAILS 
II.  SUBFAMILY  NEOPLANORBINAEi 

By  PAUL  F.  BASCH 
Department  of  Biology,  Kansas  State  Teachers  College,  Emporia 

The  subfamily  Neoplanorbinae  was  established  by  Hannibal 
(1912)  to  accommodate  5  species  in  two  unusual  genera  of  fresh- 
water mollusks.  These  species  demonstrate  an  extreme  in  endem- 
ism,  all  known  specimens  having  been  collected  from  the  lower 
Coosa  River  bordering  Coosa,  Chilton,  and  Elmore  counties  in 
central  Alabama.  As  far  as  I  am  aware,  only  t^vo  men,  H.  H. 
Smith  and  A.  A.  Hinkley,  have  ever  seen  these  animals  alive,  and 
it  is  unlikely  that  living  specimens  will  be  seen  by  anyone  else. 

The  first  mention  of  these  snails  was  by  Hinkley  in  1904,  who 
listed   both   "Planorbis   tnntillus   Pilsbry"    and   an    undescribed 


'  Supported  by  grant  G-14125  from  the  National  Science  Foundation. 


146  NAUTILUS  Vol.  75  (4) 

form,  "N.  g.,  n.  sp."  among  a  group  of  Alabama  shells  collected 
the  previous  year.  Pilsbry  (1906)  recognized  these  shells  as 
".  .  .  quite  unlike  any  fresh-water  snails  hitherto  known  in  this 
country,  having  affinities  with  the  Ancylidae."  He  established 
two  new  genera  for  them,  describing  Amphigyra  alabamensis  and 
Neoplanorbis  tantillus  as  new  species.  Two  years  later.  Walker 
(1908)  named  3  additional  species  of  Neoplanorbis:  N.  cari- 
natus,  N.  umbilicatus,  and  N.  smithii,  and  these  have  remained, 
without  change  or  addition,  to  the  present. 

In  1914  Lay  Dam,  the  first  of  a  series  of  major  hydroelectric 
power  installations,  was  completed  across  the  Coosa  River  be- 
tween Coosa  and  Chilton  counties,  13  miles  east  of  Clanton,  Ala- 
bama. This  dam  imponds  an  area  of  6,000  acres,  with  a  back- 
water extending  to  a  point  13  miles  below  Childersburg  (Pierce, 
1955) .  Sixteen  miles  downstream  Mitchell  Dam  was  completed 
in  1923,  with  5,800  acres  of  backwater  extending  to  Lay  Dam. 
In  1929,  Jordan  Dam  was  built  8  miles  north  of  Wetumpka, 
Elmore  County.  This  dam  supports  a  lake  of  4,900  acres,  extend- 
ing about  18  miles  upstream  to  Mitchell  Dam.  The  large  lakes 
formed  by  these  dams,  which  provided  power  and  recreational 
facilities  for  the  people  of  central  Alabama,  also  destroyed  the 
habitats  (and  presumably  caused  the  extinction)  of  all  known 
species  of  Neoplanorbinae.  These  tiny  mollusks  lived  on  stones  in 
the  swift  current  of  the  Coosa  (See  Goodrich,  1944) ,  in  localities 
now  covered  by  many  feet  of  impounded  lake  water.  An  attempt 
in  June,  1959,  by  the  author  and  Dr.  John  B.  Burch  to  find  living 
specimens  in  the  Coosa  was  not  successful   (Basch,  1959) . 

Through  the  kindness  of  Dr.  Henry  van  der  Schalie  of  the  Uni- 
versity of  Michigan  Museum  of  Zoology  I  have  been  given  access 
to  the  specimens  of  Neoplanorbinae  in  his  care,  with  permission 
to  extract  and  prepare  radulae  from  a  sample  of  the  specimens. 
The  radulae  were  extracted  and  mounted  using  techniques  out- 
lined in  a  previous  paper  (Basch,  1961) .  A  study  of  these  radulae 
was  undertaken  in  an  attempt  to  learn  more  about  the  systematic 
position  of  the  Neoplanorbinae  by  comparison  of  their  radulae 
with  other  groups  of  Bosommatophora. 

In  Neoplanorbis,  the  shell  is  planorboid  and  extremely  sim- 
ilar in  conformation  and  dimensions  with  shells  of  Micro- 
menetus    of    the    family    Planorbidae.    Since    Menetus    (Micro- 


April,  1962  nautilus  147 

menetus)  alabamensis  is  known  from  the  state,  1  considered  that 
possibly  Neoplanorbis  represented  a  special  group  of  Menetus- 
like  planorbid  snails.  Ecologically,  Menetus  is  an  inhabitant  of 
quiet,  plant-filled  pools  in  a  situation  vastly  different  from  the 
rocky  rapids  of  the  old  Coosa  River.  Shells  and  radulae  of 
Micromenetus  are  illustrated  by  F.  C.  Baker   (1945). 

The  following  radulae  have  been  prepared  (UMMZ  numbers): 

#102672 — A^  tantillus,  2  specimens. 
#102664 — N.  carinatus,  2  specimens. 
#102671 — ^A^.  smithii,  2  specimens. 

The  species  studied  were  all  similar  in  radular  characters.  Neo- 
planorbis tantillus,  carinatus,  and  umbilicatus  are  obviously 
closely  related  on  the  basis  of  shell  similarities  and  may  in  fact 
represent  a  single  variable  species.  The  radulae  of  A^.  tantillus 
and  N.  carinatus  are  indistinguishable  (it  was  not  possible  to 
obtain  a  suitable  specimen  of  A'^.  umbilicatus).  Neoplanorbis 
smithii,  whose  shell  is  easily  differentiated  from  the  other  three 
by  its  lack  of  spiral  sculpture,  has  a  radula  differing  in  minor 
characters,  particularly  the  angle  formed  by  the  medial  surface 
of  the  lateral  teeth  with  the  longitudinal  axis  of  the  ribbon.  In 
all  three  species  of  Neoplanorbis  the  radular  formula  is  17-1-17, 
with  the  inner  cusp  of  the  laterals  long  and  dagger-like,  bearing 
a  small  accessory  cusp  on  its  medial  surface.  The  marginal  teeth 
are  characterized  by  an  increasingly  rectilinear  upper  edge  and  a 
prominent  needle-like  cusp  located  just  medial  to  the  center  of 
the  tooth  (Page  148,  Figs.  1  and  2) .  The  extreme  smallness  of 
these  teeth  must  be  pointed  out,  for  some  of  the  finer  features  lie 
beyond  the  resolving  power  of  a  light  microscope.  The  optical 
equipment  used  in  this  study  consisted  of  a  Leitz  triocular 
Labolux  microscope  with  a  lOOX  N.  A.  1.30  oil  immersion  ob- 
jective and  paired  20X  oculars.  An  internal  prism  factor  of  1.25X 
provided  a  final  magnification  of  2500X.  Even  at  this  extreme 
magnification,  some  details  were  not  readily  distinguishable  in 
the  bacteria-sized  teeth. 

The  radula  of  Amphigyra  is  strongly  bicuspid  (cf.  Pilsbry, 
1906)  and  similar  to  that  of  Neoplanorbis.  The  formula  is 
18-1-18,  and  lateral  teeth  7  to  11  resemble  planorbid  and  some 
ancylid  laterals  rather  closely  (Page  148,  Fig.  3),  perhaps  estab- 
lishing a  point  of  homology.  Two  radulae  were  prepared  from 


148 


NAUTILUS 


Radulae  of  Neoplanorbinae.  Fig.  1,  N.  carinatus  Walker;  Fig.  2,  A^.  smithii 
Walker;  Fig.  3,  Amphigyra  alabamensis  Pilsbry. 

specimens  in  UMMZ  lot  #\Q2Q11. 

Judging  by  the  radula,  apparently  Amphigyra  and  Neoplan- 
orbis  are  closely  related  to  each  other,  but  they  have  diverged 
considerably    from    other    groups    of    Basommatophora.    With 


April,  1962  nautilus  149 

a  dearth  oi  anatomical  data  concerning  the  Neoplanorbinae,  its 

position  within  the  Ancylidac  is  not  firmly  supported  by  radular 

structure  alone,  but  no  other  evident  relationships  are  revealed. 
Literature  cited 

Baker,  Frank  C.  1945.  The  Molluscan  Family  Planorbidae. 
Urbana.  Univ.  of  Illinois  Press.  530  pp. 

Basch,  Paul  F.  1959.  The  Coosa  Revisited.  Amer.  Malac.  Union 
Annual  Rept.  for  1959:    17. 

1961.  Radulae  of  North  American  Ancylid  Snails.  I  Sub- 
family Rhodacmeinae.  Naut.  75:97-101. 

Goodrich,  Calvin.  1944.  Naut.  5.9  (1)  :11-15. 

Hanibal,  Harold.  1912.  Proc.  Maloc.  Soc.  London.  70:1 12-211. 

Hinkley,  A.  A.  1904.  Naut  18(5)  :54-57. 

Pierce,  Laurence  B.  1955.  Hydrology  and  surface-water  resources 
of  east-central  Alabama.  Geol.  Surv.  of  Alabama,  Special 
Rept.  22:1-318. 

Pilsbry,  Henry  A.  1906.  Naut.  20  {5):49-5\. 

Walker,  Bryant.  1908.  Naut.  2/ (1 1):  126-129. 


GONAD  DEVELOPMENT  AND  SPAWNING  OF 
BRACHIDONTES  RECURVUS  IN  CHESAPEAKE  BAY 

By  J.  FRANCES  ALLEN* 
National  Science  Foundation,  Washington,  D.  C. 

The  hooked  or  curved  mtissel,  Brachidontes  reciirvus  (Rafin- 
esque)  of  the  family  Mytilidae,  occurs  in  large  numbers  on  oyster 
bars  in  the  tipper  Chesapeake  Bay  and  its  tributaries.  This  species 
is  an  important  fouling  organism  on  oyster  bars  and  affects  the 
growth  habits  of  the  oyster.  It  was  reported  by  Frey  (1946)  as 
forming  much  of  the  cultch  on  oyster  bars  in  the  Potomac  River 
where  most  of  the  marketable  oysters  weie  completely  covered 
with  mussels.  As  a  result,  the  oysters  were  misshapen  from  grow- 
ing among  the  mussels.  Beaven  (1947)  states  that  this  species  is 
most  abundant  on  bars  of  the  upper  portion  of  the  Bay  and  its 
tributaries  where  the  salinity  is  low. 

Under  the  name  Mytilus  recurvus,  this  species  has  been  re- 
ported (Nelson,  1928a)  as  occurring  along  the  New  Jersey  coast. 
Chestnut  (1949)  noted  the  appearance  of  the  curved  mussel  in 
North  Carolina.  Englc  (1945)  observed  that  the  hooked  mussel, 
Mytilus  Jiaynatiis  Say,  was  present  on  oyster  bars  in  Alabama, 


*  Work  done  while  with  the  Department  of  Zoology,  University  of  Mary- 
land, College  Park,  Maryland. 


150  NAUTILUS  Vol.   75(4) 

and  later  (1948)  the  same  investigator  reported  its  presence  on 
bars  in  Mississippi  and  Louisiana.  According  to  Abbott  (1954) 
the  geographic  range  of  Brachidontes  recurvus  is  Cape  Cod  to  the 
West  Indies. 

Despite  the  wide  distribution  of  this  species  and  its  impor- 
tance as  a  fouling  organism  on  oyster  bars,  a  survey  of  the  lit- 
erature reveals  that  with  the  exception  of  the  observations  on 
distribution  previously  mentioned,  little  information  is  available 
concerning  its  biology.  This  situation  is  in  sharp  contrast  to  that 
existing  in  regard  to  other  mussels,  such  as  Mytilus  edulis  Linne; 
Mytilus  calif ornianus  Conrad;  Modiolus  demissus  (Dillwyn)  ;  and 
Modiolus  modiolus  (Linne) ,  where  considerable  data  are  avail- 
able on  numerous  aspects  of  the  biology  of  the  species. 

The  investigation  reported  here  was  part  of  a  cooperative  pro- 
gram between  the  Department  of  Zoology  of  the  University  of 
Maryland  and  the  Shellfisheries  Investigations  of  the  United 
States  Fish  and  Wildlife  Service,  located  at  Annapolis,  Maryland. 
Appreciation  is  expressed  to  Dr.  Robert  A.  Littleford  for  his 
suggestions  and  criticisms;  and  to  Mr.  James  B.  Engle  and  Staff 
of  the  Fish  and  Wildlife  Service  for  collection  of  materials  and 
for  data  on  temperature  and  salinity. 

Procedure:  Specimens  of  B.  recurvus  used  in  this  study  were 
collected  from  Hackett's  Bar,  located  on  the  western  side  of  upper 
Chesapeake  Bay  and  approximately  two  nautical  miles  north  of 
the  Severn  River.  This  bar  is  1/2  to  %  of  a  mile  offshore  from 
Hackett  Point,  in  water  varying  in  depth  from  2  to  35  feet.  Sam- 
ples were  taken  at  two  week  intervals  from  January  16,  1950' 
through  April  16,  1951,  except  when  prevented  by  weather  con- 
ditions or  mechanical  difficulties  with  the  boat.  All  the  mussels 
were  collected  with  a  standard  oyster  dredge  from  the  same  gen- 
eral location  on  the  bar  and  at  a  depth  of  16  feet. 

With  the  shell  closed  and  the  specimen  held  in  a  lateral  posi- 
tion (Newcombe  and  Kessler  1936),  shell  dimensions  were 
measured  to  the  nearest  tenth,  using  a  sliding  vernier  caliper 
calibrated  in  millimeters.  The  dimensions  used  are  defined  as 
follows:  length,  the  greatest  distance  between  the  anterior  and 
posterior  ends  of  the  shell;  width,  the  greatest  distance  between 
the  dorsal  and  ventral  edges;  and  thickness,  the  greatest  distance 
between  the  right  and  left  valves.  The  term  height  is  commonly 


I 


April,  1962  nautilus  151 

used  in  place  of  thickness  as  a  more  meaningful  designation. 

Specimens  of  the  entire  size  range  were  selected  from  each 
collection  for  microscopic  study.  The  individuals  were  fixed 
within  the  shell  by  severing  the  posterior  muscles  and  then  drop- 
ping the  entire  animal  into  Bouin's  alcoholic  fixative  (Galigher, 
1934) .  Other  fixatives  were  tried  but  Bouin's  proved  most  satis- 
factory. After  fixation  for  several  days,  they  were  removed  from 
the  shell,  washed  in  70%  alcohol  and  then  stored  in  80%  alcohol. 
They  were  dehydrated  in  alcohol  and  xylol,  embedded  in  Tissue 
Mat,  sectioned  at  10  micra,  stained  with  either  Harris's  hema- 
toxylin and  eosin  or  Mallory's  triple  stain,  and  mounted  in 
balsam. 

Plankton  samples  were  collected  simultaneously  with  the  col- 
lection of  mussels  by  pumping  100  liters  of  water  from  just  above 
the  bar  through  a  No.  20  silk  plankton  net.  Each  sample  was 
concentrated  to  100  ml.  and  preserved  with  formalin.  Examina- 
tion of  these  samples  to  determine  the  presence  or  absence  of 
mussel  larvae  was  made  with  a  Sedgewick-Rafter  Cell  (Little- 
ford,  Newcombe,  and  Shepherd,  1940) . 

Obseiuations.  The  gross  microscopic  morphology  of  the  curved 
mussel  is  identical  with  that  of  Mytihis  edulis  as  described  by 
Field  (1922) .  However,  the  alimentary  tract  is  sunounded  by 
the  visceral  mass  which  contains  the  other  organ  systems  of  the 
animal.  Immediately  surrounding  the  stomach  and  intestine  is 
the  so-called  liver,  or  digestive  diverticula.  The  latter  structure 
also  extends  into  the  mesosoma,  which  is  an  elongate,  ventral 
extension  of  the  visceral  mass,  except  during  the  period  of  gonad 
development,  when  the  mesosoma  is  filled  with  the  gonad  tissue. 

The  gonadal  tissue  of  B.  recurvus  consists  of  a  series  of  ducts 
and  canals,  lined  with  germinal  epithelium,  which  end  in  pockets 
or  follicles.  The  mature  tissue  occupies  almost  the  entire  mantle 
(Plate  15,  fig.  4)  and  the  mesosoma  to  the  edge  of  the  foot  and 
penetrates  into  the  digestive  diverticula,  where  it  is  found  ad- 
jacent to  the  so-called  liver  canals,  the  stomach,  and  the  direct 
and  recurrent  intestine.  Apparently  the  early  gonad  development 
begins  in  the  mantle  and  then  spreads  into  the  mesosoma.  The 
vesicular  tissue  which  fills  in  between  the  follicles  is  replaced  by 
the  gonad  tissue  as  it  matures.  Thus,  when  the  sex  cells  are 
mature,  nearly  the  entire  structure  of  the  organism  is  devoted  to 


152  NAUTILUS  Vol.  75(4) 

reproduction.  As  sexual  maturity  approaches,  the  mantle  and 
mesosoma  become  either  a  bright  yellow  or  a  stippled  brown 
in  color. 

The  craved  mussel  is  dioecious,  and  of  the  896  specimens  exam- 
ined, there  was  no  indication  of  the  existence  of  hermaphrodit- 
ism or  alternation  of  sex.  The  gonads  of  both  male  and  female 
show  parallel  development  in  time  and  in  the  position  of  the 
gonads.  From  those  studied,  the  sex  ratio  apparently  is  evenly 
distributed.  The  reproductive  potential  of  these  mussels  is  de- 
termined by  their  size  rather  than  by  their  age.  Studies  were  made 
of  those  forms  which  were  as  small  as  20  mm.  in  length,  and  their 
gonad  tissue  had  the  same  appearance  as  that  of  the  larger 
specimens.  Presiuiiably  they  produce  as  many  reproducti\c  cells 
as  their  anatomical  structure  will  allow.  This  also  leads  to  the 
conclusion  that  sexual  maturity  occurs  the  summer  following 
setting,  regardless  of  size  of  the  mussel. 

Seasonal  Developmerit  in  the  Male.  The  male  follicles  vary  a 
great  deal  in  size,  depending  upon  whether  the  immature  or 
mature  condition  is  observed.  From  January  until  mid-April,  the 
follicles  are  found  in  tlie  mantle,  and  to  a  lesser  decree,  in  the 
mesosoma.  They  appear  at  this  time  to  be  loosely  filled  with  cells, 
but  by  mid-April  they  become  more  dense  in  appearance  and 
they  take  a  deeper  and  a  darker  stain  (Plate  15,  fig.  1) .  As  they 
mature,  the  follicles  become  filled  with  spermatozoa  which  ra- 
diate from  the  periphery  of  the  follicle  toward  the  center  (Plate 
15,  fig  2).  These  cells  appear  as  small  oval  or  spherical  bodies 
which  are  packed  too  closely  together  for  their  detailed  structure 
to  be  clearly  visible.  They  matiue  by  June  and  remain  mature 
until  October.  Observation  on  the  density  of  the  sex  products 
in  the  follicles  leads  one  to  believe  that  spawning  may  occur 
from  the  mantle  first,  then  proceed  from  the  mesosoma.  However, 
the  appearance  indicates  that  the  animal  could  very  well  spawn 
simiUtaneously  from  both  areas.  Spawning  from  each  follicle  ex- 
tends over  a  considerable  period  of  time  so  that  some  follicles 
do  not  have  the  appearance  of  being  spent,  while  others  are 
mature.  As  the  follicles  mature,  there  are  light,  radiating  areas, 
which  under  increased  magnification  show  that  the  cells  adjacent 
to  these  areas  are  concentrated  in  short  rods,  apparently  similar 
to  the  condition  observed  in  Mytilus  edulis   (Field,  1922). 


April,  1962  nautilus  153 

While  the  follicle  increase  in  size  and  maturity,  the  supportive 
vesicular  tissue  surrounding  them  gradually  disappears  almost 
completely.  After  spawning,  the  supportive  tissue  again  appears, 
filling  the  spaces  between  the  spent  follicles. 

Seasonal  Development  in  the  Female.  The  immature  follicles 
of  the  female  are  separated  by  vesicular  connective  tissue  and 
vary  in  size.  By  the  first  week  in  May,  the  follicles  are  lined  with 
small  cells  containing  nuclei  which  with  hematoxylin  and  eosin 
stain  a  dark  blue.  By  the  third  week  in  May,  the  cells  have  in- 
creased in  size  so  that  the  prominent  nucleoli  can  be  seen,  but  the 
cells  are  still  immature.  By  mid-June,  the  mantle  and  the 
mesosoma  are  occupied  by  follicles  which  are  filled  with  mature 
and  immature  eggs  (Plate  15,  fig.  3) .  Some  of  the  vesicular  sup- 
portive tissue  is  present,  but  has  decreased  considerably  or  dis- 
appeared as  the  eggs  approach  maturity.  The  eggs  vary  a  great 
deal  in  size  and  the  nucleoli  are  large  and  prominent  (Plate  15, 
fig.  3) .  When  the  eggs  mature,  they  break  from  the  follicle  and 
are  inoved  along  by  ciliated  ducts.  The  eggs  are  not  considered 
mature  until  they  break  free.  As  is  the  case  in  the  male,  the 
female  gonad  tissue  penetrates  into  the  digestive  diverticula  and 
is  in  close  proximity  to  the  liver  canals.  (Plate  15,  fig.  3) .  The 
condition  of  the  gonad  from  July  to  October  is  similar  to  that  in 
Jime.  After  October,  however,  the  ova  are  not  as  numerous  and 
some  retain  the  same  appearance  as  those  of  June. 

On  July  3,  the  small  eggs  appeared  in  the  mantle  and  the 
larger  ones  in  the  mesosoma  which  contained  many  follicles.  In 
one  specimen  from  the  September  group,  one  lobe  of  the  mantle 
contained  only  a  few  eggs,  while  the  other  lobe  and  the  mesosoma 
were  dense  with  them.  Although  the  condition  of  the  maturity 
of  the  ova  is  such  that  the  entire  gonad  tissue  is  subject  to  spawn- 
ing at  the  same  time,  the  intensity  of  spawning  and  the  time  of 
spawning  of  these  different  areas  depends  upon  the  individual. 

Mature  eggs  are  present  in  December,  but  show  evidence  of 
degeneration  or  death,  as  shown  by  the  condition  of  the  nucleus, 
the  absence  of  nucleoli,  and  the  general  appearance  of  the  cyto- 
pla.-,m.  WHiile  the  destruction  of  the  gonadal  tissue  appears  evi- 
dent, there  is  no  similarity  to  the  comparable  mechanism  of  re- 
sorption noted  in  some  pclccypods  by  Loosanoff  and  Davis 
(1951). 


154  NAUTILUS  Vol.  75(4) 

The  development  ot  tlie  gonads  indicates  that  the  state  of  sex- 
ual maturity  exists  from  June  through  October.  Release  of  the 
sex  products  occurs  over  a  relatively  long  period  of  time,  from 
early  June  through  October,  and  in  some  cases,  through  Novem- 
ber. Individual  follicles  develop  independently  of  each  other  in 
both  sexes.  In  other  words,  the  gonads  do  not  reach  a  high  peak 
of  maturity  and  immediately  spawn  out. 

Seasonal  Distribution  of  the  Larvae.  Plankton  samples  col- 
lected from  January  16,  1950,  through  April  16,  1951,  were  ex- 
amined to  determine  the  presence,  relative  abundance,  and  stages 
of  development  of  the  larval  forms  of  B.  recurvus. 

Since  the  rate  of  development  of  the  larval  stages  shows  con- 
siderable variation,  in  part,  at  least,  because  of  the  protracted 
spawning  period,  I  decided  to  divide  the  forms  present  into  three 
groups:  pre-hinge  stage,  hinge  stage,  and  post-hinge  stage.  The 
hinge  stage  was  defined  as  "the  straight-hinge-line  embryonic 
shell"  (Field  1922),  between  the  circular  immature  larval  stage 
and  the  triangular  ovate  form  of  the  mature  larva.  In  the  area 
of  Hackett's  Bar,  the  larval  stages  of  this  species  could  be  con- 
fused only  with  the  larval  stages  of  Congeria  leucophaeta,  Cras- 
ostrea  virginica,  or  possibly  with  Tagelus  plebeius  and  Mya 
arenarin.  However,  as  is  pointed  out  by  Sullivan  (1948) ,  care- 
ful examination  will  show  numerous  diagnostic  differences  be- 
tween the  mussel  larvae  and  other  lamellibranchs.  Therefore,  the 
larval  stages  of  different  groups  may  be  easily  separated  from 
each  other,  although  some  confusion  might  possibly  arise  in 
regard  to  the  larvae  of  Brachidontes  recurvus  and  that  of  Con- 
geria leucophaeta.  However,  considering  the  rare  occurrence  of 
the  latter  form  on  Hackett's  Bar,  such  confusion  should  not 
invalidate  the  data. 

The  results  of  the  observations  on  larval  distribution  are  pre- 
sented in  Table  I.  The  relative  abundance  of  the  various  larvae 
is  expressed  as  present,  common  abundant,  and  very  abundant, 
rather  than  by  using  some  numerical  statement.  I  felt  that  such 
an  expression  of  results  had  more  validity  than  an  exact  numer- 
ical expression,  since  the  method  of  collection  was  not  subject  to 
exact  quantitative  analysis  (Littleford,  Newcombe,  and  Shep- 
herd, 1940). 

Larvae  appeared  in  the  plankton  for  the  first  time  on  June  5, 


April,  1962 


NAUTILUS 


155 


TABLE  I 

SEASONAL  OCCURRENCE  OF  URVAE  OF  BRACHIDONTES 

RECURVUS  IN  PLANKTON  SAMPLES 


Date 

Pre-hinge 

Hinge 

Post-hinge 

June  5 

Present 

Present 

Absent 

June  19 

Coramon 

Cornmon 

Present 

July  3 

Ganmon 

Ccnnion 

Common 

July  17 

Absent 

Present 

Present 

July  31 

Abundant 

Present 

Absent 

August  14 

Ccramon 

Common 

Absent 

August  28 

Abundant 

Abundant 

Abundant 

Sept.  11 

Very  Abundant 

Very  Ab'jndant 

Very  Abundant 

Sept.  25 

Common 

Present 

Cormon 

October  9 

Present 

Absent 

Common 

October  25 

Absent 

Absent 

Present 

Nov.  13 

Present 

Common 

Contnon 

Nov.  27 

Absent 

Absent 

Present 

Dec.  11 

Absent 

Absent 

Present 

1950  when  pre-hinge  and  hinge  forms  were  observed  in  the 
samples.  By  the  middle  of  June,  the  pre-hinge  and  hinge  stages 
had  quadrupled  in  number  over  the  earlier  part  of  the  month. 
The  post-hinge  stage  was  observed  for  the  first  time  on  June  19, 
1950.  During  July  and  August  there  was,  generally  speaking,  a 
continued  rise  in  abundance  of  the  larval  stages.  However,  the 
pre-hinge  stage  was  not  found  on  July  17,  1950  and  the  post- 
hinge  stage  was  not  observed  in  collections  of  July  31  and  August 
14.  All  three  stages  were  present  in  large  numbers  in  the  collec- 
tion of  August  28,  and  the  peak  of  their  numerical  abundance  in 


156  NAUTILUS  Vol.  75(4) 

the  plankton  was  ob,ser\cd  on  September  11,  1950. 

Some  idea  of  the  relative  seasonal  abundance  of  the  larval 
stages  may  he  obtained  from  the  fact  that  the  number  of  pre- 
hinge  larvae  in  June  was  27.50%  of  the  total  number  observed  on 
September  11.  Similarly,  the  number  of  the  hinge  larvae  in  June 
was  15.15%  and  the  number  of  post-hinge  was  0.54%  of  the 
total  observed  on  September  1 1 . 

(To  be  concluded) 


AUSTRALORBIS  ALBICANS  (PLANORBIDAE) 

Bv  W.  LOBATO  PARAENSE  and  NEWTON  DESLANDES' 
Institute  Nacional  de  Endemias  Rurais,  Brazil 

This  species  was  described  by  Pfeiffer  (1839,  p.  354),  on  the 
basis  of  the  shell  characters,  as  follows: 

"43.  Planorbis  albicans  Pfr. — Testa  orbiculari,  utrinque  um- 
bilicata,  solidula,  albicante  vel  pallide  fulvicante,  anfract.  3 
teretibus;  labro  subincrassato  albo;  apertura  subovata. — Diam. 
21/2,  alt.  1'". — Dem  PI.  albus   (hispidus)   am  nachsten  verwandt." 

Clessin's  desauption  (1884,  p.  119-120)  is  more  detailed  and 
extends  the  species  range  to  other  Antillean  islands: 

"87.  Planorbis  albicans  Pfeiffer.  Taf.  11.  Fig.  14.  T.  depressa, 
utrinque  centro  profundissime  immerso;  subtiliter  striatula; 
lutida,  pallide-cornea;  anfractus  4,  rapide  accrescentes,  rotundati; 
superius  valde  convexi,  sutura  profundissima  separati;  inferius 
planulati,  sutura  paulo  immersa  disjuncti;  idtimus  penultimo 
paulo  latior,  aperturam  versus  descendens;  apertura  late-lunata, 
albo-labiata,  valde  obliqua;  peristoma  acutum,  marginibus  con- 
junctis.  Diam.  5,5,  alt.   1,8   Mm. 

Planorbus  albicans  Pfeiffer  Wiegm.  Archiv  I   1839  p.  354. 

Planorbus  albicans  Reeve  conch.   Icon.   XX   f.    117.    (?) 

Gehause  gedriickt,  ober-  und  unterseits  in  de  Mitte  tief 
eigenesenkt,  sehr  fein  gestreift,  glanzend,  hellhornfarben,  Um- 
gange  4,  rasch  zunehmend,  nach  oben  sehr  gewolbt  und  durch 
eine  sehr  tiefe  Naht  getrennt;  nach  unten  flacher  und  durch 
eine  seichtere  Naht  verbunden;  der  letzte  Umgang  nur  i/gmal 
breiter  als  der  vorletzte;  gegen  die  Miindung  etwas  herabstei- 
gend;  Miindung  breit-mondformig,  weissgelippt,  sehr  schief; 
Mundsaum  scharf,  mit  verbundenen  Randern.  Vaterland:   Die 


^  Work  made  in  cooperation  with  Institnto  Oswaldo  Cruz  and  Service 
Especinl  tie  Saiulc  i'uhlica.  Aided  by  the  C'onsellio  Nacional  de  Pes(|uisas  of 
Brazil,  which  defrayed  the  expenses  of  a  trip  by  the  senior  author  to  the 
type  locality  of  A.  albicans  and  also  provided  additional  facilities  for  the 
study  of  the  material  collected  there. 


NAin  ILIIS  75(4) 


PLATE   15 


I.  Ii)(i( liidoulcs  icrun'us  in  niicl-A|)ril.  Imiiuiuirc  t'ollicles  of  (lie  male 
I^oiukI  in  the  niaiitle.  H.  and  E.  slain.  X2l)().  2.  in  June.  Male  follicles  lilled 
with  spermatozoa.  Note  the  i^adiation  from  the  perijiheiN  of  the  follide  lo 
ihe  (enter,  (iross  section  through  the  niesosoma.  Malloi\'s  Triple  slain.  \2()(). 
3.  in  mitl-Jiuie.  Matiue  female  follicles  adjaceni  lo  the  so-called  liver  canals 
in  the  di;^esti\c  di\erli(  iila.  H.  and  I-',,  stain.  X2()U.  4,  in  Jidy.  General  view 
of  cross  section  in  region  of  the  gills.  Note  the  presence  of  mature  and  im- 
mature eggs  and  the  prominence  of  the  nucleoli.  H.  and  E.  stain.  X50. 


NAUTILUS  75(4) 


PLATE   16 


Fig.  1:  Shell  of  Auslralot his  albicans  horn  Laguna  Somorostro,  Havana, 
lig.  2:  Raclula  teeth  of  Aiislralorbis  albicans  from  Lagiina  La  Canoa,  Pinai 
del  Rio  {C'.  =  central,  I  intcinicdiale,  L  ^  lateral,  M  -  marginal).  Fig.  3: 
(ieiiital  organs  of  Australorbis  albicans  from  I.agiina  La  C.anoa.  Pinar  dci 
Rio  (ag=  fragment  of  albumen  gland,  ta  ^  carrefoin-,  ng  -  nidamenlal 
gland,  od  "  proximal  segment  of  ovispermidnct,  od' =  distal  segment  of 
ovispermiduct,  ot  -  ovotestis,  o\  ^"  oviduct,  po  =  pouch  of  oviduct,  pp  = 
prepuce,  pr  ^  prostate  gland,  ps  =  penis  sheath,  rm  =  retractor  muscles  of 
penial  complex,  sd  -  spermiduct,  sp  -  spermatheca,  .sv  =  seminal  vesicle, 
ut  =  uterus,  va    -  vagina.  \tl       \as  deferens). 


April,  1962  nautilus  157 

Antillen,  liisel  Cuba,  bei  Cardenas;  Portorico,  St.  Thomas. 
(Coll.  Dunker) . 

Die  Art  gehort  zur  Sect.  Armigerus,  welche  durch  im  Innern 
der  Umgange  angebrachte  Zahne  und  Lamellen  ausgeszeichnet 
ist. — Diese  sitzen  (meist  zu  2  oder  3  an  der  Miindungswand,  und 
zu  3  an  der  gegeniiber  liegenden  Gaumenwand)  und  zwar  in 
ziemlicher  Entfernung  von  der  Miindung,  so  dass  sie  gewohnlich 
erst  beim  Zerstoren  des  Gehauses  zu  entdecken  sind,  wenn  sie 
nicht  durch  die  Umgange  durchscheinend  von  aussen  an  der 
weissen  Farbe  erkannt  werden.  Auf  der  Miindungswand  steht 
gewohnlich  in  de  Mitte  derselben  ein  grosserer  starkerer  Zahn, 
dem  nach  unten  gegen  die  Ecken  ein  kleinerer  sich  anreiht.  Auf 
der  Gaumenwand  befindet  sich  in  der  Mitte  eine  lamellen — 
oder  faltenartige  Schmelzleiste,  wahrend  zu  beiden  Seiten  dersel- 
ben in  nahezu  gleicher  Entfernung  von  ihr  und  etwa  in  der 
Mitte  des  Raumes  von  der  Mittelleiste  bis  zu  den  Anschlussecken 
zwel  Hocker — oder  zahnartige  Schmelzstiickchen  angebracht 
sind. — Ueber  den  Zweck  dieser  Gehauseverstarkungen  fehlen  bis 
jetzt   noch   Beobachtungen." 

Among  the  planorbids  collected  by  the  senior  author  in  Cuba, 
in  December,  1956,  there  were  specimens  whose  shell  agreed,  in 
shape  and  dimensions,  to  the  two  above  transcribed  descriptions, 
inclusive  as  to  the  apertural  lamellae.  Some  of  them,  dissected 
for  previous  observation,  proved  young  forms  of  a  species 
anatomically  undistinguishable  from  Planorbis  peregrinus  Or- 
bigny,  1835.  As  then  shown  (Paraense  and  Deslandes,  1958b), 
the  shape  of  the  shell  underwent  a  process  of  remodelling  along 
with  growth,  with  eventual  resorption  of  the  five  lamellae. 

A  sample  from  Puerto  Rico,  sent  us  by  Drs.  F.  F.  Ferguson  and 
Charles  S.  Richards  in  1959,  and  regarded  by  them  as  albicans, 
was  conchologically  similar  to  the  aforesaid  Cuban  material, 
but  the  specimens  stopped  growing  when  they  reached  about 
7  mm.  in  diameter,  developed  permanent  lamellae  and  showed 
remarkable   anatomical   differences. 

Those  facts  led  us  to  examine  our  remaining  material,  among 
which  we  found  1 1  specimens  anatomically  similar  to  the  Puerto 
Rican  ones.  Of  such  specimens,  7  were  collected  at  Laguna  La 
Canoa  (between  Artemisa  and  Mangas,  Candelaria,  Province 
of  Pinar  del  Rio),  and  4  at  Laguna  Somorostro  (Havana) .  The 
largest  specimens  were  5  mm.  in  shell  diameter,  and  only  two 
from  La  Canoa  showed  apertural  lamellae. 

Taking  into  account  Pfeiffer's  and  Clessin's  descriptions,  and 


158  NAUTILUS  Vol.  75(4) 

the  fact  that  we  are  dealing  with  topotypic  specimens,  we  agree 
with  Drs.  Ferguson  and  Richards  in  considering  this  species  as 
Phinorbis  albicans.  Since  the  generic  problem  of  the  group  to 
which  it  belongs  was  submitted  to  the  International  Commission 
on  Zoological  Nomenclature,  the  species  is  here  placed  in  the 
genus  Aiistralorhis,  according  to  the  reasons  presented  by  Para- 
ense    (1961). 

The  following  description  is  based  on  the  above-mentioned 
Cuban  material,  of  which  four  shells  and  one  dissected  specimen 
were  deposited  in  the  collection  of  Instituto  Oswaldo  Cruz  (No. 
7907) . 

Description:  The  empty  shell  (PI.  16,  fig.  1),  is  yellow  or 
amber,  finely  oblicjuely  striate,  has  about  31/9  whorls,  and  usually 
shows  6  internal  lamellae  deeply  situated  in  the  apertural  region. 
The  whorls  widen  rapidly,  but  at  the  region  of  the  lamellae  they 
stop  expanding  or  even  slightly  decrease,  and  then  rapidly  widen 
toward  the  opening.  The  outer  whorl  shows  various  degrees  of 
downward  deflection  at  the  apertural  region.  The  upper  (right) 
side  is  flattened  and  deeply  umbilicate.  It  shows  round  smooth- 
walled  whorls,  of  which  the  inner  one  is  obscured  at  the  bottom 
of  the  umbilicus.  The  under  (left)  side  is  concave  and  has  a 
vortex-shaped  central  depression  shallower  than  the  umbilical 
one.  On  this  side  the  whorls  are  bluntly  carinate  and  the  inner 
whorl  is  plainly  visible  at  the  bottom  of  the  central  depression. 
The  aperture  is  directed  forwards  in  young  specimens,  but  as  the 
shell  grows  larger  the  lamellae  arise  and  gradually  develop,  and 
the  aperture  gradually  bends  downwards.  In  older  specimens  a 
callous  thickening  of  the  lip  may  be  found.  A  complete  set  of 
lamellae  consists  of  two  parietal  and  four  palatal  units.  Not  only 
as  concerns  the  lamellae,  but  also  the  shell,  this  species  is  similar 
to  Australorbis  janeirensis  (see  Paraense  and  Deslandes,  1956), 
and  to  young  A.  glabratiis  and  A.  peregriniis  from  some  popula- 
tions whose  specimens  may  develop,  at  early  age,  a  transient  set 
of  apertural  lamellae  (Paraense,  1957;  Paraense  and  Deslandes, 
1958b). 

The  animal  shows  no  appreciable  difference  from  that  of  other 
small  species  of  the  genus.  The  pseudobranch  is  simple,  thin  and 
flat.  The  rectal  ridge  extends  from  the  pseudobranch  into  the 


April,  1962  nautilus  159 

pulmonary  cavity,  disappearing  at  the  level  of  the  stomach.  There 
is  no  renal  ridge.  A  dorsolateral  ridge,  facing  the  rectal  one,  runs 
parallel  to  the  left  side  of  the  renal  vein. 

The  genital  organs  are  shown  in  fig.  3. 

The  ovotestis  is  composed  of  numerous  sac-like  diverticula, 
which  are  nearly  always  simple  and  less  frequently  bifurcate.  The 
diverticula  open  into  the  ventral  collecting  canal,  which  con- 
tinues into  the  thin  proximal  segment  of  the  ovispermiduct. 
Then  follows  the  seminal  vesicle,  whose  parietal  diverticula 
are  poorly  developed,  as  in  A.  philippianus  (see  Paraense  and 
Deslandes,  1958a).  The  seminal  vesicle  gradually  merges  into  the 
distal  segment  of  the  ovispermiduct,  which  is  about  3-4  times  as 
long  as  the  proximal  segment. 

The  spermiduct,  highly  sinuous  at  the  beginning,  follows  a 
flexuous  course  in  contact  with  the  oviduct  and,  then,  traverses 
the  furrow  formed  by  the  pouch  of  the  oviduct.  At  the  point 
where  it  emerges  from  that  furrow,  it  receives  a  single  row  of 
prostrate  diverticula.  The  number  of  diverticula  varied  from  8 
to  16  in  our  material.  They  are  mainly  bi-  or  trifurcate,  less  fre- 
quently unbranched  or  arborescent.  Although  some  of  them  have 
a  short  stalk,  they  are  mostly  sessile,  their  branches  arising  al- 
most directly  from  the  prostate  duct.  The  prostate  duct  continues 
into  the  vas  deferens,  which  has  no  special  characteristics. 

The  penis  sheath  varies  from  a  little  longer  to  about  4  times 
as  long  as  the  prepuce.  It  contains  a  penis  that  shows  no  essential 
differences  from  that  of  congeneric  species.  The  prepuce  is  only 
a  little  wider  than  the  penis  sheath  and  is  internally  separated 
from  the  latter  by  a  muscular  diaphragm. 

The  oviduct  is  similar  to  that  of  other  congeneric  species,  also 
having  a  large  pouch  of  bosselated  walls.  The  nidamental  gland 
and  uterus,  taken  conjointly,  are  from  one  and  half  times  to 
thrice  as  long  as  the  oviduct.  The  vagina  is  tubular  and  smooth- 
walled.  The  spermatheca  is  pear-shaped  or  club-shaped  and  has 
a  narrow  duct  about  as  long  as  the  spermathecal  body. 

The  jaw  is  somewhat  T-shaped,  consisting  of  a  wider  upper 
piece,  vertically  striate,  and  two  narrower  lateral  pieces.  The  fol- 
lowing characteristics  were  observed  in  5  specimens  from  Pinar 
del  Rio  (fig.  2)  :  radula  formula,  14-1-14  to  15-1-15;  horizontal 
rows,  80  to  90;  central  tooth  bicuspid;  4  to  6  laterals,  2  to  3 


160  NAUTILUS  Vol.   75(4) 

intermediates,  6  to  8  marginals. 

Comparison  loitli  related  species:  In  a  conchological  diagnosis 
between  A.  albicans  and  other  congeneric  species,  there  will  al- 
ways be  place  for  uncertainty.  In  fact,  even  if  no  doubt  arises  as 
to  the  distinction  between  adult  A.  albicans  and  young  shells 
from  large  species  of  Australorbis,  there  remain  to  be  considered 
the  similarities  between  the  former  and  other  lamellate  shells.  Of 
the  Neotropical  species  studied  by  us,  the  adult  A.  janeirensis  is 
always  provided  with  a  permanent  set  of  apertural  lamellae 
(Paraense  and  Deslandes,  1956) .  Under  special  environmental 
conditions,  young  individuals  from  other  species,  such  as  A. 
glabratus  and  A.  peregrinus,  may  develop  a  transient  set  of 
lamellae  which  are  resorbed  as  the  shell  grows  larger  (Paraense, 
1957;  Paraense  and  Deslandes,  1958b) .  Thus,  a  reliable  diagnosis 
must  be  based  on  the  anatomical  characters  associated,  of  coinse, 
with  those  of  the  shell. 

A.  albicans  may  be  readily  distinguished  from  A.  glabratus  by 
the  absence  of  renal  ridge,  or  of  the  pigmented  line  that  precedes 
it  in  young  specimens,  and  of  the  vaginal  pouch  (Paraense  and 
Deslandes,  1955a,  1959)  ;  from  A.  tenagophilus  (formerly  called 
A.  nigricans  by  us),  A.  peregrinus,  A.  andecolus  and  A.  prorius, 
by  the  absence  of  tlie  vaginal  pouch  (Paraense  and  Deslandes, 
1955b,  1957,  1958b,  c)  ;  and  from  A.  stramineus  (^  centimetralis) 
by  the  absence  of  vaginal  corrugation  (Paraense  and  Deslandes, 
1955c). 

Despite  the  great  similarity  of  its  shell,  when  lamellate,  to  that 
of  A.  janeirensis,  there  are  remarkable  anatomical  differences 
bet^\'een  the  two  species  (for  comparison,  see  Paraense  and  Des- 
landes, 1956)  .  The  penial  complex,  the  vagina  and  the  sperma- 
theca  of  A.  janeirensis  are  very  long,  the  penial  complex  being 
about  half  as  long  as  the  whole  female  duct  (oviduct,  nidamental 
gland,  uterus  and  vagina  together)  ;  moreover,  the  penis  sheath 
is  from  4.7  to  7.6  times  as  long  as  the  prepuce.  In  A.  albicans,  the 
penial  complex,  the  vagina  and  the  spermatheca  are  relatively 
very  short,  the  penial  complex  being  from  14  to  1/7  (about 
1/^.5)  as  long  as  the  female  duct  and,  the  penis  sheath,  from  a 
little  longer  to  about  4  times  as  long  as  the  prepuce. 

Of  the  Australorbis  species  so  far  studied  by  us,  the  most  closely 
related  to  A.  albicans,  from  the  anatomical  standpoint,  is  A. 
philippianus    (see   Paraense  and  Deslandes,    1958a) .  The  latter 


April,  1962  nautilus  161 

also  has  a  small  penial  complex,  the  length  ot  which  is  about 
1/^  that  of  the  whole  female  duct  (range  from  1/2.5  to  1/7) .  The 
two  species  may  be  distinguished  by  the  characters  of  the  penial 
complex  and  the  prostate.  The  penis  sheath  is  from  a  little  longer 
to  about  4  times  as  long  as  the  prepuce  in  albicans;  the  two  or- 
gans are  about  the  same  length  in  philippianiis,  the  ratio  between 
the  former  and  the  latter  varying  from  0.9  to  1.5.  And  the  number 
of  prostate  diverticula  is  much  smaller  in  philippianiis:  from  0  to 
about  6,  as  against  8  to  16  in  the  present  sample  of  albicans. 

Acknowledgement:  We  are  indebted  to  the  distinguished  Cu- 
ban naturalists  Miguel  Jaume  and  Manuel  Barro,  for  their  help- 
ful assistance  to  the  senior  author  during  his  work  in  Cuba. 

Summary 
A  description  of  the  planorbid  species  Australorbis  albicans 
(Pfeiffer,  1839)    is  presented.  The  following  of  its  characters  are 
considered  as  having  diagnostic  significance: 

Shell  up  to  about  7  mm.  in  diameter  and  2.5  mm.  in  height, 
usually  with  6  apertural  lamellae  in  the  adult;  whorls  about  814, 
rapidly  widening,  bluntly  carinate  underneath.  Absence  of  renal 
ridge.  Ovotestis  diverticula  simple  or,  less  frequently,  bifurcate. 
Seminal  vesicle  with  poorly  developed  diverticula.  Prostate  di- 
verticula mainly  bifid  or  trifurcate,  less  frequently  unbranched 
or  arborescent,  and  arranged  in  a  single  row.  Penis  sheath  from 
a  little  longer  to  about  4  times  as  long  as  the  prepuce.  Well  de- 
veloped pouch  of  oviduct.  Vagina  tubular  and  smooth-walled, 
without  pouch  or  corrugation. 

References 
Clessin,  S.,    1884.  Die  Familie  der  Limnaeiden  enthaltend  die 

Genera  Planorbis,  Limnaeus,  Physa  und  Amphipeplea.  In 

MARTINI    k    CHEMNITZ:     Systematisches    Conchylien- 

Cabinet,  v.  17.  Bauer  &  Raspe,  Niirnberg. 
Paraense,  W.  L.,  1957.  Proc.  Malac.  Soc.  London,  32  (4):  175-179. 
.   1961.   Shell   versus   anatomy   in   planorbid   systematics.   I. 

Australorbis  glabratiis.  Rev.  Brasil  Biol.,  in  the  press. 
Paraense,  W.  L.  and  Deslandes,  N.,  1955a.  Mem.  Inst.  Oswaldo 

Cruz,  53(1)  :87-103. 

.   1955b.  Mem.  Inst.  Osioaldo  Cruz,  53(1)  :121-134. 

.   1955c.  Rev.  Brasil.  Biol,  15(3)  :293-307. 

.   1956.  Rev.  Brasil  Biol.,  16(1)  :81-102. 

.  1957.  Rev.  Brasil.  Biol.,  17  (2)  : 235-243. 

.   1958a.  Rev.  Brasil.  Biol.,  18(2)  : 209-2 17. 

.   1958b.  Jour.  Conchyliol.,  98  (3)  :  152-162. 

.  1958c.  Rev.  Brasil  Biol.,  18  (4):367-373. 

.  1959.  Am.  J.  Trop.  Med.  &  Hyg.,  8  (4)  : 456-472. 

Pfeiffer,  L.,  1839.  Arch.  f.  Naturgesch.,  I.  Zool.,  5  (1)  :346-358. 


162  NAUTILUS  Vol.  75(4) 

X-RAY  DIFFRACTION  EXAMINATION  OF  TWO  FORMS 
OF  OLIVA  SPICATA 

By  jerry  DONOHUE  and  KENNETH  HARDCASTLE 
Department  of  Chemistry,  University  of  Southern  CaUfornia 

Burch  (1959)  has  discussed  a  problem  concerned  with  identity 
of  two  forms  [Cf.  plate  17]  of  Oliva  spicata  (Roding,  1798)  from 
La  Paz,  Baja  California,  which  were  brought  to  his  attention  by 
Mr.  Naylor  of  San  Diego,  who  stated  that  one  form  was  rejected 
by  Indian  artisans  on  the  basis  that  they  shatter  while  being 
worked,  the  other  form  being  acceptable  in  making  various  arti- 
facts. Although  at  first  glance  both  lots  appeared  identical,  addi- 
tional study  revealed  differences  which  may  be  summarized  as 
follows: 

Used  shells:  Discarded  shells: 

Base  of  columella  Base  of  columella 

white  or  pink,  light  purple, 

more  obese,  more  slender, 

lower  spire.  higher  spire. 

For  convenience,  Burch  temporarily  identified  the  acceptable 
shells  as  Oliva  spicata  var.  venulata,  the  discarded  ones  as  var. 
ustulata,  and  concluded  that,  since  the  two  forms  were  easily 
separable,  the  lumping  of  all  these  variations  under  one  name 
was  odd.  O.  venulata  (Lamarck,  1811)  and  O.  ustulata  (La- 
marck, 1811)  are  usually  included  in  the  synonomy  of  O.  spicata, 
e.g.,  by  Johnson    (1915)    and  Keen    (1958). 

In  the  hope  of  throwing  additional  light  on  this  question,  we 
have  examined,  by  the  method  of  x-ray  diffraction,  samples  taken 
from  both  forms.  These  samples  were  taken  from  broken  frag- 
ments, and  the  locations  from  which  the  samples  were  taken  is 
indicated  in  Fig.  1,  which  presents  an  equatorial  cross  section  of 
a  characteristic  shell.  Shells  from  both  lots  showed  the  structure 
shown  in  Fig.  1,  i.e.,  a  translucent  outer  layer  which  encloses  the 
pigment,  a  somewhat  thicker,  white,  inner  layer  which  also  en- 
closes pigment  that,  however,  is  not  visible  because  this  layer  is 
thicker,  and  finally,  a  white,  opaque,  middle  layer  which  con- 
tains no  pigment.  The  outer  and  inner  layers  coalesce  at  the 
outer  lip. 

Samples  taken  from  the  outer  and  middle  layers  of  both  forms 
showed  almost  the  same  x-ray  powder  diffraction  patterns.  All 


April,   1962 


NAUTILUS 


163 


Translucent  outer  layer 
Opaque  middle  layer 


o 


I  cm 

Fig.  1.  Equatorial  cross  section  of  a  characteristic  specimen,  showing  the 
region  from  which  the  samples  were  selected.  The  dark  areas  represent  the 
pigmentation,  which  was  avoided  in  taking  the  samples  for  diffraction 
analysis. 

their  major  and  minor  features  exactly  match  the  pattern  re- 
ported by  Swanson,  Fuyat,  and  Ugrinic  (1954)  for  the  aragonite 
polymorph  of  calcium  carbonate.  An  important  difference  occurs, 
however,  in  the  case  of  both  kinds  of  samples  from  the  form 
acceptable  to  the  Indians  in  that  faint  additional  lines  character- 


164  NAUTILUS  Vol.  75(4) 

istic  of  the  powder  pattern  of  the  calcite  polymorph  of  calcium 
carbonate  (Swanson  and  Fuyat,  1953)  may  also  be  seen.  We 
estimate  that  about  5%  of  calcite  occurs  in  the  outer  layer,  and 
somewhat  more  than  half  that  much  in  the  middle  layers,  of  the 
utilized  form.  This  calcite  content  is  significantly  different  from 
that  found  in  the  samples  from  the  form  discarded  by  the  artisans 
where  none  at  all  could  be  detected  in  the  middle  layer  and  only 
a  trace  in  the  outer  layer. 

It  would  thus  appear  that,  in  addition  to  the  differences  in 
shape  and  color  between  the  two  forms,  there  is  also  a  difference 
in  chemical  constitution.  There  does  not  seem  to  be  any  a  priori 
explanation  as  to  why  this  difference  should  result  in  the  differ- 
ence in  physical  character  noted  by  the  artisans.  Although  this 
may  or  may  not  be  relevant,  calcite  is  considerably  softer  than 
aragonite  (hardness  3,  as  compared  to  4,  Mohs  scale) ,  but  pos- 
sibly the  shattering  property  may  be  the  result  of  some  other 
effect,  e.g.,  differences  in  the  orientation  of  the  aystallites  in  the 
shell. 

The  question  of  which  are  the  appropriate  names  to  assign  to 
these  forms  is  discussed  by  Burch  and  Burch   (1962) . 

Experimental  Details.  The  x-ray  diffraction  patterns  were  re- 
corded photographically  by  the  standard  techniques  in  a  Norelco 
powder  camera,  with  CuKa  radiation,  filtered  through  nickel 
foil.  All  lines  out  to  spacings  of  d  >  1.15  A  were  measured.  Rel- 
ative intensities  were  estimated  visually.  The  38  lines  in  this 
region  reported  for  aragonite  were  all  observed  at  the  expected 
spacings  and  with  the  correct  relative  intensities  for  the  four 
different  types  of  samples.  The  calcite  lines  observed  in  the  case 
of  the  samples  from  the  "accept"  form,  all  with  very  faint  in- 
tensity, were  those  at  d  =  3.86  A  and  3.035  A.  These  are  two 
of  the  strongest  lines  reported  by  Swanson  and  Fuyat  for  calcite. 
All  the  other  strong  calcite  lines  listed  by  them  are  coincident 
with  lines  due  to  aragonite  and  were  not  observed.  The  only 
calcite  line  observed  in  the  case  of  the  samples  from  the  outer 
layer  of  the  "reject"  form  was  the  one  at  spacing  3.035  A,  which 
is  by  far  the  strongest  line  of  that  pattern;  the  next  most  intense 
lines  were  too  weak  to  be  observed.  The  calcite  content  was 
estimated  by  comparison  of  the  intensities  of  the  3.035  A  line 
of  calcite  with  the  2.871  A  line  of  aragonite. 


NAUTILUS  75(4) 


PLATE  17 


Upper  and  lower,  Icli  ligs..  Iiolotxpc  ol  Ulii'a  rejecla  Burch  .^-  Burch.  Upper 
aiul  lower,  right  figs.,  Oliva  I'onilaia  Lamarck. 


NAUTILUS  75(4) 


PLATE  18 


Maxwell  Smith,   1888-1961 


April,  1962  nautilus  165 

Fig.  1  was  prepared  by  Mrs.  Maryellin  Reinecke.  We  wish  to 
thank  Mr.  John  Q.  Burch  for  bringing  this  problem  to  our 
notice  and  for  furnishing  the  specimens  of  both  forms. 

References 
Burch,  J.  Q.  1959.  Minutes  Conch.  Club  So.  Cal.  184:20. 
Burch,  J.  Q.  and  Burch,  R.  L.  1962.  [See  next  paper]. 
Johnson,  C.  W.  1915.  Nautilus  2<?.-115. 
Keen,  A.  M.  1958.  Sea  Shells  of  Tropical  West  America.  Stanford 

University  Press,  Stanford,  Calif,  p.  421. 
Swanson,  H.  E.,  and  Fuyat,  R.  K.  1953.  Nat.  Bureau  of  Standards 

Circular  539,  2;51. 
Swanson,  H.  E.,  Fuyat,  R.  K.,  and  Ugrinic,  G.  M.   1954.  Nat. 
Bureau  of  Standards  Circular  539,  3:53. 


NEW  SPECIES  OF  OLIVA  FROM  WEST  MEXICO 
By  JOHN  Q.  and  ROSE  L.  BURCH 

The  literature  on  the  history  and  many  variations  of  the 
species  commonly  known  as  Oliva  spicata  (Roding,  1798)  is 
voluminous.  The  disposition  of  most  modern  authors  has  been  to 
lump  all  varieties  as  forms  of  one  species.  In  our  opinion,  the 
last  word  has  not  been  written  on  this  assemblage. 

In  our  work,  Minutes  of  the  Conchological  Club  of  Southern 
California,  no.  184,  p.  20,  Jan.  1959,  we  discussed  the  problem  of 
the  two  species  studied  by  Donohue  and  Hardcastle.  Mr.  W.  E. 
Naylor  of  San  Diego,  a  dealer  in  commercial  shells,  referred  this 
problem  to  us.  His  trade  in  olives  is  largely  to  the  Indians  who 
use  the  shells  to  make  various  artifacts.  The  Indians  rejected  one 
variety  claiming  that  they  shatter  when  worked  and  are  unsatis- 
factory. The  acceptable  variety  is  known  to  the  trade  as  Oliva 
venulata  Lamarck,  1811.  These  shells  are  heavier,  with  lower 
spire,  and  more  obese  than  the  discarded  lot.  These  specimens 
consistently  have  the  base  of  the  columella  with  a  white  color 
and  perhaps  a  tinge  of  pink.  Shells  of  the  rejected  lot  are  more 
slender,  higher  spired,  and  the  base  of  the  columella  is  consis- 
tently a  light  purple  instead  of  white.  The  interior  of  both  is  a 
bluish  white.  At  first  glance  the  shells  in  question  seem  to  be  very 
similar,  but  after  a  little  study  they  readily  separate  into  two 
forms.  The  studies  of  Donohue  and  Hardcastle  have  shown  that 
the  two  forms  differ  in  the  chemical  analysis  of  the  shell.  Both 


166  NAUTILUS  Vol.  75(4) 

forms  are  found  on  the  same  tide  flats  at  La  Paz,  Baja  California, 
Mexico,  with  no  intergrades  in  many  hundreds  of  specimens. 
Certainly  many  species  are  recognized  upon  much  more  meagre 
grounds. 

In  our  preliminary  discussion,  for  convenience,  we  assigned 
the  discarded  form  to  Oliva  ustulata  Lamarck,  1811.  Several  very 
diff^erent  forms  have  been  recognized  by  various  authors  under 
this  name.  Tryon  in  the  Manual  of  Conchology  thought  it  to  be 
a  variety  of  the  Atlantic  Olivn  reticularis  Lamarck,  1811.  Charles 
Johnson  thought  it  to  be  a  form  in  which  the  pattern  of  the 
shell  is  obscured  by  a  dark  brown  layer.  In  our  opinion,  the  name 
Oliva  ustulata  Lamarck  cannot  be  used  for  this  form.  Seemingly 
the  species  of  Oliva  under  discussion  has  been  considered  a  fonii 
of  the  typical  by  all  authors  we  have  consulted.  Therefore,  we 
feel  justified  in  giving  it  a  name  and  the  status  of  a  new  species. 

Oliva  rejecta,  new  species.  Plate  17,  left  figs. 

Shell  cylindrically  oblong;  spire  exserted,  prominent;  color 
white  with  a  closely  reticulate  pattern  of  chocolate;  base  of 
columella  purple,  swollen,  with  4  plaits;  interior  of  aperture 
bluish  white.  Length:  35.5  mm.  Greatest  diameter:  15  mm.  Type 
locality:  tide  flats  at  La  Paz,  Baja  California. 

A  comparison  of  the  dimensions  of  the  type  of  the  new  species 
with  those  of  a  normal  specimen  of  Oliva  vemilnta  Lamarck  is 
of  interest.  Both  specimens  measure  exactly  35.5  mm.  in  length, 
but  the  greatest  diameter  of  Oliva  rejecta  is  15  mm.,  while  that 
of  Oliva  venulata  is  17.5  mm.  Also,  the  gieatest  diameter  of 
Oliva  rejecta  is  much  lower  on  the  body  whorl. 

The  holotype  is  to  be  deposited  in  the  California  Academy  of 
Sciences,  San  Francisco,  no.  12400.  Fifty  paratypes  will  be  dis- 
tributed to  various  institutions. 


NOTES  AND  NEWS 

BuLiMULUs  DEALBATUS  joNESi  Cleuch  appears  to  be  Bulimulus 
mooreanus  (Pfeiff^er)  which  has  been  introduced  into  Alabama, 
It  is  somewhat  smaller  than  is  usual  for  the  species  in  Texas,  but 
is  otherwise  typical.  Unlike  B.  dealbatus  (Say) ,  it  estivates  on 
the  weeds  where  it  can  be  found  in  large  numbers  even  in  the 
hottest  part  of  the  summer.  —  Leslie  Hubricht. 


MBI.  WHOl    I  IKKAHY 


liJH    17XP    T